Flora of Great Britain and Ireland: Volume 4, Campanulaceae - Asteraceae (Flora of Great Britain and Ireland)

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FL OR A OF GR E AT B R ITAIN A N D IRELA N D Planned in five volumes, this critical flora provides a definitive account of the native species, naturalised species, frequent garden escapes and casuals found in the British Isles. Full keys and descriptions will enable the user to name all plants occurring in the wild, plus some ornamental trees and shrubs. For the first time detailed accounts of all the large apomictic genera are given and many infraspecific variants included. Each species entry begins with the accepted Latin name, synonyms and the common English name. A detailed description follows, including information on flowering period, pollination and chromosome number. Separate descriptions are given for infraspecific taxa. Information on the status, ecology and distribution (including worldwide distribution) of the species and infraspecific taxa is also given. Clear black-and-white line drawings illustrate an extensive glossary and also illuminate the diagnostic features in a number of groups of plants. PETER S ELL joined the Herbarium in the University of Cambridge’s Department of Plant Sciences in 1944, holding the post of Assistant Curator from 1972 until his retirement in 1997. His work there on this flora continues unabated, together with almost daily visits to the University’s Botanic Garden throughout the flowering and fruiting seasons. He is co-author of Flora of Cambridgeshire (1964) and Flora of the Maltese Islands (1977), and was involved in the whole of the Flora Europaea project, also published in five volumes (1964–80). GINA MURRELL is Assistant Curator of the Herbarium in the University of

Cambridge’s Department of Plant Sciences, having previously held the post of Herbarium Technician there. She has worked with Peter Sell over a period of 30 years, and they have together collected a quarter of the British Herbarium’s 200,000 specimens.

F L O RA OF

G R E AT BRI TA I N AND

I R E L A ND VO L U ME 4 CAM PA NUL AC EAE – AST E R AC E AE

PE T E R SEL L and GINA MURRELL Herbarium, Department of Plant Sciences University of Cambridge

cambridge university press Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo Cambridge University Press The Edinburgh Building, Cambridge cb2 2ru, UK Published in the United States of America by Cambridge University Press, New York www.cambridge.org Information on this title: www.cambridge.org/9780521553384 © Cambridge University Press 2005 This publication is in copyright. Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published in print format 2006 isbn-13 isbn-10

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To our Mentors

E dred Joh n Henry Corner James Edgar Dandy Hu m ph rey Gi lbert Carter Harry Godwin William Thomas Stearn Stuart Max Walters Alexan der Stuart Watt C yri l Wes t

Contents

Foreword ix Preface xi Acknowledgements xii Introduction xiii Conspectus of families xxi Text 1 142.

C A M PA N U L A C E A E

1

146.

VA L E R I A N A C E A E

143. 144.

RU B I AC E A E

14

147.

D I P S AC AC E A E

54

C A P R I F O L I AC E A E

29

148.

A S T E R AC E A E

62

145.

A D OX AC E A E

48 New taxa and combinations 530 Abbreviations 558 Glossary 559 Index 573

48

Foreword to volume 4 by S. M. Walters ScD V.M.H.

It has been one of the continuing satisfactions of my academic career in Cambridge that the University Herbarium, of which I was Curator from 1948 to 1973, has provided an academic base for all my specialist interest in angiosperm taxonomy to develop. Indeed, I count myself doubly fortunate that, 12 years after my retirement from academic life, the Herbarium, with its staff and visitors, still provides such a base where scholarship can be pursued for its own sake. With great pleasure I welcome this volume, the first of a set of five promised to us by Peter Sell and Gina Murrell. My association with Peter goes back more than half a century: though I was ‘senior partner’ in our happy collaboration in the post-war Herbarium, ours was a symbiotic relationship from which we both greatly benefited, and I was delighted when Gina, who had been part of the team in the 1960s and 1970s, returned to the fold as Herbarium Technician in 1991. As explained in the Preface, this project to write an entirely new critical flora of the British Isles comes to fruition some 20 years after an earlier scheme, in which the late Professor David Valentine took a leading part, had failed to find any financial support. Both Clive Stace to whose New Flora of the British Isles (1997) Peter pays tribute in the Preface, and Peter himself, were enthusiastic supporters of the Valentine project, and were prepared to play major parts in writing the Flora. It is fitting that both these eminent British taxonomists should separately carry on the tradition that David Valentine so enthusiastically advocated. Two aspects of this new critical Flora seem to be especially important. One concerns the acceptance, long overdue, of the ‘alien element’ in our flora as being equally worthy of taxonomic study: in this respect Stace’s Flora represents a real change in attitude, which is to my mind unreservedly to be welcomed. The other, interestingly linked to the first by many examples, concerns the taxonomic recognition and treatment of hybrids and infraspecific variants. British botany lacks any single reference work from which the basic information about the variation of British

vascular plants can be found, yet this information is increasingly needed by ecologists, conservationists, molecular biologists and biochemists, who will, as the century closes, determine the shape of much botanical study in universities and specialised institutions. The authors of this impressive work have set themselves a colossal task. They have made an excellent start, and we can only wish them a successful conclusion. 1996 It is for me a very real pleasure to add a further word to welcome this, the second volume of ‘Sell & Murrell’, as this remarkable Flora is now widely known among British botanists. Of course, this new volume, containing in particular the genus Hieracium, must rank as Peter’s very own ‘labour of love’. One of the very special links that has grown up between Peter and me over our long-standing acquaintance in the pursuit of taxonomic botany must be our steady, persistent enthusiasm for critical apomictic genera. We do not have to explain or justify to each other our passions for, in my case, Alchemilla, and his for Hieracium. I have to admit, however, that his task, with 412 named and described species of Hieracium in this volume, casts my puny efforts with British Alchemilla into the shade! Talking to Peter and Gina about the progress of this remarkable Flora, I am encouraged by what I hear. I really believe that both Peter and I will live to see its completion, in spite of the fact that we both ‘creak a little at the joints’ – to use one of the common euphemisms we find ourselves using from time to time to describe our state of health! One final observation. How fortunate Peter is to have such a remarkable fellow-author in Gina! Writing and publishing books involves much more than producing a draft text. Some of the skill is straightforward, if laborious; but some requires real understanding at the level of human relations, and both these skills are possessed in abundance by Gina. So I conclude by saying to both Peter and Gina: keep up the good work to a successful conclusion.

Preface

For over 50 years I have worked in the herbarium at Cambridge University on the British and European floras. I have collected about 30,000 numbers consisting of some 50,000 specimens from most parts of the British Isles and made many visits to Continental Europe. Particular attention has been given to most critical genera: Cerastium, Conyza, Crepis, Dactylorhiza, Euphrasia, Fumaria, Hieracium, Limonium, Pilosella, Prunus, Rhinanthus, Salicornia, Salix, Scleranthus, Sorbus and Ulmus; and in helping friends in various ways I have considered the taxonomy of Alchemilla, Batrachian Ranunculi, Chenopodium, Potamogeton, Rubus and Taraxacum. I have also spent much time studying ecotypic and geographical variation, in particular a comparison of those variants which occur on the coasts in dunes, shingle and salt-marsh with those growing as arable weeds, and those in mountains. Special attention has also been given to trees and shrubs. It has long been my wish to publish this information in a critical flora of Great Britain and Ireland. In the 1970s a group of us tried to get a grant to carry this out, but we were unsuccessful. Clive Stace then started work on his New Flora of the British Isles, which was first published in 1991, with a second edition in 1997. In it he gives only abbreviated descriptions and omits most of the species in the large apomictic genera and many of the infraspecific variants. Numerous introduced species are included by Stace in a British and Irish flora for the first time, detailed descriptions and specimens of many of which are difficult to find. Stace’s flora is to my mind an excellent field guide, which it would be difficult to better, but it does not give the detailed descriptions that are needed to confirm the identification of a plant which is new to you. A good description in my opinion is one in which a picture of the plant unfolds before you as you read it. I considered it was possible for me to write a flora in five volumes which gave a full description of all the species in Stace’s flora and to add all the apomicts and many of the infraspecific variants, but it was too large a task to attempt

to include all the biological information envisaged by the group in the 1970s. It was necessary, however, to have the help of another author, who lived in Cambridge, to deal with the large amount of work involved. My eye fell upon Gina Murrell who had worked with me in the 1960s and 1970s, when writing accounts for Flora Europaea, Flora of Turkey and Flora of the Maltese Islands. The work of one had complemented the work of the other and we were able to criticise one another without antagonism. We started field work on this flora on 13 May 1987, by describing Ceratocapnos claviculata, which was flowering on Dunwich Heath in Suffolk, in a snowstorm. Since then we have as far as possible spent one day a week working in the field or at the Botanic Garden, Cambridge. We started writing Volume 5 in 1992, and completed it by Easter 1994. It was published on 1 April 1997, not 1996 as stated in the volume itself. I have done most of the writing and made the taxonomic and nomenclatural decisions, while Gina has done most of the measuring, sometimes sitting at the microscope dictating the description while I, surrounded by a pile of books, wrote it down, and she has set out and put the whole onto a computer. Gina has also done all the illustrations and organised our field work. Volume 4, probably the largest of the five, has taken longer than we thought due to my moving house and retiring and having to vacate my room in the Department of Plant Sciences in which I had accumulated much over 50 years. Major alterations to the whole Department have added to the difficulties. Volume 5 contained 28 families, 233 genera, 769 species, 93 subspecies, 148 varieties, 22 formae and 182 hybrids. This volume contains an introduction and full accounts of seven families, 146 genera, 1,098 species, 130 subspecies, 162 varieties, 27 formae and 51 hybrids. It deals with a whole range of taxa from very variable species which we feel cannot be further divided, to species with geographical races, ecotypes, forms and cultivars. The taxa may be outbreeding, inbreeding, apomictic or spreading vegetatively. Peter Sell

Acknowledgements

In contrast to Volume 5, the specialised knowledge that P. D. S. has of the Asteraceae has not necessitated so much outside help as regards the taxonomy, but the account would be nowhere near as complete without the very special help given by John Richards and the late Andrew Dudman in Taraxacum. They made available their first draft of the Botanical Society of the British Isles’ Handbook of Dandelions of Great Britain and Ireland and we were able to keep up to date via Philip Oswald as he was editing the final version. To the late Cyril West is also owed a very great debt. He spent the whole of his retirement working on Hieracia and came up to Cambridge regularly for over 30 years to study the large Cambridge collection. Our thanks also to the late Archie Kenneth and Mary McCallum Webster who gave almost all their numerous new collections of Hieracia to Cambridge, as have many other British and Irish botanists, particularly Allan McG. Stirling and Walter Scott. David McCosh has read and commented on the final version of Hieracium. Another special debt is due to David Tennant who wrote the very detailed account of Hieracium section Alpina. As well as growing nearly all the species of this section in his own garden he has seen them all in the field. Philip Oswald has not only translated into Latin all the new taxa in this volume, but has answered many general questions concerning Greek and Latin, and the layout of the flora. Max Walters has translated much Swedish, German and French as well as discussing many problems of taxonomy. Chris Preston has read the whole of the text and as well as much improving the accounts of the distribution of the taxa, has added their geographical classification which is set out and explained in Preston, C. D. and Hill, M. O., The geographical relationships of British and Irish vascular plants in Botanical Journal of the Linnean Society 124: 1–120 (1997). Arthur Chater has acted as a very special sort of adjudicator. Whenever we worked out infraspecific variants which were likely to occur in Cardiganshire, we telephoned him to give him the information, and he would either comment immediately or search for them at the first opportunity. His paper in Watsonia 24: 281–286 (2003) sets out the difficulties in finding information on infraspecific taxa. Charlie Jarvis has helped us while working on the Hortus Cliffortianus at the British Museum and Gina Douglas while

working on the Linnean collection at Burlington House. Barry Goddard has advised us on computer techniques. To Mrs J. E. Dandy we owe a special debt for giving us the second copy of her husband’s manuscript of his detailed work on the nomenclature of the British flora. Doug Kent continued to answer nomenclatural questions up until the time of his death. His contributions to British botany will be greatly missed. Harold Whitehouse read parts of the text and David Briggs has given us advice on breeding mechanisms. Bob Finch has helped with the account of Leontodon. To Bill and Joan Robinson we are grateful for letting us frequently raid their garden and for allowing some of their vegetables to go to flower and seed so that we were able to collect complete plant specimens. P. D. S. owes a very special debt to Brian and Rosemary Chapman. Every Friday afternoon they have led him over unfamiliar ground at Histon in Cambridgeshire and he has tried to name every plant he has come across down to forma. This has enabled him to keep his eye in with plants in the field as well as in the herbarium. In the library of the Department of Plant Sciences Richard Savage has gone out of his way to get hold of rare and obscure publications for us and somehow to keep the library going during major alterations. At the Botanic Garden, Cambridge, Professor John Parker has allowed P. D. S. to continue after his retirement to have a free run of the Garden, its library and herbarium and has endeavoured to answer some of the difficult genetical and biological questions we have put to him. Also at the Garden, James Cullen, Alexander Goodall, Caroline Hotine, Huw Jones, Peter Kerley, Clive King, Pete Michna, Sally Petit, Ann Schindler, David Stone, Tim Upson, Norman Villis and Peter Yeo have given us much help. It has been particularly helpful to know from James Cullen what was going on while Asteraceae was being written for the Garden Flora. For many years the late Jack Symons looked after the large collection of cultivated Hieracium we had in the garden. Professor Enid McRobbie and Professor Roger Leigh have allowed P. D. S. to continue to have full use of the herbarium and library after his retirement. To Clive Stace we owe a very special debt. Had he not written his New Flora of the British Isles our task would have been insurmountable. In this volume he has even saved us the onerous task of searching through the literature to sort out the correct names of the Hieracium sections.

Introduction

HISTORICAL BACKGROUND The first real flora of these islands was John Ray’s Catalogus Plantarum Angliae et Insularum Adjacentium in 1670. The first flora to use the Linnaean binomial system of nomenclature was William Hudson’s Flora Anglica nearly a hundred years later in 1762. This was followed by William Withering’s Botanical Arrangement of all the Vegetables naturally growing in Great Britain in 1776–92, the first of many floras written primarily for the amateur. James Sowerby’s English Botany, whose text was written by J. E. Smith, was first published between 1790 and 1820. It presented for the first time a complete set of coloured illustrations of our plants, illustrations which are still unsurpassed for line and colour. The third edition, published between 1863 and 1872, has inferior illustrations, but its text, rewritten by James Boswell Syme, is still important for its nomenclature and infraspecific taxa. Three especially famous floras were produced in the nineteeth century. George Bentham’s Handbook of the British Flora in 1858 was written as a before-breakfast relaxation. In it keys appeared for the first time in a British flora. It was revised by J. D. Hooker in 1886. J. D. Hooker’s Student’s Flora of the British Islands, first published in 1870 and finally revised in 1884, had very clear and concise descriptions and was the main flora used by many generations of botanists up until the 1950s. It is also important in that Hooker was one of the first authors to make frequent use of the category of subspecies. Charles Cardale Babington’s Manual of British Botany first appeared in 1843 and the tenth edition, revised by A. J. Wilmott, was published in 1922. It contains many critical species and varieties not in other floras, but the descriptions are not clear and without keys it is difficult to use. C. E. Moss’s Cambridge British Flora (1914–20), was very detailed and would have supplied a much needed critical flora, but alas only two volumes were published. The arrival of ‘C. T. & W.’, A. R. Clapham, T. G. Tutin and E. F. Warburg’s Flora of the British Isles, in 1952, heralded the beginning of a new era in the study of British plants. It was the first up-to-date treatment in the twentieth century. A much revised second edition appeared in 1962 and a third in 1987 when D. M. Moore replaced E. F. Warburg. This last edition included the information in Tutin et al., Flora Europaea 1–5 (1964–80). The nomenclature had been brought up to date by J. E. Dandy in his List of British Vascular Plants in 1958, and the work he did on this for Flora Europaea. Thus for the first time taxonomy and nomenclature had been brought in line with those of Continental Europe.

The Botanical Society of the British Isles’ publication of the Atlas of the British Flora in 1962, edited by F. H. Perring and S. M. Walters, and the Critical Supplement to the Atlas of the British Flora in 1968, edited by F. H. Perring, gave us a much better idea of the distribution of our plants. The new edition of the Atlas arrived when most of this volume had been prepared for press, but the fact that Chris Preston had checked most of our distributions meant they were not much out of date. The arrival of Clive Stace’s New Flora of the British Isles in 1991 with a second edition in 1997, and D. H. Kent’s List of Vascular Plants of the British Isles in 1992 has brought about the end of the C. T. & W. era and given us a completely up-to-date account of our flora. Major changes included the moving over of the main classification to A. Cronquist’s An integrated system of classification of flowering plants (1981) and the inclusion of almost as many alien species as native ones. The aim of our Flora is to supply full descriptions of all the species in Stace’s flora, to include all the large apomictic genera and as many infraspecific variants as practicable, and to add more information about hybrids for which extensive use has been made of Stace’s Hybridization and the flora of the British Isles (1975).

THE CONTENTS OF THE FLORA The Flora includes all the vascular plants, Lycopodiophyta (Clubmosses), Equisetophyta (Horsetails), Pteridophyta (Ferns), Pinophyta (Conifers) and Magnoliophyta (Flowering plants). The list of plants is made up of all our native species, including apomicts, and all the introduced plants given in Stace (1991), with some more added, particularly planted trees. E. J. Clement’s and M. C. Foster’s Alien plants of the British Isles arrived in 1994 after we had completed Volume 5, but we went through it and added as much information as possible. It has been used continually while preparing this volume. These alien taxa may be found to be more widespread when full attention is given to them. In his coverage of alien taxa Stace considers inclusion is merited when an alien is either naturalised (i.e. permanent and competing with other vegetation, or self-perpetuating) or, if a casual, frequently recurrent so that it can be found in most years. These criteria were applied as much to garden escapes or throw-outs as to the unintentionally introduced plants, and rarity was not taken into consideration for any of them. Cultivated species were included if they are field crops or forestry crops, or in the case of trees only, ornamentals grown on a large scale. Stace’s aim has been to include

xiv

Introduction

all taxa that the botanist might reasonably be able to find in the wild in any one year. To these we have added some ornamental trees and shrubs that are planted along streets and roadsides and in parks and estates and which we consider to be part of the landscape. Usually plants in gardens are not mentioned at all, but some species, which seed freely and spread over areas of garden and lawn where they are not planted, are included. Most of the species which Stace has mentioned, but not numbered or included in the keys, are here included, while a few have been left out altogether. We started with Volume 5 because The European Garden Flora had already covered the Monocotyledons, which has made it easier for us to deal with the garden escapes. We are following it with Volume 4, because it contains the large genera Hieracium and Taraxacum.

GEOGRAPHICAL AREA The Flora deals with the British Isles and includes England, Scotland and Wales, collectively known as Great Britain, Northern Ireland and Eire together forming Ireland, the Isle of Man, and the Channel Islands which include Jersey, Guernsey, Alderney, Sark, Herm and various small islands. In these respects it follows Stace (1991 and 1997). The smallest geographical area usually referred to is the county. This sometimes includes more than one botanical vice-county. For Great Britain we have used the boundaries adopted by H. C. Watson in 1873 in Topographical Botany and in Ireland by R. L. Praeger in 1901 in Irish Topographical Botany. These are the vice-counties used by botanists, which have the benefit of not changing at regular intervals as do the political counties. With rare or local species the actual place or area may be given. The extra-limital distributions are those given in Clapham, Tutin and Moore (1987) with as much correcting as we and Chris Preston can give them. Russia and Yugoslavia have been used in the sense of the old U.S.S.R. and Yugoslavia before recent political disruptions.

CLASSIFICATION AND NOMENCLATURE The classification follows that of Stace (1991 and 1997) and Kent (1992) which is taken from A. Cronquist, An Integrated System of Classification of Flowering Plants (1981), with the exception that the main groups are called Divisions and the second groups Classes following H. C. Bold, C. Alexopoulos and T. Deleveryas, Morphology of Plants and Fungi, ed. 4 in 1980 and A. Cronquist, A. Takhtajan and W. Zimmermann, On the higher taxa of embryobionta, in Taxon 15: 129–134 in 1966, and set out by one of us, P. D. S., in The Cambridge Cyclopedia of Life Sciences in 1985. One of us (P. D. S.) has specialised in nomenclature for many years and it is here made as accurate as possible according to the latest International Code of Botanical Nomenclature. The names of genera and species differ

little from those in Stace (1991 and 1997) and Kent (1992). Recent changes in the code of nomenclature have been used to get rid of some names which have been a long-standing source of confusion. New taxa and such changes in nomenclature and taxonomy which do occur are published at the end of the volume. No rules have been made about the number of synonyms given, as many as possible being included, but an attempt has been made to include all names used in British and Irish floras. The abbreviation auct. following a name means only that the name has not been accepted for the plant, it does not mean the type has been checked and the name rejected. Only in the case of a later homonym, which has been checked, does the word non and an author follow the name and author. This including of numerous synonyms often shows how a species has moved from one genus to another over the years, especially in the Asteraceae. The English name for the species follows Stace (1991 and 1997) as far as possible, and where they are missing from new species they have been created by us. This volume contains some very large keys. Such keys are not very easy to use. The modern idea is to break up these keys into a number of smaller ones. This is purely psychological, giving the impression it makes things easier. It does not. The initial breaks, whether in one large key or an introductory key, are the most difficult. The more one knows about the genus the more useful the key becomes and often enables one to take a short cut. Where natural groups appear in the key this has been indicated so that as users become more proficient they can move to the group that matters. In whatever way you arrive at your identified species your plant should fit exactly the detailed description in the text. If a difficult plant has any chance of being identified very detailed notes of every character should be made in the field. All too often a miserable bit of a plant rotting in a plastic bag is all that is available, as after a long day in the field the botanist put his meal and bed first. Sitting on the horizon is the so-called Code of Bionomenclature which could have a profound effect on the names of our plants in the future. Its aim is to provide a nomenclature for both botany and zoology in which there are no duplication of names. The most up-to-date account of it is by D. L. Hawksworth and J. McNeill in Taxon 47: 123–150 (1998). I have always thought there is much to be said for the zoological idea that all replacing populations have trinomials. It would mean all the subspecies and varieties in this flora would have trinomials and one would no longer have the difficulty of deciding whether a population is ecological or geographical or both, but how many name changes that would require is not known. Only forma would then be used in its present context, which is what the zoologists call morphs. Perhaps the biggest problem is priority of genera where the same epithet occurs in both botany and zoology. The genus Oenanthe for plants in 1753 and for birds in 1816 would mean over 20 familiar bird names would have to be altered. Prunella also has these same dates for plants and birds. This results in some 15 name changes. More worrying seems to be the rules for conserving names, which could result in a caucus sitting on a committee telling everyone else what to do.

Shetland Islands

100 km

Orkney Islands

GREAT BRITAIN Scottish Highlands

IRELAND Lake District River Tees Yorkshire Dales Peak District

Isle of Man Connemara

River Humber The Wash

The Burren

Snowdonia Fens Brecon Beacons

East Anglia The Broads Breckland

River Thames River Severn Lundy Island

Isles of Scilly

Alderney CHANNEL ISLANDS

FRANCE Guernsey Jersey

British Isles

112

100 km 111

109

108 107 110 106

105

95

93

94 96

104

92

Scotland

91

97 89

90

88

103 98

85

87 99 102

86

84

81

77

101 100

82

83

76

GREAT BRITAIN

78 79

75

68

80

IRELAND 72 35

40

34

67

73

74

39

33 27

61

60

24 22

23

54 52

19 14

9

49

20

58

51

50

57

56 53

10

48

13 11

7

63

59 21

18

8

62

64

Isle of Man

31

25

15

39

28

55

40

47 12

2 6 1

65

71

29

17

4

England

69

38

30

26 16

37 32

28

66

70

36

Wales

5

31

38

43

46

32

37

42

44

45

33

23

7

22

24

34 6

12

8

5

11

4 9 3 2 1

Vice-Counties of the British Isles

10

19

20

35 41

Channel Islands

25

26

30

36

3

29

27

21 17 13

18 16 14

15

ENGLAND, WALES, SCOTLAND, ISLE OF MAN 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38.

West Cornwall East Cornwall South Devon North Devon South Somerset North Somerset North Wiltshire South Wiltshire Dorset Isle of Wight South Hampshire North Hampshire West Sussex East Sussex East Kent West Kent Surrey South Essex North Essex Hertfordshire Middlesex Berkshire Oxfordshire Buckinghamshire East Suffolk West Suffolk East Norfolk West Norfolk Cambridgeshire Bedfordshire Huntingdonshire Northamptonshire East Gloucestershire West Gloucestershire Monmouthshire Herefordshire Worcestershire Warwickshire

39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76.

Staffordshire Shropshire Glamorganshire Breconshire Radnorshire Carmarthenshire Pembrokeshire Cardiganshire Montgomeryshire Merionethshire Caernarvonshire Denbighshire Flintshire Anglesey South Lincolnshire North Lincolnshire Leicestershire Nottinghamshire Derbyshire Cheshire South Lancashire West Lancashire South-east Yorkshire North-east Yorkshire South-west Yorkshire Middle-west Yorkshire North-west Yorkshire Co. Durham South Northumberland Cheviotland Westmorland Cumberland Isle of Man Dumfries-shire Kirkcudbrightshire Wigtownshire Ayrshire Renfrewshire

77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112.

Lanarkshire Peebles-shire Selkirkshire Roxburghshire Berwickshire East Lothian Midlothian West Lothian Fifeshire Stirlingshire West Perthshire Mid Perthshire East Perthshire Forfarshire Kincardineshire South Aberdeenshire North Aberdeenshire Banffshire Morayshire East Inverness-shire West Inverness-shire Main Argyllshire Dunbartonshire Clyde Islands Kintyre South Ebudes Middle Ebudes North Ebudes West Ross-shire East Ross-shire East Sutherland West Sutherland Caithness Outer Hebrides Orkney Islands Shetland Islands

IRELAND H1. H2. H3. H4. H5. H6. H7. H8. H9. H10. H11. H12. H13. H14.

South Kerry North Kerry West Cork Mid Cork East Cork Co. Waterford South Tipperary Co. Limerick Co. Clare North Tipperary Co. Kilkenny Co. Wexford Co. Carlow Laois

H15. H16. H17. H18. H19. H20. H21. H22. H23. H24. H25. H26. H27. H28.

South-east Galway West Galway North-east Galway Offaly Co. Kildare Co. Wicklow Co. Dublin Meath Westmeath Co. Longford Co. Roscommon East Mayo West Mayo Co. Sligo

H29. H30. H31. H32. H33. H34. H35. H36. H37. H38. H39. H40.

Co. Leitrim Co. Cavan Co. Louth Co. Monaghan Fermanagh East Donegal West Donegal Tyrone Co. Armagh Co. Down Co. Antrim Co. Londonderry

xviii

Introduction

VARIATION The recording of variation is most important for both ecology and conservation, and even more important for gardeners who go out of their way to both create and conserve prominent variants. Infraspecific variation is usually recorded by the recognition of subspecies, varieties, formas and cultivars. These taxa differ chiefly in ecology and distribution. A forma is a plant with a one- or two-gene difference which occurs with one or more other forms in a mixed population for most or all of its range. A variety is when one of these formas becomes more or less dominant in a particular ecological area, that is an ecotype. A subspecies is when one of these formas becomes dominant in a geographical area, that is a race. A cultivar is a forma which is selected by horticulturalists and perpetuated, usually vegetatively. Because ecotypes and races have become adapted morphologically to different conditions over a long period, it is likely that their physiology and biochemistry, and indeed their whole biology, is different. Also, as they often flower at different periods their pollinators may be different and, if climatic conditions alter, one ecotype may be better able to survive than another. Variation thus becomes very important in conservation. Because the Biological Flora of the British Isles has lumped all its information under species it can be highly misleading when applied to individual populations. It is unfortunate that many botanists tend to ignore variation completely, and they will certainly ignore it if it has no name at all; subspecies are usually more often recognised than varieties. Sometimes it is more important to conserve one variety rather than another, e.g. the Chilterns Orchis militaris var. tenuifrons is endemic, while the Suffolk var. militaris occurs in Continental Europe; Liparis loeselii var. ovata is rare in distribution, but frequent where it occurs, whereas var. loeselii is rare in Britain but occurs on the continent. Sometimes the variant will tell us whether the plant is native or not; e.g. Leucojum aestivum subsp. aestivum is native, subsp. pulchellum is a naturalised garden escape. Escaped cultivars are named wherever they can be easily recognised and are considered important. All apomicts, where possible, are treated as species, long experience showing that any sort of lumping deprives them of having an interesting ecology or distribution. Hybrids are dealt with as fully as possible, especially those that spread vegetatively. No serious attempt has been made to decide on the correct infraspecific rank as taxa are often both ecological and geographical. Uniformity of infraspecific rank is often produced in a species or genus, but usually the only important thing considered is that a morphological recognisable infraspecific taxon has a name. Where species grade gradually into one another over large distances, as the species of Larix do round the northern hemisphere, and at given points the whole population is uniformly intermediate it is regarded as a cline. Where two populations of ecotypes grow adjacent as in Geum rivale and urbanum there is often an area in which variable intermediates occur. This is also often called a cline, but it is really only so statistically, and we prefer to call it

a variable hybrid zone. If you look for these things you will be surprised how often they exist, and clear-cut species, even apomicts, are not so clear-cut as we are made to believe. Dick Brummitt and Arthur Chater writing in Watsonia 23: 161 (2000) about the genus Calystegia say: The whole genus, in which some 25 species world-wide may conveniently be recognised, is taxonomically difficult, and few if any of the species are morphologically clear-cut. They mostly vary considerably over their ranges and merge geographically one into another, and division into species and subspecies is of necessity somewhat arbitrary.

We find this true of many groups when their whole range is considered. If the origin of taxa is considered there are even more difficult problems. During the Quaternary cold stages massive glaciation from the north caused forests to retreat southwards and come up against other different floras and in some cases to hybridise with them. On climatic amelioration the forests advanced north again, often perhaps by a different path by which they went south, bringing a fresh variety of plants to join those species which survived in refugia in the cold areas. Richard West in Plant Life of the Quaternary Cold Stages: Evidence from the British Isles (2001, p. 263) writes: This overall view shows the cold stage stadial flora to have a long and complex history, originating in the latest Tertiary, occupying a major part of Quaternary time in our area, and surviving short periods of forest dominance at times of climatic amelioration. It is not surprising that the taxonomy of the species concerned is very complex.

Following the forest clearances of the Neolithic about 5,000 years ago and the agricultural revolution which followed, some of the species of open habitats, particularly coastal areas, developed ecotypes which became agricultural weeds, or were actually brought in by early Man himself. Indeed, the weeds of Cornwall, East Anglia and Scotland may have been brought in by different races of Man, at different times, from entirely different areas. To add to this state of affairs Man has brought plants from all over the world into gardens. Sometimes two species which never occur together in nature are grown together in gardens and hybridise, and may even backcross to one or both parents. These plants often escape into the countryside and sometimes our garden plants will hybridise with our native species. To confuse the issue even further Continental races of our native species are introduced to our countryside in packets of wild flower seed or are planted as trees and shrubs in our woodlands. It can thus be seen that many of our species are far from uniform genetically. Apomictic species and many ecotypic varieties are probably more uniform. DNA may help us to understand these problems, but will we ever have time and money to look in detail at all our flora?

Introduction

HERBARIA AND LITERATURE CONSULTED During the writing of the Flora the following books were consulted for every species: Boreau, A. Flore du centre de la France. 1 and 2. 1857. Paris. (This contains a large number of segregate species of A. Jordan, which he grew in cultivation for many years, and which are now recognised as infraspecific taxa or apomictic species.) Clapham, A. R., Tutin, T. G. & Moore, D. M. Flora of the British Isles. Ed. 3. 1987. Cambridge. Dandy, J. E. List of British vascular plants. 1958. London. Hegi, G. Illustrierte Flora von Mitteleuropa. Ed. 1. 1906– 1931. M¨unchen. Ed. 2. 1936– M¨unchen. Ed. 3. 1966– M¨unchen. Kent, D. H. List of vascular plants of the British Isles. 1992. London. Perring, F. H. & Walters, S. M. Atlas of the British flora. 1962. London & Edinburgh. Perring, F. H. Critical supplement to the atlas of the British flora. 1968. London. Preston, C. D. & Croft, J. M. Aquatic plants in Britain and Ireland. 1997. Colchester. Stace, C. A. Hybridization and the flora of the British Isles. 1975. London, New York & San Francisco. Stace, C. New flora of the British Isles. Ed. 1. 1991. Cambridge. Ed. 2. 1997. Cambridge. Stearn, W. T. Botanical Latin. Ed. 4. 1992. Newton Abbot. (This book carries an enormous amount of information and may well have been used more often than any other tome.) Stewart, A., Pearman, D. A. & Preston, C. D. Scarce plants in Britain. 1994. Peterborough. Tutin, T. G., Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb, D. A. Flora Europaea. 1–5. 1964–80. Cambridge. Walters, S. M., Brady, A., Brickell, C. C., Cullen, J., Green, P. S., Lewis, J., Matthews, V. A., Webb, D. A., Yeo, P. F.

xix

& Alexander, J. C. M. The European garden flora. 1–6. 1986–2000. Cambridge. Many other books and journals were consulted, mainly in the Cambridge Department of Plant Sciences, including the N. D. Simpson collection of local floras, and the Cory Library at the Botanic Garden. Where these references were considered to be important for particular plants, we have cited them under the family or genus concerned. The University herbaria at Cambridge, on which the Flora is mainly based, are ideal for the study of the British flora for the following reasons: 1 The large British collection contains specimens from most of the main collectors of British plants from 1800 onwards, including sets of published exsiccatae and specimens sent through the Botanical Exchange Clubs. Most of the critical species have been named by experts. 2 The British herbarium contains some 50,000 specimens collected by us in the last 50 years. The specimens are accompanied by detailed field notes and are often of critical species or infraspecific taxa. Often a gathering may consist of more than one sheet, particularly of trees which may have been visited three or four times. 3 There is a good herbarium of Continental European plants with which to compare the British plants. 4 The world collection contains over 50,000 sheets of John Lindley’s herbarium made when he was secretary of the Royal Horticultural Society, when plants were coming into the country from all parts of the world; and the C. M. Lemann collection, named by George Bentham, and put together at the same time. These collections are very important as regards the alien species when considered in conjunction with the Botanic Garden herbarium and recent gatherings of alien specimens. 5 The Botanic Garden herbarium contains a large collection of cultivated plants. Thus, the libraries, herbaria, our own field notes and plants grown in the Botanic Garden have enabled us to do most of the work in Cambridge. Over many years books and specimens elsewhere have been consulted.

Conspectus of families

Kingdom

P L A N TA E

Volume 1. Division

1. LY C O P O D I O P H Y TA Order 1. LY C O P O D I A L E S

1. LY C O P O D I AC E A E

Order 2. S E L A G I N E L L A L E S 2. S E L AG I N E L L AC E A E

Order 3. I S O E TA L E S 3. I S O E TAC E A E

2. E Q U I S E T O P H Y TA

Division

Order 1. E Q U I S E TA L E S 4. E QU I S E TAC E A E

Division

3. P T E R I D I O P H Y TA

Order 1. O P H I O G L O S S A L E S 5. O P H I O G L O S S AC E A E

Order 2. O S M U N D A L E S 6. O S M U N DAC E A E

Order 3. P T E R I D A L E S 7. A D I A N TAC E A E

8. P T E R I DAC E A E

Order 4. M A R S I L E A L E S 9. M A R S I L E AC E A E

Order 5. H Y M E N O P H Y L L A L E S 10. H Y M E N O P H Y L L AC E A E

xxii

Conspectus of families

Order 6. P O LY P O D I A L E S 11. P O LY P O D I AC E A E

Order 7. D I C K S O N I A L E S 12. C YAT H E AC E A E

13. D I C K S O N I AC E A E

Order 8. D E N N S TA E D T I A L E S 14. D E N N S TA E D T I AC E A E

18. DAVA L L I AC E A E

15. T H E LY P T E R I DAC E A E

19. D RYO P T E R I DAC E A E

16. A S P L E N I AC E A E

20. B L E C H NAC E A E

17. WO O D S I AC E A E (AT H Y R I AC E A E )

Order 9. S A LV I N I A L E S 21. A Z O L L AC E A E

Division

4. P I N O P H Y TA

Class 1. P I N O P S I D A Order 1. P I N A L E S 22. P I NAC E A E

24. C U P R E S S AC E A E

23. TA X O D I AC E A E

25. A R AU C A R I AC E A E

Class 2. TA X O P S I D A Order 1. TA X A L E S 26. TA X AC E A E

Division

5. M AG N O L I O P H Y TA

Class 1. M A G N O L I O P S I D A Subclass 1. M A G N O L I I D A E (DICOTYLEDONES) Order 1. M A G N O L I A L E S 27. M AG N O L I AC E A E

Order 2. L A U R A L E S 28. L AU R AC E A E

Order 3. A R I S T O L O C H I A L E S 29. A R I S TO L O C H I AC E A E

Conspectus of families

Order 4. N Y M P H A E A L E S 30. N Y M P H AC E A E

31. C E R ATO P H Y L L AC E A E

Order 5. R A N U N C U L A L E S 32. R A N U N C U L AC E A E

33. B E R B E R I DAC E A E

Order 6. PA PAV E R A L E S 34. PA PAV E R AC E A E

35. F U M A R I AC E A E

Subclass 2. H A M A M E L I D A E Order 1. H A M A M E L I D A L E S 36. P L ATA NAC E A E

Order 2. U RT I C A L E S 37. U L M AC E A E

39. M O R AC E A E

38. C A N NA BAC E A E

40. U RT I C AC E A E

Order 3. J U G L A N D A L E S 41. J U G L A N DAC E A E

Order 4. M Y R I C A L E S 42. M Y R I C AC E A E

Order 5. FA G A L E S 43. FAG AC E A E

45. C O RY L AC E A E

44. B E T U L AC E A E

Subclass 3. C A RY O P H Y L L I D A E Order 1. C A RY O P H Y L L A L E S 46. P H Y TO L AC C AC E A E

50. P O RT U L AC AC E A E

47. A I Z OAC E A E

51. BA S E L L AC E A E

48. C H E N O P O D I AC E A E

52. C A RYO P H Y L L AC E A E (I L L E C E B R AC E A E )

49. A M A R A N TAC E A E

Order 2. P O LY G O N A L E S 53. P O LY G O NAC E A E

Order 3. P L U M B A G I N A L E S 54. P L U M BAG I NAC E A E

xxiii

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Conspectus of families

Subclass 4. D I L L E N I I D A E Order 1. D I L L E N I A L E S 55. PA E O N I AC E A E

Order 2. T H E A L E S 56. E L AT I NAC E A E

57. C L U S I AC E A E (G U T T I F E R A E ; H Y P E R I C AC E A E )

Order 3. M A LVA L E S 58. T I L I AC E A E

59. M A LVAC E A E

Order 4. N E P E N T H A L E S 60. S A R R AC E N I AC E A E

61. D RO S E R AC E A E

Order 5. V I O L A L E S 62. C I S TAC E A E

65. F R A N K E N I AC E A E

63. V I O L AC E A E

66. C U C U R B I TAC E A E

64. TA M A R I C AC E A E

Order 6. S A L I C A L E S 67. S A L I C AC E A E

Volume 2. Order 7. C A P PA R A L E S 68. C A P PA R AC E A E

70. R E S E DAC E A E

69. B R A S S I C AC E A E (C RU C I F E R A E )

Order 8. E R I C A L E S 71. C L E T H R AC E A E

74. P Y RO L AC E A E

72. E M P E T R AC E A E

75. M O N OT RO PAC E A E

73. E R I C AC E A E

Order 9. D I A P E N S I A L E S 76. D I A P E N S I AC E A E

Order 10. P R I M U L A L E S 77. M Y R S I NAC E A E

78. P R I M U L AC E A E

Conspectus of families

Subclass 5. R O S I D A E Order 1. R O S A L E S 79. P I T TO S P O R AC E A E

83. C R A S S U L AC E A E

80. H Y D R A N G E AC E A E

84. S A X I F R AG AC E A E

81. E S C A L L O N I AC E A E

85. RO S AC E A E

82. G RO S S U L A R I AC E A E

Volume 3. Order 2. FA B A L E S 86. M I M O S AC E A E

88. FA BAC E A E

87. C A E S A L P I N I AC E A E

Order 3. P R O T E A L E S 89. E L A E AG NAC E A E

Order 4. H A L O R A G A L E S 90. H A L O R AG AC E A E

91. G U N N E R AC E A E

Order 5. M Y R TA L E S 92. LY T H R AC E A E

94. M Y RTAC E A E

93. T H Y M E L A E AC E A E

95. O NAG R AC E A E

Order 6. C O R N A L E S 96. C O R NAC E A E

Order 7. S A N TA L A L E S 97. S A N TA L AC E A E

98. V I S C AC E A E

Order 8. C E L A S T R A L E S 99. C E L A S T R AC E A E

100. AQ U I F O L I AC E A E

Order 9. E U P H O R B I A L E S 101. BU X AC E A E

102. E U P H O R B I AC E A E

Order 10. R H A M N A L E S 103. R H A M NAC E A E

104. V I TAC E A E

xxv

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Conspectus of families

Order 11. L I N A L E S 105. L I NAC E A E

Order 12. P O LY G A L A L E S 106. P O LY G A L AC E A E

Order 13. S A P I N D A L E S 107. S TA P H Y L E AC E A E

111. A NAC A R D I AC E A E

108. S A P I N DAC E A E

112. S I M A RO U BAC E A E

109. H I P P O C A S TA NAC E A E

113. RU TAC E A E

110. AC E R AC E A E

Order 14. G E R A N I A L E S 114. OX A L I DAC E A E

117. T RO PA E O L AC E A E

115. G E R A N I AC E A E

118. BA L S A M I NAC E A E

116. L I M NA N T H AC E A E

Order 15. A P I A L E S 119. A R A L I AC E A E

120. A P I AC E A E (U M B E L L I F E R A E )

Subclass 6. A S T E R I D A E Order 1. G E N T I A N A L E S 121. G E N T I A NAC E A E

122. A P O C Y NAC E A E

Order 2. S O L A N A L E S 123. S O L A NAC E A E

126. M E N YA N T H AC E A E

124. C O N VO LV U L AC E A E

127. P O L E M O N I AC E A E

125. C U S C U TAC E A E

128. H Y D RO P H Y L L AC E A E

Order 3. L A M I A L E S 129. B O R AG I NAC E A E

131. L A M I AC E A E (L A B I ATA E )

130. V E R B E NAC E A E

Order 4. C A L L I T R I C A L E S 132. H I P P U R I DAC E A E

133. C A L L I T R I C H AC E A E

Order 5. P L A N TA G I N A L E S 134. P L A N TAG I NAC E A E

Conspectus of families

Order 6. S C R O P H U L A R I A L E S 135. B U D D L E JAC E A E

139. G E S N E R AC E A E

136. O L E AC E A E

140. AC A N T H AC E A E

137. S C RO P H U L A R I AC E A E

141. L E N T I BU L A R I AC E A E

138. O RO BA N C H AC E A E

Volume 4. Order 7. C A M PA N U L A L E S 142. C A M PA N U L AC E A E

Order 8. R U B I A L E S 143. RU B I AC E A E

Order 9. D I P S A C A L E S 144. C A P R I F O L I AC E A E

146. VA L E R I A NAC E A E

145. A D OX AC E A E

147. D I P S AC AC E A E

Order 10. A S T E R A L E S 148. A S T E R AC E A E (C O M P O S I TA E )

Volume 5. Class 2. L I L I O P S I D A (MONOCOTYLEDONES) Subclass 1. A L I S M AT I D A E Order 1. A L I S M ATA L E S 149. BU TO M AC E A E

150. A L I S M ATAC E A E

Order 2. H Y D R O C H A R I TA L E S 151. H Y D RO C H A R I TAC E A E

Order 3. N A J A D A L E S 152. A P O N O G E TO NAC E A E

156. RU P P I AC E A E

153. S C H E U C H Z E R I AC E A E

157. NA JA DAC E A E

154. J U N C AG I NAC E A E

158. Z A N N I C H E L L I AC E A E

155. P OTA M O G E TO NAC E A E

159. Z O S T E R AC E A E

xxvii

xxviii

Conspectus of families

Subclass 2. A R E C I C A E Order 1. A R E C A L E S 160. A R E C AC E A E (PA L M A E )

Order 2. A R A L E S 161. A R AC E A E

1 6 2 . L E M NAC E A E

Subclass 3. C O M M E L I N I D A E Order 1. C O M M E L I N A L E S 163. C O M M E L I NAC E A E

Order 2. E R I O C A U L O N A L E S 164. E R I O C AU L O NAC E A E

Order 3. J U N C A L E S 165. J U N C AC E A E

Order 4. C Y P E R A L E S 166. C Y P E R AC E A E

167. P OAC E A E (G R A M I N E A E )

Order 5. T Y P H A L E S 168. S PA R G A N I AC E A E

169. T Y P H AC E A E

Subclass 4. Z I N G I B E R I D A E Order 1. B R O M E L I A L E S 170. B RO M E L I AC E A E

Subclass 5. L I L I I D A E Order 1. L I L I A L E S 171. P O N T E D E R I AC E A E

172. L I L I AC E A E (A L L I AC E A E ; A M A RY L L I DAC E A E ; T R I L L I AC E A E )

173. I R I DAC E A E

174. AG AVAC E A E

175. D I O S C O R E AC E A E

Order 2. O R C H I D A L E S 176. O R C H I DAC E A E

Division

5 . M AG N O L I O P H Y TA Cronquist, Takht. & W. Zimm.

Class

1 . M A G N O L I O P S I D A Cronquist, Takht. & W. Zimm.

Subclass 6. A S T E R I D A E Takht. Order 7. C A M PA N U L A L E S Lindl. Herbs. Leaves alternate, simple, exstipulate. Flowers actinomorphic or zygomorphic, bisexual, epigynous. Calyx 5-lobed. Corolla 5-lobed, sometimes 2-lipped, the upper 2-lobed, the lower 3-lobed. Stamens as many as the corolla lobes and alternate with them. Ovary with (1–)2–5 cells; ovules with axile placentation. Fruit a berry or capsule. Seeds with endosperm. Contains 7 families, 106 genera and about 2,500 species from all parts of the world. 142. C A M PA N U L AC E A E Juss. nom. conserv. Annual to perennial herbs, often with white latex. Leaves alternate, sometimes mostly basal, simple, sometimes petiolate, exstipulate. Inflorescence a simple or branched raceme, a corymbose raceme, or a congested spike or head, or flowers solitary. Flowers usually showy, actinomorphic or zygomorphic, bisexual, epigynous. Calyx with 5 lobes proximally fused into a tube. Corolla with 5 lobes proximally fused into a tube, the lobes more or less equal or with a 2-lobed upper and 3-lobed lower lip, often blue. Stamens 5, borne around the style-base on the receptacle, usually closely appressed around the style and free, or sometimes with the anthers or anthers and filaments fused laterally into a ring. Style 1; stigmas usually as many as ovary-cells, capitate to filiform. Ovary usually inferior, rarely superior, with (1–)2–5 cells, each cell with many ovules on axile placentas. Fruit a capsule opening variously, or a berry. Seeds with endosperm. Contains 84 genera and about 2,000 species, and occurs in most parts of the world. 1. Flowers in dense, flattish heads or globose to elongated spikes; corolla divided nearly to the base into linear lobes 2. 1. Flowers not in dense heads or globose to elongated spikes, or if so then the corolla divided not more than 3. two-thirds of the way to the base 2. Stems more or less glabrous; flowers in globose to elongated spikes without flowerless bracts at the base; 5. Phyteuma each flower with one bract; stigmas linear 2. Stems hairy; flowers in flattish heads with a conspicuous region of bracts without flowers at their base; flowers 6. Jasione without bracts; stigmas more or less globose 3. Corolla actinomorphic; filaments free, though often close together round the style and anthers sometimes fused 4. laterally 3. Corolla zygomorphic; filaments fused laterally at least 7. distally to form a tube round the style 4. Annual; ovary and capsules more than 3 times as long as 2. Legousia wide

4. Biennial to perennial; ovary and capsules less than 2(–3) 5. times as long as wide 5. Corolla tube less than 2 mm wide; styles more than 1.5 times as long as the corolla (tube and lobes) 4. Trachelium 5. Corolla tube more than 3 mm wide; style not or scarcely 6. longer than the corolla 6. At least the flowering stems erect or ascending, if all procumbent then not all the pedicels longer than the corolla; usually at least the upper leaves sessile or more or less so; capsule opening subapically or basally outside 1. Campanula the calyx 6. Stems filiform and procumbent, with solitary axillary flowers on erect pedicels much longer than the corolla; all leaves petiolate; capsule opening apically within the 3. Wahlenbergia calyx 7. Stems procumbent to decumbent, rooting at the nodes; 8. Pratia leaves broadly ovate to subrotund; fruit a berry 7. Stems erect to ascending, not rooting at the nodes; leaves 8. linear to obovate; fruit a capsule 8. Flowers and capsules pedicellate; ovary and capsules less 7. Lobelia than 15 mm, widening distally, 2-celled 8. Flowers and capsules sessile; ovary and capsules more 9. Downingia than 20 mm, cylindrical, 1-celled

Subfamily 1. Campanuloideae Flowers actinomorphic. Filaments and anthers free, or sometimes slightly laterally fused. 1. Campanula L. Marianthemum Schrank Biennial to perennial herbs. Stems ascending to erect or prostrate. Leaves alternate, the upper more or less sessile, exstipulate. Inflorescence 1- to few-flowered, in racemes or panicles, sometimes in a more or less compact head. Calyx-tube ovoid or subglobose; lobes 5, flat or folded at the sinus. Corolla usually blue, occasionally white, actinomorphic, divided up to half (to two-thirds) of the way to the base into 5 lobes, as long as or longer than calyx, rotate or campanulate. Stamens 5; filaments and anthers free. Style 1, shorter than to slightly longer than the corolla; stigmas 3–5, filiform. Ovary 3- to 5-celled. Fruit a capsule, usually less than twice as long as wide, ovoid or turbinate, dehiscing by subapical or basal pores. The flowers of Campanula species are strongly protandrous. Pollen is shed in bud and deposited on the hairs of the style, the stigmas at this stage being above the stamens. As the flower opens the stamens wither and the pollen is presented to insects which come for nectar, which is protected by the persistent, triangular bases of the stamens. The stigmas eventually separate and finally curl back so that if cross-pollination fails, self-pollination may occur as in the Asteraceae. About 300 species, chiefly in the North Temperate regions and the Mediterranean.

2

1 4 2 . C A M PA N U L AC E A E

Bailey, L. H. (1953). The garden of bellflowers in North America. New York. Crook, H. C. (1951). Campanulas. London, New York. Damboldt, J. (1976). Materials for a Flora of Turkey. XXXII. Campanulaceae. Notes Roy. Bot. Gard. Edinb. 35: 39–52. Gadella, T. W. J. (1964). Cytotaxonomic studies in the Genus Campanula. Wentia 11: 1–104. Grime, J. P. et al. (1988). Comparative plant ecology. London. [C. rotundifolia.] Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. Ser. 4, 13: 128–129, 351–352. [C. rotundifolia.] Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [C. patula.] 1. Calyx with 5 sepal-like appendages alternating with 2. 5 calyx lobes 3. 1. Calyx with 5 calyx lobes, but no extra appendages 2. Basal and lower cauline leaves cuneate to attenuate at 6. medium base 2. Basal and lower cauline leaves cordate at base 11. alliariifolia 12. glomerata 3. Flowers sessile 4. 3. Flowers with pedicels 4. Leaves not cordate at base; capsule with pores in apical 5. half 4. At least some leaves cordate at base; capsule with pores 8. at or near the base 3. lactiflora 5. Calyx lobes lanceolate to ovate, serrate 5. Calyx lobes linear to lanceolate, entire or with 1–2 small 6. basal teeth 6. Perennial, with non-flowering rosettes arising from rhizomes; corolla mostly more than 30 mm; stigma 4. persicifolia more than half as long as style 6. Usually biennial, without non-flowering rosettes; corolla mostly less than 30 mm; stigmas less than half as long 7. as style 7. Tap-root thin; basal leaves gradually narrowed to an indistinct petiole; inflorescence widely spreading 1. patula 7. Tap-root swollen and fleshy; basal leaves abruptly narrowed to a distinct petiole; inflorescence a narrowly 2. rapunculus pyramidal, racemose panicle 8. Calyx lobes linear to filiform, less than 1 mm wide at the 9. base 8. Calyx lobes linear-lanceolate, lanceolate or narrowly triangular-lanceolate to ovate-oblong or triangular11. ovate, more than 1 mm wide at the base 9. Middle cauline leaves lanceolate, ovate-oblong or narrowly so, rounded at base, crenate-serrate 16. rhomboidalis 9. Middle cauline leaves linear to linear-elliptical, very 10. gradually tapered to base, more or less entire 10. Leaves of stem mostly narrow and acute at apex; flowers several to numerous, 5–20 mm; capsules turbinate 17(a). rotundifolia subsp. rotundifolia 10. Leaves of stem blunter, the lower often like basal; flowers solitary, 20–30 mm; capsules more squat and 17(b). rotundifolia subsp. montana subrotund 11. Stems prostrate or decumbent to ascending or erect, 12. usually less than 50 cm 15. 11. Stems erect, usually more than 50 cm

13. 12. Corolla 25–40 mm 14. 12. Corolla 10–25 mm 13. Stems 15–50 cm; capsule opening by subapical pores 5. carpatica 13. Stems less than 15 cm; capsule opening by basal pores 15. fragilis 14. Leaves crenate-dentate; corolla funnel-shaped, divided one-quarter to two-fifths of the way to the base with 13. portenschlagiana erect to erecto-patent lobes 14. Leaves 2-serrate; corolla broadly funnel-shaped, divided half to three-quarters of the way to base with lobes spreading so that it appears star-shaped when you look 14. poscharskyana into its throat 15. Plant glabrous; inflorescence dense, narrowly pyramidal 10. pyramidalis or cylindrical; capsules erect 15. Plant hairy; inflorescence open and racemose; capsules 16. nodding 16. Plant forming patches, with shoots arising from extensively creeping rhizomes; calyx lobes deflexed or 9. rapunculoides patent at anthesis 16. Plant tufted, without rhizomes; calyx lobes erect or 17. erecto-patent at anthesis 17. Stems bluntly angled, glabrous or with few soft hairs; 7. latifolia leaves irregularly 1–2 serrate 17. Stems sharply angled; with few subrigid hairs; leaves coarsely dentate or irregularly crenate-serrate 8. trachelium

Subgenus 1. Rapunculus (Dumort.) Kharadze Calyx without appendages between lobes. Capsule dehiscing by 3 subapical pores or valves. 1. C. patula L. Spreading Bellflower Biennial herb with a thin, slender tap-root. Stems 20–70 cm, pale yellowish-green, erect or ascending, angled, striate, with short to medium, pale, rough simple eglandular hairs, branched in the upper half. Leaves alternate, numerous; lamina pale to bright yellowish-green on upper surface, paler beneath, the basal 2–5 × 1–2 cm, obovate or obovateoblong, more or less obtuse at apex, undulate-crenate, narrowed below to an indistinct petiole and withering before flowering, the cauline smaller, oblong, linear-lanceolate or linear, obtuse to acute at apex, shallowly crenate to entire, and narrowed to a sessile base; all glabrous or with few to numerous, short, stiff, pale simple eglandular hairs especially on the margin and midrib beneath. Inflorescence broadly paniculate, with long, spreading branches; pedicels up to 25 mm, slender, glabrous; bracteoles linear. Flowers 30–40 mm in diameter, erect. Calyx 6–22 mm, medium green, broadly funnel-shaped, glabrous, divided half to twothirds of the way to the base; lobes 5, linear to linearlanceolate, acute at apex, erect in fruit, without appendages. Corolla 17–25 mm, pale to deep bluish-violet or rarely white, broadly campanulate, divided to about half of the way to the base; lobes 5, ovate or elliptical-ovate, acute at apex, spreading, glabrous. Stamens 5; filaments 6–8 mm, yellow, greenish at base; anthers yellow. Style 1, yellow; stigmas 3, yellow, less than half as long as style, becoming separated and curled back. Capsule 8–11 mm, obconical,

1. Campanula erect, glabrous, dehiscing by 3 subapical pores; seeds 0.5– 0.6 mm, pale brown, oblong. Flowers 7–9. 2n = 20. Native. Open woods, wood-borders, rocky outcrops and hedgebanks. Locally native in southern Britain north to Shropshire, especially in the Welsh border area, formerly in Yorkshire; steadily decreasing. Often a naturalised garden escape. Most of Europe east to central Russia. A member of the European Temperate element with a continental distribution; it is a variable species which is widely naturalised outside its native range. Our plant is subsp. patula. 2. C. rapunculus L. Rampion Bellflower C. lambertiana DC.; C. calycina Boeber ex Roem. & Schult.; C. virgata DC., non Labill.; C. esculenta Salisb. nom. illegit. Biennial herb with a swollen, fleshy, napiform tap-root. Stems 20–80(–100) cm, yellowish-green, erect, angled, striate, glabrous or with short to medium, pale, rough simple eglandular hairs especially below, unbranched or with few, short branches, leafy. Leaves alternate; lamina dull medium yellowish-green on upper surface, paler beneath; the basal 3–10 × 1.5–5.0 cm, obovate to elliptical, obtuserounded to acuminate at apex, entire to crenate-dentate, the teeth rounded and abruptly narrowed to the short, glabrous or hairy petiole, the cauline linear-lanceolate to linearoblanceolate, obtuse to acute at apex, more or less entire, rounded or narrowed at base and sessile; all with few to numerous, short to medium, pale, subrigid simple eglandular hairs on both surfaces and the margins. Inflorescence a narrowly pyramidal, racemose panicle with short branches below; pedicels up to 35 mm, slender, glabrous or with a few hairs; bracteoles linear, at base of pedicel. Flowers 20–30 mm in diameter, erect and arching. Calyx 10–18 mm, medium green, funnel-shaped, glabrous, divided about twothirds to three-quarters of the way to the base; lobes 5, narrowly linear to linear-lanceolate, acute at apex, erect, without appendages. Corolla 10–25 mm, pale blue or white, funnel-shaped, divided about half of the way to the base; lobes 5, triangular-ovate, acute at apex, curved outwards. Stamens 5; filaments 7–8 mm, yellow; anthers yellow. Style 1, yellow; stigmas 3, yellow, less than half as long as style. Capsule 5–6 mm, obconical, erect, dehiscing by 3 subapical pores; seeds about 0.5 mm, pale brown, ellipsoid. Flowers 7–8. 2n = 20. Introduced. Formerly cultivated as a salad vegetable, both the carrot-like root and the shoots being edible; also grown for ornament. Naturalised in rough, grassy fields and hedgebanks, usually on gravelly soils. Scattered localities through Great Britain, especially in northern England and Scotland. Throughout Europe from Holland southwards; Asia east to Iran; North Africa. 3. C. lactiflora M. Bieb. Milky Bellflower C. biserrata K. Koch; C. celtidifolia Boiss. & A. Huet. Perennial herb with a branched, fleshy rootstock. Stems up to 150 cm, pale green, sometimes slightly tinted brownishpurple, thick, striate, erect, with rigid, pale, deflexed simple eglandular hairs which are numerous below but fewer above, branched in upper part, very leafy. Leaves alternate; lamina 2–9 × 0.7–3.5 cm, bright yellowish-green on upper

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surface, paler beneath, ovate, lanceolate, elliptical, obovate, oblong or ovate-oblong, acute at apex, serrate or slightly biserrate, the teeth narrow, narrowed or rounded at base, the basal shortly petiolate, the cauline sessile and semiamplexicaul, all sparsely clothed with short, pale simple eglandular hairs especially on the margins and veins beneath. Inflorescence broadly paniculate, leafy; pedicels up to 20 mm, pale green, with medium, pale simple eglandular hairs; bracteoles linear. Flowers 25–40 mm in diameter, erect. Calyx 11–16 mm, dull greyish-violet, divided to about half of the way to the base; lobes 5, lanceolate to ovate, gradually narrowed but obtuse at apex, serrate, without appendages; glabrous or with subrigid, medium to long, pale simple eglandular hairs on the veins of the tube and margins of the lobes. Corolla 15–30 mm, pale blue or violet, slightly paler towards the base, sometimes white or pink, broadly campanulate, divided half to one-third of the way to the base; lobes 5, ovate, acute at apex, glabrous. Stamens 5; filaments whitish; anthers yellowish. Style 1, short; stigmas 3, as long as style. Capsule 7–8 × about 6 mm, obconical, erect, glabrous or with subrigid, pale simple eglandular hairs, dehiscing by 3 subapical pores; seeds about 1.0 × 0.6 mm, pale brown, with a pale narrow margin, oblong, flattened. Flowers 7–10. 2n = 34, 36. Introduced. Grown for ornament and naturalised by streams, waysides and rough and waste ground. In scattered localities in Great Britain, especially northern England and Scotland. Native of forests, scrub and subalpine meadows in north-east Turkey, Caucasia and north-western Iran. 4. C. persicifolia L. Peach-leaved Bellflower Perennial herb with slightly creeping, slender, muchbranched rootstock and non-flowering rosettes arising from rhizomes. Stems 20–80 cm, pale yellowish-green, erect, slender and rather wiry, glabrous, usually unbranched, leafy. Leaves alternate; lamina shining medium green with pale midrib on upper surface, paler beneath and midrib prominent, the basal 7–15 × 1–3 cm, lanceolate to obovate, obtuse at apex, crenulate and narrowed at base, the petioles up to 60 mm, the cauline much smaller, linear-lanceolate to subulate, more or less acute at apex, crenulate and narrowed at base, the lower shortly petiolate, the rest sessile, all glabrous. Inflorescence a narrow, few-flowered raceme; pedicels short, slender, glabrous; bracteoles linear. Flowers 30–50(–66) mm in diameter, suberect. Calyx 10–20 mm, medium green, glabrous, divided to the base; lobes 5, linearlanceolate, acute at apex, without appendages. Corolla 25– 50 mm, blue, rarely white, broadly campanulate, divided for about one-quarter to half of the way to the base; lobes 5, broadly triangular-ovate, more or less acute at apex. Stamens 5; filaments 4–6 mm, whitish; anthers whitish or pale yellow. Style 1, whitish; stigmas 3, whitish or yellowish, more than half as long as style. Capsule 6–7 mm, ovoidconical, erect, opening by 3 subapical pores; seeds 0.8–1.0 mm, shiny brown, ovoid. Flowers 6–8. 2n = 16. Introduced. Naturalised in waste and rough ground and grassy places and banks. Scattered localities throughout Great Britain. Native of most of Europe except the extreme north, the islands and parts of the west; western Asia.

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5. C. carpatica Jacq. Tussock Bellflower Perennial herb with white, fibrous roots. Stems 15–50 cm, pale green, sometimes suffused brownish-purple, ascending or decumbent, slightly angled, glabrous or with short to medium, pale simple eglandular hairs, much branched, leafy. Leaves alternate; lamina medium bright green on upper surface, paler beneath, the basal 1–4 × 0.5–3.0 cm, ovate or subrotund, obtuse to acute at apex, irregularly serrate-dentate, cordate at base, and with long petioles, the cauline gradually decreasing in size upwards, ovate to lanceolate, acute or acuminate at apex, irregularly crenate-serrate and rounded, truncate or more or less cordate at base, the uppermost linear, all glabrous or with few, short, stiff simple eglandular hairs, all petiolate, the petioles mostly long and slender and glabrous or with few simple eglandular hairs. Flowers 35–55 mm in diameter, erect, solitary at the ends of stems and branches; pedicels long, erect, glabrous or with occasional, pale simple eglandular hairs; bracteoles linear. Calyx 16–25 mm, dull green, divided half to twothirds of the way to the base; lobes 5, linear-lanceolate, drawn out to a very narrow, acute apex, entire or with very small teeth, without appendages. Corolla 25–40 mm, pale blue, rarely white, broadly tubular-rotate, divided onethird to half of the way to the base; lobes 5, very broadly ovate, rounded-mucronulate at apex. Stamens 5; filaments 12–20 mm, pale with greenish base; anthers yellow. Style 1, greenish; stigmas 3, greenish. Capsule 6–8 mm, ovoidcylindrical, dehiscing by 3 subapical pores; seeds about 1.5 mm, pale yellow, ovate, compressed. Flowers 6–8. 2n = 34. Introduced. Garden escape established on walls and in paving at Yeoman’s Row in Middlesex and a casual elsewhere. Native of the Carpathians. Subgenus 2. Campanula Calyx with or without appendages. Capsule dehiscing by 3 basal pores. 6. C. medium L. Canterbury Bells Marianthemum medium (L.) Schur Biennial herb with a stout tap-root. Stems up to 60(–90) cm, yellowish-green, sometimes suffused brownish-purple, erect, stout, markedly striate, with numerous, pale, subrigid simple eglandular hairs, more or less branched at least in the upper part, leafy. Leaves alternate, numerous; lamina dull medium or pale yellowish-green on upper surface, paler beneath, the basal forming a rosette in the first year, 7–11 × 7–11 cm, ovate or ovate-oblong, or narrow to broadly elliptical, obtuse at apex, irregularly crenate-dentate or serrate, and attenuate at base to a short or long petiole; the cauline numerous, gradually smaller upwards, ovate-lanceolate to lanceolate or oblanceolate, obtuse at apex, crenate-dentate to subentire, and narrowed or rounded at the sessile base, all with few to numerous, stiff, pale, short simple eglandular hairs at least on the margins and veins beneath where they are sometimes dense. Inflorescence a narrow panicle of axillary and terminal flowers; pedicels up to 20 mm, with numerous medium and long, rigid, pale simple eglandular hairs; bracteoles lanceolate. Flowers 40–60 mm in diameter, sometimes more or less double, erect, solitary. Calyx 15–

25 mm, medium green, with medium to long, stiff, pale simple eglandular hairs particularly on the lobe margins, divided two-thirds of the way to the base; lobes 5, lanceolate to narrowly ovate, obtuse at apex, with broadly ovate, obtuse, deflexed appendages between the lobes. Corolla 30–60(–70) mm, violet-blue, purple, pink or white, campanulate, shortly 5-lobed, inflated in the middle, glabrous. Stamens 5; filaments 20–25 mm, whitish, ciliate; anthers yellow. Style 1, whitish; stigmas 5, cream. Capsule about 15 mm, ovoid, nodding; seeds about 2 mm, pale brown, ovoid. Flowers 5–6. 2n = 34. Introduced. Formerly much grown in gardens. Casual and naturalised in waste and rough ground and grassy places and on banks. Native of Italy and south-east France. 7. C. latifolia L. Giant Bellflower Tufted perennial herb with a short, thickened, somewhat woody rootstock with fibres. Stem 50–120 cm, pale green, erect, stout, obscurely and bluntly angled, glabrous or with few, pale, retrorse, soft simple eglandular hairs, unbranched, leafy. Leaves alternate; lamina medium green on upper surface, paler beneath, the basal 10–20 × 2.5–3.5 cm, ovate to ovate-oblong, acute at apex, irregularly 1–2 serrate, rounded or rarely cordate at the base which is usually decurrent on the petiole, withered at flowering, the cauline gradually smaller, ovate, long-acute at apex, irregularly serrate or the uppermost nearly entire and sessile or the lower petiolate, all glabrous or with stiff simple eglandular hairs especially on the veins. Inflorescence a raceme, sometimes with short branches below; pedicels up to 20 mm; bracteoles leaf-like. Flowers 35–45 mm in diameter, suberect or inclined, solitary and axillary. Calyx 20–32 mm, medium green, glabrous or puberulent at base, divided half to two-thirds of the way to the base; lobes 5, narrowly triangular-lanceolate, acute at apex, erect or erecto-patent at anthesis, without appendages. Corolla (35–)40–55 mm, bluish-purple, rarely white, campanulate, hairy inside, divided one-third to half of the way to the base; lobes 5, triangular-ovate, acute at apex. Stamens 5; filaments short, creamy-yellow; anthers creamy-yellow. Style 1, creamy-yellow; stigmas 3, yellow. Capsule 12–15 mm, ovoid, nodding, opening by 3 basal pores; seeds 1.5–2.0 mm, pale yellow, ovate, compressed. Flowers 7–8. 2n = 34, + 0–5B. Native. Rich, often damp, mainly calcareous woods and hedgebanks. Most of Great Britain, but rare to absent in southern England and northern Scotland; north-east Ireland where it may be introduced. Most of Europe except the Mediterranean region and much of the south-west; western Asia east to Iran and western Siberia. A member of the European Temperate element. 8. C. trachelium L. Nettle-leaved Bellflower C. urticaefolia Salisb., non Turra Tufted perennial herb with a short, thickened, woody rootstock with fleshy fibres. Stems 50–100 cm, pale green, erect, sometimes flecked reddish, stout, sharply angled, with sparse, subrigid, retrorse simple eglandular hairs, sometimes slightly branched, leafy. Leaves alternate, the lamina medium to deep green on upper surface, much paler beneath, the basal 8–10 × 6–7 cm, narrowly to broadly

1. Campanula ovate, narrowed to a rounded or subacute apex, coarsely dentate or irregularly crenate-serrate and deeply cordate at base, the petiole up to 120 mm and hairy, the cauline gradually becoming smaller up the stem, ovate to ovateoblong or lanceolate, more or less acute at apex, irregularly and coarsely 1- to 2-serrate and cuneate or truncate at base, the upper sessile, the lower petiolate, all with short to medium, pale, bristly simple eglandular hairs especially on the veins and margins. Inflorescence a racemose, leafy panicle with short branches bearing 1–4 flowers; pedicels up to 10 mm, recurved or erect, glabrous; bracteoles linear. Flowers 30–35 mm in diameter, erect, terminal and in the axils of leaves. Calyx 13–15 mm, medium green, the tube bristly hairy, divided one-third to half of the way to the base; lobes 5, triangular-ovate, acute at apex, erect or erecto-patent at anthesis; without appendages. Corolla 25– 35 mm, bright bluish-purple, rarely white, broadly campanulate, hairy inside, divided one-third of the way to the base; lobes 5, triangular-ovate, acute at apex, suberect. Stamens 5; filaments short, white; anthers creamy-yellow. Style 1, white; stigmas 3, yellow. Capsule 6–8 mm, ovoid, nodding, opening by 3 basal pores; seeds 0.5–0.6 mm, yellow, ovate, compressed. Flowers 7–9. 2n = 34. Native. Woods and hedgebanks, usually on clayey soils. Frequent in Great Britain north to Lincolnshire and North Wales and in south-east Ireland. It escapes from gardens and becomes well naturalised elsewhere in Britain and Ireland. Most of Europe to about 62◦ 30 N in Sweden; Turkey, Iran and western Siberia; North Africa. A member of the European Temperate element. Our plant is subsp. trachelium which occurs throughout the range of the species. 9. C. rapunculoides L. Creeping Bellflower C. rigida Stokes nom. illegit. Perennial herb forming patches, with elongated, slender, extensively creeping rhizomes producing numerous adventitious buds and fibrous roots. Stems 30–80(–100) cm, medium green, often suffused purplish, erect, rather stout, subterete, glabrous or with short, stiff simple eglandular hairs, sometimes slightly branched, leafy. Leaves alternate; lamina dull medium green on upper surface, paler beneath, the basal 5–9 × 3.0–5.5 cm, ovate to oblong-ovate, obtuse to subacute at apex, serrate or dentate, cordate or rounded at base, the petiole up to 13 cm, often purplish and glabrous, the cauline getting gradually smaller up the stem, ovate to lanceolate, more or less acute at apex, dentate and rounded at the sessile base, all with few, short, pale simple eglandular hairs, especially on the veins. Inflorescence a secund raceme or panicle; pedicels about 5 mm, recurved, with short, stiff hairs; bracteoles linear. Flowers 25–30 mm in diameter, drooping, terminal and in the axils of leaves. Calyx 10–12 mm, medium green, with short, stiff, appressed, deflexed simple eglandular hairs, divided about half of the way to the base; lobes 5, linear-lanceolate, acute at apex, deflexed or patent at anthesis; without appendages. Corolla 20–30 mm, bluish-purple, funnel-shaped, divided half to one-third of the way to the base; lobes 5, triangular-ovate, acute at apex, ciliate, spreading. Stamens 5; filaments short, white; anthers yellow. Style 1, pink; stigmas 3, pink. Capsule 6–7 mm, hemispherical, nodding, opening by 3 basal pores; seeds

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1.5–1.6 mm, yellowish-brown, ovate, compressed. Flowers 7–9. 2n = 68, 102. Introduced. Grown in gardens and naturalised in fields, woods, banks and rough ground; very persistent. In mown turf it often persists as vegetative plants. Widely scattered records in Great Britain and Ireland. Native in most of Europe but rare in the Mediterranean region and absent from the islands; western Asia; Caucasus; naturalised in North America and elsewhere. 10. C. pyramidalis L. Chimney Bellflower Perennial herb with a thick, napiform root. Stems 30– 150 cm, pale yellowish-green, erect, robust, glabrous. Leaves alternate; lamina shining yellowish-green on upper surface, paler beneath, striate, leafy, the basal 4–8 × 2– 4 cm, ovate to ovate-oblong, acute to obtuse at apex, crenate-dentate or crenate-serrate, more or less cordate at base and long-petiolate, and the cauline ovate to ovatelanceolate, obtuse to subacute at apex, crenate-dentate to crenate-serrate and cuneate to truncate at base, the lower and median petiolate, the upper sessile, all glabrous. Inflorescence a dense, numerous-flowered, narrowly pyramidal or cylindrical raceme; pedicels short, glabrous; bracteoles linear. Flowers 20–30 mm in diameter, erect. Calyx 8–9 mm, green, glabrous, divided half of the way to the base; lobes 5, narrowly triangular-lanceolate, obtuse to subacute at apex, patent or reflexed; without appendages. Corolla 10–30 mm, purplish-blue, rarely white, broadly campanulate, lobed about half of the way to the base; lobes 5, broadly ovate, subacute at apex. Stamens 5; filaments 5–6 mm, white; anthers yellow. Style 1, whitish; stigmas 3, short. Capsule 6–8 mm, subglobose, trisulcate, erect, opening by 3 basal pores; seeds almost 1 mm, ovoid, slightly compressed. Flowers 7–8. 2n = 34. Introduced. Grown in gardens and naturalised on walls. Guernsey in the Channel Islands and West Malling in Kent, a rare casual elsewhere. Native of northern Italy and the north-west part of Balkan peninsula. 11. C. alliariifolia Willd. Cornish Bellflower C. lamifolia Adans.; C. macrophylla Sims Perennial herb. Stems up to 70 cm, pale green, sometimes suffused brownish-purple, erect, robust, striate, with numerous, short to medium, pale, soft simple eglandular hairs, leafy, simple or branched. Leaves alternate; lamina medium green on upper surface, grey to whitish beneath, the basal 5–8 × 5–8 cm, triangular-ovate or ovate, obtuse to acute at apex, bicrenate, the teeth shallow and rounded, cordate at base and long-petiolate, the cauline rapidly decreasing in size and similar to basal but the lower petiolate and the upper sessile, all with numerous, short, pale, appressed simple eglandular hairs on the upper surface and numerous to dense, short, pale simple eglandular hairs below which are pectinate on the veins. Inflorescence a spike of axillary flowers on short, hairy pedicels; bracteoles linear. Flowers 20–30 mm in diameter, more or less nodding, solitary or rarely 1–3. Calyx 12–14 mm, medium green, divided for about two-thirds of the way to the base; lobes 5, lanceolate to ovate, acute at apex, alternating with the lanceolate, acute, reflexed appendages, with short to medium, pale, stiff

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hairs on both calyx and appendages particularly on the margins of lobes and appendages. Corolla 20–40 mm, white, rarely purplish, divided one-quarter to one-third of the way to the base, shortly hairy outside; lobes 5, ovate, acute at apex, long hairy on margins. Stamens 5; filaments 8–10 mm; anthers yellow. Style 1, yellow; stigmas 3, not exceeding corolla. Capsule about 8 mm, obconical, ribbed, shortly hairy, opening by 3 basal pores; seeds about 2 × 1 mm, brown with a pale margin, ellipsoid. Flowers 6–9. 2n = 34, 68, 96. Introduced. Naturalised garden escape on waste and rough ground and particularly railway banks. Mostly south and south-west England, particularly Cornwall, and increasing. Native of northern Turkey and Caucasus. 12. C. glomerata L. Clustered Bellflower Gentiana collina With. Perennial herb with a short, oblique, woody rootstock. Stem 3–25(–75) cm, pale yellowish-green, often tinted brownishred, erect, slightly angular, more or less robust, with numerous, short to medium, pale simple eglandular hairs, usually unbranched, leafy. Leaves alternate; lamina dull, medium green, paler beneath, the basal 1–6(–11) × 0.5–3.5 cm, lanceolate to ovate, obtuse at apex, irregularly crenateserrulate and cordate at base, the petioles up to 15 cm and hairy; the cauline smaller, lanceolate to ovate, obtuse to acute at apex, crenate-serrulate or the upper entire, narrowed or rounded at base and sessile and semiamplexicaul or the lower shortly petiolate, all with short to medium, pale simple eglandular hairs on both surfaces and the margins. Inflorescence subcapitate, often with several, more or less distant flowers or short, few-flowered branches below the terminal head, with large bracts. Flowers 18–22 mm in diameter, erect, sessile. Calyx 8–15 mm, medium green, more or less hairy, divided over half of the way to the base; lobes 5, triangular-ovate, acute at apex, without appendages. Corolla 12–25 mm, bright bluish-purple, rarely white, divided nearly half of the way to the base; lobes 5, triangular-ovate, acute at apex, erect, but eventually spreading. Stamens 5; filaments short; greenish-yellow, anthers greenish-yellow. Style 1, purplish-brown; stigmas 3, yellow. Capsule 6–8 mm, erect, ovoid, opening by 3 basal pores; seeds 0.5–1.0 mm, pale brown, ovate, compressed. Flowers 5–9. 2n = 30. Varies greatly in size. It is not known if the very dwarf var. nana C. Bailey, of chalk downs, would retain its habit in cultivation. Native. Chalk and limestone grassland, scrub and open woodland, cliffs and dunes by the sea; also a casual or naturalised garden escape in rough ground. Mainly in the south and east part of Great Britain, where it is locally common, north to central and east Scotland. Most of Europe and temperate Asia. A member of the Eurasian Temperate element with a continental distribution in Europe. Our native plant is subsp. glomerata which occurs throughout the range of the species. Garden escapes and some plants grown from wild flower seed are usually larger and may belong to var. dahurica Fisch. & Ker-Gawl., but there seems to be a whole range of intermediates.

13. C. portenschlagiana Schult. Adria Bellflower Perennial herb. Stems usually numerous, 15–20 cm, pale yellowish-green, sometimes tinted brownish-purple, ascending, procumbent or trailing, striate, glabrous or with very short, white simple eglandular hairs in the upper part. Leaves alternate; lamina medium green on upper surface, paler beneath, the basal 1.0–2.5 × 1.0–3.0 cm, subrotund or reniform, broadly rounded at apex, crenate-dentate and cordate at base, with petioles up to 120 mm, the cauline similar but smaller and short to long petiolate, all glabrous or with very short, pale simple eglandular hairs especially on the veins. Inflorescence lax, branched; pedicels short to rather long, glabrous or with very short, white simple eglandular hairs; bracteoles linear. Flowers 10–20 mm in diameter, numerous. Calyx 5–7 mm, green, divided nearly to base, glabrous or with very short, white simple eglandular hairs; lobes 5, linear-lanceolate or lanceolate, subacute at apex, erect to erecto-patent; without appendages. Corolla 15– 25 mm, violet-blue, funnel-shaped, divided one-quarter to two-fifths of the way to the base; lobes 5, triangularlanceolate, acute at apex, erect to erecto-patent. Stamens 5; filaments short, whitish; anthers whitish or yellowish. Style 1, purplish; stigma pale yellow. Capsule 5–7 mm, opening by 3 basal pores; seeds 1–2 mm, pale brown, ellipsoid. Flowers 6–12. 2n = 34. Introduced. Much grown on walls and rockeries in gardens; naturalised on walls and rocky banks. Scattered localities in Great Britain, mostly in the centre and south and in the Channel Islands. Probably over-recorded for C. poscharskyana. Native of Yugoslavia. Hybrids with C. poscharskyana, intermediate between the parents, occur in gardens and may escape. Named after Franz von Portenschlag-Ledermermeyer (1772–1822). 14. C. poscharskyana Degen Trailing Bellflower Perennial herb. Stems usually numerous, 15–20(–30) cm, pale yellowish-green, sometimes tinted brownish-purple, ascending, procumbent or trailing, striate, glabrous or with a few, short pale simple eglandular hairs. Leaves alternate; lamina medium green on upper surface, paler beneath, the basal 3.5–5.5 × 3.5–5.0 cm, ovate, more or less acute at apex, 2-serrate, cordate at base with petioles up to 150 mm, the cauline similar but usually getting gradually smaller upwards and short to long petiolate, all with numerous, short, pale, stiff simple eglandular hairs at first, particularly on the margins and veins, becoming glabrous. Inflorescence lax, branched; pedicels short to medium, slender, glabrous or with short, white, stiff simple eglandular hairs; bracteoles linear. Flowers 20–28 mm in diameter, erect, numerous. Calyx 5–7 mm, green, tinted reddish, divided nearly to base, with short to medium, pale, stiff simple eglandular hairs especially on the margins; lobes 5, lanceolate, acute at apex, spreading or reflexed; without appendages. Corolla 10–25 mm, dirty mauvish-blue and paler at base, broadly funnelshaped, divided about half to three-quarters of way to the base; lobes 5, triangular-ovate, more or less acute, spreading and thus appearing star-shaped as you look into the corolla. Stamens 5; filaments short, white; anthers cream. Style 1,

1. Campanula pale at base, purplish above; stigmas 3, purplish. Capsule 5–7 mm; opening by 3 basal pores; seeds 1.5–2.0 mm, pale brown, ellipsoid. Flowers 6–8. 2n = 34. Introduced. Much grown on walls and rockeries in gardens; naturalised on walls and rocky banks. Scattered localities in Great Britain, mostly in the centre and south and in the Channel Islands. Native of Yugoslavia. Named after Gustav Adolf Poscharsky (1832–1915). 15. C. fragilis Cirillo Italian Bellflower C. diffusa Vahl Perennial herb with a rhizome, woody stock and nonflowering shoots. Stems 7–15 cm, pale green, often suffused purplish, diffuse-ascending, striate, flexuous, glabrous or with few to numerous, short, white, rigid simple eglandular hairs, branched, leafy. Leaves alternate; lamina pale green on upper surface, paler beneath, the basal 0.5–2.5 × 0.5– 2.5 cm, ovate, subrotund or reniform, narrowly to broadly rounded at apex, crenate to serrate, rounded to cordate at base and with glabrous or hairy petioles up to 40 mm, the cauline numerous, similar to basal but smaller and petiolate, all glabrous or with few to numerous, short, white, rigid simple eglandular hairs. Inflorescence lax and corymbose; pedicels short, glabrous or hairy; bracteoles linear. Flowers 25–40 mm in diameter, erect, terminal or in the axils of leaves. Calyx 8–16 mm, medium green, glabrous or with few to numerous, pale, rigid simple eglandular hairs, divided nearly to the base; lobes 5, linear-lanceolate, long-acute or acuminate at apex, without appendages. Corolla 25–40 mm, pale blue to purplish-blue or white, broadly campanulate to rotate, divided about half of the way to the base; lobes 5, broadly ovate, subacute at apex, spreading. Stamens 5; filaments 4–6 mm, white; anthers yellow. Style 1, long and slender, cream; stigmas 3, cream. Capsule 6–7 mm, erect, ovoid, opening by 3 basal pores; seeds 0.2 mm, pale yellow, ovate, compressed. Flowers 6–8. 2n = 32, 34. Very variable in hairiness, leaf-shape and flower colour. Two subspecies have been recognised. Subsp. fragilis has the basal leaves subrotund-cordate, obtusely dentate to crenate, the calyx teeth 9–12 mm and the corolla 35–40 mm in diameter. Subsp. cavolinii (Ten.) Dambolt (C. cavolinii Ten.) has the basal leaves ovate, cordate and serrate, the calyx teeth 8–15 mm and the corolla 25–30 mm in diameter. Hairy and glabrous forms occur in both subspecies and gardeners have selected colour forms and large-flowered forms. Introduced. Occurs on a wall at St Peter Port in Guernsey in the Channel Islands where it is increasing. Elsewhere it is sometimes on garden paths and walls. Native of limestone rocks in central and south Italy. 16. C. rhomboidalis L. Broad-leaved Harebell C. rotundifolia subsp. rhomboidalis (L.) Bonnier Perennial herb with a napiform root and slender, sparingly branched rhizome. Stems (10–)20–40(–60) cm, yellowishgreen, often tinged with purple, erect, striate, glabrous or with short or medium, pale simple eglandular hairs, branched only above, leafy. Leaves alternate; lamina medium green on upper surface, paler beneath, the basal

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absent at anthesis, 1.5–2.5 × 0.7–2.0 cm, subrotund or reniform, rounded at apex, serrate, the teeth broad, cordate at base and petiolate, the cauline few to numerous, gradually decreasing in size upwards, ovate, lanceolate, ovateoblong or rhomboid, more or less acute or acuminate at apex, crenate-serrate, rounded or more or less cordate at base and sessile, with short, pale simple eglandular hairs, particularly on the margin and veins beneath. Inflorescence narrow and few-flowered; pedicels pale, striate, glabrous; bracteoles linear. Flowers 25–30 mm in diameter, erect. Calyx 10–12 mm, medium green, glabrous, divided about two-thirds of the way to the base; lobes 5, linear or filiform, acute at apex, spreading; without appendages. Corolla (12–) 16–25 mm, pale to bright blue, campanulate, divided onequarter to one-third of the way to the base; lobes 5, broadly ovate, obtuse-mucronulate at apex. Stamens 5; filaments short, whitish; anthers yellow. Style 1, violet-blue; stigmas 3, yellow. Capsule 6–7 mm, turbinate, opening by 3 basal pores, membranous, nodding; seeds about 1.5 mm, yellow, ovate, compressed. Flowers 6–7. 2n = 34. Introduced. A garden escape established on the wooded bank of the River Esk near Langholm in Dumfries-shire and on the bank of a sunken lane near Knock in Westmorland. Native of the south-west and central Alps and the Jura, and locally naturalised elsewhere in Europe. 17. C. rotundifolia L. Harebell C. minor Lam. nom. illegit.; C. linifolia L.; C. variifolia Salisb. nom. illegit.; C. heterophylla Gray, non L. Perennial herb with slightly creeping, very slender, elongated, much branched rootstock which produces adventitious buds. Stems 15–50(–60) cm, pale green, sometimes tinted brownish-purple, slender, wiry, erect but often decumbent at base, glabrous or with occasional hairs, simple or branched, leafy. Leaves alternate; lamina dark green, paler beneath, the basal 0.5–1.5 × 0.5–1.5 cm, with some present at anthesis, ovate to subrotund, rounded or acute at apex, crenate or crenate-dentate, truncate, rounded or cordate at base, the slender and glabrous petioles up to 6.0 cm, the cauline 15–35 × 0.7–1.5 mm, with the lower similar to basal and middle and upper linear to linear-elliptical, obtuse to acute at apex, more or less entire and narrowed to a sessile base with the middle ones intermediate, all glabrous or nearly so. Inflorescence a more or less branched panicle of few to numerous flowers, or reduced to a solitary terminal flower; pedicels very slender; bracteoles linear. Flowers 10–22 mm in diameter, faintly sweet-smelling, erect in bud, nodding in flower. Calyx 6–8 mm, medium green, glabrous, divided for about two-thirds of the way to the base; lobes 5, linear to filiform, acute at apex, spreading; without appendages. Corolla 5–30 mm, purplish-blue, rarely white, broadly campanulate, divided for one-quarter to one-third of the way to the base; lobes 5, broadly ovate, subacute at apex. Stamens 5; filaments 1.5–2.0 mm, cream; anthers cream. Style 1, bluish-purple; stigmas 3, greenish-yellow. Capsule 3–5 mm, turbinate to subglobose, nodding, opening by 3 basal pores; seeds about 0.2 mm, ovate, compressed, yellow. Flowers 7–9. Visited by beetles.

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(a) Subsp. rotundifolia Leaves of stem mostly narrow and acute at apex. Flowers several to numerous, 5–20 mm. Capsules turbinate. 2n = 68. (b) Subsp. montana (Syme) P. D. Sell C. rotundifolia var. montana Syme; C. giesekiana auct.; C. rotundifolia var. speciosa A. G. More; C. rotundifolia var. uniflora auct. Leaves of stem blunter, the lower often like basal. Flowers often solitary, 20–30 mm. Capsules more squat and subglobose. 2n = 102. Native. Grassy places, fixed dunes and rock-ledges, usually on acid, often sandy soils. In suitable places throughout Great Britain and Ireland, but absent from the Channel Islands and most of south and east Ireland. A member of a polyploid complex of taxa found in north temperate regions to over 70◦ N in Norway. As broadly defined, the species is a member of the Circumpolar Boreo-temperate element. Subsp. rotundifolia is the common subspecies and is one of several races in Continental Europe. Subsp. montana occurs mainly in Ireland, western Scotland, Isle of Man and extreme south-west England and may be endemic. 2. Legousia Durande Specularia Heist. ex A. DC. nom. illegit.; Prismatocarpus L’H´er. Annual herbs. Stems decumbent to erect. Leaves alternate, exstipulate. Inflorescence a few-flowered terminal cluster or subterminal corymbose cyme. Calyx-tube subglobose; lobes 5, erect or erecto-patent. Corolla violet-purple inside, actinomorphic, divided about half of the way to the base into 5 lobes, more or less infundibuliform-rotate, shorter than calyx. Filaments and anthers free. Style 1, shorter than corolla; stigmas 3, linear. Ovary 3-celled. Fruit a subcylindrical capsule, dehiscing by subapical pores. About 15 species in north temperate regions and South America. 1. Calyx lobes more or less erect in fruit; flowers 2–3 mm in diameter; corolla about half as long as calyx lobes 1. hybrida 1. Calyx lobes patent or recurved in fruit; flowers 15–23 mm in diameter; corolla at least as long as calyx lobes 2. speculum-veneris

1. L. hybrida (L.) Delarbre Venus’s Looking-glass Campanula hybrida L.; Specularia hybrida (L.) A. DC.; Prismatocarpus hybridus (L.) L’H´er.; L. parviflora Gray nom. illegit. Annual herb with fibrous roots. Stems 4–30 cm, decumbent to erect, pale green, sometimes tinged purplish or bronze, branched or unbranched, angled, shortly hispid. Leaves alternate; lamina 0.5–3.0 × 0.3–0.8 cm, pale greyish-green, the basal oblong-spathulate or oblong-obovate, rounded at apex and with petioles up to 10 mm, the cauline oblong or obovate, obtuse at apex, strongly undulate, sessile, sometimes semiamplexicaul and narrowly revolute, all minutely scabridulous. Inflorescence a few-flowered, terminal cluster. Flowers 2–3 mm in diameter, sessile. Calyx 10–15 mm, glabrous to minutely scabridulous, divided one-third of the

way to the base; lobes 5, 5–8 mm, lanceolate, acuminate at apex, erect or erecto-patent, slightly accrescent in fruit, glabrous or scabridulous, margins recurved. Corolla 2–3 mm, whitish outside, violet-purple within, with a wing of greenish-cream at the base, infundibuliform-rotate, divided about half of the way to the base; lobes 5, obtuse or acute at apex and mucronulate. Stamens 5; filaments 0.5–0.8 mm, cream, somewhat dilated towards the base; anthers cream or black, linear. Style 1, about 1.5 mm; stigmas 3, linear, pale yellowish-green. Capsule 15–30 × 3–4 mm, subcylindrical, strongly constricted at apex, angled, glabrous or sparsely and minutely scabridulous, dehiscing by subapical pores, crowned by the persistent calyx; seeds about 1.3 × 0.7 mm, shining brown, elliptic-ovate, strongly compressed. Flowers 5–8. Autogamous. 2n = 20. Native. Arable fields and waste land. Scattered in south, central and eastern England, mostly on calcareous soils, much decreased in the last 50 years. West and south Europe; western Asia; North Africa; Macaronesia. A member of the European Southern-temperate element. 2. L. speculum-veneris (L.) Chaix Large Venus’s Looking-glass Campanula speculum-veneris L.; Specularia speculum-veneris (L.) A. DC.; L. durandii Delarbre; Prismatocarpus speculum-veneris (L.) L’H´er. Annual herb with fibrous roots. Stems 4–40 cm, pale green, often tinged purplish, erect or sprawling, angled, glabrous or with minute hairs on the angles, usually much branched, leafy. Leaves pale or medium green, the basal 1.5–2.0 × 0.5–1.0 cm, oblong-spathulate, obtuse at apex and the petioles up to 15 mm, the cauline 0.5–2.5 × 0.3–1.0 cm, oblong or obovate, obtuse at apex, undulate and narrowly revolute and narrowed to the sessile, semiamplexicaul base, all minutely scabridulous especially towards the apex. Inflorescence of crowded, branched, terminal or subterminal corymbose cymes. Flowers 15–23 mm in diameter, sessile. Calyx 7–8 mm, green, minutely papillose-scabridulous especially towards the apex, divided for almost all of the way to the base; lobes 5, narrowly linear, acute at apex, recurved or spreading in fruit, margins recurved. Corolla 9–11 mm, violet purple with a cream area at base, rarely pale blue or white, rotate, divided for about three-quarters of the way to the base; lobes 5, broadly ovate, obtuse or acute with a small mucro at apex. Stamens 5; filaments 0.8–1.0 mm, whitish, somewhat dilated towards the base; anthers cream. Style 1, greenish-white; stigmas 3, pale yellowish-green. Capsule 10–15 mm, narrowly oblong, dehiscing from subapical pores, crowned by the persistent calyx; seeds about 1.0 × 0.8 mm, brown, broadly ovoid, strongly compressed. Flowers 5–8. Autogamous. Visited by bees. 2n = 20. Introduced. Grain casual and garden escape. Apparently persistent in arable fields near Wootton St Lawrence in Hampshire since 1916. Native of much of Europe northwards to the Netherlands; south-west Asia; North Africa. 3. Wahlenbergia Schrad. ex Roth nom. conserv. Cervicina Delile nom. rejic. Perennial herbs. Stems procumbent and filiform. Leaves alternate, exstipulate. Inflorescence of solitary, axillary

5. Phyteuma flowers. Calyx-tube hemispherical or oblong-obconical; lobes 5, erect. Corolla blue, actinomorphic, campanulate or rotate, divided one-third to half of the way to the base, lobes 5. Stamens 5; filaments and anthers free. Style 1, shorter than corolla; stigmas 3, linear. Ovary 3-celled. Fruit a turbinate capsule, dehiscing by apical pores. More than 150 species, mostly in the south temperate region, particularly South Africa, with a few in tropical America and temperate regions of the Old World. Named after Georg Wahlenberg (1780–1851). W. nutabunda (Guss.) A. DC. and W. trichogyna Stearn have been recorded as casuals. 1. W. hederacea (L.) Rchb. Ivy-leaved Bellflower Campanula hederacea L.; Cervicina hederacea (L.) Druce Perennial herb with a long, slender, creeping rootstock. Stems 10–30 cm, pale green, smooth, procumbent, interlacing, filiform, glabrous, diffusely branched, leafy. Leaves alternate; lamina 0.5–2.0 × 0.5–2.0 cm, rather shiny medium green on upper surface, paler beneath, ovate to subrotund-reniform in outline, acute at apex, angled to shallowly lobed, the lobes broadly triangular, truncate, rounded or more or less cordate at base, glabrous; petioles short to long, pale green, slender, glabrous. Flowers 5–7 mm in diameter, solitary and axillary, more or less nodding; pedicels up to 40 mm, much longer than corolla, pale green, slender, glabrous. Calyx 2–3 mm, pale green, divided up to half of the way to the base; lobes 5, subulate, acute at apex, erect. Corolla 6–10 mm, campanulate, pale blue, rarely white, divided for one-quarter to half of the way to the base; lobes 5, ovate, more or less acute at apex. Stamens 5; filaments 1.2–1.6 mm, with stiff hairs below; anthers greyish-white. Style 1, whitish; stigmas 3, greyish-white. Capsule 2.5–3.0 mm, globose or turbinate, erect, dehiscing by apical pores; seeds 0.8–1.0 mm, ellipsoid, finely reticulate. Flowers 7–8. 2n = 36. Native. Damp heathy grassland, especially by streams and flushes, and other moist acidic habitats on heaths and moors, in rocky cloughs and in woods, ascending to approximately 500 m. South and west of Great Britain north to Argyll and south and south-east Ireland, but frequent only in Wales and south-west England; formerly in the Channel Islands; rarely naturalised elsewhere in wet lawns. Western Europe from Scotland and Belgium to Spain and Portugal. A member of the Oceanic Southern-temperate element. 4. Trachelium L. Perennial herbs. Stems erect. Inflorescence of terminal, corymbose, compound cymes. Calyx 1.5–2.0 mm, 5-lobed. Corolla pale blue or whitish, actinomorphic; tube 4–6 mm, slender; lobes 5, 1–2 mm. Stamens 5; filaments and anthers free. Style 1, longer than corolla; stigmas 2–3, capitate. Ovary 2- to 3-celled. Fruit a broadly pyriform capsule, dehiscing by 2–3 sub-basal pores. Four species in the Mediterranean region. 1. T. caeruleum L. Throatwort Perennial herb with thick, fibrous roots. Stems up to 100 cm, woody at base, dark green, usually suffused brownishpurple, erect, terete, striate, shining, glabrous. Leaves all

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cauline, alternate; lamina 1.0–6.0 × 0.3–3.0 cm, dark green on upper surface, paler beneath, elliptic-ovate, ovate or lanceolate, acute at apex, serrate-dentate, the teeth narrowly and sharply triangular, cuneate or attenuate at base, glabrous or with very short, simple eglandular hairs on the margins; petioles up to 25 mm, slender, glabrous. Inflorescence of rather lax, terminal, corymbose cymes; peduncles and pedicels slender, glabrous or with very few hairs; bracts linear, acute at apex. Flowers 2–5 mm in diameter. Calyx 1.5– 2.0 mm, green, divided half of the way to the base, glabrous; lobes 5, 0.5–1.0 mm, linear-lanceolate, obtuse at apex. Corolla 7–8 mm, pale blue or whitish, shortly divided at apex; tube 4–6 mm, slender, tubular; lobes 5, narrowly triangular, acute at apex. Stamens 5; filaments 5–6 mm; anthers pale. Style 1, long-exserted, thickened towards the apex; stigmas 2–3, capitate. Capsule 2–3 mm, broadly pyriform, dehiscing by 2–3, sub-basal pores. Flowers 7–9. 2n = 32. Introduced. Naturalised on walls at St Peter Port in Guernsey since 1892; also in Jersey and at Bexley in Kent. Native of the west Mediterranean region. 5. Phyteuma L. Perennial herbs. Stems decumbent to erect. Leaves alternate, exstipulate. Inflorescence of congested, globose or elongated spikes without flowerless bracts at the base. Calyx-tube ribbed and angled, 5-lobed. Corolla violetblue or yellowish-green, slightly zygomorphic, tubular and curved or straight in bud, but split nearly to base into 5 lobes when open, more or less cylindrical. Stamens 5; filaments and anthers free, but appressed around the style. Style 1, slightly shorter than corolla; stigmas 2–3, linear. Ovary 2to 3-celled. Fruit an ovoid capsule, dehiscing by 2–3 lateral pores. About 40 species in Europe, especially the mountains of central and southern Europe and temperate Asia. The pollination mechanism is similar to that of Campanula except that the pollen is held in the tube formed by the corolla-lobes and pushed out by the elongating style. Branwell, A. E. (1872). Phyteuma spicatum. Jour. Bot. (London) 10: 307–308. Schulz, R. (1904). Monographische Bearbeitung de Gattung Phyteuma. Geisenheim am Rhein. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [P. orbiculare.] Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [P. spicata.] 1. Inflorescence oblong to cylindrical in flower; corolla 2. usually whitish to pale yellowish-green, rarely bluish 1. Inflorescence globose to very shortly ovoid in flower; 3. corolla violet-blue 2. Corolla whitish to pale yellowish-green; styles and stigma yellow to yellowish-brown 1(a). spicatum subsp. spicatum 2. Corolla bluish; stigmas yellowish-brown to blue 1(b). spicatum subsp. coeruleum 3. Bracts shorter than inflorescence; corolla strongly curved 2. orbiculare subsp. tenerum in bud 3. Bracts, at least the lower, longer than inflorescence; 3. scheuchzeri corolla nearly straight in bud

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1. P. spicatum L. Spiked Rampion Long-lived perennial herb with a somewhat fleshy rootstock and a buried fusiform enlargement. Stems 30–80(–100) cm, pale green, erect, glabrous, leafy. Leaves alternate; lamina medium green on upper surface, paler beneath, the basal present at anthesis, 3.0–5.0 × 1.5–2.5 cm, ovate or lanceolate, obtuse at apex, crenate or serrate, rounded to deeply cordate at base and the petioles up to 40 mm, the cauline gradually smaller upwards, the lower similar to basal and sometimes larger and usually petiolate, the upper lanceolate to linear-lanceolate, acute at apex, crenate, narrowed or rounded at base and sessile, all glabrous. Inflorescence 30– 80(–100) mm, a dense, oblong or cylindrical spike; bracts 4–6 mm, linear or subulate, obtuse at apex, shorter than inflorescence; bracteoles linear, obtuse at apex. Flowers 1.5–2.0 mm in diameter, tubular, more or less sessile. Calyx 3.0–3.2 mm, green, glabrous, divided about half of the way to the base; lobes 5, linear-lanceolate, acute at apex. Corolla 7–10 mm, whitish to pale yellowish-green or blue, tubular and usually curved in bud, split nearly to the base when open; lobes 5, linear, obtuse at apex. Stamens 5; filaments 3–4 mm, whitish, dilated at base; anthers greenish. Style 1, yellow to yellowish-brown, much exserted; stigmas usually 2, yellow, yellowish-brown or blue. Capsule 4.5–5.0 mm, ovoid, crowned by the calyx lobes, dehiscing by 2–3 lateral pores; seed 1.0–1.5 mm, brownish, ellipsoid, smooth. Flowers 5–6. (a) Subsp. spicatum Corolla whitish to pale yellowish-green. Style and stigmas yellow to yellowish-brown. 2n = 22 + 0–4B. (b) Subsp. coeruleum R. Schulz Corolla bluish. Stigmas yellowish-brown to blue. 2n = 22. Native and introduced. The native plant is subsp. spicatum which is found by streamsides in coppiced woods, scrub and hedgerows on acid soils in an area 20 × 10 km in the Heathfield and Hailsham areas of Sussex. It occurs from southern Norway and Estonia southwards to northern Spain and the Crna Gora. Elsewhere in Great Britain the species is a rare escape and these plants are usually referable to subsp. coeruleum, which is a native of south-central Europe and the northern part of the Balkan peninsula. The species is a member of the European Temperate element. 2. P. orbiculare L. Round-headed Rampion Perennial herb with the rootstock almost woody above and a deeply buried, fusiform, fleshy enlargement below. Stem 5– 50 cm, pale green, erect or rarely ascending, glabrous, very leafy. Leaves alternate; lamina medium green on upper surface, pale beneath, the basal 2–4 × 1.5–2.0 cm, lanceolate to ovate or ovate-oblong, obtuse to acute at apex, serratecrenate, narrowed, rounded or more or less cordate at base and the petioles up to 40 mm, the cauline becoming gradually smaller, the lower similar to basal, the upper lanceolate to linear-lanceolate, acute at apex, serrate to entire, rounded at base and sessile, all glabrous. Inflorescence subglobose to shortly ovoid; bracts narrowly triangular-ovatelanceolate, acute at apex, entire or serrate, shorter than inflorescence; bracteoles 8–10 mm, linear, acute at apex.

Flowers 12–15 mm in diameter, sessile. Calyx 4.5–5.5 mm, green, obovate, divided one-third to half of the way to the base; lobes 5, triangular-lanceolate, acute at apex. Corolla 11–13 mm, deep violet-blue, curved and cylindrical in bud and eventually divided nearly to the base; lobes 5, linearlanceolate, acute at apex, spreading or reflexed. Stamens 5; filaments 3–4 mm, white; anthers yellow. Style 1, about 10 mm, violet-blue; stigmas 2, whitish. Capsule 4.5–5.5 mm, brown, ovoid, crowned by the stiff, erect calyx lobes, opening by lateral pores; seeds 1.2–1.3 mm, brown, ellipsoid. Flowers 7–8. 2n = 22 + 0–2B. Native. Open chalk grassland. Local in southern England from Wiltshire to Sussex, formerly in Kent. From southern England and Latvia to southern Spain, Albania and southern Greece. The species is a member of the European Boreo-temperate element. Our plant is subsp. tenerum (R. Schulz) P. D. Sell (P. tenerum R. Schulz) which has a lowland distribution in west-central and south-west Europe. Subsp. orbiculare has a more easterly and more montane distribution. 3. P. scheuchzeri All. Oxford Rampion Perennial herb with a thick, fleshy stock. Stems 12–45 cm, pale green sometimes tinted brownish, erect or decumbent, slender, striate, glabrous. Leaves alternate; lamina medium green on upper surface, paler and slightly bluish beneath, the basal usually present at anthesis, 1.5–5.0 × 1.0–2.5 cm, linear-lanceolate to ovate-lanceolate, acuminate or gradually drawn out to a long-acute apex, crenate to crenateserrate, deeply cordate to rounded at base and with slender petioles up to 120 mm, the cauline gradually decreasing in size upwards, linear-lanceolate to linear, long drawn out to an acute apex, remotely crenate-serrate to entire, rounded to cuneate at base and petiolate or the uppermost sessile, all glabrous. Inflorescence 15–25 mm, shortly ovoid to globose; bracts 15–30 mm, narrowly linear, acute at apex, at least one longer than the inflorescence. Flowers 1.0–1.5 mm in diameter, more or less sessile. Calyx 4.0–5.5 mm, green, divided about two-thirds of the way to the base; lobes 5, narrowly linear-lanceolate, acute at apex, glabrous. Corolla 10–12 mm, deep violet blue, tubular, nearly straight in bud, with 5, short blunt lobes at apex. Stamens 5; filaments 3–4 mm, white; anthers yellow. Style 1, bluish, included; stigmas 3, bluish. Capsule 4–5 mm, ovoid, brown; seeds 1.0–1.3 mm, brown, ellipsoid. Flowers 5–7. 2n = 26. Introduced. Naturalised on walls and pavements in Oxford since about 1951 and in limestone cracks at Inchnadamph in Sutherland since 1992. Native of the southern Alps and northern Apennines. Named after Johannes Scheuchzer (1684–1738). 6. Jasione L. Annual to perennial herbs. Stems decumbent to erect. Leaves alternate, exstipulate. Inflorescence of congested, flattish, terminal heads of bractless flowers, the whole surrounded by one or more rows of bracts. Calyx-tube ovoid or turbinate; lobes 5, narrow and spreading. Corolla blue, actinomorphic, tubular and straight in bud, but divided nearly to the base into 5 lobes when open. Stamens 5; filaments free; anthers slightly laterally fused. Style 1, longer than corolla; stigmas 2, more or less globose. Ovary

6. Jasione inferior, 2-celled. Fruit an ovoid-globose capsule, dehiscing by 2 short, apical valves. About 10 species in Europe and the Mediterranean region. Parnell, J. (1982). Cytotaxonomy of Jasione montana L. in the British Isles. Watsonia 14: 147–151. Parnell, J. (1982). Variation in Jasione montana L. (Campanulaceae) and related species in Europe and North Africa. Watsonia 16: 249–267. Parnell, J. (1985). Jasione montana L. in Biological flora of the British Isles. Jour. Ecol. 2: 341–358. Pugsley, H. W. (1921). British forms of Jasione montana L. Jour. Bot. (London) 59: 209–216. 1. Leaves 4–10 mm wide; involucral bracts 9–11 × 4–6 mm, equalling or exceeding the flowers 1(v). montana var. latifolia 1. Leaves 1.5–5.0 mm wide; involucral bracts 2–6 × 1–5 mm, 2. not exceeding the flowers 2. Stems numerous and spreading making the total plant very wide (up to 40 cm); involucral bracts less than 1(iii). montana var. litoralis 3.5 mm 2. Stems few to numerous, outline of plant usually narrow; 3. involucral bracts less than 3.5 mm 3. Leaves densely imbricate, margins markedly revolute, with numerous to dense rigid hairs 1(ii). montana var. imbricans 3. Leaves not densely imbricate, margins not markedly 4. revolute, usually with few hairs 4. Stems 1–6 cm; involucral bracts 4–6 mm 1(i). montana var. nana 4. Stems 12–40(–60) mm; involucral bracts 5–8 mm 1(iv). montana var. montana

1. J. montana L. Sheep’s-bit J. vulgaris Gaterau; J. undulata Lam.; J. appressifolia Pau; J. espadanae Pau; J. mediterranea Rouy Usually biennial, rarely annual, overwintering herb with a slender, shining white to yellowish-white tap-root and numerous fibrous laterals. Stems 1 to numerous, (1–)5– 40(–65) cm, pale green, decumbent, ascending, spreading or erect, slender to stout, striate, with few to numerous, short to medium, white, stiff, spreading simple eglandular hairs, simple or branched, leafy below but often leafless in the upper half. Leaves alternate; lamina dull medium green on upper surface, paler beneath, the basal in a rosette, 0.3– 5.0 × 0.1–1.0 cm, linear-oblong to linear-lanceolate, obtuse to acute at apex, entire to undulate-crenate and narrowed to a sessile, slightly decurrent base, the cauline similar, but gradually decreasing in size upwards, all with very few to numerous, short to medium, pale, stiff simple eglandular hairs on both surfaces and particularly on the margins and midrib below. Inflorescence a terminal head of up to 60 or more, more or less sessile flowers surrounded by several rows of involucral bracts. Outer involucral bracts 2–11 × 2–6 mm, green, ovate or triangular-ovate, acute to cuspidate at apex, often serrate or incised, often with white, stiff simple eglandular hairs, the inner very variable but nearly always narrower and more deeply incised than the outer, the innermost linear-oblong to lanceolate. Flowers 5–7 mm in diameter. Calyx 4.0–6.5 mm, green, glabrous

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with simple eglandular hairs, divided more or less half of the way to the base; lobes 5, subulate or linear-lanceolate, acute at apex. Corolla 3–15 mm, blue, rarely pink or white, divided nearly to the base; lobes 5, linear-lanceolate, acute at apex. Stamens 5; filaments 1.5–2.0 mm, pale blue; anthers pink, turning white. Style 1, long and exserted, blue; stigmas 2, blue. Capsule 2.5–3.0 mm, oblong or ovoid-globose, dehiscing by 2 apical pores; seeds 0.6–0.8 × 0.2–0.4 mm, brown, ovoid. Flowers 4–9. Protandrous. Visited by many insects. 2n = 12. (i) Var. nana Gren. & Godr. J. montana var. maritima Br´eb., non Duby Stems few, 1–6 cm, erect. Leaves with lamina 3–10 × 1– 3 mm, linear to narrowly spathulate, more or less obtuse at apex, with papillose margins and few to numerous, stiff simple eglandular hairs. Peduncles rather thin. Heads with fewer than 40 flowers. Involucral bracts 4–6 × 2.5–3.5 mm, not exceeding the flowers. (ii) Var. imbricans J. Parn. Stems few to numerous, 6–12(–18) cm, decumbent to erect. Leaves strongly imbricate, lamina 5–10 × 1.5–5.0 mm, linear to linear-lanceolate, obtuse at apex, the margins markedly revolute, with numerous to dense rigid hairs. Peduncles thin. Heads with fewer than 40 flowers. Involucral bracts 3–5 × 2.0–2.5 mm, not exceeding the flowers. (iii) Var. litoralis Fr. J. montana var. tenella Peterm.; J. montana var. nana Boreau, non Gren. & Godr.; J. montana var. sebularia Cout. Stems 2–20 cm, numerous, decumbent to spreading making the total plant very wide. Leaves with lamina 5–12 × 1.5– 3.0 mm, oblong or linear-oblong, obtuse or acute at apex, not thickened at margin, glabrous or rarely with a few hairs. Peduncles thin. Heads with fewer than 40 flowers. Involucral bracts 2.0–3.5 × 2.0–3.5 mm, not exceeding the flowers. 2n = 12 + 1–2B. (iv) Var. montana J. montana var. major Mert. & W. D. J. Koch; J. montana var. laevis Duby; J. montana var. hirsuta Duby; J. montana var. prolifera A. DC.; J. montana var. glabra Peterm.; J. montana var. stolonifera DC.; J. montana var. gracilis Lange; J. montana var. boraei Rouy; J. montana var. timbali Rouy. Stems usually few, 12–40(–65) cm, more or less erect and forming a narrow outline. Leaves with lamina 5– 20(–30) × 2–4 mm, linear-oblong to oblong-lanceolate or lanceolate, obtuse to acute at apex, margin not thickened, glabrous or with few simple eglandular hairs. Peduncles thin. Heads with fewer than 40 flowers. Involucral bracts 5–8 × 3–4 mm, not exceeding the flowers. 2n = 12. (v) Var. latifolia Pugsley J. montana var. megaphylla Vicioso Stems few, 10–40 cm, ascending to more or less erect, forming a narrow outline. Leaves with lamina 10–50 × 4– 10 mm, lanceolate-spathulate, oblong or oblong-lanceolate, usually more or less obtuse at apex, thickened at margin and sometimes undulate-crenate, usually with numerous, stiff simple eglandular hairs especially on the margin and midrib

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beneath. Peduncles thick. Heads usually with more than 60 flowers. Involucral bracts 9–11 × 4–6 mm, equalling or exceeding the flowers. Included here are some very fleshy broad-leaved plants which may be distinct.

Wimmer, F. E. (1953). Campanulaceae – Lobelioideae. Das Pflanzenreich IV. 276b. Woodhead, N. (1951). Lobelia dortmanna L. in Biological flora of the British Isles. Jour. Ecol. 39: 458–464.

Native. Usually on acidic, shallow, well-drained, light sandy or loamy soils, on sand dunes by the sea, maritime heathland, cliff-tops and walls, hedgebanks and roadside verges inland. Predominantly western and coastal in Britain, more scattered and declining in the east, rare in northern Scotland and mainly coastal in Ireland. Most of Europe to 62◦ in Finland and 53◦ E in Russia; Morocco, Tunisia and Algeria. A member of the European Temperate element. All our plants are referable to subsp. montana which occurs throughout the range of the species except North Africa. Var. litoralis occurs on coastal dunes and also on the coasts of western Europe and Scandinavia. Var. nana is a dwarf plant of sandy shores or dwarf Calluna on seacliffs in scattered localities from Hampshire to Shetlands and at Ballyteige in Co. Wexford. It occurs also in western France. Var. imbricans occurs in Guernsey in the Channel Islands and Dingle in Co. Kerry. A similar plant occurs at Garmouth in Morayshire. It is also known from Finisterre in north-west Spain and Gironde in France. Var. latifolia grows in hedgrows and on exposed cliffs in western Great Britain and both western and eastern Ireland. It is scattered throughout Europe. The remaining plants are referable to var. montana which occurs throughout the range of subsp. montana.

1. Stem (10–)30–100(–150) cm; leaves 2–16 × 1.0–7.5 cm; 4. siphilitica corolla 15–30(–35) mm 1. Stem 20–70(–120) cm; leaves 1–9 × 0.2–1.6 cm; corolla 2. 8–20 mm 2. Leaves all basal (except for small bracts), linear, entire; plant submerged except for inflorescence, or at lakeside 3. dortmanna 2. Cauline leaves present, the lower ones serrate; plant 3. terrestial 3. Pedicels 2–3 mm; corolla lobes less than 2 mm wide 1. urens 3. Pedicels 10–30 mm; corolla lobes of lower lip more than 2. erinus 2 mm wide

Subfamily 2. Lobelioideae (Juss.) E. Wimm. Family Lobeliaceae R. Br. Flowers zygomorphic. Filaments and anthers fused laterally around the style. 7. Lobelia L. Dortmanna Hill; Rapuntium Mill.; Mecoschistum Dulac Annual or perennial herbs. Stems ascending to erect. Leaves alternate, exstipulate. Inflorescence of terminal racemes. Calyx-tube obconical; lobes 5, subulate, spreading. Corolla pale lilac to blue, zygomorphic, the tube split to the base along the back, 2-lipped, with 2 lobes in the upper lip and 3 in the lower. Stamens 5; filaments and anthers fused laterally around the style. Style 1, shorter than the corolla; stigma capitate, shortly 2-lobed. Ovary 2-celled. Capsule clavate, oblong, spherical or conical, pedicellate, dehiscing by 2 apical valves. About 250 species in all the warm and temperate parts of the world except central and east Europe and western Asia. Named after Mathias de l’Obel (1538–1616). Brightmore, D. (1968). Lobelia urens L. in Biological flora of the British Isles. Jour. Ecol. 56: 613–620. Daniels, R. E., Raybold, A. F. & Farkas, J. M. (1996). Conserving genetic variation in British populations of Lobelia urens. Biol. Conserv. 79: 15–22. Farmer, A. F. (1989). Lobelia dortmanna L. in Biological flora of the British Isles. Jour. Ecol. 77: 1161–1173. Preston, C. D. & Croft, J. M. (1997). Aquatic plants in Britian and Ireland. Colchester. [L. dortmanna.] Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [L. urens.]

1. L. urens L. Heath Lobelia Mecoschistum urens (L.) Dulac; Rapuntium urens (L.) Mill.; Dormanna urens (L.) Kuntze Perennial herb with white, acrid latex, a rhizome and brown, fibrous roots. Stems 20–60(–80) cm, pale green, erect, slender, solid, angular, glabrous or nearly so, simple or branched, leafy. Leaves alternate; lamina medium green on upper surface, paler beneath, the basal 2–10 × 1.0–1.5 cm, obovate or obovate-oblong, obtuse at apex, irregularly serrate-dentate with the teeth callous-pointed, narrowed to a subpetiolate base; the cauline gradually getting smaller upwards, the lower similar to basal, the upper linear-oblong, more or less acute, sinuate-dentate and narrowed to a sessile base, glabrous on upper surface, usually with short, pale simple eglandular hairs on the lower surface. Inflorescence a long, rather dense raceme of 40–80 flowers; pedicels 2–3 mm; bracts 10–15 mm, linear, acute at the red-tipped apex, with numerous, short, pale simple eglandular hairs; bracteoles 2, small. Flowers 8–12 mm in diameter, more or less erect or patent; pedicels 2–3 mm, scabrid. Calyx 7–8 mm, green, with short simple eglandular hairs, divided half of the way or a little more towards the base; lobes 5, linear or subulate, acute at apex, spreading. Corolla 10–15 mm, purplish-blue, paler at base, divided half of the way or a little more to the base; lobes 5, linear-lanceolate, acute at apex, recurved, 2 forming the upper lip, 3 the lower lip. Stamens 5; filaments 5.5–6.5 mm, yellow; anthers yellow. Style yellow; stigma yellow, 2-lobed. Capsule 7–8 mm, brown, oblong, dehiscing by 2 apical valves; seeds 0.3–0.4 mm, shining brown, elliptical. Flowers 6–10. Visited almost entirely by Diptera, but can self-pollinate. 2n = 14. Native. Grassy, acid heaths, rough pastures, open woods and wood borders as isolated plants or in small patches, colonizing large areas after disturbance. Very local in southern England from Cornwall to Kent; formerly in Herefordshire. Western Europe from Belgium southwards; Madeira and Azores; Morocco. A member of the Oceanic Southtemperate element. 2. L. erinus L. Garden Lobelia Rapuntium erinus (L.) Mill.; Dortmanna erinus (L.) Kuntze

8. Pratia Annual herb with fibrous roots. Stems up to 30 cm, pale green sometimes flushed brownish-purple, erect to ascending or trailing, angled, glabrous or with sparse, pale simple eglandular hairs, much branched, leafy. Leaves alternate; lamina 1–3 × 0.2–1.6 cm, pale to medium green, paler beneath, sometimes flushed purplish-brown, the lower ovate to elliptical, obtuse at apex, crenate to crenate-serrate, rounded or narrowed at base and sessile or shortly petiolate, the middle and upper oblanceolate, lanceolate, linearlanceolate or linear, obtuse to sharply acute at apex, serratedentate, narrowed at base and mostly sessile, all glabrous. Inflorescence a sparse-flowered raceme; pedicels 10–30 mm, very slender, glabrous, with 2 glands at base; bracts 10–40 mm, linear or linear-elliptical and leaf-like, acute at apex, sometimes toothed, glabrous. Flowers 10–20 mm in diameter, erect or spreading. Calyx 9–11 mm, pale green, divided up to two-thirds of the way to the base, glabrous; lobes 5, linear, acute at apex. Corolla 8–20 mm, purplishblue, often with white and yellow markings on lower lip and whitish at base, sometimes pinkish or white, divided onethird to half of the way to the base, the 3 lobes of the lower lip large, ovate, rounded and spreading, the 2 of the upper lip linear-lanceolate, obtuse and erect. Stamens 5; filaments 4–6 mm, bluish-purple or paler or whitish; anthers grey. Style 1, green suffused bluish; stigma bluish or purplish, 2-lobed. Capsule 5–8 mm, oblong-conical, dehiscing by 2 apical valves; seeds 0.2–0.3 mm, shining brown, globoseellipsoid. Flowers 5–8. 2n = 28, 42. Introduced. Much grown in gardens, hanging baskets and window boxes from which it frequently escapes and is sometimes self-sown on tips and in pavement cracks and waste places. Scattered localities through Great Britain and in the Channel Islands. Native of South Africa. 3. L. dortmanna L. Water Lobelia L. lacustris Salisb. nom. illegit.; Dortmanna lacustris Kuntze; Rapuntium dortmanna (L.) C. Presl Perennial herb with acrid, milky latex and numerous, long, pure-white, brittle, fibrous roots. Stems 20–70(–120) cm, pale green, sometimes flushed reddish, slender, terete, hollow, glabrous, unbranched, leaves few and small. Leaves alternate; lamina yellowish-green, the basal numerous, in a rosette, submerged, 2–4(–8) × 0.4–0.7 cm, linear or oblong, obtuse at apex, entire, sessile, hollow and divided into 2 longitudinal air-canals, the cauline few and bract-like, all glabrous. Inflorescence a lax, elongated, emersed raceme; pedicels 7–10 mm, glabrous; bracts much shorter than pedicels, oblong, obtuse at apex, glabrous; without bracteoles. Flowers 15–20 mm in diameter, drooping. Calyx 4.5– 5.5 mm, green, glabrous, divided about one-fifth of the way to the base; lobes 5, broadly triangular, acute at apex. Corolla 10–20 mm, pale lilac, 2-lipped, divided about half way to base; lobes oblong or oblong-oblanceolate, obtuse at apex, the 2 of the upper lip shorter than the 3 of the lower lip. Stamens 5; filaments 4–5 mm, pale; anthers brown or blackish. Style 1, pale; stigmas pale, 2-lobed. Capsule 6– 10 mm, broadly clavate, pendulous, dehiscing by 2 apical valves; seeds 0.6–0.7 mm, brown, oblong-cylindrical, rugose. Flowers 7–8(–10). Autogamous and automatically self-pollinated in bud. 2n = 28, 32.

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Native. In the margins of stony, acidic lakes up to 2 m in depth and rarely on wet ground adjacent. Locally common in north and west Great Britain south to south Wales, and in west, north and east Ireland. North and north-central Europe extending locally to south-west France and White Russia; also at similar latitudes in North America. A member of the European Boreal-montane element. Differs from Isoetes lacustris and Littorella uniflora, with which it often grows, in its broader, obtuse leaves with a milky latex. Named after a pharmacist in Groningen named Dortmann. 4. L. siphilitica L. Great Lobelia Dortmanna siphilitica (L.) Kuntze; Rapuntium siphiliticum (L.) Mill. Perennial herb. Stems (10–)30–100(–150) cm, pale yellowish-green, erect, angled, striate, glabrous or with sparse, subrigid simple eglandular hairs, simple or occasionally branching above, leafy. Leaves alternate, all cauline, largest in the middle of the stem, rapidly reduced in size upwards, less so below, lamina 2–16 × 1.0–7.5 cm, medium green on upper surface, paler beneath, lanceolate, oblanceolate, elliptical or obovate, obtuse to acute or acuminate at apex, irregularly serrate, denticulate, crenate or subentire, attenuate at base, the lower sometimes with a winged petiole, the upper sometimes semiamplexicaul, auricled and decurrent, all glabrous or sometimes with short, rigid simple eglandular hairs on the veins beneath. Inflorescence a raceme, terminating the main stem and infrequently branched at the lower end; pedicels with short simple eglandular hairs; bracts 10–30 mm, shorter than the flowers, lanceolate, acute at apex, glabrous or ciliate; bracteoles 2, minute. Flowers 10–15 mm in diameter, more or less erect. Calyx 8–20 mm, enlarging after anthesis, green, campanulate, divided for about two-thirds of the way to the base; lobes 5, lanceolate, acuminate or attenuate to a long-acute apex, entire or irregularly toothed at the base, glabrous or long-ciliate. Corolla 15–30(–35) mm, deep blue and blueand-white striped in the throat, rarely all white or pale blue, divided about one-third to half of the way to the base; lobes of lower lip 3, narrowly ovate, acute at apex and deflexed, those of the upper lip 2, narrowly triangular, acuminate at apex and erect or recurved; shortly hairy on the veins outside. Anthers blue, fused laterally to the style, 2 distinctly smaller than the rest. Style 1, shorter than corolla; stigmas capitate, 2-lobed. Capsule 6–9 × 6–10 mm, cup-shaped or spherical, dehiscing by 2 apical valves; seeds about 0.6 mm, chestnut brown, ellipsoid, more or less tuberculate. Flowers 8–10. 2n = 14. Introduced. Grown in British gardens since at least 1665 and a persistent escape on waste ground at Wisley in Surrey. Native of eastern North America. The North American Indians smoked it instead of tobacco and made a concoction from its roots which they used to treat syphilis, hence its name. 8. Pratia Gaudich. Perennial dioecious herbs. Stems creeping and rooting at the nodes. Leaves alternate, exstipulate. Inflorescence of solitary, axillary flowers; male flowers with staminal tube mostly free from the corolla, the anthers connate around

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the style, the lower pair shortly awned; female flowers with staminodes, the ovary 2-celled and the stigma 2-lobed. Calyx-tube adnate to the ovary, divided up to one-third of the way to the base; lobes 5, narrow, erect. Corolla white or cream, zygomorphic, 2-lipped, the tube split at back to the base, the upper lip 2-lobed and the lower lip 3-lobed. Stamens 5; filaments fused laterally to form a tube round the style. Fruit a berry. P. arenaria Hook. fil. (P. angulata var. arenaria (Hook. fil.) Hook. fil.) was formerly established in a lawn at Branksome in Dorset. It differs from P. angulata in its leaves being (0.4–)0.7–2.5(–3.5) × (0.4–)0.7–2.0(–2.5) cm and in its shorter peduncles. About 25 species, mostly Australasian, but also in tropical Asia, tropical Africa and South America. Named after Ch. L. Prat-Bernon (c. 1817). 1. P. angulata (G. Forst.) Hook. fil. Lawn Lobelia Lobelia angulata G. Forst.; P. treadwellii auct. Perennial dioecious herb forming large mats or diffuse patches. Stems up to 15 cm, creeping and rooting at the nodes, pale green, slender, striate, glabrous or with a few simple eglandular hairs, branched, leafy. Leaves alternate; lamina 0.2–1.2 × 0.2–0.8 cm, pale yellowish-green on upper surface, sometimes purplish beneath, slightly succulent, broadly ovate to subrotund, obtuse or acute at apex, usually more or less coarsely sinuate-dentate, rarely nearly entire, rounded at base, glabrous; petioles 1–3 mm, sometimes minutely hairy. Flowers 12–15 mm in diameter, solitary in the axils of leaves; pedicels up to 60 mm, slender, pale, glabrous. Calyx 6–8 mm, medium green, tipped purplish, glabrous, divided for one-quarter to one-third of the way to the base; lobes 5, triangular-lanceolate, acute at apex. Corolla 7–20 mm, white or cream, divided three-quarters of the way to the base; lobes 5, oblanceolate or linearoblanceolate, rounded-mucronulate at apex, the lower lip spreading, the upper lip more or less spreading. Stamens 5; filaments and anthers pale. Style 1, pale; stigma 2-lobed. Berry 7–12 mm in diameter, reddish-purple to pinkishpurple, broadly obovoid to subglobose, the calyx lobes persistent. Flowers 7–9. 2n = 70. Introduced. Grown in gardens and becoming established in lawns, mainly of parks and botanic gardens. In scattered localities in Scotland, Surrey and Kent. It has been known in turf at the Royal Botanic Garden, Edinburgh since the 1930s. Native of New Zealand. 9. Downingia Torr. nom. conserv. Bolelia Raf. nom. rejic.; Clintonia Douglas ex Lindl., non Raf. Annual herbs. Stems erect or ascending. Leaves alternate, exstipulate. Inflorescence of numerous axillary flowers on long pedicel-like ovaries. Calyx-tube very short, with 5 narrowly linear lobes. Corolla blue with a white centre, zygomorphic, the tube split to base along the back, 2-lipped, with 2 lobes in the upper lip and lower lip shallowly 3lobed. Stamens 5; filaments and anthers fused laterally round the style. Style 1 with conical stigmas surrounded by a papillose beard. Ovary long and pedicel-like, 1-celled.

Fruit a long, narrow capsule, dehiscing by 3–5 longitudinal slits. Eleven species in western North America, one extending to Chile. Named after Andrew Jackson Downing(1815–52). 1. D. elegans (Douglas ex Lindl.) Torr. California Lobelia Clintonia elegans Douglas ex Lindl. Annual herb with fibrous roots. Stems 10–25(–40) cm, pale green, sometimes tinted purplish, erect or ascending, angular, branched, sometimes to base, leafy. Leaves alternate, all cauline; lamina 0.8–3.0 × 1.0–4.0 cm, medium green on upper surface, paler beneath, linear-lanceolate, more or less acute at apex, entire, widest at the base and decurrent on stem, glabrous. Flowers 9–11 mm in diameter, on long pedicel-like ovaries which are axillary. Calyx 4.5–10.0 mm, pale green, divided almost to the base; lobes 5, narrowly linear, obtuse at apex, glabrous. Corolla 8–18 mm, 2-lipped, the upper lip 2-lobed, with the lobes blue, linear-lanceolate and acute at apex, the lower lip hemispherical and 3-lobed, the lobes and outer margin blue and the centre white with 2 yellow ridges and dark lines. Stamens 5, incurved and connate in the tube; anthers blue. Style 1, with a conical stigma and surrounded by a papillose beard. Ovary 15–40 mm, linear, twisted, minutely papillose. Capsule 25–45 mm, brown, cylindrical, dehiscing by 3–5 longitudinal slits; seeds small, numerous, shining. Flowers 6–8. 2n = 20. Introduced. Casual and perhaps naturalised in damp hollows in grassy places where it was probably introduced with grass seed. Sussex and Buckinghamshire and probably elsewhere. First found in 1978 and apparently increasing. Native of western North America.

Order 8 . RU B I A L E S Benth. & Hook. fil. Herbs, evergreen climbers or rarely shrubs. Leaves opposite, simple, with large leaf-like stipules making the leaves appear whorled. Flowers actinomorphic, bisexual or bisexual and male mixed, or more or less dioecious, epigynous. Calyx absent or 4- to 5-lobed. Corolla 4- to 5-lobed. Stamens 4–5. Ovary 2-celled; ovules with axile placentation. Fruit a capsule, berry or drupe. Contains 2 families, 607 genera and some 6,500 species from all parts of the world. 1 4 3 . RU B I AC E A E Juss. nom. conserv. Annual to perennial herbs, evergreen climbers or rarely shrubs. Leaves opposite, simple, more or less entire, usually narrow and sessile, with 1 to several stipules per leaf, the stipules usually leaf-like and as large as the leaves, so that the leaves appear to be in whorls of 4 or more. Inflorescence usually a compound, terminal and/or axillary cyme, often aggregated into a terminal panicle. Flowers small and inconspicuous, actinomorphic, bisexual or bisexual and male mixed, or more or less dioecious, epigynous. Calyx with 4–5 lobes, minute, or absent. Corolla with 4–5 lobes, fused into a long or short tube below, of various colours. Stamens 4–5, usually borne at the apex of the corolla tube, rarely near the base. Styles 1–2, if 1 often with 2 branches;

2. Nertera stigmas 1 per style or style-branch, capitate. Ovary inferior, 2-celled, each cell usually with one ovule on an axile placenta. Fruit mostly of 2, fused, 1-seeded nutlets which later separate, or succulent with 1–2 seeds; seeds endospermic, with a usually straight embryo. Contains 606 genera with some 6,500 species, cosmopolitan, but chiefly tropical. Our species of this family are mainly herbs, whereas the tropical plants are chiefly trees, shrubs or climbers. 1. Leaves opposite, usually with smaller stipules or smaller 2. leaves also at the same node 1. Leaves in whorls of 4 or more, more or less all of the 4. same size 1. Coprosma 2. Evergreen shrub 3. 2. Procumbent to ascending herb 3. Leaves ovate to subrotund or deltoid-ovate; fruit 2. Nertera succulent 5. Asperula 3. Leaves linear or nearly so; fruit of 2 nutlets 5. 4. Most or all flowers with 5 corolla lobes 6. 4. Most or all flowers with 4 corolla lobes 5. Procumbent to sprawling annual; leaves 6 or more in a 4. Phuopsis whorl; corolla deep pink; fruit dry 5. Evergreen climber or scrambler; leaves 4–6 in a whorl; 8. Rubia corolla yellowish-green; fruit fleshy 6. Calyx 0.5–1.5 mm at first, slightly enlarging in fruit; 3. Sherardia corolla pale to deep mauvish-pink 6. Calyx absent or vestigial; corolla of various colours, but 7. not mauvish-pink 8. 7. Corolla tube more than 1 mm 9. 7. Corolla tube less than 1 mm 5. Asperula 8. Ovary and fruit smooth to papillose 6. Galium 8. Ovary and fruit covered with hooked bristles 6. Galium 9. At least some whorls with more than 4 leaves 10. 9. All whorls with 4 leaves 10. Flowers in terminal panicles, white; ovary and fruit 6. Galium covered with hooked bristles 10. Flowers in dense axillary whorls, yellow; ovary and fruit 7. Cruciata smooth

1. Coprosma J. R. Forst. & G. Forst. Evergreen, usually dioecious shrubs. Branches stout, pale brown. Leaves opposite, petiolate, with small stipules. Flowers in axillary clusters with 2 partly fused bracts below them. Calyx minute. Corolla greenish, 4- to 5-lobed, with a long tube. Stamens 4–8. Styles 2, very slender; stigmas capitate, stout. Ovary 2-celled, with one ovule in each cell. Fruit a drupe, depressed-obovoid, orange-red, succulent, with 2 nuts. Over 90 species in Australasia and the Pacific Islands. 1. C. repens A. Rich. Tree Bedstraw C. baueria auct.; C. retusa Hook. fil.; C. bauriana Hook. fil.; C. stockii B. S. Williams Evergreen, usually dioecious shrub with variable habit. Stems up to 3 m, prostrate to erect; bark pale brown; branches stout; twigs striate; young shoots shortly hairy. Leaves opposite; lamina (20–)50–90 × 15–40(–50) mm, thick and rather fleshy, glossy dark green on upper surface,

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paler beneath, ovate, broadly ovate-oblong or broadly elliptical, rounded to retuse or emarginate at apex, with recurved or inrolled, entire margins, cuneate at base, glabrous; petiole 8–16 mm, stout, glabrous; stipules broadly triangular, subacute to obtuse at apex, glabrous. Inflorescence of compound, axillary clusters of flowers on branched peduncles; flowers seated in a cup formed by the partial fusion of 2 bracts. Male flowers numerous, in clusters; calyx of 4–5 minute, obtuse teeth; corolla about 5 mm, greenish, funnelshaped, divided to about half of the way to the base, with 4–5 acute lobes. Female flowers about 3 in a cluster; calyx of 4–5 short, obtuse teeth; corolla about 5 mm, greenish, subfunnel-shaped, divided for less than half of the way to the base, with 4–5 obtuse to acute lobes. Drupe 9–10 × 7–8 mm, orange-red, depressed-obovoid, succulent, with 2 nuts. Flowers 6–8. 2n = 44. Introduced. Planted as a windbreak in the Isles of Scilly and sometimes self-sown. Abundant near St Warna’s Well, St Agnes, in woodland on Tresco and on walls at Porthcressa beach, St Mary’s. Native of New Zealand. 2. Nertera Banks & Sol. ex Gaertn. nom. conserv. Perennial herbs. Stems procumbent to ascending. Leaves opposite, with small stipules. Flowers solitary, axillary and terminal. Calyx minute. Corolla greenish, 4-lobed. Stamens 4. Styles 2, slightly exserted, slender, papillose; stigma capitate. Ovary 2-celled, with 1 ovule in each cell. Fruit a drupe, globose, bright to dark reddish-orange, succulent, with 2 nuts. About 15 species in Australia, New Zealand, Malaya, Central and South America and Tristan daCunha. 1. N. granadensis (Mutis ex L. fil.) Druce Beadplant Gomozia granadensis Mutis ex L. fil.; N. depressa Banks & Sol. ex Gaertn.; N. montana Colenso Perennial herb forming small to large patches, faintly to strongly foetid when bruised. Stems up to 15 cm, procumbent and rooting to ascending, slender to stout, obscurely 4angled, glabrous. Leaves opposite; lamina (3–)5–8(–15) × 2–5(–10) mm, coriaceous or almost fleshy, bright green on upper surface, paler beneath, ovate to broadly ovate, sometimes subrotund or deltoid-ovate, obtuse to acute at apex and minutely apiculate, thickened and recurved at margin, abruptly narrowed to a rounded, subtruncate or cordate base, glabrous; petioles (1–)2–3(–4) mm, glabrous; stipules small, rather thick, narrowly to broadly triangular. Flowers very small, solitary, terminal and axillary. Calyx minute, truncate at apex. Corolla (1–)2–3 mm, greenish, subcampanulate, divided for about one-quarter of the way to the base, with 4 broadly ovate lobes. Stamens 4; filaments and anthers pale greenish. Styles 2, greenish; stigma pale greenish. Drupe about 4 mm in diameter, bright to dark reddish-orange, globose, succulent. Flowers 6–8. 2n = 44. Introduced. A garden or greenhouse escape established as a weed in turf in a few places in the Helensburgh area of Dunbartonshire and on a golf course near Largs in Ayrshire. Native of Australia, New Zealand and South America. It is grown for its show of small bead-like fruits which stay on

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the plant until late in the autumn. This very variable species needs detailed study over its whole range. 3. Sherardia L. Annual herbs. Stems procumbent to ascending. Leaves in whorls of 4–6, sessile. Flowers 4–10, in dense, terminal and axillary clusters with a whorl of 8–10 leaf-like bracts at the base, the clusters stalked. Calyx 0.5–1.5 mm at first, slightly enlarging in fruit, usually with 4–6 subulate lobes, but lobes sometimes absent. Corolla pale to deep mauvish-pink, 4lobed. Style 1, filiform, bifid, branches unequal; stigmas capitate. Ovary 2-celled, with 1 ovule in each cell. Fruit a pair of scabrid nutlets with a persistent calyx on top. One species widely distributed in Europe, the Mediterranean region and western Asia and introduced elsewhere in temperate regions. Named after William Sherard (1659– 1728). Druce, G. C. (1894). Varieties of Sherardia arvensis L. Jour. Bot. (London) 32: 240–243.

1. S. arvensis L. Field Madder Asperula sherardii H¨ock ex Hallier Annual herb with slender, reddish fibrous roots. Stems 5– 40 cm, prostrate or decumbent, spreading, 4-angled, with small, deflexed pricklets on the angles, glabrous or nearly so, simple or branched. Leaves in whorls of 4–6, the lower soon withering, lamina 5–18 × 1–5 mm, medium green on upper surface, paler beneath, obovate, elliptical or oblanceolate, cuspidate or mucronate at apex, entire, narrowed to the sessile base, more or less glabrous but the margins and underside of the midrib with forwardly directed pricklets. Flowers 2.5–3.0 mm in diameter, subsessile, 4–8 in terminal heads surrounded by an involucre of up to 10 connate bracts; bracts leaf-like, lanceolate, longer than the flowers, scabridulous. Calyx 0.5–1.5 mm at first, slightly enlarging in fruit, usually with 6 erect, subulate lobes with short, rough hairs, lobes sometimes absent. Corolla 4–5 mm, white within, deep to pale mauvish-pink outside, funnel-shaped, divided for about one-third of the way to the base; lobes 4, ovate, obtuse at apex. Stamens 4; filaments 0.4–0.5 mm, white; anthers lilac to blackish. Style 1, white, bifid; stigmas cream, capitate. Nutlet 2–3 mm, obovoid, rough with short appressed bristles, calyx persistent; seeds 0.1–0.2 mm, oblong, grooved, minutely reticulate. Flowers 5–10. Visited chiefly by flies. 2n = 22. Var. ovata Fisch.-Benz. with broader leaves, var. hirsuta Baguet a more hairy plant, and var. walravenii Wirtg. (var. maritima Griseb.; var. mutica Wirtg.), with a reduction in the calyx lobes, seem not to be worth recognising as they do not have any geographical or ecological significance. It should be noted that the calyx lobe character is that by which Sherardia is distinguished from Asperula. Native. Formerly abundant in arable fields but almost eradicated by herbicides; still found in waste places, thin grassland and in lawns where it flowers and fruits in a very dwarf form. Once frequent almost throughout Great Britain and Ireland, though local in Scotland, but now much reduced in arable regions. Through most of Europe to about

67◦ N in Norway and in the Mediterranean region; western Asia; North Africa; introduced in North America, Australia and other temperate countries, and probably also to central and northern Europe in prehistoric times. A member of the European Southern-temperate element. 4. Phuopsis (Griseb.) Hook. fil. Asperula section Phuopsis Griseb. Annual to perennial herbs. Stems procumbent to sprawling. Leaves in whorls of 6–9, sessile. Flowers numerous, in dense terminal clusters with a whorl of many leaf-like bracts at the base. Calyx minute. Corolla deep pink, 5-lobed, with a long, narrow tube. Stamens 5, included in corolla. Ovary 2-celled, with 1 ovule per cell. Style 1, long-exserted, bifid. Fruit a pair of glabrous, papillose nutlets. One species in Caucasus and Iran. 1. P. stylosa (Trin.) Benth. & Hook. fil. ex B. D. Jacks. Caucasian Crosswort Crucianella stylosa Trin.; Asperula ciliata auct. Annual to perennial herb. Stems up to 30(–70) cm, pale yellowish-green, procumbent to sprawling, 4-angled, striate, with minute prickles on the angles, branched, leafy. Leaves in whorls of 6–9; lamina 7–30 × 2–4 mm, pale greyish- or yellowish-green on upper surface, paler beneath, linear to narrowly linear-lanceolate or narrowly elliptical, acuminate at apex to a narrow point, entire and thickened on the margin, narrowed at the base, sessile, with short, forwardly directed, stiff simple eglandular hairs on the margin and sparse even shorter ones on the surface. Flowers 4–7 mm in diameter, in dense terminal clusters with a whorl of bracts at the base; bracts 12–15 × 2–4 mm, linear or linear-lanceolate, acuminate to a narrow point at apex, with stiff, forwardly directed hairs on the margin. Calyx minute. Corolla 12–15 mm, deep pink, divided about one-eighth of the way to the base, the tube long and narrow, the lobes 5, ovate, obtuse at apex and spreading. Stamens 5, enclosed in tube; anthers deep pink. Styles 1, deep pink, bifid, long exserted, slender; stigmas deep pink, clavate. Nutlets glabrous, papillose. Flowers 6–8. 2n = 20, 22. Introduced. An established garden escape. In a few scattered localities in southern and western Great Britain including a limestone bank at St Donat’s in Glamorganshire, where it has been naturalised since 1927. Native of the Caucasus and Iran. 5. Asperula L. typus conserv. Annual to perennial herbs. Stems procumbent to erect. Leaves in whorls of 4–8 and all equal, or in whorls of 4 with 2 long and 2 short, sessile. Flowers in dense, terminal clusters with a whorl of leaf-like bracts at their base, or in loose, terminal panicles. Calyx minute. Corolla white, pinkish-lilac, yellowish-pink or blue, 4-lobed. Style 1, more or less connate; stigmas capitate. Ovary 2-celled, with 1 ovule in each cell. Fruit a pair of smooth to papillose or rugose nutlets. About 200 species, mostly in Europe, the Mediterranean region and western Asia, extending to central Asia and the

5. Asperula

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Himalayas. The Australian and New Zealand species are perhaps generically distinct. A. tinctoria L., Dyer’s Woodruff, occurs as a casual garden escape.

(a) Subsp. cynanchica Laxly or densely caespitose herb. Rhizomes absent or brown. Flowers with pedicels to 1.0 mm. Corolla tube 1.0– 1.5(–2.0) times as long as lobes.

Tutin, T. G. & Chater, A. O. (1974). Asperula occidentalis in the British Isles. Watsonia 10: 170–171.

(i) Var. cynanchica Leaves 10–20(–25) mm, all linear or narrowly lanceolate.

1. 1. 2. 2. 3. 3. 4. 4. 5. 5.

2. Leaves in whorls of 6–8; corolla blue 3. Leaves in whorls of 4; corolla white or pinkish 3. arvensis Corolla 5–7 mm 4. orientalis Corolla 7–13(–15) mm Leaves equal in all whorls, elliptic-lanceolate to 2. taurina elliptic-ovate; inflorescence a dense head Leaves of at least upper whorls 2 long and 2 short, linear 4. to oblanceolate; inflorescence a diffuse panicle Rhizome orange; corolla tube about equalling lobes 1(b). cynanchica subsp. occidentalis Rhizome absent or brown; corolla tube 1.0–1.5(–2.0) 5. times as long as lobes Leaves 10–20(–25) mm, all linear or narrowly lanceolate 1(a,i). cynanchica subsp. cynanchica var. cynanchica Leaves 3–10 mm, the lower elliptical or oblanceolate, the upper linear 1(a,ii). cynanchica subsp. cynanchica var. densiflora

Section 1. Cynanchicae (DC.) Boiss. Section Cynanchica Griseb. Perennial caespitose herbs. Leaves in whorls of 4, the upper whorls with 2 long and 2 short, acute at apex, 1-veined. Inflorescence subtended by bracts and bracteoles. Corolla 4-lobed, pinkish-lilac. Nutlets ovoid, papillose to rugose, usually glabrous. 1. A. cynanchica L. Squinancywort A. minor Gray nom. illegit. Laxly to densely caespitose perennial herb with a branched stock, with or without rhizomes and numerous, slender, prostrate or ascending sterile shoots. Stems bearing flowers 3–50 cm, procumbent, ascending or erect, 4-angled, rough with short simple eglandular hairs below, more or less glabrous above, branched, leafy. Leaves in whorls of 4, the upper whorls with 2 long and 2 short; lamina 3–20(–25) × 0.5–2.0 mm, medium green on upper surface, paler beneath, linear to lanceolate, elliptical or oblanceolate, acute at apex, entire, the margin sometimes slightly revolute, slightly narrowed at base, glabrous, 1-veined. Flowers 3–4 mm in diameter, sessile or shortly pedicellate, vanilla-scented, in lax or compact, terminal and axillary, few-flowered cymes; sessile or pedicels up to 1.0 mm; bracts small, subulate; bracteoles 1.8–2.0 mm, ovate-lanceolate, acute at apex. Calyx green or flushed crimson, of 4, small lobes. Corolla 2.5–3.5 (–4.0) mm, white within, pale pinkish-lilac outside, funnelshaped, tube equalling twice as long as the lobes; lobes 4, ovate-lanceolate, acute at apex. Stamens 4; filaments 1.0– 1.2 mm, whitish; anthers purplish-black. Style 1, whitish, bifid; stigmas purplish. Nutlets 1.5–3.0 mm, ovoid, papillose to rugose, usually glabrous. Flowers 6–7. Homogamous. Visited by various small insects. 2n = 40. The three following taxa are badly in need of experimental study.

(ii) Var. densiflora Gren. & Godr. A. cynanchica var. maritima Lange nom. illegit. Leaves 3–10 mm, the lower elliptical or oblanceolate, the upper linear. (b) Subsp. occidentalis (Rouy) Stace A. occidentalis Rouy Laxly caespitose herb. Rhizomes orange. Leaves 3–10(–15) × 1–2 mm, oblanceolate to linear-lanceolate. Flowers sessile. Corolla tube about equalling lobes. Native. Limestone and chalk grassland and calcareous sand dunes with a special liking for anthills. Locally common in southern Britain and western Ireland and in scattered localities north to Westmorland and Yorkshire. Most of Europe north to about 54◦ 35 N in England and central Russia. A member of the European Temperate element. Var. densiflora occurs in maritime habitats in south and south-west England. It is also in west France and Spain. Var. cynanchica occurs through much of the range of the species. Subsp. occidentalis occurs on calcareous dunes in south Wales and western Ireland and between Santander in northern Spain and Biarritz in south-west France. Section 2. Glabella Griseb. Perennial herbs with rhizomes. Leaves in whorls of 4, equal, acute at apex, 3-veined. Partial inflorescences in dense heads; bracts forming an involucre round the flowers. Corolla 4-lobed, white tinged yellowish-pink. Nutlets ovoid, smooth, glabrous. 2. A. taurina L. Pink Woodruff A. caucasica Pobed.; A. propinqua Pobed. Perennial herb spreading by slender, orange-red rhizomes. Stems (10–)20–50 cm, pale green, erect, stout, 4-angled, with more or less numerous, pale, spreading simple eglandular hairs, leafy, sparingly branched. Leaves in whorls of 4, equal; lamina 20–60 × 10–25 mm, medium green on upper surface, paler beneath, elliptic-lanceolate to ellipticovate, acute at apex, entire, abruptly narrowed at base, with pale simple eglandular hairs especially on the margins and veins; main veins 3, with a distinct reticulum of subsidiary veins; with a very short petiole. Flowers 4.5–5.0 mm in diameter, in dense terminal heads; bracts forming a distinct involucre exceeding the flowers, 10–30 × 5–15 mm, ovate or lanceolate, acute at apex; bracteoles shorter and narrower. Calyx an indistinct annular ridge. Corolla 10–14 mm, white tinged with yellowish-pink, divided for one-third to onequarter of the way to the base; lobes 4, lanceolate, acute at apex. Stamens 4; filaments 2–3 mm, whitish; anthers pale to dark violet. Style 1, whitish, bifid; stigmas pinkish. Nutlets 1–3 mm in diameter, ovoid, smooth, glabrous.

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Flowers 5–6. Protandrous and visited by flies and bees. 2n = 22. Introduced. Garden escape or relic, naturalised by streams and in damp woods, abundantly on the banks of Turret Burn, Crieff in Perthshire. Local in central Scotland, rare and impermanent further south. Native of southern Europe, northern Turkey, Caucasia and Iran. Section 3. Asperula Annual herbs. Leaves mostly in whorls of 6–8, obtuse at apex, 1-veined. Partial inflorescences capitate, enveloped by leaf-like, long-ciliate bracts. Corolla 4-lobed, blue. Nutlets depressed-globose, smooth or finely granulose, glabrous. 3. A. arvensis L. Blue Woodruff Annual herb with fibrous roots. Stems 10–30(–50) cm, pale green, erect, slender, 4-angled, glabrous or minutely scabridulous, sometimes branched, with large, broadly oblong-obovate cotyledons often persisting until anthesis. Leaves (4–)6–8 in a whorl; lamina 6–23 × 1.5–6.0 mm, medium green on upper surface, paler beneath, narrowly oblanceolate or linear, obtuse at apex, entire, the margins sometimes narrowly revolute, gradually narrowed at base, glabrous or sparsely scabridulous especially along the midrib, 1-veined. Flowers 3.5–4.0 mm in diameter, subsessile, in crowded, terminal, hemispherical heads; bracts forming a distinct involucre exceeding or equalling the flowers, 5–18 × 1.0–2.5 mm, narrowly oblong or linear, subacute at apex, entire, ciliate with long, reddish or whitish bristles towards the base; bracteoles similar but smaller and narrower and tapered at base. Calyx an indistinct annular ridge. Corolla 5–7 mm, bright or pale blue, divided for one-quarter to one-fifth of the way to the base; lobes 4, triangular-ovate, acute at apex. Stamens 4; filaments very short, pale; anthers brownish. Style 1, pale, bifid; stigmas bluish, capitate. Nutlets 2.0–3.5 mm in diameter, dark brown, depressed-globose, more or less smooth, glabrous. Flowers 5–6. 2n = 22. Introduced. Casual on tips and in waste places, mainly from bird-seed. In scattered localities over much of Great Britain and in the Channel Islands. A widespread weed in Europe, the Mediterranean region and western Asia eastwards to Iran. 4. A. orientalis Boiss. & Hohen. Annual Woodruff A. azurea Jaub. & Spach Annual herb with fibrous roots. Stems 5–30(–40) cm, erect, 4-angled, scabrid-puberulous below, glabrous above, simple or stiffly branched, often from near the base, leafy. Leaves 4–8 in a whorl; lamina 7–25(–30) × 1.5–5.0(–10) mm, the lowest subrotund to obovate, obtuse at apex and entire, the upper oblanceolate, oblong-elliptical or linearoblong and obtuse at apex, entire, with antrorsely scabrid margins, 1-veined. Flowers 4–6 mm in diameter, subsessile, in crowded, terminal, hemispherical heads; bracts forming a distinct involucre and half to two-thirds as long as the flowers, linear-lanceolate, more or less acute at apex, antrorsely scabrid on the margins. Calyx an indistinct annular ridge. Corolla 7–13(–15) mm, bright blue, divided about

one-quarter of the way to the base; lobes 4, more or less ovate, subobtuse at apex. Stamens 4; filaments very short; anthers brownish. Style 1, pale, bifid; stigmas bluish, capitate. Nutlets 2.0–2.5 mm in diameter, dark brown, depressed-globose, finely granulate. Flowers 5–6. 2n = 22. Introduced. Casual on tips and in waste places, perhaps overlooked for A. arvensis. Native of western Syria, northern Iraq, Turkey, Caucasia and Iran. 6. Galium L. Annual to perennial herbs. Stems procumbent to erect. Leaves in whorls of 4–12 and all equal, sessile. Flowers in terminal panicles or axillary cymes. Calyx a minute annular ridge. Corolla white to yellow or greenish, rotate or funnelshaped, 4-lobed. Stamens 4. Style 1, short, bifid; stigma globose, capitate. Ovary 2-celled, with 1 ovule in each cell. Fruit a pair of smooth to bristly nutlets, the bristles sometimes hooked. About 400 species, almost cosmopolitan. G. divaricatum Pour. ex Lam., G. murale (L.) All., G. setaceum Lam. and G. virgatum Nutt. ex Torr. & A. Gray have been recorded as wool casuals. Amitage, E. (1909). Hybrids between Galium verum and G. mollugo. New Phytol. 8: 351–353. Britton, C. E. (1938). An account of the occurrence of Galium debile Desv. in Britain. Jour. Bot. (London) 76: 13–15. Dandy, J. E. (1957). Galium tricornutum Dandy. Watsonia 4: 47– 48. Grime, J. P. et al. (1988). Comparative plant ecology. London. [G. aparine, G. palustre aggr., G. saxatile, G. sterneri and G. verum.] Hult´en, E. (1958). The amphi-atlantic plants and their phytogeographical connections. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 7: 1–340. [G. boreale, G. palustre and G. saxatile.] Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 13: 1–463. [G. aparine, G. spurium and G. verum.] Klisphuis, E. K., Heringa, J. & Hogeweg, P. (1986). Cytotaxonomic studies on Galium palustre. Morphological differentiation of diploids, tetraploids and octoploids. Acta Bot. Neerl. 35: 383– 392. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [G. pumilum and G. sterneri.] Teppner, H., Ehrendorfer, F. & Puff, C. (1976). Karyosystematic notes on the Galium palustre-group (Rubiaceae). Taxon 25: 95– 97. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [G. constrictum and G. tricornatum.] 2. 1. Ovaries and nutlets with hooked bristles 1. Ovaries and nutlets smooth to rugose, tuberculate or 7. papillose, but without hooked bristles 2. boreale 2. All whorls with 4 leaves 3. 2. Most or all whorls with more than 5 leaves 3. Perennial herb with far-creeping rhizomes; tube of 1. odoratum corolla 1.0–1.5 mm 3. Annual herbs without rhizomes; tube of corolla less than 4. 1 mm. 4. Nutlets (excluding bristles) 1.5–3.0 mm, its bristles wider towards but not bulbous at their bases; corolla 0.8–1.3 12(b). spurium subsp. infestum mm in diameter

6. Galium 4. Nutlets (excluding bristles) more than 2.5 mm, its bristles with bulbous bases; corolla 2.5–3.5 mm in 5. diameter 5. Stems scrambling or ascending the stems of erect plants; internodes 40–110 mm; leaves 5–60 × 1–8 mm; 11(a). aparine subsp. aparine nutlets 4–5 mm 5. Stems prostrate, forming a round, flat patch on the ground or with spreading stems; internodes up to 35 mm; 6. leaves up to 25 × 3 mm; nutlets 2.5–4.5 mm 6. Plant not fleshy often forming a flat round patch; nutlets 3.0–4.5 mm; flowering 8–9(–3). 11(b,i). aparine subsp. agreste var. agreste 6. Plant fleshy often with spreading stems; nutlets 2.5–3.0 mm; flowering 5–7 11(b,ii). aparine subsp. agreste var. marinum 8. 7. Corolla yellow 10. 7. Corolla white to cream, greenish or pale pink 8. Leaves usually more than 3 mm wide; corolla often 7. × pomeranicum more than 3 mm in diameter 8. Leaves 0.5–1.0(–2.0) mm wide; corolla 2–3 mm in 9. diameter 9. Stems 25–120 cm; leaves 6–30(–35) mm; internodes 30–60 mm; panicle 50–200 × 30–100 mm, usually more 6(a). verum subsp. verum or less open 9. Stems 2–25 cm; leaves 1.5–8.0 mm; internodes 5–30 mm; panicle 20–50 × 10–30 mm, dense 6(b). verum subsp. maritimum 10. Leaves obtuse to acute at apex, but never apiculate, 11. mucronate or awned 14. 10. Leaves apiculate, mucronate or awned at apex 11. Leaves 0.6–1.0 mm wide; panicle obconical, widest near the top, the branches erect-ascending 4. constrictum 11. Leaves 0.8–8.0(–10) mm wide; panicle cylindrical or 12. broadly pyramidal, the branches spreading 12. Leaves 4–10(–12) × 0.8–2.0 mm; flowers 2.0–3.5 mm in diameter; nutlets 1.2–1.5 mm 5(a). palustre subsp. palustre 12. Leaves 10–30(–40) × 1.5–8.0(–10) mm; flowers 3.5–5.5 13. mm in diameter; nutlets 2.0–3.5 mm 13. Leaves 10–16(–20) × 1.5–4.0 mm; flowers 3.5–4.0 mm in diameter; nutlets 2.0–3.0 mm 5(b). palustre subsp. tetraploideum 13. Leaves 12–30(–40) × 3–8(–10) mm; flowers 3.5–5.5 mm in diameter; nutlets 2.5–3.5 mm 5(c). palustre subsp. elongatum 14. Stems glabrous or hairy, but without downwardly 15. directed pricklets on the angles 14. Stems glabrous or hairy, with downwardly directed 21. pricklets on the angles 15. Leaves with forwardly directed pricklets on the margin 16. 15. Leaves with at least some backwardly directed pricklets 19. on the margin 16. Leaves 4–11(–15) × 0.5–2.5 mm; corolla lobes acute at 10. saxatile apex 16. Leaves 8–30 × 1–7 mm; corolla lobes apiculate to 17. strongly mucronate or cuspidate at apex 17. Corolla 2–3 mm in diameter 7(a). mollugo subsp. mollugo 18. 17. Corolla 2.5–5.0 mm in diameter 18. Leaves gradually narrowed towards the apex; 7(b). mollugo subsp. erectum inflorescence narrow

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18. Leaves abruptly narrowed towards the apex; inflorescence 7(c). mollugo subsp. pycnotrichum broadly ovoid 19. Nutlets with minute, high-domed, subacute papillae 9. sterneri 20. 19. Nutlets with minute, low-domed to rounded papillae 20. Plant loosely caespitose with few stolons; stems not reddish at base; leaves 10–16(–30) × 1.4–1.6(–3.0) mm, often falcate; inflorescence lax; pedicels 1.0–1.5 mm 8(a). pumilum subsp. pumilum 20. Plant densely caespitose with numerous stolons; stems often reddish at base; leaves 4–10 × 0.5–1.0 mm, straight; inflorescence dense; pedicels under 1.0 mm 8(b). pumilum subsp. fleurotii 21. Leaves with forwardly directed pricklets on the margin 22. 21. Leaves with backwardly directed pricklets on the margin 23. 22. Flowers 2.0–2.5 mm in diameter; nutlets 4–5 mm 14. verrucosum 22. Flowers 0.5–1.0 mm in diameter; nutlets 0.8–1.0 mm 15. parisiense 23. Perennial herb with slender, rooting rhizomes; nutlets 3. uliginosum 1.0–1.5 mm 24. 23. Annual herbs with fibrous roots; nutlets 1.5–5.0 mm 24. Peduncles and pedicels divaricate at various angles in fruit, but straight; nutlets glabrous and smooth 12(a). spurium subsp. spurium 24. Peduncles and/or pedicels strongly recurved in fruit; 13. tricornutum nutlets papillose

Section 1. Hylaea (Griseb.) Ehrend. Perennial herbs with far-creeping rhizomes, smelling of hay when dried. Stems glabrous, except for a ring of hairs below the nodes, 4-angled. Leaves in whorls of 6–9, with 1 main vein, with forwardly directed marginal prickles. Inflorescence in terminal and lateral cymes, lax, few-flowered. Corolla funnel-shaped, white; lobes 4, obtuse at apex. Nutlet dry, with hooked bristles. 1. G. odoratum (L.) Scop. Sweet Woodruff Asperula odorata L. Perennial herb hay-scented when dried, with slender, branched, far-creeping rhizomes. Stems 15–45 cm, erect, pale yellowish-green, sometimes purplish below the nodes, 4-angled, the angles with a narrow, colourless margin, glabrous except for colourless hairs below the nodes, leafy, branched. Leaves in distant whorls; lamina yellowish-green on upper surface, paler beneath, the lowest of 6 in a whorl, 5–12 × 2–4 mm, oblanceolate, obtuse or subacute at apex, entire and narrowed at the base to a short, winged petiole which is often dark purple at the base; the median 6–8(–9) in a whorl, 25–45 × 6–15 mm, narrowly elliptical, oblongelliptical, oblong-lanceolate or oblanceolate, cuspidate at apex, entire and narrowed at base to a short, winged petiole, all glabrous except for the forwardly directed marginal prickles which are sometimes also on the midrib beneath and 1-veined. Flowers 4–7 mm in diameter, sweet-scented, in terminal or lateral cymes, partial inflorescence usually of 3, on long, pale yellowish-green stalks; cymes 3, forming an umbel; pedicels 1–3 mm in flower, 3–10 mm in

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G. boreale L. G. uliginosum L.

G. constrictum Chaub.

G. odoratum (L.) Scop.

G. palustre L. subsp. palustre

G. palustre L. subsp. tetraploideum A. R. Clapham ex Franco

G. palustre L. subsp. elongatum (C. Presl) Arcangeli

G. verum L.

G. pumilum Murray

G. saxatile L. G. verrucosum Huds. G. sterneri Ehrend. Fruits of Galium L.

G. mollugo L.

G. parisiense L.

6. Galium

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G. aparine L. subsp. agreste P. D. Sell

G. aparine L. subsp. aparine

G. spurium L. subsp. spurium

G. spurium L. subsp. infestum (Waldst. & Kit.) Schübler & Martens G. tricornutum Dandy Fruits of Galium L.

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fruit; bracts small, linear-lanceolate, obtuse at apex. Calyx a minute, annular ridge. Corolla funnel-shaped, white; tube 1.0–1.5 mm; lobes 4, 2.0–3.5 mm, oblong, obtuseapiculate at apex, slightly recurved. Stamens 4; filaments about 0.5 mm, white; anthers pale cream. Style 1, bifid, white; stigmas pale cream. Nutlet 2–3 mm, ovoid, with hooked, black-tipped bristles 0.5–1.0 mm. Flowers 5–6. Homogamous. Fragrant and visited chiefly by flies and bees. 2n = 44. There is difficulty in drawing the line between the genera Asperula and Galium. G. odoratum has funnel-shaped flowers, but with a corolla-tube hardly longer than the lobes. It is here included in Galium although it lacks the rotate corolla typical of that genus. Native. Locally abundant in woods on damp calcareous or base-rich soils, up to 640 m in Scotland. Throughout most of Great Britain and Ireland except the Channel Islands and the Outer Isles of Scotland but local in East Anglia, central and northern Scotland and Ireland; a garden escape in Jersey. North and central Europe and montane woods in Italy and the Balkans; North Africa; western Asia, rare in central and eastern Asia. A member of the European Temperate element. The dried leaves retain their scent for many months and were formerly used for an all-purpose domestic freshener. Section 2. Platygalium W. D. J. Koch Perennial herbs with rooting stolons. Stems shortly rigidhairy, 4-angled. Leaves in whorls of 4, with 3 parallel veins, obtuse at apex. Inflorescence many-flowered, oblongpyramidal, ultimate branches without bracts. Corolla rotate, white; lobes 4, apiculate at apex. Nutlet dry, with appressed, hooked bristles. 2. G. boreale L. Northern Bedstraw Aparine boreale (L.) Hill Perennial herb with rooting stolons. Stems 20–65 cm, pale green, erect, stout, 4-angled, very shortly rigid-hairy, at least on the angles, leafy, with erect, ascending branches. Leaves in whorls of 4; lamina 10–40 × (2–)3–5(–8) mm, bright green on upper surface, paler beneath, turning black when dried, elongate-lanceolate or narrowly elliptical, widest at or below the middle, obtuse at apex, entire, narrow or rounded at base, sessile, with very short, rigid simple eglandular hairs at least on the margins and veins, indistinctly 3veined. Flowers about 3 mm in diameter, in a dense, oblongpyramidal, terminal leafy panicle with ascending branches exceeding their bracts; pedicels (1-)2–3 mm; bracts 2–3 mm, ovate, obtuse at apex. Calyx a minute, annular ridge. Corolla rotate, white; tube very short; lobes 4, 1.0–1.5 mm, ovate, apiculate at apex. Stamens 4; filaments 0.5–1.0 mm, white; anthers pale yellow. Style 1, bifid, pale green; stigmas pale yellow. Nutlets 1.5–2.0 mm, olive-brown, oblong, with dense, whitish, appressed, hooked bristles. Flowers 7–8. Bisexual and slightly protandrous. Visited by various small insects. 2n = 44, 66. Plants described as var. stenophyllum Druce have narrower leaves, are more hairy and have the hairs on the fruits longer and more erect. They were described from sand dunes, but similar plants occur in other habitats.

Native. Rocky slopes, moraine, scree, shingle, sides of streams and lakes and stable sand dunes up to 1060 m in Scotland. Locally frequent in Great Britain north from Yorkshire, very local in north and south Wales, in scattered localities in central, west and north Ireland. North and central Europe southwards to northern Italy, Yugoslavia and Bulgaria; Caucasus; north and central Asia east to Japan; North America; Greenland. A member of the Circumpolar Boreo-temperate element. Section 3. Trachygalium Schumach. Perennial herbs with slender, rooting rhizomes. Stems 4angled, with downwardly directed pricklets on the angles. Leaves (4–)6–8 in a whorl, shortly awned at apex, with 1 main vein. Inflorescence oblong-ovoid, the ultimate branches without bracts. Corolla broadly cup-shaped, white; lobes 4, acute at apex. Nutlets rugulose and glabrous. 3. G. uliginosum L. Fen Bedstraw Perennial herb with slender, rooting rhizomes. Stems 10– 60(–100) cm, pale green, weak, decumbent or ascending, 4-angled, with downwardly directed pricklets on the angles, leafy, branched. Leaves (4–)6–8 in a whorl; lamina (8–)10–20(–25) × 2–3(–4.5) mm, yellowish-green on upper surface, paler beneath, not blackening when dry, linear-oblanceolate, obtuse with an awn at apex, entire, gradually narrowed to the sessile base, glabrous, with backwardly directed pricklets on the margin, 1-veined. Flowers 2.5–3.0 mm in diameter, sweet-scented, in small, axillary corymbs forming an oblong-ovoid panicle, the ultimate branches usually without bracts; pedicels 2–5 mm; bracts 1– 2 mm, linear-lanceolate, acute at apex. Calyx a minute annular ridge. Corolla broadly cup-shaped, white; tube 0.3–0.5 mm; lobes 4, 1.0–1.5 mm, broadly ovate, acute at apex. Stamens 4; filaments 0.5–0.6 mm, white; anthers yellow. Styles 1, bifid, pale green; stigmas pale green. Nutlets 1.0–1.5 mm, green turning dark brown, oblong, rugulose, glabrous, on deflexed stalks. Flowers 7–8. Bisexual, protandrous and visited by small insects. 2n = 22, 44. Native. Fens and base-rich, marshy places. Scattered in Great Britain and Ireland except for northern Scotland, north Ireland and Channel Islands. Europe from Portugal and north Spain, northern Italy and Greece to about 70◦ N in Scandinavia, extending eastwards to eastern Asia. A member of the Eurasian Boreo-temperate element. Section 4. Aparinoides (Jord.) Gren. Perennial herbs with slender, creeping rhizomes. Stems 4-angled, usually with downwardly directed prickles on the angles. Leaves 4–6 in a whorl, sometimes blackish when dried, obtuse to subacute but not awned at apex and 1-veined. Inflorescence cylindrical to broadly pyramidal or obconical, the partial inflorescences cymose, 1–manyflowered and the ultimate branches without bracts. Corolla shallowly funnel-shaped, white, greenish or pinkish; lobes 4, acute at apex. Nutlets more or less smooth to rugose or tuberculate, glabrous.

6. Galium 4. G. constrictum Chaub. Slender Marsh Bedstraw G. debile Desv., non Hoffmanns. & Link Perennial herb with slender, creeping, branched rhizomes and fibrous roots. Stems (5–)10–30(–60) cm, pale green, prostrate then ascending or more or less erect, slender, 4angled, the angles often whitish and smooth or with some small, downwardly directed pricklets, otherwise glabrous, leafy, much branched. Leaves 4–6 in a whorl; lamina 3– 10 × 0.6–1.0 mm, bright medium green on upper surface, paler beneath, not blackening when dry, linear or linearoblanceolate, broadest above the middle, obtuse or subacute but never mucronate, entire, margins rough with forwardly directed pricklets or revolute and smooth, narrowed at base, glabrous; 1-veined. Flowers 2.0–3.5 mm in diameter, in axillary corymbs forming an obconical panicle, those of the partial inflorescences more or less tightly together, the branches erect-ascending, the ultimate one bractless; pedicels 1–4 mm, slightly elongating after anthesis; bracts 1–3 mm, linear, obtuse or subacute at apex. Calyx a minute, annular ridge. Corolla shallowly funnelshaped, pinkish-white or greenish; tube very short; lobes 4, 0.6–1.5 mm, ovate, acute at apex. Stamens (3–)4; filaments 0.5–0.7 mm, whitish; anthers brown. Style 1, bifid, white; stigmas cream. Nutlets 2–3 mm, subglobose, tuberculate, glabrous, their pedicels not divaricate. Flowers 5–7. 2n = 24. Native. Marshy places, ditches and pond-sides. Very local in the New Forest, Hampshire, 2 sites in Wiltshire, and Channel Islands. It has been extinct in the Bovey Basin, Devon for many years. Records from Yorkshire are thought to be erroneous. South-west and south Europe from France, Spain and Portugal to southern Balkans and Crete; north-west Africa; Crimea; Caucasia. A member of the Mediterranean-Atlantic element. 5. G. palustre L. Common Marsh Bedstraw Perennial herb with slender, creeping rhizomes. Stems up to 100(–150) cm, pale green, decumbent or ascending to more or less erect, 4-angled, usually with downwardly directed pricklets on the sometimes more or less whitish angles, glabrous, leafy, branched. Leaves in whorls of 4–6; lamina 4–30(–40) × 0.8–8.0(–10) mm, bright medium green on upper surface, paler beneath, usually blackening when dry, narrowly to broadly oblanceolate, obtuse at apex, never mucronate, entire, narrowed at base, usually rough on margin and midrib; 1-veined. Flowers 2.0–5.5 mm in diameter, in a cylindrical or broadly pyramidal panicle with spreading branches, the ultimate branches without bracts; pedicels 1.0–5.5 mm; bracts 1–3 mm, linear-lanceolate, acute at apex. Calyx a minute, annular ridge. Corolla shallowly funnel-shaped, white; tube 0.4–0.6 mm; lobes 4, 1.3–2.5 mm, ovate, acute at apex. Stamens 3–4; filaments 4–6 mm, white; anthers reddish or brownish. Style 1, bifid, white; stigmas cream. Nutlets 1.2–3.5 mm, globose, almost smooth to rugose, glabrous, on spreading pedicels. Flowers 5–7. Bisexual, protandrous, visited by bees. The three subspecies are difficult to define, but have a distinct appearance.

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(a) Subsp. palustre G. witheringii Sm.; G. palustre var. serrulatum Druce; G. palustre var. witheringii (Sm.) Syme; G. palustre forma angustifolium Druce; G. palustre var. microphyllum Lange; G. palustre var. gracile auct.; G. palustre var. angustifolium (Druce) Druce Stems up to 50 cm. Leaves 4–10(–12) × 0.8–2.0 mm. Flowers 2.0–3.5 mm in diameter. Nutlets 1.2–1.5 mm. 2n = 24. (b) Subsp. tetraploideum A. R. Clapham ex Franco G. palustre var. lanceolatum Uechtr. Stems up to 80 cm. Leaves 10–16(–20) × 1.5–4.0 mm. Flowers 3.5–4.0 mm in diameter. Nutlets 2.0–3.0 mm. 2n = 48. (c) Subsp. elongatum (C. Presl) Arcangeli G. elongatum C. Presl; G. palustre var. elongatum (C. Presl) Syme Stems up to 120(–150) cm. Leaves 12–30(–40) × 3–8(–10) mm. Flowers 3.5–5.5 mm in diameter. Nutlets 2.5–3.5 mm. 2n = 96, 144. Native. Damp meadows, pond-sides, ditches, marshes and fens. Common throughout Great Britain and Ireland. Throughout most of Europe; Caucasus; western Siberia; North Africa; eastern North America. A member of the Eurosiberian Boreo-temperate element. It is probable all three subspecies occur throughout the British Isles. Subsp. palustre seems to be in drier and more acidic areas than subsp. elongatum. Subsp. elongatum may go further east on the Continent of Europe than subsp. palustre. The ecology and distribution of subsp. tetraploideum are not known, but it is widespread in Great Britain and could be the commonest subspecies in calcareous localities. Section 5. Galium Perennial herbs with slender, creeping stolons. Stems with 4 raised lines, hairy, without pricklets. Leaves 8–12 in a whorl, 1-veined. Inflorescence many-flowered with patent branches, with bracts up to the ultimate branches. Corolla rotate, yellow. Nutlets smooth, glabrous. 6. G. verum L. Lady’s Bedstraw G. floridum Salisb. nom. illegit. Perennial herb with slender, creeping stolons. Stems 2–120 cm, pale yellowish-green, sometimes faintly tinted brownish, more or less terete with 4 raised lines, decumbent to erect, with dense, very short, pale simple eglandular and glandular hairs, leafy, branched. Leaves 8–12 in a whorl; lamina 1.5–30(35) × 0.5–1.0(–2.0) mm, yellowish-green on upper surface, paler beneath, linear, obtuse-mucronate at apex, entire, sessile, 1-veined, shining and usually very shortly hairy above, densely and shortly hairy beneath, with revolute margins. Inflorescence a terminal, leafy, compound panicle with bracts up to the ultimate branches; peduncles short, densely and shortly hairy. Calyx a minute annular ridge. Corolla 2–3 mm in diameter, sweet-scented, deep yellow, glabrous, rotate; lobes 4, oblong or oblong-ovate and subacute at apex. Stamens 4; filaments deep yellow; anthers golden yellow. Style 1, bifid, cream; stigmas cream,

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shorter than anthers. Nutlets 1.0–1.5 mm, ultimately black, globose, smooth and glabrous. Flowers 6–8. Coumarin scented and visited by many small insects, especially flies. 2n = 22, 44. (a) Subsp. verum Stems 25–120 cm. Leaves 6–30(–35) mm; internodes 30–60 mm. Panicle 50–200 × 30–100 mm. (b) Subsp. maritimum (DC.) Adema G. verum var. maritimum DC.; G. verum var. littorale Br´eb. Stems 2–25 cm. Leaves 1.5–8.0 mm; internodes 5–30 mm. Panicle 20–50 × 10–30 mm, dense. Native. Dry grassy places especially on calcareous soils, waysides, hedgebanks, heaths, dunes and shingle. Throughout Great Britain and Ireland. Widespread in Europe north to northern Scandinavia; Asia east to Japan; introduced in North America. A variable species in its broader range, which is a member of the Eurasian Boreo-temperate element. Subsp. verum occurs throughout most of Europe. Subsp. maritimum is the plant of coastal dunes, shingle and cliffs and also occurs on sandy heaths inland. It is found in similar habitats in western Europe. There is in addition a large plant with interrupted inflorescence and pale flowers which does not seem to be G. × pomeranicum and may be referable to subsp. wirtgenii (F. W. Schultz) Oborny. It could have been introduced with wild flower seed. G. verum probably derived its vernacular name from the old custom of including it in straw mattresses, and especially in the beds of women about to give birth. Section 6. Leiogalium Ledeb. Perennial herbs with far-creeping rhizomes. Stems 4angled, glabrous or hairy. Leaves 5–8 in a whorl, 1-veined. Inflorescence many-flowered, the ultimate branches without bracts. Corolla white, rotate. Nutlets rugulose, glabrous. 7. G. mollugo L. Hedge Bedstraw G. vulgare Gray nom. illegit. Perennial herb, hay-scented when dried, with slender, branched, far-creeping rhizomes. Stems 15–150 cm, pale green, sometimes tinted brownish, procumbent to erect, 4-angled, glabrous or hairy especially below or beneath the nodes, the hairs short, patent, pale and simple eglandular, leafy, branched. Leaves 5–8 in a whorl, with long internodes; lamina 8–30 × 1–7 mm, medium green on upper surface, paler beneath, elliptical-obovate, oblongoblanceolate, oblanceolate or linear-oblanceolate, rounded or subacute and usually mucronate at apex, entire, glabrous on upper surface, glabrous or shortly hairy beneath, usually rough with short, forwardly directed rigid hairs on the margin, 1-veined. Inflorescence a broadly pyramidal to oblong panicle of spreading or ascending, cymose branches, the ultimate branches without bracts; pedicels 1.5–3.0(–4.0) mm, slender, glabrous. Calyx a minute annular ridge. Corolla 2–5 mm in diameter, white, strongly scented, rotate; tube very short; lobes (3–)4(–5), ovate, rounded and mucronate or cuspidate at apex. Stamens 4, exserted; filaments about 0.5 mm, white; anthers red. Style 1, bifid, white; stigmas pale green, about equalling anthers. Nutlets

1–2 mm in diameter, brown or blackish, globose, rugulose, glabrous. Flowers 6–9. Protandrous and visited chiefly by small flies. 2n = 44. Very variable. The following three subspecies occur in the British Isles with additional taxa in the total range of the species. In Continental Europe G. mollugo is considered to be a diploid (2n = 22) and G. album (2n = 44) a tetraploid, which is treated as a distinct species. Only tetraploids have been recorded from the British Isles and some botanists would regard all our plants as G. album. (a) Subsp. mollugo G. elatum Thuill.; G. mollugo subsp. elatum (Thuill.) Syme nom. illegit.; G. insubricum Gaudin; G. mollugo subsp. scabrum Hook. fil.; G. mollugo var. insubricum (Gaudin) Druce; G. mollugo var. scabrum (Hook. fil.) Druce; G. mollugo subsp. elatum var. bakeri Syme Stems weak, ascending to prostrate. Leaves mostly oblanceolate to elliptical-obovate. Inflorescence broad, with branches mostly at more than an angle of 45◦ and rather lax. Corolla 2–3 mm in diameter; pedicels strongly divaricate at fruiting. (b) Subsp. erectum Syme G. erectum Huds. 1778, non 1762; G. album Mill.; G. diffusum D. Don ex Hook.; G. aristatum auct.; G. mollugo subsp. erectum var. aristatum auct.; G. cinereum auct. Stems decumbent to more or less erect. Leaves oblanceolate to linear-oblanceolate, gradually narrowed towards the apex. Inflorescence rather narrow, with branches mostly at an angle of less than 45◦ and rather dense. Corolla 2.5–5.0 mm in diameter; pedicels weakly divaricate at fruiting. (c) Subsp. pycnotrichum (Heinr. Braun) O. Schwarz G. pycnotrichum Heinr. Braun; G. album subsp. pycnotrichum (Heinr. Braun) Krendl Stems erect, robust. Leaves oblong to broadly oblanceolate, abruptly narrowed towards the apex. Inflorescence usually broadly ovoid. Corolla 2.5–5.0 mm. Native. Grassy places, hedgerows and waysides, mainly on well-drained, base-rich soils. Throughout the British Isles, but rare in Scotland, Wales and Ireland; common in southern England and the Channel Islands. Throughout Europe; western Asia; North Africa. A member of the European Boreo-temperate element which is widely naturalised in North America and elsewhere. The distribution of the two native subspecies is not known, but subsp. erectum is usually on drier, more calcareous soils. Subsp. pycnotrichum has been recorded from Walthamstow Reservoirs. It is a native of eastern Europe which could have been introduced with grass seed or wild flower seed and should be looked for elsewhere. × verum = G. × pomeranicum Retz. G. × ochroleucum Wolf ex Schweigg.; G. × rothschildii Druce nom. nud. This partially fertile hybrid, which appears to backcross with the parents, has the stems more angled than in G. verum, the leaves usually more than 1.5 mm wide, narrowly lanceolate and usually not darkening on drying with their

6. Galium margins not prominently recurved, the inflorescence with fairly long branches not very hairy, the pedicels fairly stout, the corolla often more than 3 mm in diameter, bright yellow to whitish and its lobes more or less apiculate and the fruits smooth or rugulose. 2n = 44. Native. Frequently occurs, with the parents, along the southern coast and on inland chalk and limestone in Great Britain north to Midlothian and in the Channel Islands, and is rare in south and east Ireland. It is common in Continental Europe. There is a complicated series of nothosubspecies, only one of which, nothosubsp. hillardiae (Druce) P. D. Sell (G. hillardiae Druce), G. mollugo subsp. mollugo × verum subsp. maritimum, has a satisfactory name. Section 7. Leptogalium Lange Perennial herbs usually with stolons. Stems 4-angled, glabrous or hairy. Leaves 5–9(–10) in a whorl, 1-veined. Inflorescence comparatively few-flowered, the ultimate branches usually without bracts. Corolla white, rotate. Nutlets papillose, glabrous. 8. G. pumilum Murray Slender Bedstraw G. asperum Schreb.; G. laeve Thuill.; G. sylvestre Pollich, non Scop.; G. pusillum auct.; G. sylvestre var. nitidulum (Thuill.); G. austriacum auct.; G. pumilum var. nitidum (Thuill.) Druce; G. umbellatum Lam.; G. nitidulum Thuill. Loosely to densely caespitose perennial herb with few to numerous stolons. Stem 10–40(–70) cm, pale green, sometimes slightly reddish at base, ascending to erect, with a few, prostrate, non-flowering ones, 4-angled, glabrous or with short, pale simple eglandular hairs, leafy, little branched. Leaves in whorls of 5–9(–10); lamina 6–16(–30) × 0.5–1.6(–3.0) mm, greyish-green on upper surface, paler beneath, linear-oblanceolate to oblanceolate, sometimes falcate, obtuse-mucronate at apex, entire, narrowed to the sessile base, 1-veined, margins with backwardly directed pricklets and sometimes patent simple eglandular hairs. Flowers 2–3 mm in diameter, in a lax to dense panicle of axillary cymes exceeding the stem internodes; pedicels 0.5– 1.5 mm, glabrous. Calyx a minute annular ridge. Corolla 1.0–1.5 mm, creamy white, rotate, divided two-thirds of the way to the base; lobes 4, triangular-ovate, acute at apex. Stamens 4; filaments 0.4–0.5 mm, white; anthers yellow. Style 1, bifid, white; stigmas cream. Nutlets 1.4–1.5 mm, brown, ovoid, glabrous, with minute low dome-shaped papillae. Flowers 6–8. (a) Subsp. pumilum Plant loosely caespitose, with few stolons. Stems not reddish at base. Leaves 10–16(–30) × 1.4–1.6(–3.0) mm, narrowly oblanceolate, often falcate. Inflorescence lax; pedicels 1.0–1.5 mm. 2n = 88. (b) Subsp. fleurotii (Jord.) P. D. Sell G. fleurotii Jord. Plant densely caespitose, with numerous stolons. Stems often reddish at base. Leaves 6–10 × 0.5–1.0 mm, linearoblanceolate to narrowly oblanceolate, straight. Inflorescence dense; pedicels under 1.0 mm. 2n = 88.

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Native. Dry chalk and limestone grassland and limestone cliffs. Very local in England south-east of a line from the River Severn to the River Humber. West and central Europe from France northwards to Denmark and eastwards to the Baltic States, Poland and Roumania. A member of the European Temperate element. Subsp. fleurotii is found on cliffs at Cheddar and the surrounding areas in Somerset and similar localities in central France. The remaining plants are referable to subsp. pumilum. 9. G. sterneri Ehrend. Limestone Bedstraw G. pumilum subsp. septentrionale Sterner ex Hyl.; G. montanum auct.; G. pumilum var. glabrum auct. Perennial herb overwintering as a prostrate mat. Stems (5–)8–25(–30) cm, pale green, often red at the base, many prostrate and non-flowering at anthesis, flowering ones ascending or erect, slender, 4-angled, glabrous or with short, patent, pale simple eglandular hairs, the lower internodes short so that the leaf-whorls are crowded, the middle internodes 20–50 mm, 2.0 to 3.5 times as long as the leaves. Leaves in whorls of (6–)7–8(–10); lamina (5–)7– 14(–15) × 0.9–1.6(–2.3) mm, medium green on upper surface turning dark green when dry, paler beneath, linear to narrowly oblanceolate, mucronate at the hyaline apex, entire, narrowed to the sessile base, 1-veined, the margins with many, curved, backwardly directed pricklets. Flowers 2.3–3.5 mm in diameter, in compact, ascending cymes forming a compact, pyramidal panicle; pedicels 1–2 mm, glabrous. Calyx a minute annular ridge. Corolla 1.4–1.5 mm, creamy-white, rotate, divided two-thirds of the way to the base; lobes 4, triangular-ovate, acute at apex. Stamens 4; filaments 0.4–0.5 mm, white; anthers reddish. Style 1, bifid, white; stigmas cream. Nutlets 1.0–1.4 mm, ovoid, glabrous, with minute, high-domed, subacute papillae. Flowers 6–7. Protandrous, bisexual, self-incompatible. Visited by flies and other small insects. 2n = 22, 44. The plants from Caernarvonshire, the Durness limestone in Sutherland and western Ireland are diploid. They tend to be more slender and smaller in all their parts (flowering shoots 8–15(–20) cm, leaves 7–9 mm, flowers 2.3–3.0 mm in diameter and fruit 1.0–1.2 mm) than the tetraploid, but their measurements overlap too greatly to recognise them as two taxa. Native. Limestone or other base-rich grassland and calcareous or basic igneous rocks. Local in Britain north-west of a line from the River Severn to the River Humber where it replaces G. pumilum and is the calcicole counterpart of the calcifuge G. saxatile; western and north-eastern Ireland. North-west Europe. A member of the Suboceanic Borealmontane element. 10. G. saxatile L. Heath Bedstraw G. rupicola Bertol.; G. harcynicum Weigel; G. montanum Huds.; G. procumbens Stokes nom. illegit.; G. pulchellum Salisb. nom. illegit.; G. harcynicum var. alpestre auct.; G. harcynicum var. transiens auct. Mat-forming perennial herb with a slender tap-root and filiform, more or less rooting stolons. Stems (8–)15–35 (–40) cm, pale yellowish-green, sometimes tinted purplish, 4-angled, glabrous and smooth, the non-flowering ones

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numerous and prostrate, the flowering ones decumbent or ascending, leafy, much branched. Leaves in whorls of (5–)6–7(–8); lamina 4–11(–15) × (0.5–)1.5–2.5 mm, thin, medium green on upper surface, paler beneath, obovate to oblanceolate or oblong-oblanceolate, abruptly contracted at apex into a short, hyaline awn, the margin entire, with forwardly directed pricklets and weakly revolute, gradually narrowed at base; with one slender and distinct vein. Flowers 2.5–4.0 mm in diameter, in few-flowered, ascending cymes longer than their subtending leaves, but shorter than the stem internodes and forming a cylindrical panicle; pedicels 2–5 mm, divaricate after flowering. Calyx a minute annular ridge. Corolla 1.5–2.0 mm, pure white, rotate; lobes 4, flat, patent, ovate, acute at apex. Stamens 4; filaments incurved, white; anthers greenish-yellow. Style 1, bifid, pale green; stigmas pale greenish, shorter than anthers. Nutlets 1–2 mm in diameter, glabrous, with high-domed, subacute papillae. Flowers 5–8. Bisexual, protandrous, selfincompatible. 2n = 44. All our plants seem to be tetraploid. Diploid plants, subsp. vivianum (Kliphuis) Erend., smaller in all their parts, seem to be confined to Spain and Portugal. Native. Heaths, moors, dry grassland, rocky places and open woods on acid soils. Throughout most of the British Isles but absent in calcareous areas. West and west-central Europe, extending eastwards very locally to north-west Russia and the central Carpathians. A member of the Suboceanic Temperate element. × sterneri This hybrid is intermediate in the shape of the leaves and the patent marginal pricklets, but there are fertile plants of G. saxatile which show a few morphological characters of G. sterneri. 2n = 33, 55. Native. Recorded only in a few upland areas, where it is associated with rich ledge-vegetation although the parents grow in close proximity in many other localities. Caernarvonshire, Perthshire, Inverness-shire, Forfarshire and Sutherland. Not known elsewhere. Section 8. Kolgyda Dumort. Section Aparine W. D. J. Koch Annual herbs with fibrous roots. Stems 4-angled, glabrous or hairy. Leaves 5–10 in a whorl, 1-veined. Inflorescence pyramidal or oblong, with numerous bracts, the cymes fewflowered. Corolla rotate, white, greenish or reddish. Nutlets with hooked bristles or glabrous, papillose or smooth. 11. G. aparine L. Cleavers Aparine vulgaris Hill; G. uncinatum Gray nom. illegit. Annual herb with fibrous roots. Stems up to 3 m, procumbent to scrambling-erect, dull yellowish-green, 4-angled, with strongly downwardly directed pricklets on the angles, with more or less numerous, pale simple eglandular hairs near the nodes, leafy, branched. Leaves in whorls of 6–9; lamina 5–60 × 1–8 mm, dull, medium yellowish-green on upper surface, paler beneath, linear to linear-oblanceolate, oblong-oblanceolate or oblanceolate, abruptly contracted to an awned apex, entire, long-attenuate at base, sessile, the surfaces with very short, bulbous-based hairs and the mar-

gins with downwardly directed pricklets, with one prominent vein. Flowers 2–5 in axillary cymes, 2.5–3.5 mm in diameter; peduncles with downwardly directed pricklets, topped by 4–8 bracts which are miniature versions of the leaves; pedicels short with downwardly directed pricklets, elongating in fruit. Calyx minute. Corolla about 1.5 mm, white, cream or greenish, rotate, divided for about twothirds of the way to the base into 4 lobes; lobes triangularovate, apiculate at apex. Stamens 4; filaments white; anthers yellow. Style 1, bifid, whitish; stigmas cream, shorter than the anthers. Nutlets 2.5–6.0 mm in diameter, olive green or purplish, globose, covered with white, hooked bristles with their bases tuberculate and stalks divaricate. Flowers 5–9. Bisexual and protandrous. Sparingly visited by small insects. 2n = 42, 44, c. 66, c. 88. (a) Subsp. aparine Stems up to 3 m, scrambling and ascending the stems of erect plants; internodes 40–110 mm. Leaves 5–60 × 1–8 mm. Flowers white or cream. Nutlets 4–6 mm. Flowers 5–7. (b) Subsp. agreste P. D. Sell Stems up to 25 cm, prostrate, forming a round, flat patch on the ground or spreading; internodes up to 35 mm. Leaves up to 25 × 3 mm. Flowers often greenish. Nutlets 2.5–4.5 mm. (i) Var. agreste P. D. Sell Plant not fleshy, often forming a round flat patch, Leaves usually staying green. Nutlets 3.0–4.5 mm. Flowers 8–9 (–3). (ii) Var. marinum Fr. G. aparine forma condensata Druce Plant fleshy, stems often spreading. Leaves often turning reddish. Nutlets 2.5–3.0 mm. Flowers 5–6. Native. Throughout Great Britain and Ireland. Europe to 69◦ N in Scandinavia; north and west Asia; widely introduced elsewhere in both the northern and southern hemispheres. A member of the European Temperate element. Subsp. aparine is the plant of hedgerows, scrub, waste places and limestone scree, especially on fertile soils. Subsp. agreste var. agreste is the plant of arable land. It flowers and fruits in the stubble of cereal crops after the crop is cut, together with Aethusa cynapium subsp. agrestis. In set-aside it can go on flowering and fruiting during the winter. It also occurs in root crops. Subsp. aparine occurs on the margins of fields, growing out from the hedgerows and field margins. Subsp. agreste var. marinum is the plant of coastal dunes and shingle. 12. G. spurium L. False Cleavers Annual herb with fibrous roots. Stems 10–100 cm, diffuse or trailing or scrambling, slender, weak to rather stout, 4-angled, often with simple eglandular hairs at the nodes, with downwardly directed pricklets on the angles, leafy, branched. Leaves in whorls of 6–10; lamina (5–)30–35 × 2.5–4.0 mm, medium green on upper surface, paler beneath, linear-lanceolate, lanceolate or oblanceolate, gradually narrowed into the long-awned apex, entire, narrowed to the

6. Galium sessile base, with short, pale, bulbous-based simple eglandular hairs on both surfaces, the margins and midrib beneath with backwardly directed pricklets, 1-veined. Flowers 3–9 in oblong, axillary cymes, 0.8–1.3 mm in diameter; peduncles with backwardly directed pricklets, topped by only 2(–3) leaf-like bracts; bracteoles patent, straight, but often sharply bent just under the fruit, with backwardly directed pricklets. Calyx minute. Corolla about 1.0 mm, greenishcream, rotate, divided for about two-thirds of the way to the base; lobes 4, triangular-ovate, acute at apex. Stamens 4; filaments white; anthers yellow. Style 1, bifid, whitish; stigmas cream. Nutlets 1.5–3.0 mm, olive black, ellipsoid, glabrous and smooth or covered with hooked bristles which are wider towards, but not bulbous at, their base. Flowers 7–9. 2n = 20. (a) Subsp. spurium G. aparine var. spurium (L.) W. D. J. Koch Nutlets glabrous and smooth. (b) Subsp. infestum (Waldst. & Kit.) Sch¨ubler & Martens G. vaillantii DC.; G. aparine subsp. vaillantii (DC.) Hook. fil.; G. spurium var. vaillantii (DC.) Gren.; G. infestum Waldst. & Kit. Nutlets with hooked bristles. Probably introduced. Well naturalised as an arable weed around Saffron Walden in Essex since 1844; elsewhere as a casual or temporarily naturalised in scattered localities in Great Britain north to southern Scotland. Native of much of Europe, but rarer in the west and introduced in parts of the north; widespread in northern and central Asia; North Africa. A member of the Circumpolar Temperate element, but its native range is obscured by its frequent presence as an introduction in both the northern and southern hemispheres. Most British records, including the Saffron Walden plant, are subsp. infestum. Subsp. spurium has occurred only as a rare casual. In Continental Europe both subspecies sometimes grow together, especially in cultivated land. G. aparine subsp. agreste sometimes has greenish flowers, but has larger flowers and fruits. 13. G. tricornutum Dandy Corn Cleavers G. tricorne auct. Annual herb with fibrous roots. Stems 10–80(–100) cm, decumbent, sprawling or scrambling, stout, pale yellowishgreen, sharply 4-angled, glabrous except for the downwardly directed pricklets on the angles, leafy, usually much branched. Leaves 5–8 in a whorl; lamina 10–40 × 1–8 mm, medium green on upper surface, paler beneath, linear or narrowly oblanceolate, mucronate or cuspidate at apex, gradually narrowed at base, with backwardly directed pricklets on the margin and midrib beneath, otherwise glabrous; 1veined. Flowers 1.0–1.7 mm in diameter, mainly 1–3 in axillary cymes, scarcely longer than the leaves; peduncles (5–)8–15(–20) mm; pedicels 1–8 mm, curved distinctly inwards and downwards after flowering. Calyx a minute annular ridge. Corolla 0.9–1.1 mm, white, rotate, divided almost to the base into 4 ovate, acute lobes. Stamens 4; filaments minute; anthers yellow. Style 1, very

27

short, bifid; stigmas pale greenish. Nutlets 3–5 mm, pale green, didymous or reduced to one by abortion, glabrous, granulate with large, acute papillae. Flowers 5–9. Bisexual or lateral flowers often male. Little visited by insects. 2n = 44. Probably introduced. Arable and waste places, chiefly on calcareous soils. Formerly common, particularly in cereal crops, in south and east England with a few outlying localities, now rare and sporadic. Throughout much of Europe, but casual in the north and east; western and central Asia; introduced elsewhere. A member of the Eurosiberian Southern-temperate element. 14. G. verrucosum Huds. Southern Cleavers Valantia aparine L., non G. aparine L.; G. valantia G. Weber; G. saccharatum All. Annual herb with fibrous roots. Stems 5–50 cm, pale green, erect or ascending, sharply 4-angled, glabrous except for the downwardly directed pricklets on the angles, leafy, usually branched at the base. Leaves 5–6(–7) in a whorl; lamina 4–17 × 1.5–5.0 mm, medium green on upper surface, pale beneath, oblanceolate, mucronate at apex, narrowed at base, with forwardly directed pricklets on the margin. Flowers (1.0–)2.0–2.5 mm in diameter, usually in 3-flowered axillary cymes shorter than the leaves; pedicels 1–3 mm, deflexed after flowering. Calyx a minute annular ridge. Corolla 1.5–1.7 mm, greenish-white to white, rotate, divided almost to the base into 4, ovate, acute, spreading lobes. Stamens 4; filaments about 0.8 mm; anthers yellow. Style 1, very short, bifid; stigmas pale greenish. Nutlets 4–5 mm in diameter, didymous or commonly reduced to one through abortion, glabrous, coarsely and conspicuously verrucose-papillose. Flowers 6–9. Central flowers bisexual and lateral male. 2n = 22. Introduced. A rare bird-seed and spice casual. Native of southern Europe and western Asia. 15. G. parisiense L. Wall Bedstraw G. anglicum Huds.; G. parisiense subsp. anglicum (Huds.) A. R. Clapham; G. parisiense var. anglicum (Huds.) Hook. & Arn. Annual herb with fibrous roots. Stems 5–40 cm, procumbent to ascending, weak and slender, 4-angled, with downwardly directed pricklets on the angles, otherwise glabrous, leafy, usually much branched. Leaves in whorls of 5–7; lamina 3–12 × 0.8–3.0 mm, medium green on upper surface, paler beneath, turning black when dry, linear-oblanceolate to oblanceolate, mucronate at apex, entire, narrowed to the sessile base, 1-veined, at first spreading, then reflexed, glabrous or with short simple eglandular hairs, the margin and midrib beneath with forwardly directed pricklets, margin not or slightly revolute. Flowers 0.5–1.0 mm in diameter in few-flowered, axillary, stalked corymbs forming a long, narrow panicle; peduncles 2–7 mm, divaricate; pedicels 0.5–2.5 mm, slender, widely divaricate after anthesis; bracts up to 2 mm, solitary or in pairs, linear-lanceolate. Calyx minute. Corolla 0.5–0.6 mm, greenish inside, reddish outside, divided almost to the base; lobes 4, triangularovate, acute at apex. Stamens 4; filaments minute, whitish; anthers yellow. Style 1, bifid, white; stigmas cream. Nutlets

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0.8–1.0 mm, ellipsoid, glabrous, finely papillose. Flowers 6–7. 2n = 44, 66. Native. Walls and sandy banks, and sides of chalk and sandpits. Scattered localities in East Anglia and south-east England and decreasing. South, west and central Europe northwards to eastern England and eastwards to Bulgaria and Greece; North Africa; western Asia. A member of the Submediterranean-Subatlantic element. Our plant is var. parisiense (G. anglicum var. leiocarpum Tausch) with glabrous fruits. Both it and var. trichocarpum Tausch, with hooked bristles, are found in Continental Europe. 7. Cruciata Mill. Perennial herbs. Stems decumbent or ascending. Leaves in whorls of 4, all equal, more or less sessile. Flowers in dense axillary whorls of cymes, the terminal flower bisexual and the laterals male in each cyme; peduncles and pedicels deflexed under the leaves. Calyx absent. Corolla yellow, 4lobed, with a very short tube. Styles 2, free to base; stigmas capitate. Ovary 2-celled, with 1 ovule in each cell. Fruit of 1 or 2 smooth, more or less globose nutlets. Very close to the genus Valantia L. which has similarly deflexed peduncles and pedicels but in that genus they enlarge and coalesce during development and encircle the ripe fruit. Six species in Europe, the Mediterranean region and western Asia.

the central bisexual flowers protandrous, the laterals male. Visited by bees and flies. 2n = 22. Native. Grassy places, hedgerows, scrub and rough ground, mostly on calcareous soils. Common in Great Britain north to central Scotland; introduced in two places in east Ireland. Most of Europe northwards to Holland, northern Germany, northern Poland and central Russia; northern Turkey, Caucasus, Iran and the western Himalayas. A member of the Eurosiberian Temperate element. 8. Rubia L. Evergreen or deciduous perennial herbs. Stems trailing to scrambling, often woody at base. Leaves in whorls of 4–6, all equal, more or less sessile. Flowers in diffuse axillary and terminal panicles. Calyx a minute annular ridge. Corolla pale yellowish-green, 5-lobed, with a very short tube. Style 1, bifid, fused below; stigmas capitate. Ovary 2-celled, with 1 ovule in each cell. Fruit a subglobose, succulent, black mericarp, with 1 seed and derived from only 1 cell of the ovary, rarely didymous with the parts not separating. About 40 species with a wide distribution in Europe, temperate Asia, Africa and America. Rind, W. (1872). A history of the vegetable kingdom. London. Weston, I. (1997). Rubia tinctorum (Madder) in South Lines (v.c. 53). B.S.B.I. News 74: 42–44.

Grime, J. P. et al. (1988). Comparative plant ecology. London. [As Galium cruciata.]

1. Leaves dark green, with obscure lateral veins beneath; 1. peregrina anthers 0.2–0.3 mm 1. Leaves bright green, with prominent lateral veins 2. tinctorum beneath; anthers 0.5–0.6 mm

1. C. laevipes Opiz Crosswort C. chersonensis auct.; Galium cruciata (L.) Scop.; Valantia cruciata L.; Aparine latifolia Moench nom. illegit.; Galium valantia Gaertn., C. A. Mey. & Scherb.; Valantia ciliata Opiz ex J. & C. Presl Perennial herb with a weak primary root, slender rhizomes with extensive adventitious roots and subterranean stolons. Stems 15–60 cm, yellowish-green, slender, 4angled, decumbent to ascending, with soft, pale, spreading simple eglandular hairs 1–2 mm, usually with several ascending branches from near the base. Leaves in whorls of 4; lamina 12–25 × 4–10 mm, largest in the middle of the stem but mostly shorter than the adjacent internodes, yellowish-green on upper surface, paler beneath, lanceolate or ovate to ovate-elliptical, obtuse to acute at apex, entire, narrowed at base, with numerous, soft, short to medium simple eglandular hairs on both surfaces, 3-veined. Flowers 2–3 mm in diameter, honey-scented, in 5- to 9-flowered cymes the terminal flower bisexual and the laterals male in each cyme; pedicels with more or less numerous, soft simple eglandular hairs, elongating and recurved in fruit; peduncles with 2 bracteoles up to 8 mm which scarcely enlarge in fruit. Calyx absent. Corolla 2–3 mm, bright yellow, rotate, 4-lobed, with a very short tube; lobes ovate, acuminate at apex. Stamens 4; filaments very short, whitish; anthers cream. Styles 2, whitish; stigmas cream, capitate. Nutlets 1.8–2.7 mm in diameter, blackish, more or less globose, smooth, glabrous. Flowers 4–7. Andromonoecious,

1. R. peregrina L. Wild Madder R. anglica Huds. Evergreen perennial herb with a long, creeping stock. Stems 30–150 cm, pale green, trailing or scrambling-ascending, woody in lower part, 4-angled, the angles with downwardly directed prickles, glabrous, branched, leafy. Leaves 4–6 in a whorl; lamina 10–60 × 3–20 mm, dark green, coriaceous and shiny on upper surface, paler beneath, linear to broadly ovate-elliptical, obtuse-apiculate at apex, entire, narrowed at base, rigid and leathery, the cartilaginous margins and the underside of the midrib with backwardly directed prickles, glabrous, 1-veined, the lateral veins usually obscure. Flowers 4–6 mm in diameter, in many-flowered terminal and axillary cymes, forming a leafy panicle; pedicels 1–5 mm, glabrous; bracts 3–6 mm, ovate, cuspidate at apex. Calyx only an annular ridge. Corolla 3.0–3.5 mm, pale yellowish-green, deeply divided into 5 lobes; lobes 2–3 mm, ovate, cuspidate at apex. Stamens 5; filaments 0.4– 0.5 mm, whitish; anthers 0.2–0.3 mm, yellow, suborbicular or orbicular-ovate. Style 1, bifid; stigmas capitate. Mericarp 4–6 mm in diameter, black, subglobose. Flowers 6–8. 2n = 44. Native. Hedgerows, scrub, copses, rocky places and by walls, usually near the coast, more local inland. South and west Great Britain from Kent to the Isles of Scilly north through Gloucestershire and round the coast of Wales and in the Channel Islands, and in south, east and central Ireland. South and west Europe northwards to 53◦ 30 N in western

1. Sambucus Ireland and eastwards to Greece and Turkey; North Africa. A member of the Mediterranean-Atlantic element. 2. R. tinctorum L. Madder Deciduous perennial herb with tough, stringy, reddish rhizomes. Stems up to 1 m, sprawling or scrambling, sharply 4angled, pale green, with small, downwardly directed prickles along the angles. Leaves (2–)4–6 in a whorl; lamina 20–80(–100) × 5–30 mm, yellowish-green on upper surface, paler beneath, subcoriaceous, lanceolate or elliptical, acute or acuminate at apex, entire, cuneate at base, with backwardly directed prickles on the margin and underside of the midrib, distinctly net-veined beneath. Flowers 5–6 mm in diameter, in a much branched, lax terminal cyme; pedicels up to 5 mm, slender, glabrous; bracts 5–7 mm, lanceolate, oblanceolate or lanceolate-subulate, acute at apex. Calyx a minute annular ridge. Corolla up to 2 mm, pale greenishyellow, funnel-shaped, deeply divided; lobes 5, lanceolate or narrowly ovate, acuminate at apex. Stamens 5; filaments very short, pale; anthers 0.5–0.6 mm, linear-oblong. Style 1, bifid, pale; stigmas cream, capitate. Mericarp 5–6 mm, reddish-brown turning black, subglobose, fleshy. Flowers 6–7. 2n = 22, 44. Introduced. Formerly grown from early times for the production from its root of a range of dyes, notably Turkeyred, all of which are very stable. Later occurred as a casual and recently found established at Boothby Graffoe in Lincolnshire where it had been known for over 30 years. Probably native only of western and central Asia, but widely introduced in Europe.

Order. 9 . D I P S AC A L E S Lindl. Herbs, shrubs or small trees. Leaves opposite, simple, lobed or pinnate. Flowers actinomorphic or zygomorphic, unisexual or bisexual, epigynous. Calyx 3- to numerous-lobed. Corolla 4- to 5-lobed. Stamens 1–5. Ovary 1- to 5-celled. Fruit a berry, drupe, nut or achene; seeds with endosperm. Contains 4 families, 3 genera and about 1,200 species. 144. C A P R I F O L I AC E A E Juss. nom. conserv. Deciduous or evergreen, small trees or shrubs which are sometimes small and procumbent, or woody climbers, rarely perennial herbs. Leaves opposite, simple and sometimes lobed or pinnate, petiolate, stipulate or exstipulate. Inflorescence axillary or terminal, a corymbose or paniculate compound cyme, dense spike or head. Flowers zygomorphic or actinomorphic, bisexual or some sterile, epigynous. Calyx 5-lobed. Corolla 5-lobed, sometimes 2-lipped, with 4-lobed upper and 1-lobed lower lip, usually white to yellow, sometimes pink to reddish. Stamens 4–5, borne on the corolla tube. Styles 1, or absent; stigmas 1–5, more or less capitate. Ovary inferior, 1-to 5-celled, sometimes including 2 sterile cells, each cell with 1 apical ovule or many ovules on axile placenta. Fruit a berry, drupe or achene; seeds with a fleshy endosperm and usually a small, straight embryo.

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Consists of 13 genera and about 400 species, mostly of north temperate and boreal regions and of tropical mountains. 1. 1. 2. 2. 3. 3. 4. 4. 5. 5. 6.

6.

7. 7.

1. Sambucus Leaves pinnate 2. Leaves simple, sometimes deeply lobed 3. Flowers numerous, in corymbose compound cymes 4. Flowers 2–few, not corymbose 2. Viburnum Flowers 5-partite 8. Kolkwitzia Flowers 4-partite 4. Linnaea Main stems procumbent 5. Main stems more or less erect or climbing Bracts 12–50 mm, leaf-like, purple or green strongly 5. Leycesteria tinted purple 6. Bracts less than 15 mm, not purple Corolla actinomorphic; ovary 4-celled with 2 fertile and 2 sterile, the former each with 1 ovule; fruit a white to 3. Symphoricarpos pink berry Corolla zygomorphic to actinomorphic; ovary 2- to 3-celled, all cells fertile with one or more ovules; fruit a 7. red, blue or purple to black berry, or a capsule 6. Weigela Fruit a capsule 7. Lonicera Fruit a berry

1. Sambucus L. Deciduous shrubs, small trees or perennial herbs, with large pith. Leaves opposite, pinnate; stipulate or exstipulate. Flowers numerous, in corymbose or paniculate compound cymes, actinomorphic. Calyx 5-lobed. Corolla 5lobed, rotate. Stamens 5. Stigma sessile, 3- to 5-lobed. Ovary 3- to 5-celled. Fruit a drupe, with 3–5 seeds. About 40 species in temperate and subtropical regions, but not central and southern Africa. Grime, J. P. et al. (1988). Comparative plant ecology. London. [S. nigra.] Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 13: 134–135, 367–368. [S. racemosa.] Nelson, S. E. & E. C. (1996). Weeping Elder (Sambucus nigra L. f. pendula Dippel), wild in West Norfolk. B.S.B.I. News 71: 27. Rehder, A. (1956). Manual of cultivated trees and shrubs. Ed. 2. New York. 1. Rhizomatous perennial herb; stipules conspicuous, more 4. ebulus or less ovate; anthers reddish-purple 2. 1. Erect shrub; stipules absent or narrow; anthers cream 2. Inflorescence an ovoid or globose panicle; stipules a pair 3. of glandular appendages or linear 2. Inflorescence a flat-topped or slightly convex corymb; 6. stipules absent or subulate 4. 3. Leaves and inflorescence branches glabrous 3. Leaves and inflorescence branches hairy at least when 5. young 4. Inflorescence compact; fruits 5–6 mm in diameter 1(a). racemosa subsp. racemosa 4. Inflorescence rather lax; fruits 3–4 mm in diameter 1(b). racemosa subsp. sieboldiana

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1(c,i). racemosa var. pubens 5. Drupe bright red 1(c,ii). racemosa var. melanocarpa 5. Drupe purplish-black 6. Shrub rhizomatous with suckers; twigs with few 3. canadensis lenticels; leaflets (5–)7(–11) 6. Shrub not rhizomatous; twigs with numerous lenticels; 7. leaflets (3–)5(–7) 2(6). nigra forma pendula 7. Branches pendulous 8. 7. Branches erect or ascending 9. 8. Leaves yellow or variegated throughout life 10. 8. Leaves green until autumn 2(1). nigra forma aurea 9. Leaves golden yellow 9. Leaves variegated yellow 2(3). nigra forma luteovariegata 2(4). nigra forma laciniata 10. Leaflets divided pinnately 11. 10. Leaflets serrate only 2(1). nigra forma nigra 11. Drupe black 2(5). nigra forma viridis 11. Drupe greenish or whitish

Section 1. Sambucus Deciduous shrubs or small trees. 1. S. racemosa L. Red-berried Elder Deciduous shrub up to 4(–8) m, with a broad, rounded crown, spreading by suckers. Stems several, much branched but with straight, vigorous, erect shoots. Bark pale greyishbrown, rough and fissured; wood with thick, whitishbrown pith. Branches arching and much divided; twigs soft, pithy, pale brown; young shoots pale brown or greenish to somewhat glaucous, glabrous or with short simple eglandular hairs. Buds about 5 mm, more or less ovoid; scales dark purplish, ovate, acute at apex. Leaves foetid when bruised, yellowish-green on upper surface, paler beneath, turning yellow in autumn, sometimes variegated, imparipinnate, broadly obovate in outline; leaflets 5–9, the lamina 4–17 × 1.5–6.0 cm, narrowly elliptical, ovate, ovatelanceolate or lanceolate, the terminal sometimes oblanceolate to obovate, acuminate to caudate at apex, sharply serrate except sometimes near the apex, sometimes the lower divided again, sometimes all dissected, rounded or cuneate at the often asymmetrical base, glabrous or with very short simple eglandular hairs on the upper surface and glabrous or with short simple eglandular hairs beneath, the petiolules up to 25 mm, pale green and glabrous, the veins 4–8 pairs, not prominent; petiole 15–60 mm, pale yellowishgreen, grooved on the upper side, glabrous or with short simple eglandular hairs; stipules a pair of glandular appendages or more developed and linear. Flowers 3–6 mm in diameter, in a dense, ovoid or globose panicle up to 6 cm wide, with a definite central axis projecting well beyond the lowermost pair of branches, sickly scented. Calyx minute or absent. Corolla 3–4 mm, creamy-white, divided half of the way to the base; lobes 5, broadly elliptical, obtuse at apex. Stamens 5; filaments white; anthers cream. Stigma cream. Drupe 3–6 mm in diameter, bright red, orange or purplish-black, rarely greenish-yellow or brown, globose, juicy; nutlets more or less rugose. Flowers 5–7. Intermediate plants may occur between the following infraspecific taxa. Plants with divided or variegated leaflets may occur in any of these taxa.

(a) Subsp. racemosa Leaves and inflorescence branches glabrous. Inflorescence compact. Drupe 5–6 mm in diameter, bright red or rarely greenish-yellow. 2n = 36, 38. (b) Subsp. sieboldiana (Miq.) Hara S. racemosa var. sieboldiana Miq.; S. sieboldiana (Miq.) Schwer. Leaves and inflorescence branches glabrous. Inflorescence rather lax. Drupe 3–4 mm, bright red. 2n = 36. Named after Philipp Franz von Siebold (1796–1866). (c) Subsp. pubens (Michx) House S. pubens Michx; S. racemosa var. pubens (Michx) Koehne Leaves and inflorescence branches hairy at least when young. Inflorescence rather lax. Drupe 4–6 mm, red or purplish-black. 2n = 36. (i) Var. pubens (Michx) Koehne Drupe bright red. (ii) Var. melanocarpa (A. Gray) McMinn S. melanocarpa A. Gray Drupe purplish-black. Introduced. Well naturalised in hedges, woods and shrubberies. Frequent in Great Britain north of Cheshire and Derbyshire with scattered records further south. All the above taxa are recorded especially in Scotland. Subsp. racemosa is native of southern Europe and western Asia, subsp. sieboldiana is native of eastern Asia and subsp. pubens native of North America. Var. pubens is widespread in the north-eastern United States and adjacent Canada westwards to British Columbia. Var. melanocarpa is chiefly a plant of the Rocky Mountains. 2. S. nigra L. Elder Deciduous shrub or small tree up to 12 m, with a broad, rounded crown, often trimmed or coppiced and with several stems. Stems when mature wavy and much branched, but often with straight, vigorous, erect shoots from the base. Bark pale greyish-brown, rough, corky and fissured; wood with thick, white pith. Branches arching and much divided; twigs greyish-brown, thick, rigid and easily breaking, with numerous lenticels, rough; young shoots pale yellowishgreen, glabrous, with numerous lenticels. Buds 5–8 × 3–4 mm, more or less ovoid; scales dark purplish, ovate, acute at apex. Leaves foetid when bruised, yellowish-green on upper surface, rarely yellow or variegated, paler beneath, turning yellow and often flushed purplish in autumn, rarely bronze, imparipinnate, broadly obovate in outline; leaflets (3–) 5(–7), the lamina 3–12(–18) × 2–6(–10) cm, gradually decreasing in size downwards, lanceolate, ovate, ovateelliptical or elliptical, rarely subrotund, acuminate at apex, crenate or serrate with rather blunt, hooked teeth, rarely laciniate, cuneate, rounded or truncate at base, with short, pale simple eglandular hairs on both surfaces and the margins when young, and becoming glabrous, the petiolules up to 40 mm, pale green and shortly hairy, the veins 4–8 pairs and prominent beneath; petiole up to 80 mm, pale green, channelled above, rounded beneath, more or less shortly hairy; stipules absent or small and subulate.

1. Sambucus Flowers 5.5–7.5 mm in diameter, in a cymose, flat-topped corymb 8–24 cm in diameter, with 5 primary rays, sickly scented. Calyx about 2 mm, pale green, divided for about one-third of the way to the base; lobes 5, triangular, obtuse at apex. Corolla 2.5–3.5 mm, creamy-white, with a short tube and 5 subrotund lobes. Stamens 5; filaments 2.0–2.5 mm, white; anthers cream. Stigma cream. Drupe (4.5–)6–8 mm in diameter, globose, usually turning wine-red then black, rarely greenish, juicy. Flowers 6–7. Pollinated by small flies etc. Fruits 8–9. 2n = 36. (1) Forma nigra S. nigra var. rotundifolia DC. Branches erect or ascending. Leaves green, turning yellow and sometimes flushed purplish in autumn; leaflets lanceolate to ovate, rarely subrotund. Drupe black. (2) Forma aurea (Sweet) Schwer. Golden Elder S. nigra var. aurea Sweet; S. aurea (Sweet) Carri`ere Branches erect or ascending. Leaves golden yellow throughout the year; leaflets lanceolate to ovate. Drupe black. (3) Forma luteovariegata (Weston) Schwer. S. nigra var. luteovariegata Weston Branches erect or ascending. Leaves variegated yellow; leaflets lanceolate to ovate. Drupe black. 2n = 36. (4) Forma laciniata (L.) Zabel Parsley-leaved Elder S. nigra var. laciniata L.; S. laciniata (L.) Mill.; S. nigra var. heterophylla Endl. Branches erect or ascending. Leaves green; leaflets divided pinnately into linear, pointed lobes. Drupe black. 2n = 36. (5) Forma viridis (Weston) Schwer. S. nigra var. viridis Weston; S. nigra var. chlorocarpa Hayne; S. nigra var. virescens Sweet; S. nigra var. leucocarpa Sm. Branches erect or ascending. Leaves green; leaflets lanceolate to ovate. Drupe greenish or whitish. (6) Forma pendula Dippel Branches stiff and pendulous, hanging almost vertically downwards. Leaves green; leaflets lanceolate. Drupe black. Native. Hedgerows, woods, copses and waste and rough ground, especially on manured soils. Throughout Great Britain and Ireland but only introduced in the northern islands. Most of Europe except the extreme north; western Asia; North Africa; Azores. A member of the European Temperate element. All the forms can be seen here and there in the countryside. The berries are much favoured by birds, and are sometimes dropped by them, so that fresh bushes spring up in many new places, including cracks in drainpipes and brickwork. Both the flowers and fruits of Elder are used for making wine and cordials. It was formerly much used for making eye and skin lotions. When horses were used for agriculture, twigs were fixed in the harness to keep flies away and drovers used it in the same way for cattle. It has thousands of years of magical belief and peasant ingenuity behind it as it grows prolifically, is light in construction and if damaged can be scrapped and quickly replaced.

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3. S. canadensis L. American Elder Deciduous shrub up to 3 m, with a rounded crown, spreading by rhizomes and suckers. Stems much branched, but with straight, vigorous, erect shoots. Bark pale greyish-brown, corky and fissured; wood with thick, white pith. Branches arching and much divided; twigs greyish-brown, thick, rigid and easily breaking, with few lenticels, round; young shoots pale yellowish-green, glabrous, with numerous lenticels. Buds more or less ovoid; scales dark purplish, ovate, acute at apex. Leaves foetid when bruised, yellowish-green on upper surface, paler beneath, turning yellow, sometimes flushed purplish in autumn, sometimes variegated, imparipinnate, broadly obovate in outline; leaflets (5–)7(–11) the lowermost pair sometimes divided again, the lamina 4–15 × 2.0– 5.5(–7.0) cm, lanceolate to ovate or elliptical, acuminate to caudate at apex, serrate except at the apex with very sharp teeth, sometimes dissected, rounded and oblique to cuneate at base, glabrous on the upper surface or with very short, pale simple eglandular hairs on the midrib and with very short, pale simple eglandular hairs below especially on the veins but sometimes becoming glabrous, the petiolules up to 20 mm, pale green and glabrous or with very short simple eglandular hairs, the veins 4–8 pairs, not prominent; petiole 30–90 mm, pale yellowish-green, grooved on the upper side, glabrous; stipules absent or small and subulate, early deciduous. Flowers 4–5 mm in diameter, in a cymose, flat-topped or slightly convex corymb up to 30 cm in diameter, with 5 primary rays, sickly scented. Calyx 1.8– 2.0 mm, pale green, divided about one-third of way to base; lobes 5, triangular-ovate, obtuse at apex. Corolla 3–4 mm, creamy-white, divided half of the way to the base; lobes 5, broadly ovate, obtuse at apex. Stamens 5; filaments white; anthers cream. Stigma cream. Drupe 4–6 mm in diameter, globose, usually dark purple, rarely greenish-yellow or reddish. Flowers 7–9. Pollinated by small flies. 2n = 36. Introduced. Naturalised in scrub, rough ground and on railway banks where it has been widely planted. Scattered localities in Scotland, northern England and Surrey. Native of eastern North America. Section 2. Ebulus Spach Ebulus Garcke Perennial herbs with creeping rhizomes. 4. S. ebulus L. Dwarf Elder S. humilis Mill.; S. herbacea Stokes nom. illegit.; Ebulus humilis Garcke Foetid perennial herb with a creeping rhizome forming clumps. Stems several to numerous, 60–200 cm, medium green with paler ridges and often faintly speckled, glabrous, pith wide and white, simple or little branched. Leaves opposite, broadly ovate in outline, pinnate; leaflets 5–13, the lamina 5–16 × 1.0–4.5 cm, medium green with paler midrib on upper surface, only slightly paler beneath, oblong to oblong-lanceolate, long-acute or acuminate, serrate with curved, ascending, mucronate teeth, rounded or cuneate and sometimes with leaf-like lobes or segments at the base, sessile or petiolulate, with some very short simple eglandular hairs above, more numerous beneath, with 12–16 pairs of

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prominent lateral veins; rhachis channelled above, ridged below, with few hairs; petiole up to 80 mm, like rhachis; stipules 5–30 × 3–15 mm, ovate, acute at apex, toothed like the leaves. Flowers 9–11 mm in diameter, in corymbs, with 3(–4) primary rays; pedicels and branches glabrous. Calyx small, pale green, with 5 triangular lobes. Corolla 3–4 mm, cream, sometimes pink-tinged; lobes 5, elliptical or obovate, pointed at apex. Stamens 5; filaments white; anthers reddish-purple. Stigmas reddish. Drupe 4–5 mm in diameter, black, globose. Flowers 7–8. Pollinated by insects. 2n = 36. Probably introduced, but possibly native. In clumps by waysides, rough and waste ground, old banks and hedges and churchyards, often upon a mediaeval site. By English legend it grew spontaneously out of the blood of the slaughtered Danes. It was much used as a medicinal herb for purging, dropsy, gout and piles. It gives off a blue dye for staining cloth. It was known to the Romans, Gauls and Anglo-Saxons and there is every possibility it was introduced. It is scattered over much of Great Britain and Ireland. Most of Europe from the Netherlands and northern Ukraine southwards; western Asia to the Himalayas; North Africa; Madeira. A member of the European Southern-temperate element. 2. Viburnum L. Shrubs or small trees. Leaves deciduous or evergreen, simple, sometimes lobed; stipulate or exstipulate. Flowers numerous, in corymbose, compound cymes, actinomorphic, sometimes some sterile. Calyx 5-lobed. Corolla 5-lobed, rotate to campanulate. Stamens 5. Style 1, short; stigma 3lobed. Ovary 3-celled, with 1 fertile with a single ovule and 2 sterile. Fruit a drupe, with 1 seed. About 200 species in the north temperate zone, extending south to Central America and Java. Bean, W. J. (1980). Trees and shrubs hardy in the British Isles. Vol. 4. Ed. 8. London. Hult´en, E. (1958). The amphi-atlantic plants and their phytogeographical connections. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 7: 74–75. [V. opulus.] Kollmann, J. & Grubb, P. J. (2002). Viburnum lantana L. and Viburnum opulus L. (V. lobatum Lam., Opulus vulgaris Borkh.) in Biological flora of the British Isles, Jour. Ecol. 90: 1044–1070. Rehder, A. (1956). Manual of cultivated trees and shrubs. Ed. 2. New York. Stearn, W. T. (1950). × Viburnum bodnantense. Bot. Mag. 167: t. 113.

3.

4. 4. 5. 5. 6. 6. 7. 7. 8. 8. 9. 9. 10. 10. 11. 11. 12. 12. 13.

13.

14. 14. 15.

The segregates of both V. lantana and V. opulus are very difficult to key out but the taxa are much easier to recognise in the field, where you can see the whole range of leaves on the bush.

15.

1. Leaves palmately lobed; outer flowers of cymes larger 2. than inner and sterile or all large and sterile 7. 1. Leaves not lobed; all flowers similar in size and small 2. All flowers of cymes large and sterile 9(2). opulus forma roseum 2. Outer flowers of cymes sterile and larger than the fertile 3. inner 3. Upper leaves of branches with central lobe often longer than broad, the teeth of the lobes sharp and up to 14 per

16.

16.

lobe, the apical tooth of each lobe often very long, the 11. trilobum glands of the petiole shortly stalked Upper leaves of the branches with central lobe usually wider than long, the teeth of the lobes less sharp and up to 11, the apical teeth usually longer than the rest but not 4. very long, the glands of the petiole sessile Leaves pale to medium yellowish-green; anthers yellow; 5. drupe 8–11 × 8–11 mm Leaves dull medium to rather dark yellowish-green; 6. anthers yellow to purple; drupe 9–12 × 7–11 mm 9(1). opulus forma opulus Drupe red 9(3). opulus forma flavum Drupe yellow Drupe yellow, turning red 10(1). sargentii forma sargentii 10(2). sargentii forma flavum Drupe yellow Flowering in winter; without leaves, with last year’s 8. leaves hanging on, or evergreen 11. Flowering in spring, with young leaves or evergreen 9. Evergreen Usually without leaves when flowering or last years 10. leaves hanging on Leaves medium yellowish-green and wrinkled 5. × rhytidiophylloides 7. tinus Leaves dark green and smooth 1. farreri Corolla tube 6.5–9.0 mm 2. × bodnantense Corolla tube 8–11 mm 8. davidii Leaves with 3 main longitudinal veins 12. Leaves with 1 main longitudinal vein 13. Leaves mostly broadly elliptical, deciduous Leaves mostly oblong or elliptical-oblong, evergreen, or 16. partly evergreen and partly deciduous Leaves with minute stellate hairs on the veins of the lower surface at anthesis, and sometimes a few on the surface, becoming glabrous or nearly so later in the year at least on the under surface making it look green 3(iii). lantana var. glabratum Leaves with minute stellate hairs on the veins of the lower surface and numerous to dense more open and longer stellate hairs all over the lower surface which remain until the shrub is in fruit, making the under 14. surface look grey Leaves with stellate hairs on under surface stalked; 4. veitchii calyx with dense stellate hairs Leaves with stellate hairs on under surface sessile or shortly stalked; calyx glabrous or with few stellate hairs 15. Leaves scarcely rugose on upper surface 3(i). lantana var. lantana Leaves strongly rugose on upper surface 3(ii). lantana var. rugosum Leaves mostly elliptic-oblong with a few oblong and a few elliptical, partly evergreen and partly deciduous, 5. × rhytidiophylloides glabrous on the upper surface Leaves all oblong or narrowly elliptical-oblong, evergreen, with few to numerous stellate hairs on the 6. rhytidiophyllum upper surface

Section 1. Thyrsosma (Raf.) Rchb. Thyrsosma Raf. Deciduous shrubs or small trees. Leaves unlobed, glabrous or with simple or forked hairs. Drupes oblong, bright red.

2. Viburnum 1. V. farreri Stearn Farrer’s Viburnum V. fragrans Bunge, non Loisel. Deciduous shrub or small tree. Stems numerous, up to 3.5 m, erect then arching. Bark pale greyish-brown to reddishbrown, shredding in papery flakes. Branches numerous, arching; twigs pale brown, longitudinally ridged; young shoots reddish-brown, glabrous. Buds 5–7 × 2–3 mm, narrowly ovoid; scales reddish-brown, ovate, acute at apex, with very short hairs. Leaves opposite; lamina 5–9 × 2.0– 5.5 cm, dull medium to dark green on upper surface, paler and sometimes touched purplish beneath, oblong-elliptical, elliptical, obovate or oblanceolate, acute to acuminate or cuspidate at apex, serrate-dentate with small mammiform teeth, cuneate or rounded at base, with scattered hairs on both surfaces, becoming glabrous except for small tufts in the vein axils; veins 5–7 pairs, the midrib sometimes reddish-purple; petiole 10–35 mm, usually reddishpurple, channelled above, rounded beneath, glabrous; stipules absent. Inflorescence terminal or axillary, in dense corymbs; pedicels pale yellowish-green, with short, white simple eglandular and very short glandular hairs. Flowers 7.5–10.0 mm in diameter, rotate, scented. Calyx 2.5–3.0 mm, yellowish-green, glabrous; tube about 2 mm; lobes 5, about 1.0 mm, broadly ovate, rounded at apex. Corolla 9–10 mm, white tinted pale pink, becoming white, rose in bud; tube 6.5–9.0 mm; lobes 5, 1–3 mm, ovate, obtuse at apex. Stamens 5, 2 fertile and 3 sterile; filaments 5–7 mm, white; anthers purplish-red. Style 1, 3–4 mm, pale green. Drupe 10–13 mm, oblong, bright red. Flowers 11–4. 2n = 16, 32. Introduced. This hardy shrub, whose flowers can stand up to 12◦ of frost, is grown in gardens and sometimes by roads, in parks and in landscaped areas. It is established in scrub near Parkgate House on Ham Common, Surrey. Native of China. Named after Reginald John Farrer (1880–1920). 2. V. × bodnantense Aberc. ex Stearn Bodnant Viburnum V. farreri × grandiflorum Wall. Deciduous shrub. Stems numerous, up to 3 m, erect, thin, arching. Bark pale dull brown or orange-brown, shallowly and irregularly longitudinally ridged. Twigs pale brown, irregularly longitudinally marked; young shoots rather shiny reddish-brown, glabrous. Buds 2–5 × 2–3 mm, narrowly ovoid; scales reddish-brown, ovate, subobtuse to acute at apex, with very short hairs, becoming glabrous. Leaves opposite; lamina 6–10 × 4–6 cm, dull medium green on upper surface, paler beneath, ovate, obovate or lanceolate, more or less acute at apex, serrate-dentate, the teeth mammiform, cuneate or rounded at the base, with few to numerous simple or forked eglandular hairs on both surfaces; veins 6–9 pairs, impressed above, prominent beneath, sometimes purplish-red; petioles 15–25 mm, usually purplish-red, channelled above, rounded beneath, glabrous. Inflorescence terminal or axillary, 8- to 35flowered, in cymose corymbs; pedicels pale yellowishgreen, with numerous, short, white simple eglandular hairs. Flowers 8–10 mm in diameter, rotate, strongly scented. Calyx 2.5–3.0 mm, yellowish-green, glabrous; tube about

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2 mm; lobes 5, 0.5–1.0 mm, broadly ovate, rounded at apex. Corolla 9–13 mm, white tinted pale pink, denser rose in bud; tube 8–11 mm; lobes 5, 1–3 mm, subrotund. Stamens 5, 2 fertile and 3 sterile; filaments 6–10 mm, white; anthers chestnut. Style 1, 3–4 mm, pale green; stigmas yellowishgreen, 3-lobed. Drupe 11–13 × 7–8 mm, bright red, oblong. Flowers 11–4. 2n = 16. The most common cultivated variety is cv. Dawn, which has the flowers rich rose-red in bud. Although the characters which distinguish it from V. farreri are small, it can usually be recognised at a glance by its more handsome, more deeply coloured flowers and broader outline. Introduced. A hybrid artificially produced in Bodnant Gardens, Denbighshire, now widely grown in gardens, and sometimes in parks, landscaped areas and by roads and streets. Section 2. Viburnum Section Lantana Spach Deciduous or evergreen shrubs. Leaves unlobed, with stellate hairs at least beneath. Drupe subglobose, oblong or elliptical, turning red, then black. 3. V. lantana L. Wayfaring Tree V. farinosum Stokes nom. illegit. Deciduous shrub or small tree up to 5 m, with a single trunk or several slender stems, regular and compact or somewhat straggling. Bark reddish or greyish, rough. Branches irregular, erect or sometimes spreading. Twigs pale, dull brown, with numerous stellate hairs; young shoots reddishbrown, with dense, greyish stellate hairs. Buds 8–10 × 6–7 mm, ovoid, pointed at apex, stellate hairy. Leaves opposite, simple; lamina 5–11 × 3–7 cm, deep yellowish-green on upper surface, paler beneath with paler veins, slowly suffused dull brownish-red in autumn, lanceolate, ovate, obovate, elliptical or oblong, subacute to rounded at apex, undulate-spinulose-denticulate, rounded to cordate at the sometimes slightly unequal base, firm and thick, glabrous or with numerous stellate hairs on the upper surface, with small stellate hairs on the veins beneath, sometimes also larger stellate hairs on the surface; veins 4–7 pairs, curved upwards; petioles 3–30 mm, pale green, more or less stellate-hairy. Inflorescence a dense umbel-like cyme 60–100 mm in diameter. Flowers 6–7 mm in diameter, all alike and all fertile, with a rather unpleasant smell. Calyx 0.8–1.0 mm, deeply divided; lobes 5, triangularovate, obtuse at apex. Corolla about 6 mm, cream, deeply divided; lobes 5, oblong or obovate, rounded at apex. Stamens 5; filaments 3–4 mm, white; anthers pale yellow. Styles 1, short and thick; stigma capitate. Drupe 8.5–10.0 × 8–9 mm, turning deep shining red, then black, ellipsoidal. Flowers 5–6. Pollinated by insects or self-pollinated. Fruits 7–9. 2n = 18. (i) Var. lantana Leaves hardly rugose on upper surface, with minute stellate hairs on the veins of the lower surface and numerous to dense more open and larger stellate hairs all over the lower surface which remain until the plant is in fruit, making the under surface look grey.

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V. × rhytidiophylloides J. V. Suringar (faintly rugose above)

V. rhytidiophyllum Hemsl. ex Forbes & Hemsl. (rugose above)

V. lantana L. var. lantana (faintly rugose above) Leaves of Viburnum

2. Viburnum

V. lantana var. glabratum Chab. (faintly rugose above)

V. lantana var. rugosum Lange (very rugose above)

V. veitchii C.H. Wright (rugose above) Leaves of Viburnum

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(ii) Var. rugosum Lange Leaves strongly rugose on upper surface, with minute stellate hairs on the veins of the lower surface and numerous to dense, more open and larger stellate hairs all over the lower surface, making the undersurface look grey. (iii) Var. glabratum Chab. V. lantana var. viride A. Kern. Leaves not rugose on upper surface, with minute stellate hairs on the veins of the lower surface and some on surface at anthesis, becoming glabrous or nearly so at least on the lower surface later in the year making it look green. Native. Scrub, woods and hedges especially on base-rich soils, and planted by roadsides and new woods. Common in southern England north to Nottinghamshire and Derbyshire, rare in Wales; occasionally introduced elsewhere in Great Britain and Ireland. Central and south Europe extending from Great Britain and northern Spain eastwards to the central Ukraine; Caucasus; northern Turkey; Iran; Morocco and Algeria. A member of the European Temperate element. The complete distribution of the three varieties is not known, but in Great Britian var. glabratum seems to be confined to south-west England and Wales east to the Isle of Wight and it may be the only variety in the far west of its range. It is also widely planted in Cambridgeshire and extends into France. Var. lantana probably applies to the rest of the native plants. Var. rugosa is much planted in Cambridgeshire and may be widespread. It could be part of the V. × rhytidiophylloides complex. 4. V. veitchii C. H. Wright Asian Wayfaring Tree Deciduous, fairly open shrub. Stems up to 4 m, numerous, erect, much branched. Bark pale greyish-brown, rough. Branches rather long, ascending, glabrous. Twigs pale greyish-brown, with some stellate hairs; young shoots pale green, densely clothed with small, yellowish stellate hairs. Buds 8–9 × 3.0–3.5 mm, ovate, pointed at apex. Leaves opposite, simple; lamina 7–15 × 5–9 cm, rather dark green on upper surface, paler yellowish-green beneath, with even paler veins, slowly suffused brownish-red in autumn, broadly ovate, broadly elliptical or nearly subrotund, rounded or shortly acute at apex, more widely dentate with blunter teeth than in V. lantana, rounded to subcordate at base, thick, with scattered stellate hairs on upper surface and rugose, with dense long-stalked stellate hairs on lower surface and smaller stellate hairs on the veins; veins 7–9 pairs, curved upwards; petioles up to 20 mm, densely stellate-hairy. Inflorescence a dense umbel-like cyme up to 120 mm in diameter. Flowers 6–7 mm, all alike and all fertile, with a rather unpleasant smell. Calyx 0.8–1.0 mm, deeply divided, more or less densely stellately hairy; lobes 5, triangular-ovate, obtuse at apex. Corolla about 6 mm, cream, deeply divided; lobes 5, oblong or obovate, rounded at apex. Stamens 5; filaments 3–4 mm, white; anthers yellow. Style 1; stigma capitate. Drupe 8.5–10.0 × 6–7 mm, turning deep shining red, then black, narrowly ellipsoidal. Flowers 5–6. Introduced in 1901. Planted in new woodlands in Cambridgeshire and probably elsewhere. Native of central China.

5. V. × rhytidiophylloides J. V. Suringar Hybrid Viburnum V. lantana × rhytidiophyllum Dense, much branched, partly evergreen and partly deciduous shrub up to 6 m. Stems numerous, up to 6 m, erect, then arching. Bark pale brown, rather smooth. Branches arching and much divided. Twigs pale reddish-brown, glabrous, with scattered lenticels; young shoots with dense stellate hairs. Buds naked or with a few scales. Leaves opposite; lamina 6–12 × 3–6 cm, shining yellowish-green to medium green and much rugose on upper surface, greyish-green to yellowish-green beneath, sometimes turning reddish in autumn, oblong to elliptical or ovate-elliptical, obtuse or subacute at apex, sinuate-denticulate or subentire, rounded to subcordate at base, with few to numerous stellate hairs on the upper surface, with dense stellate hairs on the veins and less dense ones on the surface beneath; veins 6–8 pairs, impressed above, prominent beneath; petioles 10–40 mm, pale yellowish-green, densely stellate-hairy. Inflorescence a dense, umbel-like cyme 30–60 mm in diameter. Flowers 6–9 mm in diameter, all alike, with a rather unpleasant smell. Calyx 3.0–4.5 mm, pale yellowish-green, divided about one-sixth of the way to the base, with stellate hairs; lobes 5, ovate, more or less obtuse at apex. Corolla 4–7 mm, white, divided one-third of the way to the base; lobes 5, obovate, broadly rounded at apex. Stamens 5; filaments 2.5– 3.5 mm, white; anthers pale yellow. Style 1, green; stigma green, much shorter than anthers. Drupe 8–10 mm in diameter, subglobose, turning red, then black. Flowers 11–6. 2n = 18. The normal plant looks much more like the V. rhytidophyllum parent, but there are planted trees of V. lantana on roadsides that have a much more wrinkled upper surface to the leaves and that are much less rounded than in that species, and they may belong to this hybrid as may V. lantana var. rugosa. Introduced. Grown in gardens and sometimes planted on roadsides in parks and landscaped areas. Two trees have been found in the wild in Kent, in Darenth Wood and on a railway bank at Eynsford. The original garden clone was raised in 1925. 6. V. rhytidiophyllum Hemsl. ex Forbes & Hemsl. Wrinkled Viburnum Dense, much branched, evergreen shrub. Stems usually numerous, up to 6 m, erect. Bark pale dull brown, rough, scaly. Branches spreading and much divided. Twigs pale reddish-brown, very rough, glabrous or with the remnants of stellate hairs; young shoots pale green, with dense, manyarmed stellate hairs. Buds naked or with a few scales. Leaves opposite; lamina 5–20 × 2–5 cm, dark shiny green and much wrinkled on upper surface, pale greenish-white beneath, oblong or narrowly elliptical-oblong, obtuse or subacute at apex, entire, narrowed to a usually asymmetrically rounded base, glabrous above, with dense stellate hairs beneath; veins 7–9 pairs, impressed above, prominent and slightly reddish-brown beneath; petioles 10–35 mm, with dense, sometimes reddish-brown stellate hairs. Inflorescence 10–16 cm in diameter, a terminal, domed,

2. Viburnum compound-corymbose cyme; peduncles and pedicels with dense, reddish-brown stellate hairs. Flowers 4–7 mm in diameter, all fertile, rather nasty smelling. Calyx 3.0–3.5 mm, greenish, divided about one-third of the way to the base; lobes 5, ovate, obtuse at apex. Corolla 3.0–3.5 mm, divided two-thirds of the way to the base, yellowish-white; lobes 5, oblong-ovate, obtuse at apex. Stamens 5; filaments 2.0–2.5 mm, white; anthers yellow. Style 1, white; stigma whitish, much shorter than anthers. Drupe 8–10 × 5–7 mm, oblong, compressed, turning shiny red, then black. Flowers 5–6. Fruits 9–10. 2n = 18. Introduced. Much grown in shrubberies and naturalised or a relic in old woodland and parkland. Also planted by roadsides and in landscaped areas. First introduced in 1900. Native of central and western China. Section 3. Tinus Maxim. Evergreen shrubs. Leaves unlobed, glabrous except for axillary tufts in the axils of veins beneath. Drupe subglobose, ellipsoid or ovoid, deep blue. 7. V. tinus L. Laurustinus Dense, much branched, evergreen shrub up to 6 m, often as wide or wider than high. Bark dull brown, rough. Branches ascending, sometimes arching. Twigs dull, rather pale brown, with numerous lenticels, glabrous; young shoots yellowish-green or brown on upper side, with numerous stiff hairs. Buds narrowly ovoid, obtuse at apex, covered with stellate hairs. Leaves opposite; lamina 3–12 × 1–7 cm, dark green and smooth on upper surface, much paler beneath, lanceolate, ovate or subrotund-ovate, obtuse to subacute at apex, entire, rounded or cuneate at base, glabrous or with scattered simple or stellate hairs particularly in the vein axils; veins 4–7 pairs, curving upwards; petioles 5–15(–25) mm, pale green often tinted purplish-brown, often with long simple hairs. Inflorescence 4–9 cm in diameter, a terminal cyme; peduncle and pedicels pale green, angled, glabrous. Flowers 5–9 mm in diameter, uniformly fertile, sweet-smelling. Calyx about 1 mm, purplish, divided about two-thirds of the way to the base; lobes 5, triangular. Corolla 3–4 mm, pinkish outside, white within, divided to halfway to the base; lobes 5, broadly rounded-ovate, spreading. Stamens 5; filaments about 2.5 mm, white; anthers cream. Style 1, whitish; stigma whitish, much shorter than anthers. Drupe 5–6 × c. 5 mm, deep blue, finally bluishblack, ovoid. Flowers 11–4. 2n = 36. Introduced. Cultivated since at least the sixteenth century. Naturalised on cliffs, banks and in rough ground and planted in parks and cemeteries and along roads. Widespread in southern England and south Wales. Native of southern Europe, south-west Asia and North Africa. Our plant is subsp. tinus. 8. V. davidii Franch. David’s Viburnum Dense evergreen shrub often wider than high. Stems up to 2 m, numerous, erect or ascending, but very twisted and flexuous. Bark dull, dark greyish-brown, rough. Branches numerous, ascending, thick; twigs dark reddish-brown, with numerous lenticels; young shoots shining yellowishgreen with prominent brown lenticels, glabrous. Buds up

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to 7 mm, lanceolate or oblong-lanceolate, acute at apex. Leaves opposite; lamina 6–13 × 4–8 cm, dull dark green on upper surface with paler veins, pale yellowish-green beneath with even paler veins, broadly or narrowly elliptical or ovate, rounded to an acute or shortly acuminate apex, subentire to very shallowly undulate-denticulate, rounded at base, glabrous; with 3 main longitudinal veins, impressed above, prominent beneath, lateral veins numerous; petioles 5–30 mm, reddish-purple and channelled above, green and rounded beneath, glabrous. Inflorescence a terminal corymbose cyme; peduncles compact, 10–35 mm, yellowish-green, glabrous; pedicels 2–4 mm, yellowishgreen, glabrous. Flowers 4.5–5.5 mm in diameter, uniformly fertile. Calyx 1.5–2.5 mm, pale green, divided nearly to the base; lobes 5, triangular-ovate, obtuse to acute at apex. Corolla 2.8–3.2 mm, white, divided for two-thirds of the way to the base; lobes 5, rounded-ovate. Stamens 5; filaments about 2 mm, white; anthers blackish-red. Style 1, whitish; stigma whitish. Drupe 5–6 × about 4 mm, blue, ellipsoid or ovoid, slightly compressed. Flowers 6. 2n = 18. Although the shrubs appear to have bisexual flowers, the plants appear to be functionally dioecious. Introduced. Planted in parks, landscaped areas and shrubberies. Native of western China. Named after Abb´e Armand David (1826–1900). Section 4. Opulus (Mill.) DC. Opulus Mill. Deciduous shrubs. Leaves palmately lobed, with simple eglandular hairs beneath. Drupes subglobose, bright red, soft. 9. V. opulus L. Guelder Rose Opulus palustris Gray nom. illegit. Deciduous, fairly dense, broad shrub. Stems up to 4 m, numerous, erect, much branched. Bark pale greyish-brown, smooth but faintly wrinkled, slightly longitudinally ridged. Branches rather short; twigs pale greyish-brown, faintly wrinkled, slightly angled, glabrous; young shoots pale green, often tinted brownish-purple. Buds 5–7 × 3.0–4.5 mm, ovoid, acute at apex; scales reddish-brown. Leaves opposite, simple; lamina 3–8(–12.5) × 4.5–7.0(–9) cm, dull medium yellowish-green on upper surface, paler beneath, with even paler veins, sometimes touched purplish, turning deep reddish-purple in autumn, obovate in outline, palmately lobed, the lobes 3(–5), the middle lobe usually wider than long, ovate, acute or acuminate at apex, irregularly dentate, the teeth more or less mammiform, rounded or subcordate at base, glabrous on the upper surface, with numerous, short, pale simple eglandular hairs beneath, especially on the veins; veins impressed above, prominent beneath, with a main vein to each lobe and up to 7 pairs of side veins; petioles 10–25 mm, pale green, with a few discoid glands; stipules subulate or filiform, sometimes laciniate; leaf-scar V-shaped with 3 bundle traces. Flowers numerous, with a sickly scent, in compound, flat-topped cymes 50–120 mm in diameter, the outer large and sterile, the inner smaller and fertile, sometimes all large and sterile; peduncles and pedicels pale yellowish-green, sometimes tinted red, glabrous or with minute glandular hairs. Outer flowers

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1 4 4 . C A P R I F O L I AC E A E

V. sargentii Koehne V. opulus L. Leaves of Viburnum (Taken from non-flowering shoots below to flowering shoots above)

2. Viburnum

V. trilobum Marsh. Leaves of Viburnum (Taken from non-flowering shoots below to flowering shoots above)

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1 4 4 . C A P R I F O L I AC E A E

(9–)15–25 cm in diameter; calyx absent or of minute lobes; corolla white, divided up to two-thirds of the way to the base, the 5 lobes equal in size, more or less obovate with a rounded apex; sexual organs absent. Inner flowers 4–7 mm in diameter; calyx small, very pale green the 5 lobes triangular-ovate, acute and sometimes dark at tips; corolla white, divided about halfway to the base, the 5 lobes ovate, rounded at apex and curved outwards; stamens 5, the filaments 3–4 mm, white, the anthers yellow; stigmas cream with a red spot at apex. Drupe 8–11 mm in diameter, bright red, rarely yellow, subglobose, soft. Flowers 6–7. Pollinated by insects or selfed. 2n = 18.

to each lobe and up to 7 pairs of side veins; petiole 15– 35 mm, pale green, with discoid glands; stipules subulate. Flowers numerous, with a sickly scent, in compound, flattopped cymes, the outer large and sterile, the inner smaller and fertile; peduncles and pedicels pale yellowish-green, sometimes tinted, glabrous. Stamens 5; filaments pale; anthers purple. Stigma cream. Drupe 9–12 × 7–11 mm, yellow or yellow turning dark red, subglobose, soft. Flowers 6–7.

(1) Forma opulus Outer flowers of inflorescence large and sterile, inner smaller and fertile. Drupe red.

(2) Forma flavum Rehd. Drupe yellow.

(2) Forma roseum (L.) Hegi Snowball Tree V. opulus var. roseum L.; V. opulus var. sterilis DC. All flowers of inflorescence large and sterile, at first white turning deep rose as they fade. Drupe absent. (3) Forma flavum (Horwood) P. D. Sell V. opulus var. flavum Horwood Outer flowers large and sterile, inner smaller and fertile. Drupe yellow. Native. Woods, scrub and hedgerows, especially on damp soils. Frequent throughout Great Britain and Ireland except in northern Scotland. Europe except for the north and most of the Mediterranean region; central and west Asia; doubtfully recorded from Algeria. The wild tree is forma opulus. Forma roseum is commonly grown in gardens and is sometimes seen planted on roadsides and in parks; it has been known in European gardens since the sixteenth century. Forma flavum is sometimes bird-distributed from gardens. A very large number of those planted by roads, in estates and in planted woods and infills in woods and hedgerows may well belong to one or other of the following two species, especially if they originate from nurseries. 10. V. sargentii Koehne Asian Guelder Rose Deciduous, fairly open, broad shrub. Stems up to 4 m, numerous, erect, much branched. Bark medium greyishbrown, fissured and rather corky. Branches rather long, ascending; twigs pale greyish-brown, faintly wrinkled, slightly angled, with numerous lenticels, entire, glabrous; young shoots pale green, often tinted reddish, markedly striate. Buds 5–7 × 3–4 mm, ovoid, reddish-brown, scaly. Leaves opposite, of thicker texture than V. opulus, simple; lamina 5–9 × 5–9 cm, dull medium to rather dark yellowishgreen on upper surface, paler beneath, with even paler veins, turning reddish-purple in autumn, obovate in outline, palmately lobed, the lobes 3, the central lobe as wide or wider than long, ovate, rounded at apex and dentate, the teeth more or less mammiform, the lateral lobes broad, shallow and rounded with rather shallow teeth, rounded and entire for much of lower part, glabrous on upper surface, with numerous simple eglandular hairs on the veins beneath and sometimes on the surface; with a main vein

(1) Forma sargentii Drupe turning yellow, then red.

Introduced in 1892. It has been planted in plantations and as an infill in hedges in Cambridgeshire and is almost certainly widespread, being mistaken for V. opulus. It is native of north-east Asia. Forma sargentii is the common plant, but forma flavum may be the garden escape recorded in several places. 11. V. trilobum Marsh. American Guelder Rose V. opulus subsp. trilobum (Marsh.) Hult´en; V. opulus var. americanum Aiton; V. opulus var. pimina Michx; V. edule Pursh; V. oxycocus Pursh; V. americanum auct. Deciduous, fairly open, broad shrub. Stems up to 4 m, numerous, erect, much branched. Bark grey, fissured and rather corky. Branches rather long, ascending or spreading; twigs pale greyish-brown, longitudinally ridged; young shoots green, longitudinally ridged, glabrous. Buds 3.5– 5.0 × 2–3 mm, ovoid, obtuse at apex; scales green, ovate, obtuse at apex. Leaves opposite, simple; lamina 5–11 × 4– 10 cm, medium yellowish-green on upper surface, paler beneath, with even paler veins, turning reddish-purple as early as July, mostly ovate in outline, palmately lobed, the lobes 3, the central lobe at least of the larger leaves usually longer than wide, coarsely dentate, sometimes throughout sometimes only at apex with 4–14 teeth, the apical tooth often rather long, the lateral lobes dentate with 4–12 teeth, the apical tooth sometimes long, base of leaf rounded or subcordate, mostly entire, glabrous or nearly so on the upper surface, with short simple eglandular hairs beneath, pectinate on the veins; with a main vein to each lobe and up to 8 pairs of side veins; petiole up to 35 mm, green or brownish, glabrous, shallowly grooved above; with a few, small, disclike glands near the apex; stipules subulate. Flowers numerous, with sickly scent, in compound flat-topped cymes 7– 10 cm in diameter, the outer large and sterile, the inner smaller and fertile; peduncles and pedicels pale yellowishgreen, with minute, scattered glandular hairs. Stamens 5; filaments pale; anthers yellow. Stigma cream. Drupe 7– 9 × 7–8 mm, orange or red, subglobose, soft. Flowers 6–7. Introduced in 1812. Planted in plantations and as infills in hedgerows in Cambridgeshire and is almost certainly widespread, being mistaken for V. opulus. It is native of North America. There is a yellow-fruited form of it in gardens.

3. Symphoricarpos 3. Symphoricarpos Duhamel Symphoria Pers.; Anisanthus Willd. ex Roem. & Schult.; Magaris DC. Deciduous shrubs. Leaves simple, entire or sometimes deeply lobed, exstipulate. Flowers in dense terminal spikes, actinomorphic. Calyx (4–)5-lobed, campanulate. Corolla 5-lobed. Stamens (4–)5. Style 1, slender; stigma capitate. Ovary 4-celled, with 2 fertile cells each with a single ovule, and 2 sterile cells. Fruit a drupe with 2 seeds. About 17 species in North America and one in China. Gilbert, O. L. (1995). Symphoricarpos albus (L.) S. F. Blake (S. rivularis Suksd.; S. racemosus Michx) in Biological flora of the British Isles. Jour. Ecol. 83: 159–166. Jones, G. N. (1940). A monograph of the genus Symphoricarpos. Jour. Arnold Arbor. 21: 201–252. 1. 1. 2. 2. 3. 3. 4. 4.

2. Drupe 9–15 mm 3. Drupe 4–9 mm 1. albus var. laevigatus Leaves glabrous; drupe white Leaves hairy beneath; drupe white, pink or pinkish-purple 4. × doorenbosii 2. orbiculatus Drupe coral pink to purplish all over Drupe white or tinged pink or pink on one side and white 4. on the other Style hairy; drupe 4–7 mm in diameter, purplish-red on 3. × chenaultii exposed side, whitish on other side Style glabrous; drupe 7–9 mm, white or tinged with pink 5. microphyllus

1. S. albus (L.) S. F. Blake Snowberry Vaccinium album L.; S. racemosus Michx; S. rivularis Suksd.; Xylosteon album (L.) Moldenke; Lonicera racemosa (Michx) Pers.; Symphoria racemosa (Michx) Pursh Deciduous shrub spreading underground and suckering. Stems numerous, 1–3 m, erect, slender, branched. Bark brown, shredding on older branches. Branches arching; twigs slender, brown; young shoots yellowish-brown, glabrous. Buds 1.5–2.0 × 1.2–1.5 mm, brown, ovate, acute at apex; scales ovate, acute at apex. Leaves opposite; lamina (2.0–)2.5–8.0(–9.0) × 1.2–5.0 cm, dull green on upper surface, paler beneath, glabrous; those of the twigs elliptical or ovate, obtuse at apex, entire or a few sinuately lobed, cuneate or rounded at base; petiole up to 5 mm; leaves of the sucker shoots often conspicuously lobed; veins 4–6 pairs. Flowers 3–7, 5–6 mm in diameter, in terminal spike-like racemes; bracts and bracteoles 1.0–1.5 mm, ovate, acuminate at apex, entire, glabrous. Calyx 2.5–3.0 mm, green, glabrous, divided about one-fifth of the way to the base; lobes (4–)5, triangular-ovate, acute at apex. Corolla 5–6 mm, white and hairy within, pink outside, divided halfway to the base; lobes 5, triangular-ovate, subobtuse at apex. Stamens (4–)5; filaments 2.0–2.5 mm, white; anthers yellow. Style 1, slender, white; stigma green, capitate. Drupe 9–15 mm in diameter, white, subglobose; seeds 2, 4–6 × 3.0–3.5 mm, white, ellipsoid, plano-convex. Flowers 6–9. Pollinated by bees, wasps and syrphids. Fruits 9–11. 2n = 36, 54, 72.

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Introduced. First successfully introduced to gardens in 1817. Naturalised in woods, scrub, cliffs, scree slopes, river shingle and rough ground where it spreads by suckers and forms large dense thickets. It is outside the area where it can be established by seed. Frequent throughout Great Britain and Ireland. Native of western North America from Alaska and Alberta to California and Colorado. Our plants, like those of Continental Europe, are referable to subsp. laevigatus (Fernald) P. D. Sell (S. racemosus var. laevigatus Fernald; S. albus var. laevigatus (Fernald) S. F. Blake) and are distinguished by their glabrous twigs and leaves and smaller fruits. 2. S. orbiculatus Moench Coralberry Lonicera symphoricarpos L.; S. vulgaris Michx; Symphoria glomerata Pursh; Symphoria rubra Raf.; S. imberbis Tausch; S. spicatus Engelm. Deciduous shrub spreading underground and suckering. Stems 0.5–2.0 m, erect or ascending, slender, branched. Bark grey and shredding. Branches erect or ascending, grey; young shoots pale brown or purplish, sparsely to densely short-hairy. Buds small and scaly. Leaves opposite; lamina (1.5–)2–5(–6) × 1.0–3.5 cm, dull green on upper surface, bluish beneath, turning crimson in autumn, elliptical, ovate or subrotund, obtuse to acute at apex, entire or undulate, rounded or slightly cuneate at base, glabrous on upper surface, variously shortly hairy beneath; veins impressed; petiole 1–4 mm, shortly hairy. Flowers 2–3 mm in diameter, in densely crowded, short, axillary, spike-like racemes on the branches of the season; bracts and bracteoles small. Calyx 2.0–2.5 mm, divided half of the way to the base; lobes (4–)5, triangular, ciliate. Corolla 2.5–4.0 mm, pinkish, broadly campanulate, divided half of the way to the base; lobes 5, rounded; glabrous on outer surface, hairy within. Stamens 5; filaments white; anthers yellow. Style 1, hairy; stigma green, capitate. Drupe 4–6 mm in diameter, coral-pink to purplish all over, ellipsoid, fleshy; nutlets 2, 2.5–3.5 mm, flattened-ellipsoid, obtuse at each end. Flowers 7–8. Fruits 9–11. 2n = 18. Introduced. Naturalised on Dartford Heath in Kent since at least 1982 and perhaps elsewhere. Native of North America from southern New York to Florida, westwards to Texas, northern Mexico, Colorado and east South Dakota. 3. S. × chenaultii Rehder Pink Snowberry S. microphyllus × orbiculatus Deciduous shrub spreading underground and suckering. Stems up to 2 m, erect to procumbent, rooting at tips, much branched. Bark grey and shredding. Branches erect or ascending and arching, grey; young shoots reddish-brown, with dense, short hairs. Buds small and scaly. Leaves opposite; lamina 1–2 × 0.6–1.7 cm, dark green on upper surface, bluish-green beneath, elliptical or broadly elliptical, obtuse to acute and mucronate at apex, rounded or cuneate at base, glabrous on upper surface, densely short hairy beneath; veins 3–5 pairs, impressed; petioles 1–2 mm, shortly hairy. Flowers 2–3 mm in diameter, in axillary and terminal fascicles and spikes; bracts small. Calyx 2.0–2.5 mm, divided half of the way to the base; lobes (4–)5, triangular-ovate,

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ciliate. Corolla 3–6 mm, whitish, tinted pink, divided onethird of the way to the base, glabrous; lobes 5, elliptical. Stamens 5; white, glabrous; anthers yellow. Style 1, longhairy; stigma greenish, capitate. Drupe 4–7 mm in diameter, subglobose, bright purplish-red on the upper exposed side with numerous, minute, pale dots, passing gradually into pinkish-white on the lower side sprinkled with purplish spots. Flowers 7–8. Fruits 9–11. 2n = 18. Introduced. Originated in the gardens of L´eon Chenault et Cie, at Orl´eans, France about 1910. Naturalised in woods and scrub and rough ground, planted by roadsides and in landscaped areas. Scattered records in Great Britain north to central Scotland. 4. S. × doorenbosii Kr¨ussm. Doorenbos Snowberry S. albus subsp. laevigatus × microphyllus × orbiculatus Deciduous shrub spreading underground and suckering. Stem up to 2 m, stiffly erect or ascending, much branched. Bark grey and shredding. Branches erect, pendulous or ascending, grey; young shoots reddish-brown, with dense, short hairs. Buds small and scaly. Leaves opposite; lamina up to 4.0 × 2.5 cm, dark green on upper surface, paler beneath, elliptical to broadly obovate, obtuse at apex, entire, rounded to cuneate at base; glabrous on upper surface, shortly hairy beneath; veins 3–5 pairs; petioles up to 2 mm, with short hairs. Flowers 2–3 mm in diameter, in short racemes; bracts small. Calyx 2–3 mm, divided half of the way to the base; lobes (4–)5, triangular, acute at apex. Corolla up to 7 mm, campanulate, pink, divided one-third of the way to the base; lobes 5, elliptical or ovate, glabrous. Stamens 5; filaments white, glabrous; anthers yellow. Style 1, glabrous; stigma greenish, capitate. Drupe 12–13 mm, white tinted pink, white or purplish-pink, globose. Flowers 7–8. Fruits 9–11. This group of hybrids was first raised in Holland by G. A. Doorenbos, after whom it was named, from a mixed planting of S. albus subsp. laevigatus, S. orbiculatus and S. × chenaultii. Introduced. Persistent garden escapes have been recorded from a meadow west of Borough Green, and a hedge near Hextable Nature Reserve, both in Kent. They will almost certainly be found elsewhere. 5. S. microphyllus Kunth Small-leaved Snowberry S. montanus Kunth; S. glaucescens Kunth; Anisanthus microphyllus (Kunth) Willd. ex Roem. & Schult.; Symphoria microphylla (Kunth) Spreng.; Symphoria glaucescens (Kunth) Spreng.; Symphoria montana (Kunth) Spreng.; S. mexicanus K. Koch; Margaria nudiflora DC. Deciduous shrub spreading underground and suckering. Stem 2–3 m, erect, much branched. Bark smooth, scarcely shredding. Branches erect or spreading, smooth; young shoots dark brownish-red, crisp-puberulent with curved hairs. Buds small and scaly. Leaves opposite; lamina 1.0– 2.5 × 0.7–1.5 cm, dull green on upper surface, greyishgreen beneath, broadly elliptical to subrotund, rounded acute or apiculate at apex, entire, rounded or cuneate at base, glabrous on upper surface, shortly hairy beneath especially on the veins; veins 3–5 pairs, impressed; petioles

1–2 mm, shortly hairy. Flowers 2–3 mm in diameter, in pairs or solitary and axillary or in a short terminal spike; bracts 1.0–1.2 mm, ovate, obtuse at apex. Calyx 2.0–2.5 mm, divided half of the way to the base; lobes 5, triangular, acute at apex. Corolla 9–10 mm, white flushing pinkish, narrowly campanulate, divided one-third of the way to the base; lobes (4–)5, ovate; glabrous outside, hairy inside. Stamens 5; filaments white; anthers yellow. Style 1, glabrous; stigma greenish, capitate. Drupe 7–9 mm in diameter, white or tinged with pink, translucent, globose; nutlets 2, about 3 mm, flattened-ellipsoid, obtuse at each end. Flowers 7–8. Fruit 9–11. Introduced. Recorded from a disused railway yard at Woofferton in Shropshire and in a wood at Earlsmill, Darnaway in Morayshire. Probably no longer in cultivation. Native of New Mexico, Mexico and Guatemala. 4. Linnaea L. Procumbent, evergreen, dwarf shrubs. Leaves opposite, simple, shallowly crenate-serrate, exstipulate. Flowers in pairs on long peduncles which are terminal on short lateral branches, with one bract each at base and 2 bracteoles at the apex of the peduncle, actinomorphic. Calyx 5-lobed. Corolla 5-lobed, campanulate. Stamens 4, 2 shorter, inserted towards the base of the tube. Style 1, filiform; stigma capitate, bilobed. Ovary 3-celled, 1 fertile cell with a single ovule, 2 sterile cells. Fruit an achene. One species with several races in Europe, Asia and North America. Named after Carl von Linn´e (Linnaeus) (1707– 78). Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 13: 156–157, 369. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough.

1. L. borealis L. Twinflower Evergreen, dwarf shrub forming large mats. Stems up to 40 cm, procumbent, pale brown, slender, with short and very short, pale simple eglandular hairs, with leafy branches. Leaves opposite; lamina 0.5–1.5 × 0.5–1.5 cm, deep yellowish-green on upper surface, paler beneath, subrotund or broadly ovate, rounded at apex, crenate-serrate with a few shallow teeth, rounded at base, with few, short, pale simple eglandular hairs particularly on the margin and midrib beneath; petioles 2–4 mm, pale green, with short, pale simple eglandular hairs. Flowers 8–12 mm in diameter, in pairs; peduncles 30–70 mm, pale green, with numerous, very short to short, pale, unequal glandular hairs; pedicels 7–20 mm, with short, unequal glandular and some longer simple eglandular hairs; bracts 2, lanceolate, obtuse to acute at apex; bracteoles 2, lanceolate, obtuse at apex. Calyx 2.5– 3.0 mm, divided almost to the base, with glandular and simple eglandular hairs; lobes 5, linear-lanceolate, obtuse to acute at apex. Corolla 7.5–10.0 mm, pink to mauvishpink, campanulate, divided half way to the base; lobes 5, broadly ovate, rounded at apex, hairy inside. Stamens 4; filaments 2.0–2.5 mm. Style 1, 6–7 mm; stigma capitate, bilobed. Achene 2.5–3.0 mm, ellipsoid, glandular-hairy,

7. Lonicera now rarely produced. Flowers 6–8. Pollinated by insects. 2n = 32. Native. Rather bare ground in the shade of rocks or trees, mostly in woods, ascending to 730 m. Local to rare and decreasing in eastern Scotland north to Sutherland, formerly south to Yorkshire. Northern Europe and Asia, from 71◦ 10 N in Norway extending southwards to the Alps and eastern Carpathians, the Caucasus and Japan. Our plants are subsp. borealis which extend eastwards through Asia to westernmost North America. Cultivated plants are often subsp. americana (Forbes) Hult´en (L. americana Forbes; L. borealis var. americana (Forbes) Rehder) which has the corolla 10–16 mm, and is the widespread American race. The species is a member of the Circumpolar Borealmontane element. 5. Leycesteria Wall. Deciduous shrubs, often semi-herbaceous. Leaves opposite, simple, entire to serrate, stipulate or exstipulate. Flowers in crowded, terminal spikes with large purple or purplishgreen bracts, more or less actinomorphic. Calyx unequally 5-lobed. Corolla regularly 5-lobed, funnel-shaped. Stamens 5. Style 1, slender; stigma capitate. Ovary 5(–8)celled, each cell with several ovules. Fruit a several-seeded berry. Six species in the Himalaya and China. Named after Wm. Leycester, a judge in Bengal. 1. L. formosa Wall. Himalayan Honeysuckle Deciduous shrub, often semi-herbaceous. Stems numerous, up to 2 m, erect or arching, bluish-green with a bloom, hollow, smooth. Branches spreading or arching; young shoots green, with appressed, pale simple eglandular hairs. Leaves opposite, lamina 6–19 × 2–12 cm, dull medium yellowish-green on upper surface, paler and more creamygreen beneath, narrowly to broadly ovate, long-acute to acuminate at apex, sometimes almost caudate, entire or shallowly serrate, shortly cuneate, subtruncate or subcordate at base, minutely hairy on both surfaces; stipules when present linear, acute; petioles 5–35 mm, pale green or sometimes flushed purplish, channelled above, with long, pale, semi-appressed simple eglandular hairs. Flowers 13– 15 mm in diameter, in crowded terminal spikes or in the axils of the upper cauline leaves; bracts 12–50 × 5–23 mm, deep reddish-purple, ovate to ovate-lanceolate, acuminatecuspidate at apex, entire, cordate at base. Calyx 5–7 mm, purplish, persisting, purple glandular-hairy, with 5, very unequal and subulate lobes. Corolla 15–20 mm, white, funnel-shaped, with a small glandular dilation at its base, glabrous, the 5 regular lobes ovate, obtuse at apex and spreading, with 5 nectariferous glands at the base of the tube. Stamens 5, as long as the corolla; filaments white, glabrous; anthers pale yellow. Style 1, longer than corolla, white; stigma capitate, pale yellow. Berry 5–12 mm, globose or broadly ellipsoid, reddish-purple to blackish-purple, glandular-hairy. Flowers 7–9. Fruits 9–11. 2n = 18. Introduced. An established garden escape naturalised in woods, shrubberies and rough ground, by railways and sometimes planted for Pheasant cover. Probably distributed by birds. Scattered localities throughout Great Britain and

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Ireland and probably increasing. Native of the Himalayas and Burma. 6. Weigela Thunb. Deciduous shrubs. Leaves opposite, simple, crenate-serrate, exstipulate. Flowers in axillary clusters of 1–4(–6) on short lateral twigs, actinomorphic; bracts present. Calyx 5-lobed. Corolla 5-lobed. Stamens 5. Style 1, long; stigma peltatecapitate. Ovary 2-celled. Fruit a capsule, opening by 2 valves. About 12 species in temperate Asia. Named after Christian Ehrenfried Weigel (1748–1811). 1. W. florida (Bunge) DC. Weigela Calysphyrum florida Bunge; Diervilla florida (Bunge) Siebold & Zucc.; W. rosea Lindl. Deciduous shrub up to 2.5 m, with spreading habit. Branches arching; twigs brownish, slightly hairy; young shoots pale green, with 2 lines of hairs. Leaves opposite, simple; lamina 3.5–9.0 × 2–4 cm, medium green on upper surface, paler beneath, ovate or obovate, acute to acuminate at apex, crenate-serrate, rounded at base, sparsely to densely white-hairy on the veins of both surfaces and the margins; veins 5–6 pairs, curved upwards; petiole 2–4 mm, very hairy. Inflorescence an axillary cluster of 1–6 flowers. Bracts small, opposite, subulate, appressed. Calyx 8–9 mm, divided to about halfway to the base, glabrous; lobes 5, linear-lanceolate, obtuse at apex. Corolla 25–35 mm, deep rose on the outside, paler and almost white within, funnelshaped, hairy, divided for about one-third of the way to the base; lobes 5, broadly oblong, rounded at apex, spreading. Stamens 5, slightly exserted; filaments 10–15 mm, white; anthers yellow. Style as long as the stamens; stigma peltatecapitate, slightly 2-lobed. Capsule 15–20 × 1.5–2.0 mm, linear. Flowers 5–6. Introduced to gardens in 1844. Garden escape persistent on an overgrown area by the railway at Redhill in Surrey and on a disused tip at Bedminster Down, Somerset; casual elsewhere. Native of northern China and Korea. 7. Lonicera L. Deciduous or evergreen shrubs or climbers. Leaves opposite, simple, sometimes lobed, or entire, exstipulate. Flowers sessile, in pedunculate axillary pairs or in terminal heads; bracts and bracteoles usually present. Calyx 5-lobed. Corolla 5-lobed and more or less actinomorphic, or zygomorphic with a 4-lobed upper lip and 1-lobed lower lip. Stamens 5. Style 1, long; stigma capitate or slightly lobed. Ovary 2- to 3-celled, with several ovules per cell. Fruit a several-seeded berry. About 200 species in the north temperate zone, extending south to Mexico and Java. Barrett, R. (1995). Aliens on Hayling Island. B.S.B.I. News 68: 37–39. [L. tatarica.] Grime, J. P. et al. (1988). Comparative plant ecology. London. [L. periclymenum.] Rehder, A. (1903). Synopsis of the Genus Lonicera. Annual Rep. Missouri Bot. Gard. 14: 27–232. Yeo, P. F. (1964). Lonicera pileata and L. nitida in cultivation. Baileya 12: 56–66.

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1. Flowers and berries sessile in terminal and subterminal 2. whorls 1. Flowers and berries in pairs, sessile at the apex of a common axillary stalk and sometimes crowded at the 5. ends of branches 3. 2. All leaves separate, not fused in pairs 2. At least most of the apical pair of leaves on each branch 4. fused around the stem at their base 3. Leaves glabrous or with an occasional simple eglandular hair; corolla usually with reddish colouring, 8(i). periclymenum var. periclymenum strongly fragrant 3. Leaves with long, pale simple eglandular hairs, especially beneath; corolla usually without reddish colouring, only 8(ii). periclymenum var. hirsuta slightly fragrant 4. Bracteoles at base of each flower minute or absent 9. caprifolium 4. Bracteoles about 1.5–2.0 mm, obscuring the base of the 10. × italica ovary 6. 5. Stems twining 7. 5. Stems not twining 6. The 2 bracts at the base of each flower-pair subulate; flowering nodes often clustered into terminal spikes; 6. henryi corolla glabrous on outside 6. The 2 bracts at the base of each flower-pair ovate or elliptical and leaf-like; flowering nodes not clustered near the branch apex; corolla glandular-hairy on outside 7. japonica 7. The 2 bracts at the base of each flower-pair (and the bracteoles within) ovate, obscuring the base of the 3. involucrata flower, purple and enlarging in fruit 7. The 2 bracts at the base of each flower-pair subulate to linear-lanceolate, not obscuring the ovaries and scarcely 8. enlarging in fruit 9. 8. Leaves deciduous; corolla distinctly 2-lipped 10. 8. Leaves evergreen; corolla actinomorphic 4. tatarica 9. Twigs and leaves more or less glabrous 5. xylosteum 9. Twigs and leaves hairy 10. Plant rarely more than 1 m; leaves (6–)12–32 mm, 1. pileata narrowed at base 10. Plant often more than 1 m; leaves (4–)6–16(–20) mm, 2. nitida truncate, rounded or subcordate at base

Section 1. Lonicera Xylosteon Mill. Erect or procumbent deciduous shrubs. Flowers in axillary pairs. Corolla actinomorphic or 2-lipped; tube short. 1. L. pileata Oliv. Box-leaved Honeysuckle L. ligustrina var. pileata (Oliv.) Franchet; Caprifolium pileatum (Oliv.) Kuntze Evergreen or partially deciduous shrub of low, neat, dense, spreading habit. Stems up to 2 m, often procumbent or spreading. Bark pale brown, fissured. Branches often horizontal; twigs pale greyish-brown with pale fissures, with few, pale simple eglandular hairs; young shoots often dark purple above, pale beneath, slender, with numerous to dense, pale simple eglandular hairs. Buds scaly, brownish. Leaves distichous; lamina (0.6–)1.2–3.2 × 0.4–1.4 cm, bright dark green on upper surface, paler beneath, thick,

narrowly elliptical, narrowly lanceolate, oblong or oblonglanceolate, narrowed but more or less obtuse at apex, entire with a thickened margin, the midrib prominent beneath, glabrous, narrowed at base; petiole very short. Flowers about 6 mm in diameter, in sessile pairs in the leaf axils, fragrant; bracts lanceolate to subulate. Calyx about 1.5 mm, pale green, unequally 5-lobed, the lobes lanceolate or ovate and obtuse at apex, with a collar-like appendage, subtended by bracts and bracteoles. Corolla 6–8 mm, yellowish-green or cream, hairy, actinomorphic, the 5 lobes about one-third the length of the tube, triangular-ovate, obtuse at apex and more or less spreading. Stamens 5, exceeding the corolla; filaments 5–7 mm, hairy, white; anthers cream. Style 1, greenish, not exceeding the corolla. Berry 5–8 mm in diameter, purplish-violet, globose. Flowers 4–5. Much visited by bees. 2n = 18. Introduced to gardens in 1900. Much-grown in shrubberies and road-borders and sometimes self-sown. Scattered localities in central and southern Britain. Native of the West Hupeh and West Szechwan Provinces of China. 2. L. nitida E. H. Wilson Wilson’s Honeysuckle L. ligustrina var. yunnanensis Franch.; L. pileata forma yunnanensis (Franch.) Rehder Evergreen shrub of dense, leafy habit. Stems up to 1.8 cm, erect, much branched. Bark greyish, flaking. Branches arching; twigs pale brown, striate; young shoots often purplish, slender and erect, minutely hairy. Buds scaly, greenish or brownish. Leaves close together; lamina (0.4–)0.6– 1.6(–2.0) × 0.4–1.0(–1.3) cm, bright green on upper surface, paler beneath, lanceolate, ovate or elliptical, more or less obtuse at apex, entire, truncate or rounded at base or sometimes subcordate, glabrous or with a few short hairs; veins conspicuous beneath; petiole very short, often reddish, shortly hairy. Flowers in pairs in the leaf axils, about 6 mm in diameter, fragrant; bracts lanceolate to subulate. Calyx green, sometimes violet-purple, with a thickened, downwardly overlapping rim at the base and 5 shallow, obtuse lobes above. Corolla 5.5–6.0 mm, creamy-green, actinomorphic, tubular, gibbous at the base, 5-lobed above, glandular on the outside, with very short, simple eglandular hairs inside. Stamens 5, inserted near the middle of the corolla; filaments white, hairy towards the base; anthers pale yellow, exserted. Style 1, greenish; stigma shorter than anthers, about as long as the corolla. Berries in 2s, 4–7 mm in diameter, shining purple when ripe, more or less globose. Flowers 4–5. 2n = 18. Introduced. Much grown for hedging and then rarely flowering. Also grown in gardens from which it escapes and is self-sown in scrub, hedges, woodland, banks and rough ground. Scattered records throughout Great Britain and Ireland. Native of China. 3. L. involucrata (Richardson) Banks ex Spreng. Californian Honeysuckle Xylosteon involucratum Richardson; L. ledebourii Eschsch.; L. mocinianan DC.; Caprifolium involucratum (Richardson) Kuntze; Distegia nutens Raf. Dense deciduous shrub with broad, rounded habit. Stems numerous, up to 2 m, often intertwined. Bark pale

7. Lonicera greyish-brown, furrowed. Branches numerous, long, erect, arching or spreading; twigs pale brown, ridged, glabrous; young shoots pale green, to purplish-brown, ridged. Buds scaly, pale brown. Leaves decussate; lamina 4–12 × 3–6 cm, dark green on upper surface, paler beneath, elliptical, elliptic-oblong, ovate or oblong, acute-cuspidate at apex, entire, cuneate or rounded at base, glabrous on upper surface, with numerous, short, pale simple eglandular hairs beneath and on the margins; veins impressed on upper surface, prominent beneath; petiole up to 10 mm, pale green, with unequal, pale simple eglandular hairs. Flowers in pairs in the leaf axils, 8–10 mm in diameter, not or barely fragrant, on stalks up to 30 mm which are clothed with numerous, unequal hairs, subtended by 2 reddish-purple, rounded bracts, up to 12 × 20 mm and 2 bracteoles much smaller, purple, ovate and enlarging in fruit, all the bracts and bracteoles with numerous unequal glandular hairs and longer simple eglandular ones. Calyx absent. Corolla 8–10 mm, actinomorphic, infundibuliform, saccate at base, dull orange to red above, yellow below, 5-lobed, the lobes broadly ovate and rounded, with short, glandular hairs. Stamens 5, inserted above the middle of the tube; filaments red; anthers yellow. Style 1, orange-red; stigmas green. Berry 7–8 mm in diameter, shining black. Flowers 5–6. 2n = 18. Introduced. Frequently grown in gardens and sometimes bird-sown in marginal and rough ground. Scattered records in England and Ireland. Relics of old hedges have survived for many years in Ireland. Native of western North America. 4. L. tatarica L. Tartarian Honeysuckle Xylosteon cordatum Moench; Xylosteon tataricum (L.) Dum.-Cours.; Chamaecerasus tataricus (L.) Billiard; Caprifolium tataricum (L.) Kuntze Deciduous shrub with rather narrow outline. Stems numerous, up to 4 m, erect, much branched. Bark pale brown, rough and shredding. Branches erect, arching over above; twigs pale brown, slender, spreading, glabrous, with hollow pith; young shoots pale brown, striate, glabrous. Buds scaly, pale brown. Leaves opposite; lamina 2.5–8.0 × 1.5– 4.5 cm, matt medium green on upper surface, paler and greyish beneath, lanceolate, ovate or elliptical, obtuse to acute at apex, entire, rounded, subtruncate or cordate at base, glabrous; petioles up to 10 mm, pale green, glabrous. Flowers 10–15 mm in diameter, in stalked pairs in the leaf axils; bracts 2–5 mm, subulate, glabrous; bracteoles elliptical, obtuse at apex, free or nearly so. Calyx 2.0–2.5 mm, pale green, with 5 rounded lobes. Corolla 15–22 mm, white to deep pink, zygomorphic, deeply 5-lobed with a short tube, the lobes elliptical to obovate, rounded at apex, 2 small and 3 large, glabrous. Stamens 5, exceeding the tube; filaments 5–6 mm, white, glabrous; anthers yellow. Style 1, white; stigma green, protruding from tube. Berry 6.5– 7.5 mm in diameter, red, flattened-globose. Flowers 5–6. 2n = 18. Very variable in flower colour and size. Introduced. A persistent garden escape or relic. Recorded from Hayling Island in Hampshire, West Grinstead in Sussex, Hextable in Kent and Oulton Broad in Suffolk. Native of western and central Asia.

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5. L. xylosteum L. Fly Honeysuckle Caprifolium dumetorum Lam. nom. illegit; Caprifolium xylosteum (L.) Gaertn.; Lonicera dumetorum Moench; Xylosteon dumetorum Moench; Xylosteon vulgare Borkh. Lonicera vulgaris (Borkh.) R¨ohl.; Lonicera pubescens Stokes nom. illegit.; Chamaecerasus xylosteum (L.) Billiard; L. ochroleuca St-Lag.; Euchylia villosa Dulac; L. villosa (Dulac) K. Koch Deciduous shrub of vigorous growth and rounded, dense, leafy habit. Stems numerous, up to 3 m, erect, much branched. Bark pale greyish-brown, rough and furrowed. Branches spreading; twigs pale greyish-brown, furrowed, glabrous; young shoots usually purplish on upper surface, very pale beneath, with numerous, short, pale, wavy simple eglandular hairs. Buds scaly, greenish. Leaves opposite; lamina 3–7 × 2–4 cm, dark greyish- or bluish-green on upper surface, paler beneath, soft, elliptical, broadly elliptical, oblong-elliptical or almost subrotund, rounded to subacute at apex, entire with a thickened margin, rounded to subcordate at base, covered with short, curled, pale simple eglandular hairs on both surfaces and the margins, becoming less so above; petiole up to 7 mm, pale green or purplish, shortly hairy. Flowers 6–8 mm in diameter, in stalked pairs in the leaf axils, not or barely fragrant, subtended by 2 subulate to linear-lanceolate bracts; bracteoles about half as long as ovaries. Calyx about 1 mm, green, with 5 unequal lobes more or less ovate and acute at apex. Corolla 8–15 mm, yellowish-green or cream, hairy, zygomorphic, divided for two-thirds of the way to the base into 5 oblong, rounded lobes and markedly 2-lipped, with a short tube. Stamens 5, exceeding the tube; filaments 5–6 mm, white, hairy at base; anthers greenish. Style 1, white; stigma green, protruding from tube. Berry 6–8 mm in diameter, globose but flattened, dark red. Flowers 4–5. 2n = 18. Native in Ash, Beech and Tilia platyphyllos woodland in north-facing coombes just above the spring-line on the South Downs of West Sussex; also widely bird-sown in hedges, woods and scrub throughout much of Britain and Ireland. Europe from Scandinavia to Spain, Sicily, Macedonia and Caucasus; northern and western Asia. A member of the Eurosiberian Temperate element, but variable and may need dividing into subspecies. Section 2. Nintooa (Spach) Maxim. Woody climbers. Flowers in axillary pairs. Corolla 2lipped; tube long. 6. L. henryi Hemsl. Henry’s Honeysuckle Caprifolium henryi (Hemsl.) Kuntze Evergreen climber of spreading habit. Stems up to 5(–10) m, sinuate, ascending. Bark pale brown, strongly ridged, peeling. Branches ascending or spreading. Twigs chestnutbrown, slender, peeling, with numerous lenticels; young shoots yellowish-green, with dense, short, curled, creamy brown, ascending simple eglandular hairs. Buds scaly. Leaves distichous; lamina 4–13 × 1.0–3.5 cm, medium to dark, matt green with a paler midrib on upper surface, glossy, paler yellowish-green beneath, lanceolate, oblonglanceolate or ovate, acute or acuminate at apex, entire,

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1 4 4 . C A P R I F O L I AC E A E

rounded to subcordate at base, with pale, appressed hairs on the margin and midrib above, otherwise glabrous; veins 5–10 pairs, impressed on upper surface, prominent beneath; petiole 5–10 mm, yellowish-green, sometimes curved round stem, sometimes hairy. Flowers in sessile, axillary pairs, but clustered in terminal groups, zygomorphic, 15–25 mm in diameter, fragrant; bracts 3–4 × about 1 mm, yellowishgreen suffused reddish, subulate, obtuse or acute at apex; bracteoles partly obscuring the ovaries. Calyx 5.0–6.5 mm, yellowish-green, divided for about one-quarter of the way to the base into 5 lobes, the lobes triangular-ovate, more or less acute at apex, sparsely hairy. Corolla 15– 25 mm, pale creamy-yellow, tinged purplish-red, 2-lipped, the upper lip with 4, oblong, obtuse lobes, the lower lip undivided, oblong and obtuse at apex, glabrous outside, appressed-hairy inside. Stamens 5, slightly exserted; filaments cream, sparsely hairy; anthers yellow, suffused red. Style 1, yellowish-green, sparsely hairy; stigma green, about as long as the anthers. Berry 7–8 × 9–10 mm, more or less globose, blackish-purple with a pale blue bloom. Flowers 6–7. Fruits 9–10. 2n = 54. Introduced. Grown in gardens and naturalised from throw-outs or bird-sown. In a few localities in Surrey and from Sherrardspark Wood in Hertfordshire. Native of western China. 7. L. japonica Thunb. ex Murray Japanese Honeysuckle Nintooa japonica (Thunb. ex Murray) Sweet; Caprifolium japonicum (Thunb. ex Murray) Dum.-Cours.; L. cochinchinensis G. Don; L. repens Miq. Vigorous, rampant evergreen climber. Stems up to 5(–10) m, dark orange brown, hollow, twining or trailing, with numerous, greyish simple eglandular hairs. Bark smooth. Branches spreading; twigs medium brown, paler beneath, with numerous, pale simple eglandular hairs; young shoots pale brownish-green, with dense, unequal simple eglandular hairs. Buds scaly, brownish-green. Leaves opposite; lamina 2–8 × 0.8–6.0 cm, dull olive or yellowish-green on upper surface, paler beneath, ovate, elliptical or oblong-elliptical, rounded, abruptly acuminate or subacute at apex, entire, narrowed to rounded at base, hairy on both surfaces to nearly glabrous; veins prominent beneath; petiole 3–8 mm, hairy. Flowers 10–15 mm in diameter, in pairs in the leaf axils, zygomorphic, very fragrant; peduncles 9–18 mm, hairy; bracts leaf-like, ovate or elliptical; bracteoles one-third to half as long as ovaries. Calyx about 4–5 mm, divided for about one-third of its length; lobes 5, triangular-ovate, acute at apex, hairy mainly on the margin. Corolla 30–50 mm, zygomorphic, white to pale yellow, sometimes purplish, covered with reddish glandular hairs; limb 2-lipped, the upper lip varying from shorter to longer than the tube. Stamens 5; filaments white; anthers cream to yellowish. Style 1, white; stigma green, exceeding anthers. Berry 7–8 mm in diameter, black, globose. Flowers 6–11. 2n = 18. Introduced. Often grown in gardens and naturalised in hedges, scrub, banks and rough ground. In scattered localities in central and south Britain and in the Channel Islands. Has been known at Bere Ferrers in Devon since the 1930s. Native of eastern Asia.

Section 3. Caprifolium (Mill.) DC. Caprifolium Mill.; Periclymenum Mill. Woody climbers. Flowers in terminal heads or whorls. Corolla 2-lipped; tube long. 8. L. periclymenum L. Honeysuckle Periclymenum vulgare Mill. Deciduous woody climber, but often low and trailing or scrambling. Stems up to 6 m, flexuous, more or less striate, glabrous or slightly hairy. Bark pale brown. Branches spreading; twigs pale brown; young shoots pale yellowishgreen, sometimes flushed purplish, with sparse to dense, unequal, pale brownish-yellow glandular hairs, sometimes also with few to numerous, subrigid, pale, long simple eglandular hairs. Leaves distichous; lamina 2–7 × 1.5–4.5 cm, dull medium yellowish-green to dark green on upper surface, paler and often bluish beneath, narrowly to broadly elliptical, sometimes subrotund or oblong, rounded-obtuse to shortly acute at apex, entire, sometimes lobed in shade, rounded or cuneate at base, glabrous or with long, subrigid, pale simple eglandular hairs especially beneath, shortly petiolate or the uppermost sessile; petioles up to 10 mm, usually long-hairy. Flowers 15–25 mm, in diameter, in sessile terminal whorls, more or less fragrant, especially at night; bracts 15–25 × 15–18 mm, ovate, shortly acute at apex, entire, rounded at base, glandular-hairy; bracteoles 2– 3 × 2–3 mm, broadly ovate, acute at apex, glandular-hairy, partly obscuring ovary. Calyx 3.0–4.5 mm, pale green, often tinted reddish, markedly lobed, the lobes narrowly triangular and obtuse to subacute at apex, glandular-hairy. Corolla 30–50 mm, cream flushed purplish, or cream outside and creamy white within, zygomorphic, the tube 20–30 mm, slender, with 4 narrowly elliptical or linear upper lobes, and a single lower lobe divided into 4, ovate, round lobules, with numerous glandular hairs and sometimes some longer simple eglandular hairs also. Stamens 5, exceeding corolla; filaments white; anthers yellow. Style 1, pale green; stigma green, exceeding the anthers. Berry 5–8 mm, globose, dark red. Flowers 6–9. Pollinated by hawk-moths and bumble-bees etc. Fruits 8–9. 2n = 18, 36, 54. (i) Var. periclymenum L. periclymenum var. clarkei Hesl.-Harr.; L. periclymenum var. quercifolia Aiton Young shoot usually only with glandular hairs. Leaves glabrous or with an occasional simple eglandular hair. Corolla usually with reddish colouring mixed with the cream, strongly fragrant. (ii) Var. hirsuta (Rouy) P. D. Sell L. periclymenum subvar. hirsuta Rouy Young shoot with long simple eglandular hairs mixed with the glandular hairs. Leaves with long, pale simple eglandular hairs especially beneath. Corolla usually without reddish colouring, only slightly fragrant. Native. Woods, hedges, scrub, shady rocks and shingle, often not flowering in the shade. Common throughout Great Britain and Ireland. Europe extending northwards to Norway and eastwards to southern Sweden and the Balkans;

3. Knautia north and central Morocco. A member of the Suboceanic Southern-temperate element. Var. periclymenum is the common plant. Var. hirsuta occurs in scattered localities, especially near the coast. It also occurs rarely in France. Its relationship to L. periclymenum subsp. hispanica (Boiss. & Reut.) Nyman; (L. hispanica Boiss. & Reut.; L. periclymenum var. glaucohirta Kunze) from south Spain and Portugal, which hardly differs in its more glaucous underside of the leaf and in being even more hairy, needs further consideration. Both our plant and the plants from south Spain and Portugal lose their hairs slowly with age. More needs to be found out about the distribution of our plant. 9. L. caprifolium L. Perfoliate Honeysuckle Periclymenum italicum Mill. nom. illegit.; Caprifolium hortense Lam.; Caprifolium rotundifolium Moench; L. suavis Salisb.; L. rotundifolia (Moench) Medik.; Caprifolium italicum (Mill.) Medik.; Caprifolium vulgare Medik.; Caprifolium germanicum Delarbre; Caprifolium perfoliatum (R¨ohl.) Gray; L. pallida Host; L. italica (Mill.) Wood; L. perfoliata (R¨ohl.) Edwards; Caprifolium pallidum (Host) Schur; Periclymenum perfoliatum Gray nom. illegit. Deciduous woody climber with spreading habit. Stems up to 6(–10) m, sinuous, with numerous lenticels, glabrous. Bark pale brown. Branches spreading; twigs pale brown; young shoots pale green, glabrous. Leaves distichous; lamina, 3– 10 × 2–8 cm, dull medium green with pale midrib on upper surface, much paler and rather glaucous beneath, broadly elliptical or ovate, rounded or rounded-retuse at apex, entire, the upper pair connate at base, the lower rounded and shortly petiolate, all glabrous. Flowers 25–35 mm in diameter, in sessile, terminal and subterminal whorls, fragrant; bracts 10–25 × 10–30 mm, rounded, entire; bracteoles absent or minute. Calyx 2.5–3.0 mm, green, sometimes tinted reddish, very shallowly lobed, the 5 lobes broadly triangular and more or less acute at apex. Corolla 30–50 mm, pinkish-purple and cream outside, creamy-yellow inside, zygomorphic, the tube 25–40 mm and slender, the upper 4 lobes linear, the lower lobe divided into 4 ovate, rounded lobules, with sparse minute glandular and an occasional simple hair. Stamens 5, not exceeding corolla; filaments pale greenish-white; anthers pale browny-cream. Style 1, greenish-white; stigma green, exceeding the anthers. Berry 7–8 mm, ovoid, red or orange-red. Flowers 5–6. Pollinated by hawk-moths. Fruits 8–9. 2n = 18. Introduced. An established garden escape naturalised in hedges, plantations and rough ground. Scattered localities in Britain north to central Scotland. Known at Cherry Hinton, Cambridgeshire since 1763 and Bagley Wood, Berkshire since 1700. Native of east-central and south Europe westwards to Italy; Caucasus and Turkey. 10. L. × italica Schmidt ex Tausch Garden Honeysuckle L. caprifolium × etrusca Sant. L. × americana auct. Deciduous woody climber with spreading habit. Stems up to 6(–10) m, often procumbent or spreading. Bark pale brown, almost smooth. Branches spreading; twigs pale

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brown, spreading, glabrous; young shoots pale green, sometimes brownish-purple on upper side, with few to numerous, fine simple eglandular hairs. Leaves distichous; lamina 3–7 × 1.7–5.5 cm, dark, rather shiny green on upper surface with a pale midrib, much paler and glaucous beneath, oblanceolate, obovate, elliptical or broadly elliptical, subacute to broadly rounded at apex, entire or sometimes wavy, the upper connate at base, the lower cuneate and shortly petiolate, the veins numerous, all glabrous. Flowers 35–45 mm in diameter, in sessile, terminal and subterminal whorls, fragrant; bracts 10–20 × 10– 30 mm, ovate, obtuse to subacute, entire, connate; bracteoles 1.5–2.0 × 1.0–1.5 mm, minute versions of the bracts and obscuring the base of the ovary. Calyx about 3 mm, yellowish-green, sometimes tinted reddish, unequally 5-lobed, the lobes triangular-ovate and more or less acute at apex. Corolla 45–65 mm, pinkish-purple and cream outside, creamy-yellow inside, zygomorphic, the tube 24–40 mm, slender, with glandular and longer simple eglandular hairs on outside, upper 4 lobes linear, lower lip broadly ovate and divided for one-third of the way to the base into 4 lobules. Stamens 5, exceeding the corolla; filaments white, glabrous; anthers cream. Style 1, white; stigma greenish, exceeding the anthers. Berry 7–8 mm, ovoid, red. Flowers 5–6. 2n = 18. Introduced. One of the most common garden honeysuckles, having mostly replaced L. caprifolium. Naturalised in marginal and rough places. Recorded from Surrey and several places in East Anglia and should be looked for elsewhere as it is overlooked for L. caprifolium. Of garden origin. 8. Kolkwitzia Graebn. Deciduous shrubs. Leaves opposite, simple, sinuate-serrate in upper half, exstipulate. Flowers in a corymbiform cyme at the end of short, leafy twigs; bracts present. Calyx 4lobed. Corolla 4-lobed, actinomorphic. Stamens 4. Style 1, filiform; stigma minutely 3-lobed. Ovary 3-celled, 1 usually empty. Fruit a pair of beaked nutlets, densely strigose, each with 1 seed. A genus of few species in eastern Asia and the Fiji Islands. Named after Richard Kolkwitz (1873–1956). 1. K. amabilis Graebn. Beauty Bush Deciduous shrub. Branches numerous. Twigs pale brown, glabrous; young shoots reddish-brown, with dense, unequal simple hairs and very short glandular hairs. Leaves opposite; lamina 1.5–4.5 × 0.8–2.5 cm, yellowish-green on upper surface, paler beneath, ovate or elliptical, acuminate at apex, subentire to sinuate-serrate in upper half, rounded at base, shortly hairy on both surfaces, particularly on the veins and margins; veins 3–5 pairs, curving upwards; petioles 2–3 mm, hairy. Inflorescence a corymbiform cyme at the end of short, leafy twigs, with the flowers in pairs; bracts about 3 mm, linear or subulate. Calyx 4–5 mm, purplish, divided nearly to the base; lobes 4, linear, acute at apex, often flexuous, hairy. Corolla 11–17 mm, white, flushed with rosepink, actinomorphic, tubular-campanulate, 4-lobed. Stamens 4, the anterior pair adnate for one-third of the tube. Style 1, filiform; stigma minutely 3-lobed. Fruit a pair of

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1 4 6 . VA L E R I A NAC E A E

beaked nutlets, densely strigose, each with 1 seed. Flowers 5–6. 2n = 32. Introduced to gardens in 1901. A persistent garden escape, self-sown on an old wall at Rochester, Kent. Native of Hupeh, China. 145. A D OX AC E A E Trautv. nom. conserv. Perennial rhizomatous herbs. Leaves opposite, 1- to 3ternate, petiolate, exstipulate. Inflorescence a compact, cubical terminal head, 4 flowers lateral and 1 terminal. Flowers actinomorphic, bisexual, half-epigynous. Calyx with 2 lobes in terminal flowers and 3 lobes in lateral flowers. Corolla with 4 lobes in terminal flowers and 5 lobes in lateral flowers, yellowish-green. Stamens 4 in terminal flowers and 5 in lateral flowers, but appearing to be 8 and 10 due to longitudinal division into half-stamens, borne at apex of corolla tube; nectar is secreted in a ring round the base of the stamens. Styles as many as ovary cells; stigmas capitate. Ovary 2- to 5-celled, mostly 4 in terminal and 5 in lateral cells, with 1 ovule per cell. Fruit a rather dry drupe; seeds with copious endosperm. Contains 3 monotypic genera of obscure relationships. 1. Adoxa L. As family. One species only. Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 13: 112–113.

1. A. moschatellina L. Moschatel A. tuberosa Gray nom. illegit. Perennial herb with far-creeping, somewhat swollen rhizomes which have fleshy, white scales at their apices and long, slender stolons. Stems 5–15 cm, dull, pale yellowishgreen, sometimes reddish-tinted below, erect, glabrous. Leaves dull pale green on upper surface, paler and rather glossy beneath, glabrous; basal ternate, 15–35 × 15–30 cm, broadly ovate in outline, the leaflets with lamina 10–30 × 8– 16 mm, ovate in outline, of 3 segments, the terminal obovate, obtuse at apex, with 1–2 obtuse lobes on each side and narrowed at the base, the lateral ovate or obovate with 1–2 obtuse lobes which are cuneate or rounded at base; petiolules 5–25 mm; petioles up to 70 mm, pale green, sometimes pink-tinted; cauline leaves 2, opposite, ternate, the lobes obovate or lanceolate, toothed or trisect, with short petioles expanded at base. Inflorescence 6–10 mm, a compact, cubical, terminal head. Flowers 5, 1 terminal and 4 lateral, 6–8 mm in diameter, faintly smelling of musk. Calyx 1.0–1.5 mm, pale green, glabrous, divided almost to the base with 2 lobes in the terminal flower and 3 in the lateral; lobes lanceolate, obtuse at apex. Corolla 5–6 mm, pale greenish, rotate, divided almost to the base; lobes ovate, obtuse at apex. Stamens 4 in terminal flower, 5 in lateral flowers; filaments short, yellow, divided nearly to base, each fork with a yellow anther. Styles as many as ovary cells, whitish; stigma greenish, capitate. Fruit a rather dry drupe, 4.5–5.5 mm in diameter, green, subglobose, half-enclosed by the adnate calyx which becomes fleshy, rarely produced. Flowers 4–5.

Visited by various small insects; homogamous or slightly protogynous; self-pollination possible. 2n = 36, 54, 72. Easily recognisable by its small, yellowish-green ‘townhall clock’ (one of its vernacular names) flower-head. Native. Woods and hedgebanks, also shady rocky places on mountains, mostly on damp, humus-rich soil alluvium by streams and rivers. Frequent throughout Great Britain except northern Scotland, Co. Antrim and introduced in Co. Dublin in Ireland. Europe, from Scandinavia to the mountains of Spain, Italy, Bulgaria and north Greece; mountains of north-west Morocco apparently extinct; north, central and eastern Asia, Caucasus, Himalaya and North America. A member of the Circumpolar Boreo-temperate element. 146. VA L E R I A N AC E A E Batsch nom. conserv. Annual to perennial herbs. Leaves opposite, simple or pinnate, petiolate or sessile, exstipulate. Inflorescence a terminal, paniculate cyme, often more or less corymbose. Flowers more or less actinomorphic to zygomorphic, unisexual or bisexual, epigynous. Calyx represented by none to many teeth, very small in flower, similar in fruit or developing long feathery appendages. Corolla with 5 lobes, equal or those on abaxial side slightly longer, fused into the tube proximally, the tube straight or slightly pouched or with a long backwardly directed spur at the base. Stamens 1 or 3, borne on the corolla tube. Style 1; stigma 1 and capitate or 3 and linear-oblong. Ovary 3-celled; 1 adaxial cell fertile with 1 ovule; 2 abaxial cells sterile and equally large to vestigial. Fruit a 1-seeded nut; seeds with endosperm. Contains 15 genera and about 400 species nearly cosmopolitan in distribution but best developed in the North Temperate Region and the Andes. 1. Stems forked into 2 at each node; calyx remaining minute 1. Valerianella at fruiting 2. 1. Main stem simple or with lateral branches 2. Leaves green, upper cauline pinnate, pinnatisect or 2. Valeriana sharply and irregularly serrate; stamens 3 2. Leaves glaucous, upper entire or irregularly dentate; 3. Centranthus stamen 1

1. Valerianella Mill. Annual herbs. Stems repeatedly forked, leafy. Leaves opposite, simple, entire to serrate or sparsely lobed. Flowers in rather lax to dense, compound cymes, bisexual. Calyx more or less absent or small, persistent but remaining small on top of fruit, asymmetrical. Corolla tube not pouched or spurred. Stamens 3. Style 1; stigmas 3. Ovary with 1 fertile cell and 2 distant, but sometimes small sterile empty cells. Fruit a 1-seeded nut. About 50 species, one group centred in the Mediterranean and Middle East, extending to north-west Europe, central Asia, Pakistan and Kenya, the other North American. The Old World species all seem to produce a strong and characteristic stale odour when dried which attracts cats, the New World species seem to be odourless. The young leaves in spring and autumn were gathered for salads and are now included in salad mixtures sold in supermarkets. Our species are very similar vegetatively and ripe fruits are needed for identification.

1. Valerianella V. kotschyi Boiss. has been recorded as an esparto casual. FitzGerald, R. (1990). Rare plant survey of south-West England. IV. Dorset. Nat. Conserv. Council Rep. no. 1061 [V. eriocarpa.] Pearman, D. A. & Edwards, B. (2002). Valerianella eriocarpa Desv. in Dorset, and a reassessment of its status as a presumed introduction in Britain. Watsonia 24: 81–89. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [V. dentata.] Wigginton, M. J. (Edit.) British red data books. Vol. 1. Vascular plants. Peterborough. [V. eriocarpa, rimosa.] 1. Calyx in fruit absent or as a very small tooth less than 2. one-tenth as long as the rest of the fruit 1. Calyx in fruit conspicuous, from a quarter as long as to 4. nearly as long as the rest of the fruit 2. Fruit about as thick as wide, much longer than wide or thick, with a deep groove separating the sterile 2. carinata cells 2. Fruit about twice as thick as wide, scarcely longer than 3. thick, with a shallow groove separating the sterile cells 3. Stems usually single or few, (5–)10–40 cm; more or less erect; internodes 35–100 mm; leaves up to 7 cm 1(a). locusta subsp. locusta 3. Stems usually numerous, 2–10 cm, much branched, the stems spreading so that the whole plant forms a ball; internodes up to 35 mm; leaves up to 3.5 cm 1(b). locusta subsp. dunensis 4. Calyx in fruit usually with 6 teeth, more than two-thirds as 5. long as the rest of the fruit and nearly as wide as the fruit 4. Calyx in fruit with less than 6 teeth, often only one, 6. usually less than half as long as the rest of the fruit 4 (i). eriocarpa var. eriocarpa 5. Fruit hairy 5. Fruit glabrous or nearly so 4 (ii). eriocarpa var. glabrescens 6. Main tooth of calyx in fruit scarcely or not toothed; fruit more or less smooth on all faces, with 2– 6 fine grooves and or longitudinal ridges, with easily broken walls 3. rimosa 6. Main tooth of calyx in fruit usually with 2 or more distinct subsidiary teeth; fruit with 2 distinct ribs 7. delimiting an ovate more or less flat area 5(i). dentata var. dentata 7. Fruit glabrous or nearly so 5(ii). dentata var. mixta 7. Fruit hairy

1. V. locusta (L.) Laterr. Common Cornsalad Valeriana locusta L.; Valeriana locusta var. olitoria L.; V. olitoria (L.) Pollich; Fedia olitoria (L.) Gaertn.; Valeriana olitoria (L.) Willd.; Fedia ecalyculata Stokes nom. illegit. Annual herb with fibrous roots. Stems 2–40 cm, pale yellowish-green, rather brittle, weakly angled, erect, ascending or prostrate, repeatedly forked, glabrous or with short simple eglandular hairs in the lower half, leafy. Leaves opposite; lamina 0.6–7.0 × 0.3–1.0 cm, bright medium green on upper surface, paler beneath, the lower broadly spathulate to ovate, upper lanceolate to linear, obtuse to acute at apex, entire to remotely sinuate-dentate, narrowed to a sessile or shortly petiolate base, glabrous. Flowers 1.5– 1.8 mm in diameter, in terminal, capitate, cymose heads and solitary in the fork of branches; bracts green, oblongspathulate, obtuse at apex, entire, with scarious auricles, shortly ciliate. Calyx reduced to a minute tooth above each loculus. Corolla 2.5–3.0 mm, pinkish or bluish, divided

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one-third of the way to the base; lobes 5, subrotund. Stamens 3; filaments about 0.5 mm, white; anthers yellow. Style 1, slender, white; stigmas 3, cream. Fruit 1.8–2.5 mm long, 1.0–1.5 mm wide and 1.0–1.5 mm thick, brown, in dense, hemispherical terminal clusters and solitary ones in the forks of branches, subrotund in side view, compressed and narrowed elliptical in end view, glabrous to minutely hairy, smooth or transversely rugose, fertile cell with outer wall spongy, more or less as thick as rest of cell; sterile cells about as large as fertile cell without the spongy wall, scarcely grooved between them; calyx inconspicuous. Flowers 4–6. 2n = 16. (a) Subsp. locusta Stems usually single or few, (5–)10–40 cm, more or less erect; internodes 35–100 mm. Leaves up to 7 cm. (b) Subsp. dunensis (D. E. Allen) P. D. Sell V. locusta var. dunensis D. E. Allen Stems usually numerous, 2–10 cm, much branched, the stems spreading so that the whole plant often forms a ball; internodes up to 35 mm. Leaves up to 3.5 cm. Native. Arable and rough ground, bare places in grassland, banks, walls, rocky outcrops, dunes and shingle. Throughout Great Britain and Ireland but much decreased in arable land because of the use of herbicides. Subsp. dunensis is common and sometimes abundant on coastal dunes and shingle. It occurs elsewhere in Europe in similar habitats. The remaining plants in Great Britain and Ireland are referable to subsp. locusta. Other variants occur in Continental Europe including var. oleracea (Schltr.) Breistr., which is a large plant with an obtusely angled fruit of about 4 mm, with prominent spongy thickening on the sterile loculi, and is cultivated for salads.The species occurs in most of Europe, but is rare in the north; Madeira; North Africa; western Asia; introduced in North America. It is a member of the European Temperate element. 2. V. carinata Loisel. Keel-fruited Cornsalad Fedia carinata (Loisel.) Steven; V. carinata var. pubescens auct. Annual herb with fibrous roots. Stems 7–40 cm, pale yellowish-green, rather brittle, weakly angled, erect or ascending, repeatedly forked, glabrous or with short, simple eglandular hairs in the lower half, leafy. Leaves opposite; lamina 0.5–3.0 × 0.3.–0.7 cm, bright medium green on upper surface, paler beneath, obovate-spathulate, lanceolate, oblong or linear, obtuse to acute at apex, entire or the upper one sometimes with a few teeth, narrowed or rounded to the sessile or shortly petiolate base, glabrous or with some simple eglandular hairs. Flowers 1.5–1.8 mm in diameter, in terminal, capitate, cymose heads and solitary in the forks of branches; bracts green, oblong-spathulate, obtuse at apex, entire, with scarious auricles, shortly ciliate. Calyx reduced to an indistinct tooth above the fertile loculus. Corolla 2.5–3.0 mm, pinkish or bluish, divided onethird of the way to the base; lobes 5, subrotund. Stamens 3; filaments about 0.5 mm, white; anthers yellow. Style 1, slender, white; stigmas 3, cream. Fruit 2.0–2.7 mm long, 0.8–1.4 mm wide and 0.8–1.4 mm thick, brown, in dense

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1 4 6 . VA L E R I A NAC E A E

V. locusta (L.) Laterr.

V. carinata Loisel.

V. rimosa Bastard

V. eriocarpa Desv. var. eriocarpa

V. dentata (L.) Pollich var. mixta Dufour

V. dentata (L.) Pollich var. dentata Fruits of Valerianella

1. Valerianella hemispherical terminal clusters and solitary ones in the fork of branches, oblong-ovoid, nearly quadrangular in section, glabrous or minutely hairy, smooth or transversely rugose; fertile cell without spongy wall; sterile cells slightly smaller than fertile one, with a deep groove between them; calyx inconspicuous. Flowers 4–6. 2n = 16. Native. Arable land, waste ground, banks, old walls and rocky outcrops. Widespread in southern England and Wales, and the Channel Islands; introduced in south and east Ireland. Central and south Europe and south-west Asia extending eastwards to Iran; North Africa. A member of the European Southern-temperate element. It has apparently increased in recent years and in some counties (e.g. Dorset) it is now the commonest species in the genus. 3. V. rimosa Bastard Broad-fruited Cornsalad V. auricula DC.; Fedia auricula (DC.) Mert. & W. D. J. Koch; V. rimosa var. dasycarpa auct. Annual herb with fibrous roots. Stems 7–30(–50) cm, pale yellowish-green, rather brittle, weakly angled, erect or ascending, repeatedly forked, rather rough on the angles, leafy. Leaves opposite; lamina 1.0–5.0 × 0.3–1.0 cm, bright medium green on upper surface, paler beneath, the lower ovate-spathulate, the median elliptical-oblong, the upper oblong or linear, obtuse to acute at apex, entire or the upper with narrow teeth near the base, narrowed to a sessile or shortly petiolate base, glabrous or with short simple eglandular hairs. Flowers 1.5–1.8 mm in diameter, in terminal, rather lax, cymose heads and solitary in forks of branches; bracts green, linear, obtuse at apex, entire. Calyx usually scarcely toothed the tooth over the fertile loculus usually obtuse and entire. Corolla 2.5–3.0 mm, pinkish or bluish, divided one-third of the way to the base; lobes 5, subrotund. Stamens 3; filaments about 0.5 mm, white; anthers pale yellow. Style 1, slender, white; stigmas 3, cream. Fruit 1.5–2.5 mm long, 1.5–2.5 mm wide and 1.0–1.5 mm thick, brown, in numerous, small, terminal fasciculate clusters and solitary ones in the forks of branches, broadly ovoid, more or less smooth on all faces, with 2–6 fine grooves and/or longitudinal ridges, with easily broken walls, glabrous; fertile cell without a spongy wall; sterile cells slightly larger than fertile cell, scarcely grooved between them; calyx conspicuous, 0.5–1.0 mm. Flowers 6–8. 2n = 16. Native. Cornfields and rough ground on chalk and limestone. Very local in southern England and Ireland; formerly widespread in Great Britain north to central Scotland and Ireland. The decrease is presumably due to the use of herbicides. Europe from Great Britain and Ireland and Denmark southwards, but scarce in the Mediterranean region and absent from many of the islands; Caucasia; North Africa. A member of the European Temperate element. 4. V. eriocarpa Desv. Hairy-fruited Cornsalad Fedia eriocarpa (Desv.) Rchb. Annual herb with fibrous roots. Stems 7–45 cm, pale yellowish-green, rather brittle, weakly angled, erect or ascending, repeatedly forked, rather rough on the angles, leafy. Leaves opposite; lamina 1.0–3.5 × 0.3–0.6 cm, bright medium green on upper surface, paler beneath, linear, linear-lanceolate, lanceolate or oblong, obtuse to acute at

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apex, entire or occasionally with 1–2 teeth towards the base, narrowed to a sessile or shortly petiolate base, glabrous. Flowers 1.0–1.3 mm in diameter, in terminal, dense, cymose heads and solitary in the forks of branches; bracts green with narrow, scarious margins, the lower linear-spathulate and obtuse at apex, the upper narrowly triangular and acute at apex. Calyx obliquely coroniform, almost as wide and long as the fruit, with 6, usually subequal teeth. Corolla 2.0–2.5 mm, pale pink or lilac, divided about one-third of the way to the base; lobes 5, obovate, rounded at apex. Stamens 3; filaments about 0.3 mm, white; anthers yellow. Style 1, slender, white; stigmas 3, cream. Fruits 1.5–2.0 mm long, 0.7–1.2 mm wide and 0.6–0.8 mm thick, in numerous, small, fasciculate terminal clusters and solitary ones in the forks of branches, narrowly ovoid, usually more or less hairy, strongly net-veined; fertile cell without spongy wall; sterile cells reduced to ridges separated by a flat, ovate area; calyx very conspicuous. Flowers 4–7. 2n = 16. (i) Var. eriocarpa Fruit hairy. (ii) Var. glabrescens Cosson Fruit glabrous. Native or introduced. Locally frequent in thin soils over limestone and chalk in coastal habitats in Dorset; naturalised on banks, old walls and rough ground elsewhere in a few, scattered localities in southern England and the Channel Islands. Native of southern Europe and North Africa. The two varieties have been included mainly to show that V. eriocarpa and V. dentata cannot be distinguished on fruit hairiness. 5. V. dentata (L.) Pollich Narrow-fruited Cornsalad Valeriana locusta var. dentata L.; Fedia dentata (L.) Mirb.; Valeriana dentata (L.) All.; Fedia morisonii Spreng.; V. morisonii (Spreng.) DC. Annual herb with fibrous roots. Stems 7–30(–50) cm, pale yellowish-green, rather brittle, weakly angled, erect or ascending, repeatedly forked, rather rough on the angles and sometimes with short, deflexed simple eglandular hairs, leafy. Leaves opposite; lamina 1.0–5.0 × 0.3–2.0 cm, bright medium green on upper surface, paler beneath, ovatespathulate, lanceolate, oblong or linear, rounded-obtuse to acute at apex, entire to sinuate or slightly lobed near the base, narrowed to a sessile or shortly petiolate base, glabrous or with short simple eglandular hairs. Flowers 1.0–1.3 mm in diameter, in terminal, rather lax, cymose heads and solitary in the forks of branches; bracts green, with narrow, scarious margins, the lower linear-spathulate and obtuse at apex, the upper narrowly triangular and acute at apex. Calyx obliquely truncate, much shorter and narrower than the fruit, with unequal teeth, the tooth over the fertile cell acute. Corolla 2.0–2.5 mm, pale pink or lilac, divided about one-third of the way to the base; lobes 5, obovate, rounded at apex. Stamens 3; filaments about 0.3 mm, white; anthers yellow. Style 1, slender, white; stigmas 3, cream. Fruits 1.5– 2.0 mm long, 0.7–1.2 mm wide and 0.6–0.8 mm thick, in numerous small, fasciculate terminal clusters and solitary in the forks of branches, narrowly ovoid, glabrous or hairy,

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1 4 6 . VA L E R I A NAC E A E

scarcely veined; fertile cell without spongy wall; sterile cells reduced to ridges separated by a flat, ovate area; calyx conspicuous. Flowers 6–7. 2n = 16. (i) Var. dentata Fruit glabrous or nearly so. (ii) Var. mixta Dufour Fruit hairy. Native. Cornfield weed germinating in both spring and autumn, but surviving best in less competitive spring-sown crops. Scattered localities in south and east England and Wales and Ireland, formerly much more common and north to central Scotland. The decline is probably due, as perhaps also in other species, to the regular use of herbicides and the application of nitrogenous fertilisers to highly competitive modern crop varieties. Throughout most of southern and central Europe and through Turkey to the western shores of the Caspian Sea; North Africa. A member of the European temperate element. Var. mixta is probably the commoner variant in Britain and Ireland. 2. Valeriana L. Perennial dioecious or monoecious herbs, mostly with a bitter taste and peculiar smell, particularly evident when dry. Stems not repeatedly forked. Leaves opposite, simple and pinnate on same plant. Flowers in rather dense compound cymes, bisexual or unisexual. Calyx inrolled in flower, developing long, feathery projections when fruiting. Corolla tube not or slightly pouched at the base; 5-lobed. Stamens 3. Style 1; stigmas 3. Fruit a unilocular, 1-seeded nut, the sterile cells of ovary scarcely discernible. About 200 species in Europe, Asia, Africa and America. Alston, A. G. H. (1950). Abstract from Walther, F. (1949) see below. Watsonia 1: 379–380. Grime, J. P. et al. (1988). Comparative plant ecology. London. [V. officinalis.] Hult´en, E. (1958). The amphi-atlantic plants and their phytogeographical connections. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 7: 68–69. [V. dioica.] Walther, E. (1949). Zur Morphologie und Systematik des Arzneibaldrians in Mitteleuropa. Mitt. Th¨ur. Bot. Ges. Beihelt. 1: 7–105. 1. Basal and cauline leaves pinnate with several lateral 2. leaflets as large as terminal one 4. 1. At least some of the basal leaves simple 2. Plants without stolons; stems hairy below; middle cauline leaves with 15–27, linear to oblong, entire leaflets, the terminal not wider than the middle lateral leaflets 1(a). officinalis subsp. collina 2. Plant with both epigeal and hypogeal stolons; stems glabrous; middle cauline leaves with 5–9, lanceolate to ovate, dentate leaflets, the terminal distinctly wider than 3. the middle lateral leaflets 3. Stems up to 50 cm; leaflets up to 10 mm wide 1(b). officinalis subsp. dunensis 3. Stems up to 200 cm; leaflets usually much wider, up to 1(c). officinalis subsp. sambucifolia 60 mm 3. pyrenaica 4. Basal leaves dentate, cordate at base 5. 4. Basal leaves entire, rounded or cuneate at base

5. Stem up to 1.5 m, rhizomatous, monoecious; lamina of 2. phu basal leaves more than 3 cm; corolla white 5. Stem up to 40(–50) cm, rhizomatous and stoloniferous, dioecious; lamina of basal leaves less than 3 cm; corolla 4. dioica pink or white

1. V. officinalis L. Common Valerian Perennial herb with a short, simple, not very stout rhizome, sometimes with stolons. Stems 20–200 cm, medium green, erect, robust, striate, channelled, sometimes with short, pale simple eglandular hairs in the lower half or glabrous, leafy, branched above. Leaves opposite; lamina 5–20 × 3–15 cm, fresh medium green on upper surface, paler beneath, gradually decreasing in size upwards, the lower pinnate, obovate, oblanceolate or elliptical in outline and acute at apex, the leaflets 3–25, with the lamina 18–90 × 3–40 mm, ovate to lanceolate, linear-lanceolate or oblong, more or less acute at apex, coarsely dentate to entire and with petioles up to 25 cm, the uppermost linear or lanceolate, acute at apex, sparsely dentate or entire and sessile, all glabrous. Flowers 4–5 mm in diameter, bisexual, in compact cymes, clustered at the top of the stems and from branches in the axils of the upper leaves; pedicels short, glabrous; bracts lanceolate or linear-lanceolate, acute at apex. Calyx involute during flowering, spreading out to form a deciduous pappus in fruit of numerous, erect, incurved, plumose rays. Corolla 3–8 mm, white or tinged pink or lilac, divided halfway to the base; lobes 5, obovate, rounded at apex. Stamens 3; filaments 2.5–3.0 mm, white; anthers yellow. Style 1, white; stigmas 3, cream. Nut 2–5 mm, brown, ovoid, hairy or glabrous; pappus 3.5–4.0 mm, plumose. Flowers 6–8. The following 3 subspecies are easier to understand if populations are examined rather than individual plants, as they seem to occur in different habitats. The diploid subsp. officinalis has not been recorded for Great Britain and Ireland. (a) Subsp. collina (Wallr.) Nyman V. collina Wallr.; V. officinalis var. mikanii auct. Plants without stolons. Stems hairy below. Middle cauline leaves with 15–27, linear to oblong, entire leaflets, the terminal not wider than the middle lateral leaflets. Inflorescence sometimes with lower branches. Corolla 3–6 mm. Nut 2–4 mm. 2n = 28. (b) Subsp. dunensis P. D. Sell Plants with stolons. Stems hairy or glabrous, up to 50 cm. Middle cauline leaves with 7–13 narrowly lanceolate, denticulate leaflets, the terminal slightly wider than the middle lateral leaflets. Inflorescence rarely with lower branches. Corolla 5–6 mm. Nut 3–4 mm. (c) Subsp. sambucifolia (J. C. Mikan ex Pohl) Hayw. V. sambucifolia J. C. Mikan ex Pohl; V. officinalis var. latifolia auct. Plant with both epigeal and hypogeal stolons. Stems hairy or glabrous, up to 200 cm. Middle cauline leaves with 5–9, lanceolate to ovate, dentate leaflets, the terminal distinctly wider than the middle lateral leaflets. Inflorescence often with lower branches. Corolla 4–8 mm. Nut 4–5 mm. 2n = 56.

2. Valeriana Native. Dry and damp, grassy places, rough ground and dune slacks. Frequent throughout much of Great Britain and Ireland. Subsp. collina occurs in the drier habitats, particularly boulder clay woods, subsp. sambucifolia in the wetter ones. Subsp. dunensis occurs in dune hollows between Birkdale and Ainsdale in Lancashire and on the Bar at the Culbin Sands in Nairn. D. E. Allen has written on a herbarium sheet that the Lancashire plant is a dwarf form uniform over several miles of dunes. It should be looked for in other dune localities. Its chromosome number is not known, but it is likely to be tetraploid. A variable polyploid complex, found in northern and central Europe, south to the mountains of Spain, Italy and north Greece and eastwards in Asia to Japan; naturalised in North America. The complex is a member of the Eurasian Boreo-temperate element. 2. V. phu L. Turkey Valerian Perennial herb with a thick, woody, cylindrical, usually simple rhizome producing cylindrical roots, from the lower surface. Stem up to 1.5 m, hollow, subterete to smooth, glabrous except for short hairs at the base and nodes, branched above, leafy. Leaves opposite; lamina medium green on upper surface, paler beneath, glabrous to very shortly hairy; basal (3–)6–8(–10) × 1.0–2.5 cm, simple or compound, the simple ones oblong-ovate, subacute at apex, entire, rounded or cuneate at base, the compound ones if present variously pinnatisect, the terminal lobes similar to simple leaves and subpetiolulate, the lateral lobes 2–5 × 1–5 cm, oblonglanceolate, sessile and frequently decurrent, on long petioles 5–15 cm; lower cauline leaves similar to compound basal ones and amplexicaul; upper cauline leaves pinnatisect to subpinnatifid, the lobes lanceolate to filiform, amplexicaul. Flowers 4–5 mm in diameter, in compact cymes, clustered at the top of the stems and in the axils of the upper leaves; pedicels short, glabrous; bracts 2–3 mm, triangularlanceolate, acute at apex. Calyx involute during flowering, spreading out to form a deciduous pappus in fruit of numerous, incurved, plumose rays. Corolla 8–9 mm, white, divided about halfway to the base; lobes 5, oblanceolate, rounded at the apex. Stamens 3; filaments 4–5 mm, white; anthers yellow. Style 1, white; stigmas 3, cream. Nut about 4–5 mm, brown, lanceolate, hairy between the veins; pappus 3–4 mm. Flowers 6–8. Introduced. Long naturalised on roadsides at Mydroilin in Cardiganshire, where it still persists. Native of Turkey where it is very rare; widespread in Europe but thought to be introduced. 3. V. pyrenaica L. Pyrenean Valerian Perennial herb with a short, stout, simple rhizome. Stems 70–110 cm, dark green, striate, erect, with short, pale simple eglandular hairs at the nodes, leafy. Leaves opposite; lamina dark green on upper surface, paler beneath, glabrous; basal with lamina 8–20 × 8–20 cm, ovate or subrotund, rounded at apex, deeply and irregularly dentate, cordate at base, the petioles pale green and up to 20 cm; cauline in opposite pairs, the lower similar to basal, the middle and upper ovate, acute at apex, sharply and irregularly serrate, often with 2 pairs of linear or lanceolate, acute, toothed lobes at the base, the petioles short and glabrous. Flowers bisexual,

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4–5 mm in diameter, in compact cymes clustered at the top of the stem and from the axils of the uppermost leaves; pedicels short, glabrous; bracts linear or linear-lanceolate, acute at apex. Calyx involute during flowering, spreading out to form a deciduous pappus in fruit of numerous, erect, incurved, plumose rays. Corolla 6–7 mm, pink, divided about half way to the base; lobes 5, elliptical or obovate, rounded at apex. Stamens 3; filaments 2.0–2.5 mm, white; anthers yellow. Style 1, white; stigmas 3, cream. Nut 4.5– 6.0 mm, ovoid, glabrous; pappus 4.5–5.0 mm. Flowers 6–7. 2n = 16. Introduced. Damp woods and shady hedgebanks. Naturalised in north and west Britain from east Cornwall to Aberdeenshire, frequent in south and central Scotland, rare in west and north-east Ireland. Native in the Cordillera Cantabrica, Pyrenees and one station in south-east Spain. 4. V. dioica L. Marsh Valerian V. sylvestris Gray, non Groschke Dioecious perennial herb with a creeping, usually unbranched rhizome and long stolons. Stems 10–40(–50) cm, pale green, erect, striate, with short, pale simple eglandular hairs at the nodes, leafy, branched above. Leaves opposite, medium yellowish-green on upper surface, paler beneath, glabrous; basal and those of stolons with lamina 2–5 × 1–3 cm, ovate, oblong or elliptical, more or less obtuse at apex, entire, cuneate or rounded at base and petiolate; cauline in opposite pairs, gradually decreasing in size, obovate or oblanceolate in outline, obtuse at apex, pinnate, the terminal leaflet ovate or lanceolate, entire and cuneate at base, the lateral leaflets linear or linear-lanceolate, obtuse at apex and entire or with a few teeth and sessile. Male flowers 4–5 mm in diameter, female flowers 1.8–2.2 mm in diameter, in a compound, compact cyme, terminal or at the end of short branches in the angles of the upper leaves, those of female smaller and tighter than those of the male; pedicels short, glabrous; bracts linear or linear-lanceolate, acute at apex. Calyx involute during flowering, spreading out to form a deciduous pappus in fruit with numerous spreading plumose rays. Corolla of male flower 4–6 mm, pink, divided about halfway to the base with lobes 5, obovate and rounded at apex; the female flowers 4–5 mm, white, divided about half way to the base; the lobes 5, obovate and rounded at apex. Stamens 3; filaments 2.0–2.5 mm, white; anthers yellow. Style 1, white; stigmas 3, cream. Nut 2.5–3.0 mm, brown, ellipsoid, glabrous; pappus 4.5–5.0 mm. Flowers 5–6. 2n = 16, 32. Native. Marshes, fens and bogs, often in shade. Frequent in Great Britain north to central Scotland. From southern Norway and southern Sweden to north Spain eastwards to Macedonia and the western borders of Russia; central Asia. Also in North America as subsp. sylvatica (Rich.) F. G. Mey. Our plant is subsp. dioica which occurs throughout the European range of the species. A member of the European Temperate element. 3. Centranthus DC. Kentranthus orthogr. var. Annual or perennial herbs. Stems not repeatedly forked. Leaves opposite, simple, entire to deeply pinnately lobed

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1 4 7 . D I P S AC AC E A E

or more or less pinnate. Flowers in dense, compound cymes, bisexual. Calyx inrolled in flower, developing long, feathery projections when fruiting. Corolla tube with a backwardly directed spur. Stamen 1. Style 1; stigma 1. Fruit a 1-seeded nut, the sterile cells of the ovary scarcely discernible. Nine species in Europe and the Mediterranean. 1. C. ruber (L.) DC. Red Valerian Valeriana rubra L.; C. marinus Gray nom. illegit. Erect, rhizomatous perennial herb with a branched rootstock and fleshy roots. Stems 30–80 cm, weakly branched above, hollow, pale glaucous, finely striate, usually somewhat thickened at the nodes, glabrous. Leaves opposite; lamina 3–8(–12) × (0.5–)1–5(–6) cm, glaucous, rather fleshy, lanceolate to ovate, obtuse to acute or sometimes acuminate at apex, entire or the uppermost irregularly dentate, glabrous, the lower narrowed to a short, flattened petiole, the upper broadly cuneate or rounded at the more or less sessile, amplexicaul base. Flowers 4–5 mm in diameter, arranged in 2 or more approximate or distant pairs of crowded cymes, subtended by bracts; bracts 2–10 × 1– 3 mm, subulate or narrowly oblong, glabrous or sparsely hairy. Calyx about 1.5 × 0.5 mm, tubular, the teeth involute and hardly visible. Corolla 7–10 mm, scented, pale or deep reddish-pink or white, with a very slender tube produced basely into a slender, pointed spur (2–)5–10(–12) mm; lobes 5, spreading, subequal, oblong, obtuse at apex. Stamen 1, inserted at the mouth of the corolla tube; filaments about 3 mm; anthers about 2 × 1.3 mm, yellow, oblong. Style 1, about 12 mm, slender; stigma 1, subclavate, obscurely lobed. Nut about 3.5 × 1.5 mm, dull brown, narrowly ovoid; pappus about 4.5 mm, of white, plumose hairs. Flowers 6–8. Protandrous and cross-pollinated by long-tongued insects, mainly Lepidoptera. 2n = 32. Introduced. Frequently cultivated and naturalised on old walls, cliffs, dry banks, chalk pits and waste places. Common in the Channel Islands and south and south-west Great Britain extending to the northern part of Ireland and central Scotland. Native of the Mediterranean region and western Asia; cultivated widely and now well naturalised in western Europe outside its native range. All our plants seem to be referable to subsp. ruber. 147. D I P S AC AC E A E Juss. nom. conserv. Biennial or perennial herbs. Leaves opposite, simple to pinnate, petiolate or sessile, exstipulate. Inflorescence a terminal or axillary capitulum, subtended by involucral bracts, the ovary and fruit enclosed in a tubular epicalyx (involucel). Flowers weakly zygomorphic, bisexual or gynodioecious, epigynous. Calyx small, cup-shaped or divided into 4–8 teeth or bristles, arising from the top of the ovary outside the corolla and often less conspicuous than the epicalyx (involucel) which arises at the base of the ovary but encloses the latter in a tubular structure and often expands into lobes around the calyx. Corolla of 4–5 lobes, fused into a tube proximally, the lobes usually larger on the abaxial side, especially in the outer flowers. Stamens 4, free, more or less exserted, borne on the corolla tube. Style 1; stigma

2-lobed, or simple and capitate or oblique. Ovary 1-celled, with 1 ovule. Receptacle usually with a scale associated with each flower within the capitulum. Fruit an achene, which when ripe remains enclosed in the epicalyx; seeds with endosperm. Contains 7 genera and about 350 species native to Eurasia and Africa. 1. Stems and usually the midribs on the underside of the 1. Dipsacus leaves prickly 2. 1. Stem and leaves not prickly 2. Corolla 5-lobed; epicalyx (involucel) expanded at the 5. Scabiosa apex into a membranous, veined funnel 2. Corolla 4-lobed or 4-angled; epicalyx (involucel) 3. variously expanded at apex but not membranous 3. Calyx without bristles and scarcely toothed; corolla cream 2. Cephalaria 3. Calyx with 4–8 teeth or bristles; corolla blue to purple or 4. violet, rarely white or pinkish 4. Receptacle without scales at the base of each flower; calyx with 8 bristles; outer flowers much longer than 3. Knautia inner ones 4. Receptacle bearing scales, one subtending each flower; calyx with 4–5 bristles; flowers all more or less equal in 4. Succisa size

1. Dipsacus L. Virga Hill; Galedragon Gray nom. illegit. Biennial herbs. Stems erect, prickly, leafy. Leaves opposite, simple or pinnate, underside of midrib usually prickly. Capitula solitary at the ends of stems or branches, surrounded by very long spiny bracts. Receptacle with spinetipped scales subtending each flower. Flowers all more or less of one size. Epicalyx (involucel) 4-angled, scarcely toothed at the apex. Calyx cup-shaped, scarcely toothed. Corolla 4-lobed. Stamens 4. Style 1; stigma oblique, entire. Fruit a 4-angled achene, crowned by the calyx at least in the early stages of maturity. About 15 species in Europe, western Asia and North Africa. The name Dipsacus originates from Greek words associated with thirst, referring to the cups made by the connate cauline leaves in which water accumulates. Campbell, W. D. (1993). Dipsacus laciniatus. B.S.B.I. News 63: 30–31. Reynolds, S. C. P. (1997). The putative hybrid between two teasels, Dipsacus fullmum L. and D. sativus (L.) Honck. (Dipsacaceae) in Dublin (v.c. 21). Watsonia 21: 285–289. Ryder, M. L. (1996). Is the Fuller’s Teasel (Dipsacus sativus) really a distinct species. The Linnean 11(4): 21–27. 1. Upper cauline leaves shortly petiolate, not fused in opposite pairs; capitula globose; involucral bracts and 2. receptacular scales similar 1. Upper cauline leaves sessile, connate in pairs; capitula ovoid to ovoid-cylindrical; involucral bracts and 3. receptacular scales dissimilar 2. Capitula 12–25 mm in diameter; receptacular scales 4. pilosus 10–12 mm 2. Capitula 20–50 mm in diameter; receptacular scales 5. strigosus 15–25 mm

1. Dipsacus 3. Receptacular scales equalling the flower, with a recurved, 2. sativus rigid apical spine 3. At least the lower receptacular scales exceeding the 4. flowers, with a straight, flexible apical spine 4. Cauline leaves crenate-serrate to entire; involucral bracts 1. fullonum long-linear or narrowly linear-lanceolate 4. Cauline leaves laciniate or pinnatifid; involucral bracts 3. laciniatus lanceolate-subulate

1. D. fullonum L. Wild Teasel D. sylvestris Huds.; D. fullonum subsp. sylvestris (Huds.) P. Fourn. Biennial herb with a stout, yellowish tap-root. Stems 50– 200 cm, bright pale yellowish-green, sometimes brownishpurple, erect, stout, angled, furrowed, with many hooked prickles on the angles, glabrous, branched, leafy. Leaves opposite; rugose and medium yellowish-green with pale margins on upper surface, paler and sometimes tinted reddish beneath; basal lying flat on the ground in a rosette, dying early in the second season, the lamina 15–60 × 3– 6 cm, oblong to elliptical-oblanceolate, obtuse or acute at apex, entire to crenate, narrowed at base and subpetiolate, with scattered, short, markedly swollen-based, pale prickles, veins very prominent beneath, glabrous; cauline in pairs, each pair connate at the sessile base into a water-collecting cup; lamina 5–34 × 1–13 cm, ovate-lanceolate, lanceolate or linear-lanceolate, acute at apex, crenate-serrate or the uppermost entire, narrowed at base, often ciliate, with sharp prickles only on the prominent midrib beneath. Capitula (15–)40–90 mm, ovoid-cylindrical, at the ends of stems or branches; peduncles long, angled, with numerous prickles on the angles. Involucral bracts 10–130 mm, of unequal length, with the longest equalling or exceeding the capitulum, long-linear or narrowly linear-lanceolate, strongly curved upwards, acute at apex, with numerous prickles. Receptacle narrowly ellipsoid; scales 7–30 mm, exceeding the flowers, pale green, oblong-oblanceolate or oblongobovate, long spine-tipped at apex, and long-ascending ciliate. Involucel 4.5–5.5 mm, pale green, 4-angled, 8-ribbed, with strong ribs at the corners and weaker ribs up the centre of each face, scarcely toothed at apex. Calyx about 2 mm, with a short tube and a wide, 4-angled, very shortly hairy cup which has a ciliate, shallowly lobed margin, usually falling before the fruit matures. Corolla 12–13 mm, pinkishpurple to lilac, with 4, short, unequal, ovate, obtuse lobes at apex. Stamens 4; filaments 4.5–5.0 mm, pale pink or lilac; anthers pink or lilac, exserted. Style 1, pale pink or purple; stigma pink or lilac. Achene about 3.7 × 1.1 mm, pale brown, 4-angled, with one longitudinal ridge in the middle of each side, appressed-hairy, crowned by the calyx when young. Flowers 7–8. Visited by bees and long-tongued flies. 2n = 18. Native. Rough ground, grassland, waysides, railway banks, stream banks, woods, thickets, field margins and banks by the sea, especially where heavy soil has been disturbed. Frequent in the Channel Islands and Great Britain north to central Scotland, local in Ireland. South, west and central Europe eastwards to central Russia and Turkey; western Asia; North Africa. A member of the European Temperate element.

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× laciniatus = D. × fallax Simonk. In this hybrid the leaf-cutting and involucral bracts are halfway between those of the parents. Growing with both parents in Oxfordshire. Also occurs in central Eruope. × sativus This hybrid has fewer stem prickles that D. fullonum, cauline leaves with prickles on the underside of the midrib, the longer involucral bracts shorter than the inflorescence, receptacular scales slightly recurved at the tip and achenes about 4.6–1.8 mm, with 1, rarely 2 longitudinal ridges on the middle of each side and dark brown with appressed hairs. Recorded rarely on waste ground in Bedfordshire, Somerset and Co. Dublin. Occurs widely in Continental Europe. 2. D. sativus (L.) Honck. Fuller’s Teasel D. fullonum var. sativus L.; D. fullonum subsp. sativus (L.) Thell.; D. fullonum subsp. fullonum auct. Biennial herb with a stout, yellowish tap-root. Stems 50– 200 cm, pale green, erect, stout, angled, furrowed, with fewer, blunter prickles on the angles than in D. fullonum, glabrous, branched, leafy. Leaves opposite, medium green with pale veins on upper surface, paler beneath; basal lying flat on the ground in a rosette, dying early in the second season, the lamina 10–38 × 3–7 cm, oblong-elliptical, oblanceolate-elliptical or oblanceolate, obtuse to acute at apex, entire to crenate, narrowed at base and subpetiolate, glabrous but with scattered, swollen-based prickles; cauline in pairs, each pair connate at the sessile base into a watercollecting cup, the lamina 5–20 × 1–5 cm, linear-lanceolate to oblong, obtuse to acute at apex, usually distinctly and irregularly crenate-dentate, without prickles on the prominent midrib beneath. Capitula 30–90 mm, ovoid, at the ends of stems and branches; peduncles long, angled, with numerous prickles on the angles. Involucral bracts 10–50 mm, of unequal length and spreading so that none overtop the capitulum, lanceolate-subulate, acute at apex, without prickles. Receptacle narrowly ellipsoid; scales 12–14 mm, equalling the corolla, pale green, oblong-lanceolate or oblong-ovate, spinose-ciliate, with a rigid, recurved apical spine. Involucel 3.5–4.5 mm, pale green, 4-angled, 8-ribbed, with strong ribs at the corners and weaker ribs up the centre of each face, scarcely toothed at apex. Calyx about 2 mm, with a short tube and a wide, 4-angled, very shortly hairy cup which has a ciliate, shallowly lobed margin, usually falling before the fruit matures. Corolla 12–14 mm, pinkish-purple, with 4, short, unequal, ovate to oblong, obtuse lobes at apex. Stamens 4; filaments 4.5–5.0 mm, pale pink or purple; anthers pale pink or purple, exserted. Style pale pink or purple; stigmas pink. Achene about 4.3 × 2.0 mm, pale brown, 4angled, silvery-white appressed-hairy, crowned by the calyx when young. Flowers 7–8. 2n = 18. Introduced. Still grown for fulling in Somerset for which it was formerly more widely grown; now a frequent casual from bird-seed on tips and waste ground. Scattered records in the Channel Islands and Great Britain. Origin uncertain, but possibly derived from D. ferox Loisel., a species of Corsica, Sardinia and central Italy. The dried, ripe capitula were used in the dressing of woollen cloths (i.e. by fullers),

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the nap of which was raised by means of the recurved apical spines of the receptacular scales. 3. D. laciniatus L. Cut-leaved Teasel Biennial herb with a stout, yellowish tap-root. Stems 50– 200 cm, pale green, erect, angled, furrowed, covered with slender prickles, glabrous, branched, leafy. Leaves opposite, medium green with pale veins on upper surface, paler beneath; basal lying flat on the ground in a rosette, dying early in the second season, the lamina 10–30 × 3–5 cm, oblong-elliptical, oblanceolate-elliptical or oblanceolate, obtuse to acute at apex, entire to crenate, narrowed at base and subpetiolate, glabrous but with scattered swollen-based prickles; cauline in pairs, each pair connate at the sessile base into a water-collecting cup, the lamina 5–26 × 2–11 cm, ovate in outline, long acute or acuminate at apex, laciniate or pinnatifid, the lobes lanceolate or oblong-lanceolate, acute at apex, toothed or again lobed, with prickles on the midrib beneath. Capitula 40–90 mm, ovoid-cylindrical, at the ends of stems or branches; peduncles long, angled, with numerous prickles. Involucral bracts 20–79 mm, of unequal length with the longest equalling or exceeding the capitulum, lanceolate-subulate, acute at apex, curved upwards, with numerous prickles. Receptacle narrowly ellipsoid; scales 16–20 mm, exceeding the flowers, pale green, lanceolate, with a long spine at the apex. Involucel 4.5–5.5 mm, pale green, 4-angled, 8-ribbed, with strong ribs at the corners and weaker ribs up the centre of each face, scarcely toothed at apex. Calyx about 2 mm, with a short tube and a wide, 4-angled, very shortly hairy cup, which has a ciliate, shallowly lobed margin, usually falling before the fruit matures. Corolla 13–15 mm, pale pink, with 4, short, unequal, ovate, obtuse lobes at apex. Stamens 4; filaments 4.5–5.0 mm, pale pink or purple; anthers pink or purple. Style 1, pale purple or mauve; stigma pink or purple. Achene 4.5–5.0 mm, pale brown, 4-angled, silvery-white appressedhairy, crowned by the calyx when young. Flowers 7–8. 2n = 16, 18. Introduced. By a track at Charlbury Station and tips in Oxfordshire; also reported from roadsides in Middlesex, Surrey and Lancashire, perhaps originating from wild flower seed mixtures. Native of Europe northwards to central France, northern Germany and northern Ukraine; western Asia. 4. D. pilosus L. Small Teasel Virga pilosa (L.) Hill; Galedragon pilosus (L.) Gray Biennial herb with a tap-root. Stems 30–150 cm, bright pale green, erect, angled, furrowed, with sparse, weak prickles on the angles, glabrous, branched, leafy. Leaves opposite medium to dark bright green on upper surface, paler and yellowish-green beneath; basal lying flat on the ground in a rosette, the lamina 4–9 × 2.0–4.5 cm, ovate or oblongovate, acute or shortly acuminate at apex, crenate or crenateserrate, narrowed to subtruncate at base with a long petiole, with pale simple eglandular hairs particularly on the margins and veins beneath and sometimes with prickles on the midrib beneath; cauline 4–20 × 1–10 cm, simple and ovate to narrowly elliptical, acute at apex, entire to crenate-serrate and shortly petiolate, or ternate with a pair of unequal,

elliptical, acute, crenate-serrate leaflets completely free at the base and with a short petiole, with scattered, pale simple eglandular hairs. Capitula 12–25 mm in diameter, at first drooping, then erect, at the end of stems or branches; peduncles long, with numerous weak prickles. Involucral bracts 10–12 mm, unequal, shorter than the flowers, triangularlanceolate, acute at apex and abruptly contracted into a spine-tip, spreading, slightly reflexed or slightly curved upwards, sparsely covered with long, silky simple eglandular hairs. Receptacle narrowly ellipsoid; scales 10–12 mm, equalling the flower, pale green, obovate, abruptly contracted into a straight, spiny point, ciliate with long, silky hairs. Involucel 4.5–5.5 mm, pale green, 4-angled, 8-ribbed, with strong ribs at the corners and weaker ribs up the centre of each face, with numerous minute teeth at apex, hairy becoming glabrous. Calyx about 2 mm, with a short tube and a wide, 4-angled, shortly hairy cup which has a ciliate, shallowly lobed margin, strongly contracted between tube and cup, usually falling before the fruit matures. Corolla 6–9 mm, whitish, with 4, short, unequal, ovate, obtuse lobes at the apex. Stamens 4; filaments 4–5 mm, creamywhite; anthers dark violet. Style 1, pale; stigma lilac. Achene 4–5 mm, brown, 4-angled, oblanceolate, appressed-hairy, crowned by the calyx when young. Flowers 7–8. Sparingly visited by small bees and flies. 2n = 18. Native. Damp places in woods, hedgebanks and by streams and ditches. In scattered localities in England and Wales, especially on chalk or limestone, up to 260 m in Derbyshire. Central Europe from northern Spain, northern Italy, north Balkan peninsula and south Russia, north to Holland, Denmark, Sweden and Hungary; western Asia. A member of the European Temperate element. 5. D. strigosus Willd. Yellow-flowered Teasel Biennial herb with a tap-root. Stems 50–200 cm, pale green, erect, angled, deeply furrowed, the angles with distinctly upward-pointing prickles, glabrous, widely branched, leafy. Leaves opposite, medium green with a paler midrib on upper surface, paler beneath; basal lying flat on the ground in a rosette, or ascending, the lamina 5–17 × 3–8 cm, ovate, acute or shortly acuminate at apex, crenate or crenateserrate, sometimes with 2 pairs of distant leaflets, narrowed at base into a petiole, with pale, stiff simple eglandular hairs on the upper surface and often prickles on the veins beneath and petioles; cauline with lamina 4–21 × 1–8 cm, simple and ovate to narrowly elliptical, acute at apex, entire to crenate-serrate and shortly petiolate or ternate with a pair of unequal, elliptical, acute, crenate-serrate leaflets completely free at the base and with a short petiole, with scattered, pale simple eglandular hairs. Capitula 20–50 mm in diameter, globose, at the ends of stems or branches; peduncles long, with numerous, weak prickles. Involucral bracts (10–)15–25 mm, unequal, shorter than the flowers, triangular-lanceolate, tapering gradually into a spiny point, more or less spreading, sparsely covered with long, silky simple eglandular hairs. Receptacle narrowly ellipsoid; scales 15–25 mm, broadly obovate, tapering more or less gradually into a long, spine-like appendage, sparingly ciliate with long, silky hairs. Involucel 4.5–5.5 mm, pale green, 4-angled, 8-ribbed, with strong ribs at the corners

3. Knautia and weaker ribs up the centre of each face, with numerous minute teeth at apex, hairy at first but becoming glabrous. Calyx about 2 mm, with a short base and a wide, 4-angled, shortly hairy cup which has a ciliate, shallowly lobed margin, usually falling before the fruit matures. Corolla 6–9 mm, pale yellow, with 4, short, unequal, ovate, obtuse lobes at the apex. Stamens 4; filaments 5–6 mm, white; anthers dark violet. Style 1, pale; stigma lilac. Achene 4.0–4.5 mm, greyish-brown with black streaks, 4-angled, oblanceolate, appressed-hairy, crowned by the calyx when young. Flowers 7–8. Introduced. Recorded in the Cambridge area for many years and possibly naturalised in a chalk pit at Cherry Hinton since at least 1828; a rare casual elsewhere in England. Native of southern Russia, the Ukraine, Turkey and Iraq, and naturalised elsewhere in temperate Europe. In Great Britain it was long confused with D. pilosus. 2. Cephalaria Schrad. nom. conserv. Perennial herbs. Stems erect, usually more or less hairy, leafy. Leaves opposite, simple and deeply pinnately lobed to pinnate. Capitula solitary at the end of branches or stems. Involucral bracts in several rows. Receptacle with rather leathery scales subtending each flower. Flowers unequal, outer longer than inner. Epicalyx (involucel)8-ridged, with 8 apical teeth. Calyx cup-shaped, scarcely toothed. Corolla 4-lobed. Fruit an achene, 8-angled, with 4 long, rigid hairs and 4 shorter ones. About 65 species, mainly in southern Europe, the Mediterranean and western Asia, but also in South Africa. C. syriaca (L.) Roem. & Schult. is a rare bird-seed alien or garden escape. 1. C. gigantea (Ledeb.) Bobrov Giant Scabious Scabiosa tatarica M. Bieb., non L.; Scabiosa gigantea Ledeb.; Scabiosa elata Hornem.; C. elata (Hornem.) Schrad.; C. tatarica Roem. & Schult. Perennial herb with a stout tap-root. Stems up to 3.5 m, erect, striate, stout, pale green, with short simple eglandular hairs above and long retrorse ones below, leafy. Leaves opposite, medium yellowish-green on upper surface, paler beneath, lower with lamina 20–40 × 15–20 cm, broadly oblong in outline, lyrate to pinnatisect, divided nearly to midrib, the terminal segment narrowly to broadly elliptical or lanceolate, narrowed to a short acute or acuminate apex, crenateserrate and narrowed at base, the lateral segments 5–7 pairs, oblong or lanceolate, shortly acute at apex, crenate-serrate and narrowed at base, the upper at least connate and decurrent on the midrib, with the petiole very long, with long, pale deflexed simple eglandular hairs; upper pinnatisect to pinnate, similar to lower but smaller and with shorter petioles; with numerous, short to medium, pale, appressed simple eglandular hairs to nearly glabrous on both surfaces. Capitula 40–60 mm in diameter, globose, solitary at the ends of stems or branches; peduncles very long, furrowed, with very short, pale simple eglandular hairs. Involucral bracts in several rows, 10–12 × 3–5 mm, green, ovate, obtuse to acute at apex, with long, white, rather shaggy simple eglandular hairs. Flowers lightly scented, the outer longer than inner. Receptacle with rather leathery, oblanceolate, acuminate,

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hairy scales subtending each flower. Involucel 4–7 mm, with 8 teeth, densely long-hairy. Calyx 1.0–1.5 mm, cupuliform, fimbriate, very hairy. Corolla up to 20 mm, 4-lobed, pale yellow or cream. Stamens 4; filaments and anthers yellow. Style 1, yellow; stigma 2-lobed, yellow. Achene 3–4 mm, 8-angled, with 4, long, rigid hairs and 4 shorter ones at apex, hairy. Flowers 7–8. 2n = 36. Introduced. Grown in gardens and naturalised in rough, grassy places and waste ground. Scattered records in south and central England. Native of stream banks, wet meadows and rocky slopes in northern Turkey and the Caucasus. 3. Knautia L. Trichera Schrad. ex Roem. & Schult. Perennial or biennial herbs. Stems erect, hairy, leafy. Leaves opposite, simple, crenate to pinnate. Capitula solitary or up to 3, at the ends of stems and branches; surrounded by an involucre. Receptacle hairy, but without scales subtending each flower. Flowers unequal, the outer longer than the inner. Epicalyx (involucel)4-ridged, scarcely toothed, densely hairy. Calyx with 8 long bristles. Corolla 4-lobed. Fruit an ovoid to cylindrical achene, with a basal appendage; dispersed by ants. About 50 species in Europe, western Asia and North Africa. 1. K. arvensis (L.) Coult. Field Scabious Scabiosa arvensis L.; Trichera arvensis (L.) Schrad. ex Roem. & Schult.; Scabiosa arvensis var. pinnatisecta auct. Perennial or biennial herb with a sympodial stock and usually with rhizomes. Stems (15–)25–75(–100) cm, pale green, often tinted or spotted purplish especially towards the base, erect, faintly striate, slightly angled, rough with numerous, short, medium and rather long, subrigid, downwardly directed simple eglandular hairs, simple or branched, leafy. Leaves opposite, dull medium green on upper surface, paler beneath; basal in an overwintering rosette from the stock or the base of an old flower stem, the lamina ovate or elliptical, obtuse or acute at apex, entire to crenate or sometimes lyrate-pinnatifid and shortly petiolate; cauline widely spaced and often confined to the lower half of the stem, the lamina 4–25 × 2–8 cm, oblanceolate, obovate or broadly elliptical in outline, acute or obtuse at apex, usually deeply pinnatifid or the upper sometimes, rarely all only crenate-dentate, the terminal segments linear or linear-lanceolate, obtuse to acute at apex and entire or with an occasional tooth, the lateral segments (2–)4–12(–16), linear or linear-lanceolate, obtuse to acute at apex and with an occasional tooth or small lobe and shortly petiolate or sessile; midrib impressed above, prominent beneath; all with more or less numerous, short to long, pale simple eglandular hairs. Plants gymnodioecious, capitula in bisexual plants 30–40 mm in diameter, in female plants (15–)20–30 mm in diameter, the capitula solitary or up to 3 at the ends of branches; peduncles pale green with numerous, long, subrigid, bulbous-based simple eglandular hairs and dense, very short, curled or glandular hairs. Involucral bracts in 2 rows, 9–10 × 4–5 mm, yellowish-green, ovate to ovatelanceolate, subacute at apex, with dense, long, pale, ascending, subappressed simple eglandular hairs. Flowers scented,

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the outer larger than inner. Receptacle with long white hairs, but no scales subtending the flowers. Involucel 3–6 mm, pale green, 4-ridged, scarcely toothed, densely hairy. Calyx 1–2(–3) mm, green, cupuliform, hairy, with about 8, suberect, setaceous teeth about 2–3 mm, which are ciliate below. Corolla pale bluish-lilac, rarely purple, pink or white, those of the marginal flowers 14–15 mm, larger and more unequally 4-lobed than those of the central flowers. Stamens free; filaments 6–8 mm, whitish; anthers lilac. Style 1, whitish; and stigma whitish, tinged lilac. Achene 5–6 mm (without calyx), ovoid to cylindrical, more or less hairy, with a basal appendage. Flowers 6–9. Protandrous, and visited by bees and butterflies. 2n = 20, 40. Very variable in the size of the plant, dissection of the leaves and colour of the flower. Plants with entire or slightly dentate leaves have been called var. integrifolia (Roth) DC. (Scabiosa integrifolia Roth; Scabiosa arvensis var. integrifolia (Roth) Peterm.; Trichera arvensis var. subintegerrima Lange) but intermediates occur and they are found in mixed colonies. Native. Dry rough pasture, especially over limestone and chalk, but also on sand; along hedgebanks, field margins and waste places, and locally on limestone rock-ledges, up to 366 m in Derbyshire. Throughout Great Britain and Ireland, but very local in the north of Ireland and northern and western Scotland. Europe northwards to northern Norway, and in western Siberia; North Africa. A member of the Eurosiberian Temperate element. 4. Succisa Haller Perennial herbs. Stems erect, ascending or prostrate, sparsely hairy, leafy. Leaves opposite, simple, entire or distantly toothed. Capitula solitary at the ends of stems and branches. Flowers all more or less the same size. Receptacle with herbaceous bracts at base and smaller scales subtending each flower. Epicalyx (involucel)4-angled, with 4 teeth. Calyx with 4–5, bristle-tipped teeth. Corolla 4angled. Fruit an elliptical achene, crowned by the persistent calyx. Contains only one species. Grime, J. P. et al. (1988). Comparative plant ecology. London. 1. Stem leaves laciniate-dentate at base 1(v). pratensis var. grandifolia 2. 1. Stem leaves entire or crenate-dentate 1(i). pratensis var. subacaulis 2. Stem up to 7 cm 3. 2. Stem much more than 7 cm 3. Stems prostrate or spreading and turned up at the ends 1(ii). pratensis var. arenaria 2. 3. Stems ascending or erect 4. Large elliptical basal leaves and few small cauline ones; 1(iii). pratensis var. ovalis flowering May to July 4. Large leaves at base getting gradually smaller up the stem; flowering July to October 1(iv). pratensis var. pratensis

1. S. pratensis Moench Devil’s-bit Scabious Scabiosa succisa L.; Scabiosa borealis Salisb. nom. illegit.; S. fuchsii Gray nom. illegit.; S. praemorsa Asch. Perennial herb with short, erect premorse rhizome with long, stout roots. Stems 6–100 cm, prostrate, decumbent,

ascending or erect, pale yellowish-green, often purplish near the base, striate, with short to medium, pale, ascending-appressed simple eglandular hairs. Leaves opposite, medium green with a pale midrib on upper surface, paler and greyish beneath, sometimes slightly suffused purplish; basal and lowest cauline with lamina 1–30 × 1–5 cm, elliptical, oblong, ovate or lanceolate, rounded to subacute at apex, entire or distinctly toothed, rounded, cuneate or attenuate at base, the petioles up to 15 cm, glabrous or with long, pale simple eglandular hairs; remaining cauline few and smaller, in pairs, the lamina ovate or lanceolate or the upper linear, more or less acute, entire, shortly petiolate or sessile; all with medium to long, pale, appressed simple eglandular hairs on both surfaces and the margins. Capitula 2–25 mm in diameter, solitary or few at the ends of branches; peduncles very long, with numerous, short, pale simple eglandular hairs. Involucral bracts in 2–3 rows, 4–10 × 1.5–2.2 mm, green, sometimes suffused dark red, lanceolate or ovate, acute at apex, with numerous medium to long, pale simple eglandular hairs on the margin and minute glandular hairs on the surface. Receptacle elongated, with green, darktipped, oblanceolate, acute, ciliate scales. Flowers scented. Involucel 1.5–3.0 mm, green, 4-angled, with 2 furrows on each face, ending above in 4, erect, green, triangular lobes, densely hairy. Calyx 1.5–2.0 mm, brownish, 4-lobed, cupular, with 4 setaceous teeth. Corolla 4–7 mm, mauve, pink or rarely white, the inner about equalling those of the marginal flowers, but usually smaller in female capitula, mauve to dark bluish-purple, rarely white or pink, hairy. Stamens 4; filaments whitish; anthers reddish-purple. Style 1, cream; stigma cream. Achene 4.5–5.0 mm, elliptical, crowned by the persistent calyx. Flowers 5–10. Protandrous; visited by butterflies and various bees. 2n = 20. There is a variable degree of abortion of anthers in ‘female’ capitula. (i) Var. subacaulis (Bernardin.) P. D. Sell S. subacaulis Bernardin.; Scabiosa succisa var. subacaulis (Bernardin) Rouy Stem up to 7 cm. Leaves numerous, elliptical, obovate or lanceolate. Capitula 10–20 mm in diameter. Flowering July to August. (ii) Var. arenaria (Rouy) P. D. Sell Scabiosa succisa var. arenaria Rouy Stems prostrate or spreading and turned up at the ends. Leaves all similar, 2–5 × 0.2–2.0 cm, elliptical, entire, more or less acute at apex. Capitula 12–15 mm in diameter. Flowering August to October. (iii) Var. ovalis (Rouy) P. D. Sell Scabiosa succisa var. ovalis Rouy Stems ascending or erect. Leaves numerous in a basal rosette, elliptical, entire, cauline few and small. Capitula 15–20 mm in diameter. Flowering May to July. (iv) Var. pratensis Scabiosa succisa var. hispidula auct.; Scabiosa succisa var. glabrata auct. Stems more or less erect. Leaves large at base and gradually getting smaller upwards; entire or distantly toothed. Capitula 15–25 mm in diameter. Flowering July to October.

5. Scabiosa (v) Var. grandifolia (Rouy) P. D. Sell Scabiosa succisa var. grandifolia Rouy Stem very tall, up to 100 cm. Leaves up to 11 × 4 cm, the basal with petioles up to 9 cm, the cauline serrate-dentate and laciniate near the base. Capitula numerous, up to 25 mm in diameter. Flowering July. Native. Acid or calcareous marshes, fens, meadows and pastures, damp woods and cliff-tops. Common over most of Great Britain and Ireland. Europe northwards to 68◦ 21 N; Caucasus; western Siberia; North Africa. A member of the Eurosiberian Temperate element. The common plant is var. pratensis. Var. arenaria is known only from cliff-tops in Cardiganshire and the Mid Etudes. It should be looked for elsewhere in western Britain. It also occurs on the coast of France. Var. ovalis is known only from hay meadows at Pixie Mead in Oxfordshire. It is grown by one wild flower seed company and may be planted elsewhere. It also occurs in France. Var. subacaulis is found in Cornwall, the Outer Hebrides, Shetland and Fair Isle. It is similar in habit to Jasione montana var. nana and has the same sort of distribution. It also occurs in France. Var. grandifolia is known only from a peaty, woodland swamp at Ashurstwood in Sussex. It also occurs in France. 5. Scabiosa L. Columbaria J. & C. Presl. Perennial herbs. Stems erect, hairy, leafy. Leaves opposite, simple, serrate or lobed to pinnate. Capitula solitary at the ends of stems or branches surrounded by an involucre. Receptacle with narrower scales subtending each flower. Flowers unequal, the outer longer than the inner. Epicalyx (involucel)8-ridged, expanded at apex into a membranous funnel. Calyx with 5 long bristles at the top of the stalk. Corolla 5-lobed. Fruit an ellipsoid achene, crowned by the persistent calyx and wind dispersed. About 100 species in temperate Europe and Asia, the Mediterranean region, the mountains of eastern Africa and South Africa. Grimes, J. P. et al. (1988). Comparative plant ecology. London. [S. columbaria.] 1. Flowers dark purple; with scarious, funnel-shaped apex (2a). atropurpurea subsp. atropurpurea with 8 veins 2. 1. Flowers pale lilac, pinkish, bluish, mauve or white 2. Involucel with scarious funnel-like apex with 16–24 veins 1. columbaria 2. Involucel with scarious funnel-like apex with 8 veins (2b). atropurpurea subsp. maritima

1. S. columbaria L. Small Scabious Columbaria minor Gray nom. illegit.; Asterocephalus columbarius (L.) Rchb. Perennial herb with a long tap-root, prolonged upwards into an erect, branching stock. Stem 10–100 cm, pale yellowishgreen, sometimes purplish below, slender, with numerous, pale, short, downwardly directed, appressed simple eglandular hairs. Leaves opposite, 2–12 × 1–6 cm, yellowishgreen on upper surface; paler beneath, basal elliptical, lanceolate or ovate, rounded-obtuse at apex, crenate to pinnatisect and gradually narrowed at base or pinnatifid

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to the midrib with oblong or oblanceolate, obtuse and entire or dentate lobes, shortly petiolate; cauline mainly in the lower half, usually in pairs, the lamina pinnatisect to the midrib, with linear, oblong or very narrowly elliptical, obtuse lobes, the lobes sometimes themselves divided into narrow lobelets, the lower petiolate, the upper sessile, rarely the lower cauline like the undivided basal; all with few to numerous, pale simple eglandular hairs. Capitula solitary to fairly numerous, 15–30 mm in diameter, shallowly domed in flower, ovoid in fruit, solitary at the ends of stems or peduncles; peduncles very long, up to 25 cm, pale yellowish-green, with numerous, pale, downwardly directed, appressed simple eglandular hairs. Involucral bracts in 2 rows, 7–20 × 0.3–1.5 mm, yellowish-green, linear, subacute at apex, with numerous to dense, short, pale simple eglandular hairs. Flowers faintly scented. Receptacle elongated, with pale green, brown-tipped, spathulate or linear-lanceolate scales. Involucel 3.0–4.5 mm, pale green, appressed-hairy, with 8 furrows ending in a more or less scarious, pleated 16–24 veined, funnel-shaped cup. Calyx 0.5–0.8 mm, green, cup-shaped, with 5 brown setaceous teeth 3–6 mm which are minutely hairy. Corolla pinkish, bluish, mauve or white, the outer up to 15 mm, the inner up to 7 mm, hairy, with a short tube and 5 unequal lobes, 2 short and 3 long. Stamens 4; filaments white; anthers white. Style 1, mauve; stigma mauve. Achene 3.0–3.5 mm, pale brown, ellipsoid, appressed-hairy. Flowers 7–8. Homogamous to protandrous and visited by bees and Lepidoptera. 2n = 16. There seems to be every graduation between coastal plants up to 35 cm with pinnate leaves and a solitary capitulum to plants up to 1 m with bipinnate leaves and numerous capitula. Although many subspecies and varieties have been described we cannot equate them to either geography or ecology. Very tall plants with narrow leaf segments planted with wild flower seed in Cambridgeshire appear to be referable to var. patens (Jord.) Car. & St-Lag., described from France. Native. Dry calcareous pastures and banks, chalk and limestone grassland, sandy places near the coast and rocky places up to 610 m in northern England. Locally common in Great Britain north to southern Scotland, but absent from large non-calcareous areas in Wales and the southwestern peninsula. Europe to southern Scandinavia; west Asia; North Africa. A member of the European Temperate element. Plants of Great Britain are referable to subsp. columbaria. 2. S. atropurpurea L. Sweet Scabious Sixalix atropurpurea (L.) Greuter & Burdet Annual or biennial herb with fibrous roots. Stems often numerous, 20–60 cm, pale yellowish-green, sometimes purplish below, with few to numerous, pale, downwardly directed, appressed simple eglandular hairs. Leaves opposite 2–10 × 1–5 cm, dull medium green on upper surface, paler beneath; basal and lower cauline with lamina oblanceolate or oblong-oblanceolate, obtuse to acute at apex, crenate-dentate or incise-dentate to lobed, the terminal lobe large, the lateral narrow and petiolate; middle and upper cauline pinnatifid, with linear or oblanceolate-linear, obtuse, entire or dentate segments, sessile or nearly so; all with few to numerous, short, pale simple eglandular hairs

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1 4 8 . A S T E R AC E A E

inner

ligule 3-lobed

ligule 5-lobed

style

outer

style involucral bracts pappus achene style

ligulate flower

ligulate flower tubular flower

receptacular scale

style

achene

rows of involucral bracts

receptacle

(after Davis et al. Flora of Turkey Vol. 5. 1975) tubular flower

Anthemis Calendula Anthemis

Echinops

Carthamus Carlina Carlina Inula

Eupatorium Echinops

Eupatorium Styles

(after Kåre Bremer 1994) Flowers of Asteraceae

Stamens

148.ASTERACEAE

pappus of simple eglandular hairs

Tragopogon pratensis subsp. minor (Mill.) Wahlenb.

pappus of plumed hairs

Sonchus asper (L.) Hill

pappus a small rim

Arctium lappa L. subsp. minus

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Matricaria recutita L.

pappus of scales

Achillea millefolium L. Galinsoga parviflora L. Achenes of Asteraceae

Leontodon hispidus L.

pappus a rim

Anthemis arvensis L.

pappus of downwardly barbed awns

Bidens tripartita L.

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1 4 8 . A S T E R AC E A E

or nearly glabrous. Capitula solitary to fairly numerous, 30–40 mm in diameter, domed in flower, oblong-ellipsoid in fruit, solitary at the ends of stems or peduncles; peduncles very long, up to 25 cm, pale yellowish-green, with few to numerous, pale, downwardly directed, appressed simple eglandular hairs. Involucral bracts in 2 rows, 5–16 × 0.3– 3.0 mm, yellowish-green, linear, more or less acute at apex, with numerous, short, pale, appressed simple eglandular hairs. Flowers faintly scented. Receptacle elongated, with green-tipped, white-based, oblong-lanceolate scales. Involucel 3.0–5.0 mm, pale green, hairy at base and apex, with 8 furrows, ending in a more or less scarious funnel-like apex with 8 veins. Calyx 0.3–0.7 mm, green, cup-shaped, with 5, red, setaceous teeth 6–8 mm, which are minutely hairy. Corolla dark purple, pinkish or lilac, the outer 12–18 mm, slightly longer than the inner, hairy, with 5 unequal lobes, 2 short and 3 long. Stamens 4; filaments and anthers whitish. Style 1, mauve to purple; stigma mauve to purple. Achene 4.0–4.5 mm, pale brown, ellipsoid. Flowers 6–9. (a) Subsp. atropurpurea Flowers dark purple. (b) Subsp. maritima (L.) Arcangeli S. maritima L.; S. ambigua Ten.; S. ateridoi Pau; S. cupanii Guss.; S. grandiflora Scop.; S. paui Sennen; Sixalix atropurpurea subsp. maritima (L.) Greuter & Burdet Flowers pinkish or lilac. Introduced. Subsp. atropurpurea is much grown in gardens and is naturalised on chalk cliffs and railway banks by the sea in Cornwall, and at Folkestone in Kent where it has been known since 1862. It is a rare casual elsewhere. It is native of Spain and Portugal. The specimen on which C. C. Babington based his record of S. columbaria in Jersey is in fact S. atropurpurea subsp. atropurpurea. Subsp. maritima was present at Les Hanni`eres, St Ouen, Jersey from the 1870s to the 1920s, where it was probably planted. It is native throughout the Mediterranean region.

Order 10. A S T E R A L E S Lindl. Annual to perennial herbs, rarely shrubs. Leaves usually alternate, sometimes opposite, simple and entire to pinnately or palmately lobed or variously pinnately compound, exstipulate. Flowers in dense, terminal heads (capitula), zygomorphic or actinomorphic, bisexual or unisexual, epigynous. Calyx absent or represented by a pappus of scales, teeth, bristles, hairs or a membranous ring. Corolla of (4–) 5 petals fused into a tube with 4–5 teeth at apex (tubular flowers) or with a strap-like ligule (ligulate flowers). Stamens 5, borne on the corolla tube. Ovary 1-celled with 1 ovule. Fruit an achene. Seeds without endosperm. Contains 1 family occurring worldwide. 148. A S T E R AC E A E Dumort. nom. conserv. Family COMPOSITAE Giseke nom. alt. Annual to perennial herbs, often woody at base, rarely shrubs, often with latex or oil-canals. Leaves usually alternate, sometimes opposite, simple and entire to pinnately or

palmately lobed or variously pinnately compound; petiolate or sessile; exstipulate. Inflorescence of a solitary capitulum, or with few to numerous capitula arranged cymosely (sometimes called a capitulescence), often cymose-corymbose. Receptacle of capitulum expanded and concave, flat, convex or elongated-conical. Flowers usually numerous, small, borne on the receptacle in dense terminal heads (capitula), the outer opening first, zygomorphic or actinomorphic, often both in the same capitulum, bisexual or unisexual, epigynous. Capitula bearing round the outside of the flowering zone a series of sepal-like organs (involucral bracts), which also sometimes occur mixed with the flowers (often 1 per flower) on the receptacle (receptacular scales or bristles); flowers may be all tubular (discoid capitulum), or all ligulate (ligulate capitulum), or with a central region of tubular flowers (disc flowers) and an outer region of ligulate flowers (ray flowers forming a radiate capitulum); sometimes the capitulum has very few flowers, rarely only 1, when the capitula are aggregated into large spherical heads (Echinops). Sepals absent or represented by a pappus of scales, teeth, bristles, hairs or a membranous ring, often enlarging in fruit. Petals 5, rarely 4 or absent, fused into a tube with a distal limb of either 5, rarely 4, actinomorphic lobes (tubular flowers) or with a unilateral strap-like ligule often with 3 or 5 apical lobes (ligulate flowers), the tube often extremely short in the latter. Stamens 5, epipetalous, borne on the corolla tube; anthers often sagittate or tailed below and often with terminal appendages and are fused laterally into a cylinder round the style. Ovary inferior, 1-celled, with 1 basal, anatropous ovule. Style 1, usually branched, each branch with a linear, stigmatic surface. Fruit an inferior achene (cypsela), often with a persistent pappus. Over 1,500 genera and 25,000 species (excluding apomicts) occurring throughout the world. The largest family of flowering plants, comprising examples of almost every ecological type and life-form. The combination of the flowers being borne in capitula, 5 stamens with laterally fused anthers and the fruit an achene makes it a unique family. The most common type of flower is the tubular flower, which is usually bisexual, but sometimes male or sterile. The ligulate flower is usually female or sterile, or bisexual. Capitula may have all the flowers bisexual or the inner flowers usually bisexual and the outer flowers female or sterile. Rarely the flowers of the capitulum are all female or all male. Unisexual capitula may be borne on the same plant (monoecious condition) or on different plants (dioecious condition). The arrangement of the anthers and style arms is associated with the mode of pollen presentation. The anthers ripen before the stigmas, and discharge their pollen into the tube formed by the cylinder of fused anthers. At the time this happens the style is short and the style arms pressed together. The style then elongates up the anther tube, from which the pollen is swept by the hairs of the style arms and presented at the apex of the anther tube to any visiting pollinator. Later the style arms separate to expose the stigmatic surfaces. Thus each flower goes through a staminate then a pistillate stage, before finally the style arms may recurve sufficiently for the stigmas to make contact with pollen from the anthers

148. ASTERACEAE of their own flower. Thus self-compatible species may selfpollinate should cross-pollination fail. Measurements of open capitula are usually only approximate as they have to be made in most cases from pressed specimens. Measurements of involucral bracts are made from the base of the involucre to the tip of the involucral bracts and are mostly made from pressed specimens. Baagøe, J. (1978). Taxonomical application of ligule microcharacters in Compositae. Bot. Tidsskr. 72: 125–147. Bentham, G. (1873). Notes on the classification, history, and geographical distribution of Compositae. Jour. Linn. Soc. London (Bot.) 13: 355–458. Bremer, K. (1994). Asteraceae: cladistics and classification. Portland, Oregon. Burtt, B. L. (1961). Compositae and the study of functional evolution. Trans. & Proc. Bot. Soc. Edinb. 39: 216–232. Carlquist, S. (1976). Tribal interrelationships and phylogeny of the Asteraceae. Aliso 8: 465–492. Cronquist, A. (1955). Phylogeny and taxonomy of the Compositae. Amer. Midl. Naturalist 53: 478–511. Cronquist, A. (1977). The Compositae revisited. Brittonia 29: 137– 153. Czapik, R. (1996). Problems of apomictic reproduction in the Families Compositae and Rosaceae. Folia Geobot. Phytotax. 31: 381– 387. Heywood, V. H., Harborne, J. B. & Turner, B. L. (Edits). (1977). The biology and chemistry of the Compositae. 2 vols. London, New York, San Francisco. Hind, D. J. N., Jeffrey, C. & Pope, G. V. (1995). Advances in Compositae systematics. Kew. King, R. M. & Dawson, H. W. (1975). Cassini on Compositae collected from the Dictionnaire des sciences naturelles with an index by the authors. A facsimile reprint of Cassini’s articles on Compositae published in Cuvier, Dict. Sci. Nat. in 3 volumes with precise dates and indication of original pagination. Morton, J. K. (1977). A cytological study of the Compositae (excluding Hieracium and Taraxacum) of the British Isles. Watsonia 11: 211–223. Reveal, J. L. (1997). Early suprageneric names in Asteraceae. Compositae Newsletter 30: 29–45. Wagenitz, G. (1975). Systematics and phylogeny of the Compositae (Asteraceae). Pl. Syst. Evol. 125: 29–46. 1. Shrubs, with new growth arising each year from older, 2. woody stems 15. 1. Herbs, stems sometimes woody at base 3. 2. Outer flowers of capitulum conspicuously ligulate 2. Flowers of capitulum 1, few or many, all tubular or the outer sometimes with very short, ligule-like lobes not or scarcely exceeding the involucral bracts, in Helichrysum 6. beware ligule-like inner involucral bracts 3. Leaves pinnate or pinnately divided nearly to midrib, or if 82. Senecio not then the ligules purple 3. Leaves entire to toothed less than halfway to midrib; 4. ligules white or yellow 4. Leaves dying in winter; clothed with dense, very sticky 51. Dittrichia glandular hairs 5. 4. Leaves evergreen; not sticky with glandular hairs 63. Olearia 5. Ligules white 86. Brachyglottis 5. Ligules yellow 6. Leaves divided more than halfway to midrib; very 7. fragrant when bruised

63

6. Leaves entire to shallowly lobed or toothed; usually not 8. fragrant when bruised 7. Capitula less than 5 mm in diameter, clustered 67. Artemisia 7. Capitula more than 5 mm in diameter, solitary on long 68. Santolina peduncles 8. Stems procumbent, rooting, less than 20 cm; inner involucral bracts spreading in flower, white and ligule-like 48. Helichrysum 8. Stems erect to scrambling, often more than 20 cm; inner 9. involucral bracts not spreading nor ligule-like 9. Leaves with a revolute margin and sometimes 49. Cassinia glandular-hairy 10. 9. Leaves not so 10. Young stems and lower side of leaves densely white- to 11. buff-tomentose 13. 10. Stem and leaves glabrous or nearly so 11. At least the larger leaves remotely sinuate-lobed 86. Brachyglottis 12. 11. Leaves entire or sharply toothed 48. Helichrysum 12. Leaves white-tomentose on upper side 63. Olearia 12. Leaves green on upper side 59. Chrysocoma 13. Leaves linear, less than 5 mm wide 13. Leaves lanceolate to subrotund, more than 10 mm wide 14. 14. Erect shrub; leaves linear to ovate, cuneate at base; 64. Baccharis flowers white 14. Scrambler; leaves palmate, cordate to hastate at base; 85. Delairea flowers yellow 15. Capitula each with 1 flower, the capitula aggregated into 1. Echinops tight, globose heads 15. Capitula each with more than 1 flower, not forming 16. tight, globose heads 16. Male and female flowers in separate capitula on the same plant, the male more apical and with several 17. flowers, the female lower down and with 1–2 flowers 16. Male and female flowers not in separate capitula, or if in separate capitula then on different plants, both with 18. several to many flowers 17. Leaves mostly opposite; fruiting capitula without terminal processes, with short straight spines or with 94. Ambrosia tubercles 17. Leaves alternate; fruiting capitula with 2 prominent terminal processes covered with stiff, hooked bristles 96. Xanthium 18. Capitula with all ligulate flowers, the ligules usually 19. 5-toothed at apex; milky latex usually present 18. Capitula with flowers all tubular or tubular and ligulate, the ligules when present usually 3-toothed, milky latex 45. absent 20. 19. Pappus absent or of scales or a raised rim 19. Pappus, at least in the central flowers of a capitulum of 26. hairs 15. Scolymus 20. Leaves spiny; plant thistle-like 21. 20. Leaves not spiny; plant not thistle-like 22. 21. Pappus of scales 21. Pappus absent or the achene terminating in a raised rim 24. 22. Capitula less than 20 mm in diameter; ligules yellow 20. Hedypnois

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1 4 8 . A S T E R AC E A E

22. Capitula more than 20 mm in diameter; ligules blue or 23. rarely white 23. Most or all leaves at flowering time cauline; capitula sessile or with a short peduncle; receptacle without 16. Cichorium scales 23. Most or all leaves at flowering time basal; capitula solitary on long peduncles; receptacle with scales 17. Catananche 24. Leaves all basal; peduncles much swollen below the 18. Arnoseris capitula 24. Cauline leaves present; peduncles not or scarcely 25. swollen below the capitula 25. Capitula 10–30 mm in diameter; outer involucral bracts 19. Lapsana not enfolding achenes, nor patent in fruit 25. Capitula 4–5 mm in diameter; outer involucral bracts enfolding the achenes, patent and making a stellate 21. Rhagadiolus involucre in fruit 26. Pappus of plumose hairs on at least some of the achenes, 27. that is the hairs having long slender side-branches 26. Pappus hairs simple and smooth or denticulate, that is 33. having minute side-branches 27. Cauline leaves usually absent although scale-like bracts 28. are often present 30. 27. Obvious cauline leaves present 22. Hypochaeris 28. Receptacular scales present 29. 28. Receptacular scales absent 29. Stems with scale-like bracts, often branched; hairs on stem and leaves simple; capitula often more than 1; involucral bracts merging into the scales of the stem and 23. Scorzoneroides peduncles 29. Stems without bracts, unbranched; hairs on stem forked; 24. Leontodon capitula solitary; scales on stem absent 30. At least some of the abundant hairs anchor-shaped 31. (glochidiate) 32. 30. Glabrous or if hairy none of the hairs anchor-shaped 31. Outer involucral bracts lanceolate and similar to inner 25. Picris 31. Outer involucral bracts ovate with a cordate base and 26. Helminthotheca much wider than inner 32. Involucral bracts more than 10, in several rows 27. Scorzonera 28. Tragopogon 32. Involucral bracts about 8, in 1 row 33. Leaves with strong spines; receptacular scales present, 15. Scolymus enclosing achenes; pappus hairs 2–4 33. Leaves not or weakly spiny; receptacular scales usually 34. absent; pappus hairs numerous 35. 34. Achenes distinctly compressed 38. 34. Achenes not or scarcely compressed 35. Achenes with a distinct, narrow beak at apex, or at least 36. markedly narrowed distally 35. Achenes without a beak and scarcely narrowed distally 37. 36. Involucral bracts in several rows; pappus hairs in 2 equal 31. Lactuca rows 36. Involucral bracts in 2 distinct, but unequal rows; pappus 33. Mycelis hairs in 2 unequal rows 30. Sonchus 37. Ligules yellow 32. Cicerbita 37. Ligules blue to mauve 38. Stems without leaves or scales or stolons; capitulum 34. Taraxacum 1 per stem

38. Stems with leaves or scales or if neither with stolons; 39. capitulum often more than 1 39. Pappus of inner achenes of rigid hairs somewhat 36. Tolpis expanded at base 40. 39. Pappus usually of soft hairs not expanded at base 40. Receptacle with long, silky hairs equalling or exceeding 37. Andryala the flowers 41. 40. Receptacle glabrous or with short hairs 42. 41. Pappus hairs pure white 43. 41. Pappus hairs yellowish-white to pale brown 42. Plant with long, thin rhizomes, often bearing whitish, 29. Aetheorhiza subglobose tubers; capitulum 1 per stem 42. Plant without rhizomes; usually more than 1 capitulum 35. Crepis per stem 43. Involucral bracts in 2 rows, the outer shorter than the 35. Crepis inner; achenes narrowed or beaked at apex 43. Involucral bracts in several rows; achenes narrowly 44. obconical and truncate at apex 44. Plant usually stoloniferous; leaves never distinctly petiolate; ligules often with reddish stripe on outer face, sometimes red; achenes less than 2.5 mm, each rib shortly projecting above to form a crenulate apex; pappus hairs mainly in 1 row with a few shorter than the 38. Pilosella rest 44. Plant never stoloniferous; leaves usually distinctly petiolate; ligules yellow or rarely green; achenes 2.5–5.0 mm, the ribs apically confluent into an obscure 39. Hieracium ring; pappus hairs in 2 rows 45. Leaves and involucral bracts often spiny; anthers usually with long tails at base; flowers mostly pink or blue to 46. purple, rarely white, all tubular 45. Leaves and involucral bracts rarely with spines; anthers mostly without basal tails; flowers of various colours, 60. often yellow, the outer ones often ligulate 46. Involucral bracts subulate, stiff and at least some 3. Arctium strongly hooked at apex 47. 46. Involucral bracts not hooked at apex 48. 47. Corolla pale yellow to reddish-orange 51. 47. Corolla pink or blue to purple or mauve, rarely white 6. Cirsium 48. Pappus of plumose hairs 48. Pappus absent, or of narrow scales, or of simple or 49. denticulate hairs 49. Capitula not surrounded by large leaves or large outer 12. Centaurea involucral bracts 49. Capitula surrounded by large leaves or large outer 50. involucral bracts 13. Cnicus 50. Pappus of 2 rows of rigid hairs 14. Carthamus 50. Pappus absent or of linear scales 52. 51. Pappus hairs plumose 51. Pappus absent, of narrow scales, or of simple or 55. denticulate hairs, rarely barbed 52. Inner involucral bracts with a distinct apical portion, sometimes only a spine, but always abruptly delimited 8. Cynara from basal portion 52. Involucral bracts without distinct apical and basal portions, if with apical spine, then gradually narrowed 53. into it 53. Leaves more or less entire to distantly toothed, the teeth not spinose or bristle-like; involucral bracts obtuse to 4. Saussurea rounded at apex

148. ASTERACEAE 53. Leaves spiny or at least with fine bristle-like teeth; at least the outer involucral bracts spinose, mucronate or 54. acuminate at apex 54. Outer involucral bracts leaf-like, the inner ones scarious and spreading in dry weather, pale yellow on inner face 2. Carlina and appearing like ligules 54. All involucral bracts linear-lanceolate to more or less 6. Cirsium ovate 56. 55. Leaves with sharp spines 58. 55. Leaves without spines 56. Receptacle glabrous, but achene pits fringed with teeth 7. Onopordon 57. 56. Receptacle densely hairy or bristly 57. Stem leaves decurrent down the stem in a spiny wing; involucral bracts all entire, with a terminal spine 5. Carduus 57. Stem leaves not decurrent down the stem; stems not spiny; outer involucral bracts with spine-tipped lateral 9. Silybum lobes or teeth 10. Serratula 58. Involucral bracts simple and entire 58. At least the inner involucral bracts with a distinct apical 59. portion which is scarious, toothed or spiny 59. All flowers bisexual and of the same size; apical portion of involucral bracts scarious, not separated from the 11. Acroptilon main part by a constriction 59. Outer flowers of capitulum functionally female though sometimes with sterile stamens, often longer than the inner flowers; apical portion of involucral bracts usually dentate or spiny, if just scarious then separated from the 12. Centaurea main part by a constriction 61. 60. At least the lower leaves opposite 78. 60. All leaves basal or alternate 62. 61. All flowers in a capitulum tubular 66. 61. Outer flowers of capitulum ligulate, the inner tubular 62. Terminal capitula male only, in elongated, bractless 94. Ambrosia racemes 63. 62. All capitula bisexual, variously arranged 111. Eupatorium 63. Pappus of hairs 64. 63. Pappus absent, or of scales, or of stout bristles 64. Leaves pinnate or simple with a narrow cuneate base; 103. Bidens pappus of barbed bristles 64. Leaves simple, broadly cuneate to cordate at base; 65. pappus absent or of scales 65. Flowers greenish-white; receptacular scales present; 95. Iva pappus absent 65. Flowers blue; receptacular scales absent; pappus of 112. Ageratum scales 108. Schkuhria 66. Capitulum with only 1 ligulate flower 67. 66. Capitulum each with 3–numerous ligulate flowers 68. 67. Plant with large underground tubers 69. 67. Plant without large underground tubers 100. Helianthus 68. Leaves simple; ligules yellow 68. Leaves pinnately divided; ligules red to pink or purple 106. Dahlia or white 70. 69. Ligules pink or white 72. 69. Ligules yellow to greenish- or brownish-yellow 70. Leaf-lobes linear to filiform; ligules usually pink, rarely 105. Cosmos white 70. Leaf-lobes lanceolate to ovate; ligules white, rarely 71. purplish

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71. Capitula less than 7 mm in diameter excluding ligules; 102. Galinsoga pappus of scales 71. Capitula more than 7 mm in diameter excluding ligules; 103. Bidens pappus of strong, barbed bristles 103. Bidens 72. Pappus of strong, barbed, persistent bristles 72. Pappus absent or of weak, deciduous bristles and/or 73. scales 74. 73. Most leaves divided to the base or midrib 73. Leaves simple, subentire to toothed or shallowly lobed 75. 74. Pappus absent, or minute, or of 2 small scales; 104. Coreopsis receptacular scales present 74. Pappus of conspicuous scales; receptacular scales absent 107. Tagetes 75. Outer involucral bracts linear with dense glandular 98. Sigesbeckia hairs; ligules less than 10 mm 75. Outer involucral bracts lanceolate to ovate, not or 76. scarcely glandular-hairy; ligules more than 10 mm 97. Guizotia 76. Annual; leaves sessile and amplexicaul 76. Annual or perennial; leaves petiolate or sessile but not 77. amplexicaul 77. Pappus of 2 deciduous setae and rarely some small 100. Helianthus scales 77. Pappus absent or a small, more or less toothed rim 101. Heliopsis 78. Capitulum with flowers all tubular or if outer flowers 79. ligulate, ligules not exceeding inner involucral bracts 78. Capitulum with outer flowers ligulate, the ligules 108. obviously exceeding the inner involucral bracts 80. 79. Pappus absent, or of small scales, or a few bristles 94. 79. Pappus of hairs, sometimes apically expanded 81. 80. Leaves divided less than halfway to midrib 85. 80. Leaves divided more than halfway to midrib 81. Plant densely white-woolly; receptacular scales present 69. Otanthus 82. 81. Plant not densely woolly; receptacular scales absent 82. Pappus of 1–8 stiff, barbed bristles and sometimes also 56. Calotis some small scales 83. 82. Pappus absent or a small, membranous ring 83. Capitula very numerous, in racemes or panicles 67. Artemisia 84. 83. Capitula 1–numerous, if numerous then in corymbs 84. Stems procumbent to weakly erect; pappus absent 81. Cotula 84. Stems more or less erect; pappus a membranous ring 66. Tanacetum 86. 85. Capitula very numerous, in racemes or panicles 85. Capitula 1–few, or if more or less numerous in 87. corymbs 43. Berkheya 86. Involucral bracts spinose 67. Artemisia 86. Involucral bracts not spinose 87. Corolla hairy; stem and leaves densely hairy 67. Artemisia 88. 87. Corolla glabrous; stem and leaves glabrous to hairy 89. 88. Receptacular scales present 90. 88. Receptacular scales absent 89. Corolla with a small pouch at base, obscuring top of 71. Chamaemelum ovary in one plane 89. Corolla not pouched at base, not obscuring top of ovary 72. Anthemis

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90. Stems procumbent to weakly erect; achenes 81. Cotula compressed 90. Stems usually stiffly erect to ascending; achenes not or 91. scarcely compressed 92. 91. Perennial; capitula flat, in flat corymbs 93. 91. Capitula conical to convex, not in corymbs 66. Tanacetum 92. Achenes 5- to 12-ribbed 78. Oncosiphon 92. Achenes 4-ribbed 93. Achenes with 4–5 ribs, without resin glands near the 79. Matricaria apex 93. Achenes strongly 3-ribbed, with 2 resin glands near the 80. Tripleurospermum apex 94. Leaves broadly ovate to subrotund, cordate at base, 95. petiolate 94. Leaves linear to ovate, cuneate at base, petiolate or 97. sessile 95. Plant with scrambling stems and palmately lobed leaves 85. Delairea 95. Plant with aerial stems bearing only reduced bract-like 96. leaves, the main leaves all basal 91. Petasites 96. Capitula in inflorescences 92. Homogyne 96. Capitula solitary 51. Dittrichia 97. Plant densely glandular-hairy, very sticky 97. Plant glabrous to woolly, not obviously 98. glandular-hairy, nor sticky 99. 98. Plant glabrous to hairy, but not woolly 98. Plant woolly at least in part, especially near the tops of 104. stems 99. Leaves conspicuously serrate to deeply lobed or more or less pinnate; involucral bracts in 1 main row with much shorter supplementary ones near the base of the 82. Senecio capitulum 99. Leaves entire to slightly and remotely serrate; 100. involucral bracts in several rows 100. Involucral bracts totally scarious, coloured or white 48. Helichrysum 101. 100. Involucral bracts at least partly herbaceous 102. 101. Stems glabrous or sometimes weakly scabrid 103. 101. Stems hairy 102. At least the lower leaves more than 2.5 cm, either succulent or flat; pappus straw-coloured to pale reddish 58. Aster or brownish 102. All leaves less than 2.5 cm, with margins rolled under 59. Chrysocoma 103. Flowers yellow; outer involucral bracts with spreading 50. Inula to recurved tips 103. Flowers white to cream or pinkish; involucral bracts 61. Conyza appressed to flowers 105. 104. Annual with a simple root system 104. Perennial with rhizomes or stolons, though often short 106. 105. Outer involucral bracts herbaceous, woolly beyond halfway to apex; outer flowers with receptacular 44. Filago scales 105. All involucral bracts more or less scarious, glabrous or woolly in lower half; receptacular scales absent 47. Gnaphalium 47. Gnaphalium 106. Capitula bisexual, in elongate panicles 106. Capitula unisexual and more or less dioecious, in more 107. or less crowded, terminal, subcorymbose clusters

107. Plant stoloniferous; stems rarely more than 20 cm; largest leaves forming basal rosettes, the cauline much 45. Antennaria narrower 107. Plant rhizomatous; stems rarely shorter than 25 cm; leaves not forming a basal rosette, the largest ones 46. Anaphalis cauline 108. Pappus of at least the inner flowers of hairs, sometimes 109. also of scales 108. Pappus absent, or of scales, and/or a few bristles, or a 127. rim or tube 110. 109. Ligules white, or pink, or blue to purple, or mauve 117. 109. Ligules yellow to orange 110. Main leaves all basal, broadly ovate to subrotund, cordate at base; flowering stems with only reduced 91. Petasites bract-like leaves 110. Stems with leaves, or if all more or less basal then not 111. cordate at base 111. Outer involucral bracts broader than inner, green and 62. Callistephus more or less herbaceous 111. Outer involucral bracts similar to or smaller than the 112. inner ones 113. 112. At least the larger leaves truncate to cordate at base 115. 112. Leaves all cuneate at base 58. Aster 113. Involucral bracts in a graded series of rows 113. Involucral bracts in 1 main row, sometimes with smaller 114. supplementary ones near the base of the capitulum 114. Leaves pinnately veined; involucral bracts in 1 main row, with smaller supplementary ones at base of 82. Senecio capitulum; ligules white 114. Leaves palmately veined; capitula without supplementary involucral bracts; ligules white or 83. Pericallis coloured 115. Involucral bracts all green or more green in apical half than basal half; ligules linear to narrowly elliptical, 58. Aster more than 1 mm wide 115. Involucral bracts more green in basal half than apical half; ligules filiform to linear, usually less than 0.6, but 116. sometimes up to 1.5 mm wide 116. Inner tubular flowers more numerous than outer filiform ones, or the filiform ones absent; ligules more 60. Erigeron than 1 mm wide 116. Inner tubular flowers fewer than outer filiform ones; 61. Conyza ligules less than 1 mm wide 117. Main leaves all basal, cauline reduced to bracts; 90. Tussilago flowering stems with 1 capitulum 117. Leaves present on flowering stem, or if all more or less 118. basal, then capitula more than 1 per stem 118. Involucral bracts in 1 main row, often with smaller 119. supplementary ones near the base of the capitulum 123. 118. Involucral bracts in 2 or more, often indistinct rows 119. Capitula numerous in cylindrical or pyramidal panicles, 120. with 2–4 tubular flowers and 2–4 ligulate flowers 119. Capitula not in dense pyramidal panicles, with more than 5 tubular flowers and more than 4 ligulate flowers 121. 120. Lower leaves deeply pinnately lobed; petioles not 87. Sincalia sheathing stem; ligulate flowers usually 3 120. Lower leaves deeply palmately lobed; petioles 88. Ligularia sheathing stem; ligulate flowers usually 2 121. Basal and cauline leaves with petioles having broad, 88. Ligularia sheathing bases

148. ASTERACEAE 122. 121. Basal and cauline leaves without sheathing petioles 122. Involucral bracts in 1 main row with supplementary 82. Senecio smaller ones near the base of the capitulum 122. Involucral bracts in 1 row, with no supplementary small 84. Tephroseris ones near the base of the capitulum 123. Involucral bracts in 2 distinct rows of equal length 89. Doronicum 123. Involucral bracts in 2 or more indistinct rows, 124. becoming progressively longer towards the centre 124. Plant densely glandular-hairy and very sticky 51. Dittrichia 125. 124. Plant not or slightly glandular-hairy, but not sticky 125. Pappus of inner row of hairs and outer row of small, 52. Pulicaria often laterally fused scales 126. 125. Pappus entirely of hairs 126. Capitula more than 1.5(–2.0) cm in diameter including ligules; ligules more than 10 mm; anthers with 50. Inula branched tails 126. Capitula less than 1.5(–2.0) cm in diameter including ligules; ligules less than 10 mm; anthers without tails 57. Solidago 128. 127. Receptacular scales or abundant bristles present 127. Receptacular scales and bristles absent, but sometimes 136. short fringes round the achene pits 128. Receptacle with abundant bristles only; pappus of 5–10 scales which have long apical bristles 109. Gaillardia 128. Receptacle scales present; pappus absent, a rim, or 129. small scales not bristle-tipped 129. Achenes strongly compressed, more than twice as wide 130. as thick, with strong lateral ribs or narrow wings 129. Achenes angular, not or only slightly compressed, less than twice as wide as thick, without lateral ribs or 131. wings 130. Capitula less than 20 mm in diameter including ligules; 70. Achillea ligules less than 10 mm 130. Capitula more than 20 mm in diameter including 104. Coreopsis ligules; ligules more than 10 mm 132. 131. Ligules white 133. 131. Ligules yellow to orange 132. Corolla of tubular flowers with a small pouch at base, 71. Chamaemelum obscuring top of ovary in one plane 132. Corolla of tubular flowers not pouched nor obscuring 72. Anthemis the top of the ovary 133. Involucral bracts with broad scarious margins and tips 72. Anthemis 134. 133. Involucral bracts green and entirely herbaceous 134. Lower leaves cordate at base; ligules less than 2 mm 54. Telekia wide 134. Lower leaves cuneate at base; ligules more than 2 mm 135. wide 53. Buphthalmum 135. Plant not scabrid; receptacle convex 99. Rudbeckia 135. Plant scabrid; receptacle strongly conical 108. Schkuhria 136. Capitulum with 1 ligulate flower 137. 136. Capitulum with at least outer flowers ligulate 77. Chrysanthemum 137. All flowers ligulate 138. 137. Inner flowers tubular

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138. At least some achenes strongly curved, very warty on 93. Calendula outer face 138. Achenes not or slightly curved, if warty then not just 139. on outer face 139. Pappus of 1–8 bristles, sometimes with minute scales 140. as well 139. Pappus absent or a minute rim, or of scales, or tubular 141. 140. Fruiting capitula more than 10 mm in diameter, very resinous; pappus bristles smooth or forwardly serrated 55. Grindelia 140. Fruiting capitula less than 10 mm in diameter, not resinous; pappus bristles barbed and persistent 56. Calotis 142. 141. Pappus absent 143. 141. Pappus of scales or tubular 142. Achenes of the ligulate flowers 3-angled with the ribs often winged, those of the tubular flowers more or less 73. Xanthophthalmum cylindrical 142. Achenes all similar, cylindrical-obconical, 5- to 77. Chrysanthemum 8-ribbed 43. Berkheya 143. Leaves more or less spiny 144. 143. Leaves not spiny 144. Involucral bracts fused into a cup-like structure 42. Gazania 145. 144. Involucral bracts not fused into a cup-like structure 145. Cauline leaves decurrent on the stem as wings 110. Helenium 146. 145. Cauline leaves not decurrent 147. 146. Pappus of scales or tubular 149. 146. Pappus absent or a minute rim 75. Coleostephus 147. Pappus tubular 148. 147. Pappus of scales 148. Outer flowers of capitulum female; achenes with 3 40. Arctotis wing-like ridges on 1 side 148. Outer flowers of capitulum sterile; achenes without 41. Arctotheca ridges 149. Rosette plant; capitula solitary on leafless stems 65. Bellis 150. 149. Flowering stems bearing leaves 150. Stem leaves simple, shallowly to deeply toothed but not 151. to midrib, the teeth simple 150. Stem leaves pinnately lobed to midrib or nearly so, the 153. lobes further lobed 66. Tanacetum 151. Ligules less than 10 mm 152. 151. Ligules more than 10 mm 152. Tubular part of ligulate flower not winged 74. Leucanthemella 152. Tubular part of ligulate flower with 2 narrow 76. Leucanthemum transparent wings 153. Ultimate leaf segments lanceolate to ovate, flat 66. Tanacetum 153. Ultimate leaf segments linear to filiform, not or 154. scarcely flattened 154. Achenes with 4–5 ribs, without resin glands 78. Matricaria 154. Achenes strongly 3-ribbed, with 2 resin glands near the 80. Tripleurospermum apex

Subfamily 1. Cichorioideae (Juss.) Chev. nom. conserv. Subfamily Lactucoideae (Cass.) Lindl.; Subfamily Liguliflorae; Subfamily Cichoriaceae Juss.; Subfamily

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Arctotideae Cass.; Subfamily Carduoideae Cass. ex Sweet; Subfamily Carlinoideae (Cass.) Lindl.; Subfamily Centaureoideae (Cass.) Lindl.; Subfamily Cynaroideae (Durande) Chev.; Subfamily Echinopsoideae (Cass.) Lindl.; Subfamily Hieracioideae Burmeist.; Subfamily Hyoseridoideae Burmeist.; Subfamily Hypochaeridoideae Burmeist.; Subfamily Mutisioideae (Cass.) Lindl.; Subfamily Nassaurioideae (Cass.) Lindl.; Subfamily Scorzoneroideae Burmeist.; Subfamily Vernonioideae (Cass.) Lindl.; Subfamily Xeranthemoideae Burmeist. Plants often producing white latex. Stem leaves usually spiral, sometimes absent. Capitula with flowers all ligulate, all tubular or rarely the inner tubular, the outer ligulate. Tubular flowers if present with long, narrow lobes, usually blue or red, rarely yellow. Filaments joining anthers on back. Style branches 2, usually each with 1 broad stigmatic surface on the inner face. Pollen grains ridged, spiny or both. Contains 6 tribes, 25 subtribes, 391 genera and about 6,700 species, plus a large number of apomictic microspecies. Cosmopolitan. Tribe 1. Cynareae Lam. & DC. Tribe Cardueae Cass.; Tribe Echinopseae Cass.; Tribe Carlineae Cass.; Tribe Centaureae Cass.; Tribe Xerantheminae Kostel; Tribe Carthaminae DC.; Tribe Serratuleae (Less.) W. D. J. Koch; Tribe Silybeae Kitt.; Tribe Crupineae Godr. Plants not producing white latex, often spiny. Capitula with the flowers all tubular, the outermost often longer and with larger lobes and thus pseudo-radiate, usually red to blue or white, rarely yellow. Anthers tailed, the apical appendage elongate; filaments glabrous or papillose-hairy; pollen generally spiny. Style with long, linear branches which are dorsally papillose-hairy and with a ring of hairs on the thickening beneath the branches. Contains 4 subtribes, 83 genera and about 2,500 species. Mainly Eurasia and North Africa with a few genera elsewhere. 1. Echinops L. Biennial to perennial herbs with fleshy tap-roots and fibrous side-roots. Stems erect, sulcate. Leaves alternate, deeply pinnately lobed, white to grey arachnoid-hairy on lower side, with lobes and teeth ending in rather weak spines. Capitula each with 1 flower, grouped on a receptacle into 1 or more globose heads more than 25 mm in diameter, with laciniate basal bracts. Involucral bracts in more or less 3 rows, herbaceous, long-pointed at apex, with long, fine rigid hairs on the margin and a row of white hairs or narrow scales outside them. Flowers bisexual. Corolla tubular, white to blue, deeply 5-lobed. Anthers tailed; filaments glabrous. Style slender and with rather long, linear style branches, with a ring of hairs beneath the branches. Achenes oblong, angled, densely appressed-hairy; pappus of partially connate scales. About 120 species in Eurasia and North and East Africa, introduced elsewhere.

Kruse, J. & Meusel, H. (1969). Zur Blattfolge und Blattbildung einiger. 1. Echinops. Feddes Rep. 80: 339–356. 1. Plant without glandular hairs; involucral bracts with a 2. exaltatus long recurved or bent apex 1. Plant with glandular hairs present at least on the upper surface of the leaf; involucral bracts erect or very slightly 2. curved at apex 2. Plant often more than 150 cm; outer involucral bracts with numerous glandular hairs; corolla pale greyish-blue 1. sphaerocephalus 2. Plant not more than 120 cm; outer involucral bracts without glandular hairs; corolla more pure blue 3. bannaticus

1. E. sphaerocephalus L. Glandular Globe Thistle E. multiflorus Lam.; E. paniculatus Jacq.; E. viscosus Wierzb. Perennial herb with fleshy tap-root and fibrous sideroots. Stems 50–160(–250) cm, pale yellowish-green, often flushed purplish-brown, erect, ridged, with dense, short, reddish-brown glandular hairs and scattered to fairly numerous, pale, longer simple eglandular hairs, leafy, branched in the upper part. Leaves all cauline at anthesis, 3–20 × 2– 10 cm, dull, medium yellowish-green with a pale midrib on upper surface, greenish-grey beneath, ovate-oblong, oblong or lanceolate in outline, spinose-acute at apex, pinnatifid, the lobes of the lower leaves triangular-ovate, with a spine at the apex and irregular, unequal, yellowish-brown spines along the margins, those of the upper leaves smaller and narrower, the interlobes short and broadly winged, more or less amplexicaul at the sessile base, with numerous, short and occasional medium, pale glandular hairs on the upper surface and margin, densely arachnoid hairy beneath and with numerous, pale short glandular and eglandular hairs on the prominent veins. Inflorescence 30–60 mm in diameter, globose, solitary at the ends of branches; branches pale green often suffused purplish-brown, arachnoid-hairy and densely glandular-hairy. Involucral bracts in 3 rows, 14– 17 × 1.0–2.5 mm, dull, medium green with a narrow, scarious margin and apex, oblanceolate or linear-oblanceolate, the outer erect, gradually drawn out to a very fine, sometimes slightly curved apex, with short glandular hairs on the outer face and medium to rather long, white hairs on the margin of the upper half, the inner similar but without glandular hairs and with long, white hairs at the obtuse apex. Flowers about 5 mm, all tubular, pale greyish-blue, shortly 5-lobed at apex. Receptacle of inflorescence pitted, the individual capitulum of 1 flower with involucral bracts sitting in the pit and surrounded by long, white hairs or narrow scales. Achenes 7–8 mm, oblong, densely appressed-hairy; pappus 1.0–1.5 mm, of partially connate, hairy scales. Flowers 8–9. Visited by bees. 2n = 32. Introduced. A garden escape naturalised in waste places, rough ground, waysides and railway banks. There are scattered records throughout much of Great Britain, but this plant is probably over-recorded as it was assumed to be the only species in our area. Native of central and southern Europe and western Asia. Our plant is subsp. sphaerocephalus.

2. Carlina 2. E. exaltatus Schrad. Globe Thistle E. commutatus Jur.; E. strictus Fisch. ex Sims; E. mollis Schur Perennial herb with fleshy tap-root and fibrous side-roots. Stems 40–250 cm, pale yellowish-green, sometimes tinted brownish-purple, erect, strongly ridged and channelled, arachnoid-hairy, leafy, simple or branched in the upper part. Leaves all cauline at anthesis, 10–35 × 4–30 cm, dull medium green on upper surface, grey beneath, broadly ovate to oblong in outline, spinose-acute at apex, pinnatifid, the lobes triangular-ovate to oblong, spinose-acute at apex and shallowly pinnatifid, the lobules triangular-ovate and spinose-acute, smaller spines occurring in the interlobular area and in the interlobe area which is short and broadly winged, with short, bent or curved, rigid simple eglandular hairs on the upper surface, densely arachnoidhairy beneath; veins prominent beneath; lower leaves with a short, winged, toothed petiole, all leaves amplexicaul at base. Inflorescence 35–60 mm in diameter, globose, solitary at the end of branches; peduncles strongly ridged, densely arachnoid-hairy. Involucral bracts in 3 rows, 16–22 × 1.5– 2.0 mm, pale brown or whitish, the outer spathulate, with a long recurved or bent spine at apex and ascending fimbriae on the margin, the inner oblanceolate-spathulate and longer, but otherwise similar, all glabrous on the surface. Flowers 5–7 mm, all tubular, pale greyish-mauve or white, shortly 5-lobed at apex. Receptacle of inflorescence pitted, the individual capitulum of 1 flower with involucral bracts sitting in a pit and surrounded by white hairs or scales as long as the outer involucral bracts. Achenes 7–8 mm, oblong, densely appressed-hairy; pappus about 2 mm, of partially connate, hairy scales. Flowers 8–9. 2n = 30. Introduced. Grown in gardens and naturalised in waste places, rough ground, waysides and railway banks. There are scattered records throughout much of Great Britain and this species is probably commoner than E. sphaerocephalus. Native of east-central Europe, north-east Italy, the northern part of the Balkan peninsula and western Asia. 3. E. bannaticus Rochel ex Schrad. Blue Globe Thistle E. ritro auct.; E. ruthenicus Rochel, non M. Bieb.; E. rochelianus Griseb. Perennial herb with fleshy tap-root and fibrous side-roots. Stems 50–120 cm, pale yellowish-green, sometimes suffused brownish-purple, erect, strongly ridged and channelled, white arachnoid-hairy, leafy, simple or branched near the top. Leaves all cauline at anthesis, 10–20 × 2–10 cm, dull medium green on upper surface, whitish beneath, broadly ovate or elliptical in outline, spinose-acute at apex, deeply pinnatifid, the lobes lanceolate or ovate and obtuse or cuneate and spine-tipped at apex, the lobules mammiform or oblong, the interlobe area broadly winged, sessile, with numerous, short, pale glandular hairs on the upper surface and the revolute margins, glandular and densely, whitish arachnoid-hairy beneath; veins prominent beneath; usually yellowish, with slender spines along the margin. Inflorescence 25–50 mm in diameter, globose, solitary at the ends of branches; branches pale green, densely arachnoidhairy. Involucral bracts in 3 rows, 14–17 × 1.4–1.6 mm,

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pale green and mauve-blue towards the apex, the outer triangular-lanceolate, the inner lanceolate, erect, long-acute or acuminate at apex, the outer with long, erect cilia but without glandular hairs. Flowers 11–14 mm, all tubular, bluish, deeply divided into 5, narrow lobes at apex. Receptacle of inflorescence pitted, the individual capitulum of 1 flower with involucral bracts sitting in the pit and surrounded by white, rigid hairs connate at the base and equalling or shorter than the outer bracts. Achenes 5–6 mm, oblong, densely appressed-hairy; pappus 2.0–2.5 mm, of partially connate, hairy scales. Flowers 8–9. 2n = 30. Introduced. A garden escape naturalised in waste places, rough ground, railway banks and roadsides. There are scattered records throughout Great Britain. Native of south-east Europe and west Asia. 2. Carlina L. Biennial herbs with a tap-root. Stems erect, not spiny. Leaves alternate, pinnately lobed, more or less arachnoidhairy, very spiny. Capitula solitary or in terminal groups of 2–3(–6) forming a corymb. Involucral bracts in several rows, spiny, the outer more or less leaf-like, the innermost linear, entire, scarious, patent and ligule-like in dry weather. Flowers all tubular and bisexual. Corolla tubular, purple, shallowly 5-lobed. Anthers tailed; filaments glabrous. Style shortly bilobed, glabrous or papillose to minutely hairy apically and dorsally on the lobes, generally with a subapical ring of longer hairs beneath the lobes. Receptacle flat, with laciniate scales more or less connate to a honeycombed structure covering the receptacle and enclosing the achenes. Achenes oblong, hairy; pappus a ring of plumose hairs often united proximally. Contains 28 species in Europe and south-western Asia, North Africa and the Canary Islands. Greig-Smith, J. & Sagar, G. R. (1981). Biological causes of local rarity in Carlina vulgaris in Synge, H. (Edit.) The biological aspects of rare plant conservation, pp. 389–400. Chichester. Grime. J. P. et al. (1988). Comparative plant ecology. London. 1. Leaves irregularly lobed, rather sparsely arachnoid-hairy 1(i). vulgaris var.vulgaris 1. Leaves simple with a regular, spiny margin and densely 1(ii). vulgaris var. maritima arachnoid-hairy

1. C. vulgaris L. Carline Thistle Biennial herb with a tap-root. Stems 10–60 cm, pale green, often suffused reddish-purple, stiffly erect, markedly striate, more or less appressed arachnoid-hairy, usually branched above, persisting for at least a season after the plant dies. Leaves numerous, condensed at base but sometimes dying before flowering, the cauline gradually decreasing in size or abruptly smaller just above the base, 2–13 × 0.5– 2.0 cm, greyish-green, linear, oblong, oblong-lanceolate or oblong-oblanceolate, acute at apex, undulate to irregularly pinnately lobed, rarely simple, with numerous, short, pale rather weak spines at the margin, attenuate at base, the basal petiolate, the cauline semiamplexicaul, more or less arachnoid-hairy especially beneath. Capitula solitary or in terminal groups of 2–3(–6) forming a corymb, 15–30 mm in diameter; peduncles usually short, striate,

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arachnoid-hairy. Involucral bracts in several rows; outer more or less leaf-like, 10–20(–25) × 3–7 mm, pale green, linear-oblong to lanceolate, spiny-acute at apex, spiny on the margin, sparsely arachnoid-hairy; inner 12–20 × 1.0– 1.5 mm, straw-coloured and tinted purplish in the lower part, linear, long-acute at apex, entire, glabrous, becoming patent and ligule-like in dry weather. Flowers all bisexual. Corolla tubular, 5-lobed, purple. Receptacle flat, with scales more or less connate to a honeycombed structure covering the receptacle, the scales linear, scarious and deeply laciniate at apex. Achenes 2–4 mm, dark brown, oblong, covered with long, reddish-brown, appressed hairs; pappus 5–8 mm, a ring of pale brownish cream plumose hairs in one row, united at the base in groups of 2–4, deciduous. Flowers 7–10. Visited chiefly by bees and hoverflies. 2n = 20. (i) Var. vulgaris Taller plant. Leaves irregularly lobed, rather sparsely arachnoid-hairy. (ii) Var. maritima (Rouy) P. D. Sell C. vulgaris forma maritima Rouy Squat robust plant. Leaves simple with a regular spiny margin and densely arachnoid-hairy. Native. Calcareous grassland, open basic rocky habitats and stabilised sand dunes; a poor competitor which in closed turf is favoured by heavy grazing. Frequent in suitable habitats in most of Great Britain and Ireland north to Orkney, but only coastal in the north of Ireland and Scotland and absent from the Outer Hebrides and Shetland. Europe northwards to about 62◦ N in Scandinavia; Siberia; Turkey; Caucasus; very variable. A member of the Eurosiberian Temperate element. All our plants are referable to subsp. vulgaris. Var. vulgaris is the widespread plant. Var. maritima in known only from the Mull of Galloway and Arran. It was described from maritime rocks on Finist`ere, France. 3. Arctium L. Lappa Scop. nom. illegit.; Bardana Hill nom. illegit. Biennial herbs with long, stout tap-roots. Stems erect, not spiny. Leaves alternate, large, simple, entire to remotely denticulate or dentate, arachnoid-hairy. Capitula solitary at ends of branches or in corymbose to racemose clusters, dispersed as a unit with the achenes enclosed. Involucral bracts numerous, in several rows, rigid, subulate, mostly strongly hooked at apex, spreading to form a subglobose capitulum. Flowers all tubular and bisexual. Corolla tubular, usually purple, rarely white, deeply 5-lobed. Anthers tailed; filaments glabrous. Style with long, linear branches, dorsally hairy and with a ring of hairs beneath the branches. Receptacle flat, with dense rigid hairs. Achenes obovoid-oblong, compressed, rugose; pappus of scabrid, yellow hairs, free to the base. About 8 species throughout most of Europe and temperate Asia and widely introduced elsewhere. Ar`enes, J. (1950). Monographie du genre Arctium L. Bull. Jard. ´ Bruxelles 20: 67–156. Bot. Etat Babington, C. C. (1856). On the British species of Arctium. Ann. Mag. Nat. Hist. ser. 2, 17: 369–377.

Beeby, W. H. (1908). The British species of Arctium. Jour. Bot. (London) 46: 380–382. Duistermaat, H. (1996). Monograph of Arctium L. (Asteraceae). Gorteria Suppl. 3, 1–143. Evans, A. H. (1913). The British species of Arctium. Jour. Bot. (London) 51: 113–119. Grime, J. P. et al. (1988). Comparative plant ecology. London. [A. minus aggr.] Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 13: 208–209 [A. lappa sens. strict.] Perring, F. H. (1960). Report on the survey of Arctium L. agg. in Britain, 1959. Proc. B.S.B.I. 4: 33–37. 1. 1. 2. 2. 3.

3.

4.

4. 5. 5.

2. tomentosum Inner involucral bracts not hooked at apex 2. Inner involucral bracts hooked at apex 3. Petioles solid 4. Petioles with a central hollow Capitula 30–43 mm in diameter; involucral bracts green, glabrous or slightly arachnoid-hairy and with minute 1(a). lappa subsp. lappa glandular hairs Capitula 20–35 mm in diameter; involucral bracts usually tinted brownish-red, often very arachnoid-hairy 1(b). lappa subsp. pubens Capitula 15–25 mm in diameter, subsessile or on peduncles up to 8 mm; involucral bracts 7–15 mm 1(c). lappa subsp. minus Capitula 20–40 mm in diameter, sessile or on peduncles 5. up to 40(–120) mm; involucral bracts 6–19 mm 1(b). lappa subsp. pubens Peduncles 5–40(–120) mm Peduncles up to 8 mm or capitula sessile 1(d). lappa subsp. nemorosum

1. A. lappa L. nom. conserv. Burdock Biennial herb with a long, stout tap-root. Stems 90–150 cm, stout, pale green, often suffused purplish-red, furrowed, more or less clothed with arachnoid hair, with many, spreading ascending branches. Basal and lower cauline leaves with lamina 15–58 × 15–45 cm, broadly ovate, apiculate to acute at apex, entire or distantly denticulate to crenatedentate and often undulate, cordate at base; middle and upper leaves gradually becoming smaller, narrower, less cordate and with shorter petioles, the uppermost often broadly lanceolate to ovate-lanceolate and not all cordate; all green and more or less arachnoid-hairy on upper surface and with pale veins, paler and more or less arachnoid-hairy beneath with prominent veins; petioles 5–550 mm, pale green, grooved above and round and ridged below, solid or with a central hollow, slightly arachnoid-hairy. Capitula 15–42 mm in diameter, 1–4 together in loosely corymbose groups terminating the main branches or in racemes, globose in bud, hemispherical and widely open in fruit; peduncles 3–100 mm or subsessile, pale green, slightly arachnoid-hairy. Involucral bracts numerous, 6–21 × 0.7– 2.6 mm, yellowish-green or straw-coloured to darker green and purplish-tinged, narrowly linear-lanceolate, flat and imbricate below, narrowing above into a long, rigid, subulate, more or less spreading point, all hooked at apex, glabrous to densely arachnoid-hairy and often with minute glandular hairs. Flowers all tubular and bisexual, deeply 5lobed, usually reddish-purple, rarely white. Receptacle flat,

3. Arctium with numerous, rigid hairs 5–15 mm. Achenes 5–9 × 2– 3 mm, pale to dark brown, often with blackish blotches, obovoid-oblong, compressed, ribbed, more or less rugose; pappus 1.0–4.9 mm, of scabrid, golden-yellow hairs in several rows, free to the base. Flowers 7–9. Visited by bees and Lepidoptera, but usually self-pollinated before the flowers open. There is continuous variation in size of capitulum between the large-headed subsp. lappa and the smallheaded subsp. minus. In this account subspp. lappa, nemorosum and minus are narrowly defined and the remainder put in subsp. pubens. It is probable that subsp. pubens originated from hybridisation between subsp. lappa and subsp. minus with possible backcrossing with both parents. Thus, subsp. pubens is extremely variable and tends to form distinct local populations which are kept constant by predominantly self-pollination. Much of subsp. nemorosum is outside the range of the other subspecies but comes in contact with subsp. minus where intermediates are difficult to separate from subsp. pubens. Experiments at Cambridge in the 1950s showed that seeds were produced if the capitula were bagged, but not if the flowers were emasculated. Intermediates were as fertile as the subspecies. With this in mind populations have been examined in the field over much of Great Britain with the following conclusions. We do not understand the classification of Duistermaat (1996), whose conclusions differ radically from ours. (a) Subsp. lappa Greater Burdock Lappa vulgaris Hill; Bardana lappa (L.) Hill; Lappa major Gaertn. nom. illegit.; A. ruderale Salisb. nom. illegit.; A. majus Bernh. nom. illegit.; A. vulgare (Hill) Druce Leaves with lamina entire or distally dentate; petioles solid. Capitula 30–42 mm in diameter on peduncles 3–120 mm, in loosely corymbose groups terminating the main inflorescence branches. Involucral bracts 8–21 × 1.0–2.6 mm, green, entire, glabrous or slightly arachnoid-hairy and with few to numerous minute glandular hairs. Achenes 6.0–7.7 mm; pappus 1.0–4.9 mm. Receptacular hairs 5–9 mm. 2n = 36. (b) Subsp. pubens (Bab.) P. D. Sell Hairy Burdock A. pubens Bab.; Lappa pubens (Bab.) Boreau; A. minus subsp. pubens (Bab.) Ar`enes; Lappa notha Ruhmer; A. notha (Ruhmer) J. Weiss; Lappa subracemosa Simonk.; A. subracemosum (Simonk.) Nyman; Lappa mixta E.G. Camus; Lappa ruhmerio Gu´etrot; A. debrayi Senay; A. batavum Ar`enes Leaves with lamina mostly rather sharply dentate; petioles usually hollow, but sometimes more or less solid. Capitula 20–35 mm in diameter on peduncles 5–40(–120) mm, in a lax raceme in groups of 1–4. Involucral bracts 9–17 × 0.8– 2.2 mm, pale green, often tinted brownish-red, entire or slightly toothed, usually very arachnoid-hairy at first, less so later. Achenes 5–8 mm; pappus 1–3 mm. Receptacular hairs 5–11 mm. 2n = 36. Subsp. pubens includes a whole range of fertile intermediates between subsp. lappa and subsp. minus and it is impossible to exclude most intermediates between all three of these subspecies and subsp. nemorosum. All the

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subspecies can grow on their own in pure populations without other taxa being present. (c) Subsp. minus (Hill) Hook. fil. Lesser Burdock Lappa minor Hill; A. minus (Hill) Bernh.; Bardana minor (Hill) Hill; A. personata Mill., non L.; Lappa glabra Lam. Leaves with lamina mostly rather sharply dentate; petiole hollow. Capitula 15–25 mm in diameter, subsessile or on peduncles up to 8 mm, in racemes with the terminal capitulum usually solitary. Involucral bracts 7–15 × 0.7–2.0 mm, pale green, suffused brownish-purple, minutely toothed, slightly arachnoid-hairy, sometimes with minute glandular hairs, or glabrous. Achenes 5–6 mm; pappus 1–3 mm. Receptacular hairs 5–8 mm. 2n = 36. (d) Subsp. nemorosum (Lej.) P. D. Sell Northern Burdock A. nemorosum Lej.; Lappa major var. racemosa (Lej.) G. Mey.; Lappa macrosperma Wallr.; Arctium intermedium Lange; Lappa intermedium (Lange) Rchb. fil.; Lappa nemorosa (Lej.) K¨orn.; A. newbouldii A. Benn. nom. provis.; A. austriacum Teyber; A. minus subsp. nemorosum (Lej.) Syme; A. majus subsp. nemorosum (Lej.) Rouy; A. majus subsp. intermedium (Lange) Nyman Leaves with lamina distantly crenate-dentate; petiole hollow. Capitula (25–)30–40 mm in diameter, sessile or on peduncles up to 8 mm, in racemes but with the top 2–4 in a subcorymbose cluster. Involucral bracts 6–19 × 1.2–2.6 mm, green, often tinted brownish-purple especially at the tips, slightly toothed, slightly arachnoid-hairy. Achenes 6– 9 mm; pappus 1.0–3.5 mm. Receptacular hairs 9–15 mm. 2n = 36. Native. The species occurs throughout Great Britain and Ireland except for the Scottish Highlands. It also occurs throughout much of Europe, North Africa and south-west Asia, and is widely naturalised outside its native range. It is a member of the Eurasian Temperate element. All subspecies occur on waysides and in field borders and waste places. Subsp. lappa is scattered through south and central Britain and is absent from the north and much of the west. It is scattered through most of Europe but rare in the Mediterranean and in Asia through Iran, Pakistan and northern India eastwards to Japan, and is introduced in North and South America and Australasia. Subsp. minus is the only subspecies which frequently grows in woods and shady places as well as in open habitats. It is the most common subspecies in central and south Britain, central and southern Ireland and the Channel Islands, and is the predominant taxon in southwest England. It is absent from Scotland, north England and north Ireland. It also occurs in most of Europe to about 66◦ N in Scandinavia and in south-west Asia east to Iran and in North Africa. It is introduced in North and South America and New Zealand. Subsp. nemorosum is the only taxon in much of north and west Britain and north Ireland, but is absent from south-west Britain. Subsp. nemorosum to a considerable extent replaces subsp. minus in these areas, but there is an overlap in south-east England and in a belt stretching from north and west Wales to Lincolnshire and Yorkshire. In Europe it is in mainly central but extends to 64◦ N in Scandinavia and south to southern Italy and northern

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Greece, and eastwards to western Russia. Subsp. pubens is widespread in central and southern Britain with a few records in Ireland and further north. It occurs in west, central and south Europe, but seems to be absent from the north, and is introduced in North and South America. It probably originates from hybridisation between subsp. lappa and subsp. minus, but subsp. nemorosum may also be involved. Other subspecies occur in Europe, central Asia and North Africa. 2. A. tomentosum Mill. Woolly Burdock Lappa arctium Hill; Lappa tomentosa (Mill.) Lam.; A. bardana Willd.; A. crispum Cretz.; Bardana arctium (Hill) Hill Biennial herb with a long, stout tap-root. Stems 50–150 cm, stout, erect, pale green often tinted brownish-red, furrowed, more or less clothed with arachnoid hair and numerous very short glandular hairs, with many erecto-patent branches above. Basal and lower cauline leaves with lamina 15–50 × 15–40 cm, broadly ovate, apiculate to acute at apex, entire or distantly denticulate and often undulate, cordate at base; middle and upper leaves gradually becoming smaller, narrower, less cordate and with shorter petioles, the uppermost often broadly lanceolate to ovatelanceolate and not at all cordate; all green on upper surface and glabrous or with some hairs, paler and more or less arachnoid-hairy beneath and with prominent veins; petioles up to 500 mm, pale green, grooved above, rounded and ridged below, solid, slightly arachnoid-hairy. Capitula 15– 30 mm in diameter, in groups of 1–4, in corymbs or rarely on elongated panicles, more or less globose; peduncles 30– 100 mm, pale green, with dense, minute glandular hairs and more or less arachnoid-hairy. Involucral bracts 3–16 × 1.0– 3.2 mm, densely arachnoid-hairy to more or less glabrous; outer yellowish-green, linear-lanceolate below, abruptly contracted at the middle to a linear, hooked apex; inner oblong-lanceolate, with an acute or obtuse, not hooked, membranous, purplish apex. Flowers all tubular, deeply 5-lobed, usually reddish-purple, rarely white. Receptacle flat, with dense, rigid hairs 1.4–3.0 mm. Achenes 5–6 × 2– 3 mm, pale to dark brown, often with blackish blotches, obovoid-oblong, compressed, ribbed, more or less rugose; pappus 1.0–3.5 mm, of scabrid, golden-yellow hairs in several rows, free to the base. Flowers 7–9. 2n = 36. This species may be confused with A. lappa subsp. pubens which sometimes has a more or less solid petiole. Its heads are sometimes almost free of arachnoid hairs, var. glabrum (K¨orn.) Ar`enes (Lappa tomentosa var. glabra K¨orn.; Lappa palladinii Marcow.; A. palladinii (Marcow.) Grossh.), but this variant does not seem to be recorded from Great Britain and Ireland. Introduced. A grain casual which may be under-recorded. Native of west and central Europe southwards to northern Italy, eastwards to central and south Russia, Crimea and Transcaspia and naturalised in North America. It can hybridise with A. lappa. 4. Saussurea DC. nom. conserv. Bennettia Gray Perennial herbs with short stolons. Stems erect, solitary. Leaves alternate, simple, subentire to denticulate; or rarely

dentate, densely white or greyish arachnoid-hairy on lower surface, not spiny. Capitula in more or less contracted terminal corymbs or a corymbiform panicle. Involucral bracts in many rows, obtuse at apex, more or less hairy. Flowers all tubular and bisexual. Corolla tubular, purple, deeply 5-lobed. Anthers with long, acute, terminal apendages and basal feathery tails; filaments glabrous. Style with long linear branches which are dorsally hairy and with a ring of hairs beneath the branches. Receptacle flat, with scales. Achenes cylindrical, glabrous, smooth or rugulose; pappus of 1 row of plumose hairs, often united proximally, with an outer row of simple, shorter hairs. About 30 species, mostly alpine herbs in central and eastern Asia, with others in Europe and North America and one in south-east Asia extending to northern Australia. Named after Horace B´en´edict de Saussure (1740–99). 1. S. alpina (L.) DC. Alpine Saw-wort Serratula alpina L.; Cirsium montanum Hill nom. illegit.; Bennettia alpina (L.) Gray Perennial herb with a more or less scaly stock producing short stolons which end in leaf rosettes. Stem 7–45 cm, solitary, yellowish-green, sometimes tinted purplish, leafy, ridged, arachnoid-hairy. Leaves yellowish-green on upper surface, greyish-white beneath; basal and lower cauline with lamina 2–18 × 1.5–4.0 cm, narrowly lanceolate to ovate, obtuse to acute at apex, entire to sinuate-denticulate or more rarely sinuate-dentate, rounded or cuneate at base, with a narrowly winged petiole; upper cauline gradually decreasing in size, lanceolate to linear-lanceolate, shortly to long acute at apex, entire to sinuate-denticulate, sessile; all sparsely arachnoid-hairy above, more densely so beneath. Capitula solitary to compactly corymbose, sometimes with a long, lower branch, 10–20 mm in diameter; peduncles very short, densely arachnoid-hairy. Involucral bracts in many rows, 4–6 × 1.5–2.5 mm, green or purplish, narrowly to broadly ovate or lanceolate, obtuse at apex, the outer sparsely arachnoid-hairy, the inner villous with long, grey hairs. Flowers all tubular and bisexual, purple, fragrant, deeply 5-lobed. Receptacle flat; scales numerous, paleaceous. Achenes 3–4 mm, brown with 4 pale ribs, cylindrical, glabrous; pappus of 1 row of whitish plumose hairs, often united proximally, and an outer row of simple, shorter hairs. Flowers 7–9. Protandrous. Visited by flies and bees. 2n = 52, 54. Native. Rocky streamsides, mountain cliff ledges and scree to 1190 m, descending to the maritime zone in north Scotland where it is sometimes in reach of sea spray; very susceptible to grazing. Usually found on calcareous or ultrabasic rocks but sometimes on acidic rocks including granite. Local in north Wales, Craven Pennines, Lake District, western and northern Scotland, Inner and Outer Hebrides, Orkney and Shetland Islands and the highest mountains of Ireland. Northern Europe and the mountains of central Europe; northern and central Asia, reaching 75◦ N in Siberia. A member of the Eurasian Arctic-montane element. Our plant is subsp. alpina which occurs through much of the range of the species.

5. Carduus 5. Carduus L. Polycantha Hill; Ascalea Hill Annual or biennial herbs. Stems erect, with spiny wings. Leaves alternate, variously lobed, sharply and densely spiny, decurrent down the stem. Capitula solitary or in clusters at the ends of the main stems and branches. Involucral bracts in numerous rows, simple, linear-subulate to narrowly ovate, spine-tipped. Flowers all tubular and bisexual. Corolla tubular, usually purple or pink, rarely white, deeply 5-lobed, the lobes sometimes unequal. Anthers tailed; filaments hairy. Style with long, linear branches which are dorsally hairy and with a ring of hairs beneath the branches. Receptacle flat, densely bristly. Achenes obovoid-oblong or oblong, smooth or glabrous, with an apical rim; pappus of many rows of deciduous simple rigid hairs, united proximally in a ring. About 90 species in Eurasia and North and East Africa and widely introduced elsewhere. C. argentatus L. and C. thoermeri Weinm. have been recorded as bird-seed and grain casuals. C. macrocephalus Desf. is grown in gardens and was formerly naturalised in southern England. It differs from C. nutans in having a prominent (not obscure) midrib to its involucral bracts and a more densely hairy stem. Hybrids recorded between species of Carduus and Cirsium are very doubtful. Kasmi, S. M. A. (1964). Revision der Gattung Carduus (Compositae). Mitt. Bot. Staatssamml. (M¨unchen) 5: 279–550. Moore, R. J. & Mulligan, G. A. (1956). Natural hybridisation between Carduus acanthoides and Carduus nutans in Ontario. Canad. Jour. Bot. 34: 71–85. Mulligan, G. A. & Moore, R. J. (1961). Natural selection among hybrids between Carduus acanthoides and C. nutans in Ontario. Canad. Jour. Bot. 39: 269–279. 1. Capitula more or less cylindrical; corolla with 5, more or 2. less equal lobes 1. Capitula more or less globose to hemispherical; corolla 3. distinctly 2-lipped with one entire and one 4-lobed 2. Stems with spiny wings right up to the base of the capitula; capitula in clusters of 3–10(–20); involucral bracts thin and transparent on the margins, without a strongly thickened midrib except sometimes near the 1. tenuiflorus apex 2. Stems with interrupted spiny wings and at least some peduncles unwinged distally; capitula in clusters of 1–3; involucral bracts with strongly thickened margins and 2. pycnocephalus midrib for at least the distal half 3. Capitula (20–)30–65 mm in diameter, usually solitary, more or less drooping; involucral bracts lanceolate, 4. narrowed just above the base; corolla 15–25(–28) mm 3. Capitula 15–35 mm in diameter, solitary or in small clusters, more or less erect; involucral bracts linear-lanceolate or linear-subulate and not narrowed just 5. above the base; corolla 12–18 mm 4. Stems usually less than 50 cm; capitula 1–3 on short 5(i). nutans var. litoralis branches 4. Stems up to 100(–150) cm; capitula usually numerous on 5(ii). nutans var. nutans long branches 5. Spines of stem wings up to 5 mm; capitula 25–35 mm in diameter; inner involucral bracts 1-veined in the distal 3. acanthoides quarter

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5. Spines of stem wings up to 3 mm; capitula 15–25 mm in diameter; inner involucral bracts faintly 3-veined in the 6. distal one-fifth 6. Stems 30–45 cm; capitula few and congested 4(i). crispus subsp. multiflorus var. glareicola 6. Stems up to 150 cm; capitula on long branches 4(ii). crispus subsp. multiflorus var. multiflorus

Section 1. Leptocephali Rchb. fil. Capitula cylindrical, deciduous when ripe. Corolla tube widened above into an ellipsoid cup 1.0–1.5(–2.0) mm, with 5 more or less equal lobes. 1. C. tenuiflorus Curtis Slender Thistle Annual or biennial herb with a stout tap-root and fibrous side-roots. Stems 1-several, (3–)7–75(–120) cm, yellowishgreen, solid, triangular, with a continuous toothed wing on each angle up to the base of the capitulum, the teeth up to 12 mm, triangular, with a creamy-green apical spine up to 5 mm and a triangular base, more or less arachnoid-hairy especially in the upper part, leafy, usually only branched in the upper one-third, but sometimes in upper two-thirds, the branches erect-ascending. Leaves yellowish- to medium green with a broad, pale midrib on the upper surface, slightly paler beneath, basal in a rosette, dying early, the lamina 5–18 × 2–8(–9) cm, elliptical or oblanceolate in outline, acute-spinose at apex, pinnatifid, the lobes broadly ovate, acute at apex, with the spine up to 5 mm, pinnately divided, and each lobule with a spine, narrowed to the base; cauline similar but gradually decreasing in size upwards and decurrent at the sessile base; all sparsely hairy on upper surface, arachnoid-hairy beneath. Capitula 4–17 mm in diameter, cylindrical, sessile, solitary or in compact clusters of 3– 10(–20). Involucral bracts in several rows, 14–16 × 1.5–3.0 mm, yellowish-green, with a darker, purplish-green centre and the inner with a thin scarious margin, triangularovate-lanceolate, narrowed into a creamy-green spine up to 1.5 mm which is outwardly curved at the apex, with the midrib sometimes raised near the apex, glabrous or the outer arachnoid-hairy. Flowers 10–15 mm, all tubular, corolla usually mauve or pinkish-rose, sometimes white, deeply divided into 5 equal linear lobes. Receptacle flat, deeply pitted, with dense bristles. Achenes 4–5 mm, pale brown, shining, obovoid-oblong, swollen and smooth, the apical prominence shortly stipitate, clavate and entire; pappus 10–13 mm, dirty white, of many rows of scabrid (not plumose) hairs united proximally and deciduous. Flowers 6–8. Visited by bees. 2n = 54. Native. Waysides, rough and open ground and shingle. Locally frequent round the coasts of Great Britain and Ireland except in north and west Scotland; a few scattered localities inland where it is often a wool alien. Western Europe eastwards to Holland and Italy, including islands of the western Mediterranean; naturalised in Scandinavia. A member of the Suboceanic Southern-temperate element. 2. C. pycnocephalus L. Plymouth Thistle Annual or biennial herb with a stout tap-root. Stems 15–120 cm, yellowish-green, solid, triangular, with a discontinuous toothed wing on each angle which is absent from just below

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at least some of the capitula, the teeth up to 5 mm, triangular, with a creamy-green apical spine up to 5 mm and a triangular base, more or less arachnoid-hairy and greyish-tomentulose above, leafy, usually only branched in the upper one-third, but sometimes lower, the branches erect-ascending. Leaves yellowish- to medium green, with a broad, pale midrib on the upper surface, slightly paler beneath; basal in a rosette, dying early, the lamina 7–20 × 1.5–8.0 cm, oblanceolate, oblong-lanceolate or oblong in outline, spinose-acute at apex, pinnatifid, the lobes ovate, acute-spinose at apex with the spine up to 12 mm, pinnately divided and each lobule with a spine, narrowed to the base; cauline similar but gradually decreasing in size upwards, decurrent at the sessile base; sparsely hairy on the greenish upper surface, grey arachnoid-lanuginose beneath. Capitula 7–15 mm in diameter, cylindrical, solitary or in clusters of 2–3, sessile or on peduncles up to 100 mm. Involucral bracts in several rows, 3–20 × 1–3 mm, greenish or purplish, the inner with scarious margins, lanceolate-subulate, spine-tipped at apex, glabrous or arachnoid-hairy, with thickened margins and midrib for at least the distal half. Flowers 10–14 mm, all tubular, the corolla rose-purple, divided into 5, equal, linear lobes. Receptacle flat, deeply pitted, with dense bristles. Achenes 4–5 × 1.8–2.0 mm, oblong, strongly compressed, the apical prominence small, shortly stipitate, clavate and entire; pappus 10–14 mm, tinged brownish, of many rows of scabrid (not plumose) hairs united proximally and deciduous. Flowers 6–8. 2n = 54. Introduced. Established on a cliff at Plymouth Hoe in Devon since 1868 but now in very small quantity; elsewhere a wool and bird-seed casual. Native of southern Europe from Spain to the Balkans; Canary Islands; North Africa; western Asia. Our plant is subsp. pycnocephalus which occurs throughout much of the range of the species. Section 2. Carduus Capitula depressed-globose or more or less globose, persistent. Corolla tube widened above into an oblong cup (2–)2.5–5.0 mm, distinctly 2-lipped with one entire and one 4-lobed. 3. C. acanthoides L. Broad-winged Thistle C. crispus auct. et CTM (1987); C. polyacanthos Schreb. nom. illegit.; Polycantha acanthoides (L.) Hill Biennial herb with a slender tap-root. Stem 30–150 cm, pale yellowish-green, with continuous, undulate, spinous wings up to 8 mm wide, up to or nearly up to the capitula, the broadly triangular teeth with an apical spine up to 5 mm, arachnoid-hairy, leafy, with long branches. Leaves yellowish-green, with a pale midrib on upper surface, paler beneath; basal with lamina 20–60 × 6–20 cm, oblong to oblong-lanceolate in outline, acute at apex, pinnatifid, the terminal segment lanceolate, long-acute at apex and dentate, the lateral segments 6–8 pairs, broadly ovate, spinose-acute at apex, spinose-dentate with shorter, more slender spines round the rest of the margin, without spaces between the segments, all veins prominent beneath, narrowed at base to short, winged petioles; cauline gradually becoming smaller up the stem, lanceolate or elliptic-lanceolate in outline, the segments often 3-lobed with the middle lobe longest, or in

the upper leaves with segments much narrower, narrowed to a sessile, decurrent base; all sparsely arachnoid-hairy. Capitula 25–35 mm in diameter, erect, more or less globose, sessile, more or less erect, shortly peduncled and solitary or in small clusters; peduncles arachnoid-hairy. Involucral bracts 15–18 × 1.5–2.0 mm, the outer and middle imbricate, or patent to slightly deflexed, linear-lanceolate and long-acute at apex, the inner middle obtuse and contracted into the spine at the apex, the midrib raised throughout its length, arachnoid-ciliate and minutely verruculose on the back, the inner longer than the middle and 1-veined in the distal quarter. Flowers 16–18 mm, all tubular, the corolla usually reddish purple, sometimes white, 2-lipped with one entire and one 4-lobed. Receptacle flat, deeply pitted, with dense bristles. Achenes 3–4 mm, pale brown, oblong, compressed, with fine wrinkles and a small, sessile, sinuatelobed apical prominence; pappus 11–13 mm, pale brownish, of many rows of simple but scabrid hairs. Flowers 6–8. 2n = 16, 20, 22. ?Introduced. The present status is not known, but there are old records, probably as casuals and it has occurred recently in several Cambridgeshire sites. Much of Europe, but absent from most of the south-west and only an introduction in the north; Caucasus; widely naturalised elsewhere. 4. C. crispus L. Welted Thistle C. acanthoides auct. et Dandy (1958) et CTM (1987). Biennial herb with a slender tap-root. Stem 30–150 cm, pale yellowish-green, with a continuous, undulate, spinous wing up to 6 mm wide up to, or nearly up to the capitula, the triangular teeth with a spine at the apex up to 3 mm, sparsely arachnoid-hairy, leafy, usually with long branches. Leaves yellowish-green with a pale midrib on upper surface, paler beneath; basal with lamina 20–65 × 6–20 cm, elliptical or oblong-elliptical in outline, acute at apex, pinnatifid, the terminal segments lanceolate, long-acute at apex and dentate, the lateral segments 6–8 pairs, broadly ovate, spinous-acute at apex, spinous-dentate with shorter, more slender spines round the rest of the margin and without spaces between the segments, all veins prominent beneath, narrowed at base to a short, winged petiole; cauline gradually becoming smaller upwards, lanceolate to oblong in outline, but segments often 3-lobed with the middle lobe longest, or in the upper leaves with segments much narrower, narrowed to a sessile, decurrent base. all glabrous or nearly so, or with short hairs on the veins beneath, or sparsely arachnoid-hairy. Capitula 15–25 mm in diameter, erect, more or less globose, usually in dense clusters of 3–5, occasionally solitary; peduncles very short, arachnoid-hairy. Involucral bracts 7–10 × 1.0–1.2 mm, imbricate, or the outer and middle slightly recurved at apex, the outer and middle linear-subulate, verruculose in the proximal one-third, with the midrib raised in the distal two-thirds and arachnoid-ciliate, the inner longer than the outer, faintly 3-veined in the distal one-fifth, often purplish, with a weak, slender spine at apex and arachnoid-hairy. Flowers 12–15 mm, all tubular, the corolla usually reddishpurple, sometimes white, 2-lipped with one entire and one 4-lobed. Receptacle flat, deeply pitted, with dense bristles. Achenes 3–4 mm, pale brown, oblong, with fine wrinkles and a small, sessile, sinuately lobed apical prominence;

6. Cirsium pappus 8–12 mm, pale brownish, of many rows of simple but scabrid hairs. Flowers 6–8. Visited by many bees, hoverflies and Lepidoptera. 2n = 16. (i) Var. glareicola P. D. Sell Stems 30–45 cm. Capitula few and congested. (ii) Var. multiflorus (Gaudin) DC. Stems up to 150 cm. Capitula on long branches. Native. Hedgerows, waysides, grassy and waste places, cultivated ground, especially set-aside, dunes and shingle. Frequent in Great Britain north to central Scotland, scattered records in Ireland and western and northern Scotland. Europe northwards to about 71◦ N in Scandinavia; Siberia; Caucasus; naturalised in eastern Asia; North America and elsewhere. A member of the Eurosiberian Temperate element. Our plant is subsp. multiflorus (Gaudin) Franco (C. multiflorus Gaudin; C. crispus subsp. occidentalis Chass. & Ar`enes; C. acanthoides subsp. multiflorus (Gaudin) Nyman) which occurs in west and north-west Europe. The common plant is var. multiflorus. Var. glareicola occurs on shingle and sand by the sea. The two names C. acanthoides L. and C. crispus L. were correctly transposed according to their types by Dandy (1958), and this was accepted by CTW (1987). We have followed Flora Europaea (1976) and Stace (1997) and believe the best solution would be to conserve the names in this sense with new types, as that is the sense in which they have been used in the great bulk of the literature. × nutans = C. × dubius Balb. C. × orthocephalus auct.; C. polyacanthus Schleich., non Lam. This hybrid has the capitula intermediate in size, often 2–4 on a branch and narrowly drooping, the involucral bracts are narrower and less strongly spinous than in C. nutans but more spreading than in C. crispus and the achenes are mostly abortive. Good achenes are sometimes found and in Canada hybrid swarms occur, plants showing a complete range of variation from one species to the other. 2n = 16–22. Native. Scattered records in Great Britain north to Yorkshire. Widespread in Continental Europe. 5. C. nutans L. Musk Thistle Ascalea nutans (L.) Hill A usually biennial herb. Stems 20–100(–150) cm, yellowish-green, sometimes striated brownish-purple or whitish, erect, with 4 wings which are interruptedly spinosetoothed, the teeth 5–10 mm, greyish-green, triangular or palmate and with spines up to 8 mm, arachnoid-hairy, densely so under the capitula where it is devoid of wings, leafy, simple or branched from above halfway, the branches long and sometimes longer than the main stem. Leaves with the lamina of basal elliptical, sinuate and narrowed into a stalk-like base, all cauline at anthesis, widely spaced and spreading, 2–30 × 0.8–8.0 cm, greyish-green with a pale midrib on the upper surface, slightly paler beneath, elliptical, oblong or oblong-lanceolate in outline, spinousacute at apex, pinnately divided almost to the midrib, the lobes 2–11 pairs, triangular-ovate or palmate with an apical brownish spine up to 8 mm, the interlobe area with a broad wing, sessile, sparsely arachnoid-hairy on both surfaces and

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woolly on the prominent vein beneath. Capitula (20–)30– 65 mm in diameter, hemispherical, usually solitary at the ends of branches or rarely 2–4 in a loose corymb; peduncles more or less drooping, densely arachnoid-hairy. Involucral bracts 5–32 × 2–3 mm, green, flushed deep purplish-brown, lanceolate, the outer and median spreading or reflexed, the inner erect, all narrowed just above the base to a spinousacute, arachnoid-hairy apex. Flowers 15–25(–28) mm, all tubular, the corolla reddish-purple, with a musky scent, 2lipped with one entire and one 4-lobed. Receptacle flat, with dense rigid hairs. Achenes 3–5 mm, pale brown, obovoidoblong, grooved, glabrous; pappus 13–24 mm, pale brown or whitish, of many rows of denticulate hairs. Flowers 5–8. Visited by bees, hoverflies and Lepidoptera. 2n = 16. (i) Var. litoralis P. D. Sell Stems usually less than 50 cm. Capitula 1–3, on short branches. (ii) Var. nutans Stems up to 100(–150) cm. Capitula usually numerous, on long branches. Native. Grassy and bare places, waysides, rough ground, arable fields, heaths, dunes and shingle up to 500 m, mostly on calcareous gravelly or sandy soils. Locally frequent in the Channel Islands and Great Britain north to southern Scotland, rare elsewhere in Scotland and Ireland and sometimes only casual. Europe northwards to about 67◦ N in Fennoscandia; Siberia; Caucasus; Turkey; North Africa; naturalised in North America, Australasia and elsewhere. A member of the Eurosiberian Temperate element. Our plant is subsp. nutans. Var. nutans is the common plant. Var. litoralis is found on maritime shingle and sand. (Carduus nutans L. × Cirsium arvensis (L.) Scop. has been tentatively recorded from Herefordshire and Carduus nutans L. × Cirsium vulgare (Savi) Ten. tentatively recorded from Lincolnshire, but both are almost certainly errors.) 6. Cirsium Mill. Ixine Hill; Tetralix Hill, non Griseb. nom. conserv. Biennial to perennial herbs. Stems erect, with or without spiny wings. Leaves alternate, denticulate to deeply lobed, spiny or at least with bristle-pointed teeth. Capitula solitary or in corymbs or dense clusters. Involucral bracts in many rows, simple, linear-subulate to ovate, spine-tipped to mucronate or acuminate at apex, generally with a more or less distinct resin duct. Flowers all tubular, bisexual or female. Corolla tubular, usually purple, rarely white, deeply 5-lobed, the lobes sometimes unequal. Anthers tailed; filaments hairy. Style with long, linear branches which are dorsally hairy and with a ring of hairs beneath the branches. Receptacle flat, with numerous bristly hairs. Achenes oblong, or obovoid-oblong, compressed, the truncate apex with an annular margin surrounding a subconical central projection; pappus of several rows of plumose hairs. Contains about 250 species in Eurasia, North and East Africa and North America, and is widely introduced elsewhere. C. rivulare (Jacq.) All. has been recorded as a casual garden escape.

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Dandy, J. E. (1969). Cirsium acaule Scop. Watsonia 7: 167. Druce, G. C. (1914). Der Formenkreis des Cirsium eriophorum (L.) Scop. in Europa by F. Petrak. Rep. Bot. Soc. Exch. Cl. Brit. Isles 3: 361–362. Everet, S. (1993). Cirsium tuberosum (L.) All. Tuberous Thistle in Gillam, B. (Edit.) The Wiltshire flora, pp. 83–90. Newbury. Grime, J. P. et al. (1988). Comparative plant ecology. London. [C. arvense, C. palustre and C. vulgare.] Hult´en, E. (1958). The amphi-atlantic plants and their phytogeographical connections. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 7: 102–103, 162–163. [C. heterophyllum and C. palustre.] Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 13: 290–291. [C. arvensis.] Klinkhamer, P. G. L. & DeJong, T. J. (1993). Biological flora of the British Isles. Cirsium vulgare (Savi) Ten. (Carduus lanceolatus L.; Cirsium lanceolatum (L.) Scop., non Hill). Jour. Ecol. 81: 177–191. Pigott, C. D. (1968). Biological flora of the British Isles. Cirsium acaulon (L.) Scop. (C. acaule (L.) Weber, Carduus acaulos L.; Cnicus acaulis (L.) Willd.). Jour. Ecol. 56: 597–612. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [C. tuberosum.] 1. 1. 2. 2. 3. 3. 4. 4. 5. 5. 6. 6. 7. 7. 8.

8.

9. 9. 10. 10. 11. 11.

2. Leaves with rigid hairs on upper surface 4. Leaves glabrous or with soft hairs on upper surface 1. eriophorum Stem not winged 3. Stem with discontinuous spiny wings Stems up to 50 cm; capitula few, congested on short 2(i). vulgare var. litorale branches Stems up to 150(–300) cm; capitula numerous, on 2(ii). vulgare var. vulgare spreading branches 5. Corolla yellow 6. Corolla purple or rarely white Upper part of stem with only very reduced distant 5. erisithales leaves; capitula drooping Upper part of stem with large yellowish-green leaves 7. oleraceum exceeding the erect capitula Biennial with a tap-root; stems continuously spiny 7. winged Perennial with at least short rhizomes; stems not winged 8. or base of leaf shortly decurrent Stems up to 150 cm; capitula numerous, on long 9(i). palustre var. palustre branches Stems up to 50 cm; capitula few, congested, with very 9(ii). palustre var. ferox short peduncles Capitula few to numerous; distal part of corolla about half as long as proximal narrow part, lobed for more 9. than three-quarters of the way to the base Capitula usually solitary, rarely up to 3; distal broad part of corolla about as long as proximal narrow part, lobed 13. about halfway to the base Leaves with a flat lamina which is subentire or 10. shallowly undulate lobed 11. Leaves 3-dimensional and deeply lobed Leaves green, but sometimes arachnoid-hairy beneath 10(iv). arvense var. integrifolium Leaves white or grey tomentose beneath 10(v). arvense var. vestitum Stems up to 50 cm; capitula few, in a condensed, 10(i). arvense var. maritimum unbranched inflorescence Stem up to 120(–150) cm; capitula numerous, in a 12. branched inflorescence

10(ii). arvense var. arvense 12. Leaves deeply obtuse-lobed 12. Leaves deeply and narrowly acute-lobed 10(iii). arvense var. horridum 13. Stem absent or up to 10 cm, or if up to 30 cm then more 8. acaule than 1 well-developed leaf near the top 13. Stem more than 10 cm with only distant much reduced 14. leaves in upper quarter 14. Cauline leaves widened to an amplexicaul base; 6. heterophyllum involucre mostly more than 20 mm 14. Cauline leaves narrowed to base, not or scarcely 15. amplexicaul; involucre less than 20 mm 15. Roots not swollen; lower cauline leaves not or rather 3. dissectum shallowly lobed, white or grey on underside 15. Some roots swollen into tubers; lower cauline leaves 4. tuberosum deeply lobed, green on underside

Section 1. Eriolepis (Cass.) Dumort. Eriolepis Cass. Leaves with spine-like appressed hairs on upper surface. Flowers bisexual or the outermost functionally male or sterile. Corolla limb 5-fid to about halfway to base. Pappus shorter than or equalling the corolla. 1. C. eriophorum (L.) Scop. Woolly Thistle Carduus eriophorus L.; Cnicus eriophorus (L.) Roth.; Tetralix eriophora (L.) Hill; Cynara eriocephala Stokes nom. illegit. Biennial herb with a tap-root and fibrous side-roots. Stems (40–)60–150(–200) cm, pale yellowish-green, erect, markedly striate, robust, arachnoid-hairy, the hairs with swollen bases, not winged, branched above, leafy to the top. Leaves with lamina 5–40 × 4–15 cm, medium green on upper surface, greyish beneath, broadly oblong or oblonglanceolate in outline, acute at apex, deeply pinnatisect, the segments linear or oblong, with a rigid, yellow, apical spine (1–)5–12(–25) mm, the interlobe areas wide and near the midrib winged, sometimes additional long spines on the margin and veins beneath, with numerous, yellow, appressed, spine-like hairs on the upper surface and margin and dense grey tomentum on the lower surface, but with the midrib clear. Capitula few, 40–90 mm in diameter, in a loose racemose panicle or corymb; peduncle densely arachnoidhairy. Involucral bracts in numerous rows, 10–50 × 2–3 mm, pale yellowish-green, linear-lanceolate to lanceolate, acute or acuminate at apex with a spine (0.5–)1–4(–5) mm, the outer patent to recurved, the median ones with or without an apical appendage, all arachnoid-hairy. Flowers all tubular, the corolla 25–44 mm, reddish-purple, deeply 5lobed. Receptacle flat, pitted, with dense bristly hairs. Achenes 4.5–6.0 mm, buff mottled with black, obovoid-oblong; pappus 20–33 mm, of many rows of whitish plumose hairs. Flowers 7–9. Visited by long-tongued bees and Lepidoptera. 2n = 34. Native. Grassland, open scrub, roadsides and banks on calcareous soils. Locally frequent in Great Britain north to Co. Durham. Central Europe from France, Belgium and Holland to the north Balkan peninsula and Upper Volga. A member of the European Temperate element. A very variable species in which many subordinate taxa have been described. Our plant has been named subsp. britannicum Petr., but not enough good material from

6. Cirsium elsewhere in Europe has been seen to make a decision on its validity. × vulgare = C. × grandiflorum Kitt. C. × gerhardtii Sch. Bip. This hybrid has shortly decurrent, slightly clasping leaves, capitula intermediate in size between those of the parents and less arachnoid-hairy than C. eriophorum. It is partially fertile and apparently backcrosses with C. eriophorum when the offspring are difficult to distinguish from that species. It occurs rarely in England north to Yorkshire. It is also recorded widely in Continental Europe. 2. C. vulgare (Savi) Ten. Spear Thistle Carduus vulgaris Savi; Carduus lanceolatus L.; Cirsium lanceolatum (L.) Scop., non Hill; Ascalea lanceolata (L.) Hill; Cnicus lanceolatus (L.) Willd.; Cynara lanceata Stokes nom. illegit. Biennial herb with a long tap-root. Stem (20–)50– 150(–300) cm, pale yellowish-green, sometimes tinted purplish, winged, the wings medium green, interrupted, and with unequal, lanceolate to ovate, spine-tipped teeth, furrowed, sparsely arachnoid-hairy, leafy, with long branches above. Leaves dull medium green on upper surface with a pale midrib which is sometimes tinted brownish-purple, greyish-green beneath; basal with lamina 15–30 × 6–10 cm, lanceolate-obovate or elliptic-oblong in outline, longacute at apex, deeply pinnatifid, the terminal segment long linear-lanceolate or lanceolate, acute at apex with a yellow spine, entire and with very fine spinules on the margin, the lateral segments pointing in various directions, broadly ovate, trilobed, each lobe with a yellow spine at the apex and the margins finely spinulose, area between the lobes near the midrib narrowly winged and finely spinulose, narrowed into a short, winged petiole; cauline becoming gradually smaller, similar to basal but sessile and decurrent for the whole internode; all arachnoid-hairy, sparsely above, more so beneath. Capitula 30–50 mm in diameter, ovoidoblong, solitary or 2–3 in a pedunculate cluster; peduncles short or long, pale green, arachnoid-hairy. Involucral bracts in numerous rows, 20–40 × 0.5–1.5 mm, green, linearlanceolate, long acute to acuminate at apex to a yellowish spine 2.0–3.5 mm, more or less arachnoid-hairy. Flowers 26–36 mm, all tubular, reddish-purple, deeply divided into 5 linear lobes at apex. Receptacle flat, pitted, with numerous bristly hairs. Achenes 3.5–5.0 mm, yellow streaked with black, oblong, compressed, the truncate apex with a distinct, annular margin surrounding a subconical central projection; pappus 20–30 mm, pale brownish-white, of several rows of plumose hairs. Flowers 7–10. Visited chiefly by longtongued bees, hoverflies and Lepidoptera. 2n = 68, 102. (i) Var. litorale P. D. Sell Stems up to 50 cm. Capitula few, congested on short branches. (ii) Var. vulgare Stems up to 150(–300) cm. Capitula numerous, on spreading branches. Native. Field margins, grassy and waste places, waysides and gardens, dunes and shingle. Common through-

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out the British Isles. Var. vulgare is the widespread plant. Var. litorale occurs on maritime shingle and dunes. Very widespread in Europe, north to the Arctic circle, and western Asia; North Africa; naturalised in North and South America, South Africa and Australasia. A member of the Eurasian Temperate element. Section 2. Cirsium Section Chamaeleon DC. Leaves without spine-like hairs on upper surface. Flowers bisexual or the outermost male or sterile. Corolla limb 5fid for about halfway to base. Pappus shorter than, rarely equalling the corolla. 3. C. dissectum (L.) Hill Meadow Thistle Carduus dissectus L.; Carduus pratensis Huds.; Cnicus pratensis (Huds.) Willd.; Carduus anglicus Lam.; Cirsium anglicum (Lam.) DC.; Cirsium lanceolatum Hill nom. illegit.; Cnicus anglicus (Lam.) C. C. Gmel.; Cnicus britannicus Druce nom. illegit.; Cirsium pratense (Huds.) Druce, non DC. Perennial herb with a short, obliquely ascending stock, cylindrical roots and short stolons. Stems 15–18 cm, pale yellowish-green, erect, terete, striate, unwinged, arachnoidhairy, usually unbranched, usually with few, small, bractlike leaves above the middle. Leaves green and sparsely arachnoid-hairy on upper surface, sparsely to densely white arachnoid-lanate-tomentose beneath; basal with lamina 12– 25 × 1.5–3.0 cm, oblong-elliptical or elliptic-lanceolate, acute at apex, sinuate-dentate or slightly pinnatifid, with soft prickles, the longest on the teeth or lobes which are broadly triangular, narrowed to a long petiole; cauline 3–5, like the basal but more oblong and semiamplexicaul with basal auricles. Capitula 20–30 mm in diameter, usually solitary, rarely up to 3; peduncles arachnoid-hairy, short. Involucral bracts in numerous rows, 10–20 × 2–3 mm, pale green, lanceolate, appressed, the outer spine-tipped, the inner acuminate at apex, sparsely arachnoid-hairy. Flowers all tubular, bisexual, the corolla 15–25 mm, dark reddish-purple, deeply 5lobed. Receptacle flat, pitted, with dense bristles. Achenes 3–4 mm, pale brown, obovoid, smooth; pappus 10–20 mm, pure white, of several rows of plumose hairs. Flowers 6–8. 2n = 34. Native. Fens, bog margins and wet meadows on peaty soil up to 500 m. Local in England and Wales north to Yorkshire and Merionethshire; very local in south-west Scotland; throughout Ireland. Western Europe from Spain to Holland and north-west Germany. A member of the Oceanic Temperate element. × palustre = C. × forsteri (Sm.) Loudon Cnicus forsteri Sm.; Carduus forsteri (Sm.) Bab. This hybrid has the leaves decurrent, sinuate-pinnatifid or pinnatipartite with lobed, weakly spinose segments and is arachnoid-hairy beneath. Capitula are solitary or 2–3 together and bear shortly spinose, arachnoid-hairy involucral bracts. Not uncommon throughout the range of C. dissectum. It has also been recorded for France and Holland.

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4. C. tuberosum (L.) All. Tuberous Thistle Carduus tuberosus L.; Cnicus tuberosus (L.) Roth Long-lived perennial, non-stoloniferous herb with an obliquely ascending stock and swollen, fusiform roots. Stems 20–100 cm, medium green heavily suffused brownish-purple, erect, grooved, unwinged, arachnoidhairy, leafy chiefly below the middle, branched above. Leaves 4–25 ×1–8 cm, greyish-green on upper surface, slightly paler beneath, basal large and petiolate, the cauline gradually decreasing in size and sessile, elliptical in outline, acute at apex, deeply pinnatifid to pinnatisect, the terminal segment small, the lateral oblong or ovate, with 2–3(–5), oblong to elliptic-lanceolate, sometimes dentate lobes, the basal narrowed to a winged petiole, the upper semiamplexicaul; all sparsely arachnoid-hairy on upper surface, densely so beneath, intermixed with short simple eglandular hairs. Capitulum 25–30 mm in diameter, solitary at the ends of branches; peduncles with 1–2 bracts, long, arachnoid-hairy. Involucral bracts in several rows, 4–9 × 1.5–2.5 mm, the outer greyish-green, the inner greyishgreen heavily suffused purplish, triangular-ovate to oblonglanceolate, obtuse at apex, some mucronate, appressed, slightly arachnoid-hairy. Flowers all tubular, slightly sweet scented, the inner about 15 mm, the outer about 23 mm, all pinkish-purple, deeply divided into 5 narrowly linear lobes. Styles pinkish-purple. Receptacle convex, with numerous, white, bristly hairs. Achenes 3–4 mm, pale brown, obovoid; pappus 11–16 mm, greyish-white, of several rows of plumose hairs. Flowers 6–8. Visited by bumblebees and butterflies. 2n = 34. Native. Dry, ungrazed or lightly grazed ancient calcareous grassland. Very local in Wiltshire, where it grows with C. × medium at almost all its localities, Dorset and Glamorganshire. It became extinct in Cambridgeshire in 1974, following ploughing, but was planted in the same area from local stock in 1987. West and central Europe, eastwards to southern Sweden, Saxony, Tirol and northern Italy. A member of the Suboceanic Southern-temperate element. 5. C. erisithales (Jacq.) Scop. Yellow Thistle Carduus erisithales Jacq.; Cnicus erisithales (Jacq.) L.; Cirsium ochroleucum DC.; Cirsium glutinosum Lam. Perennial herb with obliquely ascending stock. Stem (30–) 60–120(–150) cm, pale yellowish-green, ridged, erect, arachnoid-hairy, usually sparingly branched, leafless above the middle or with a few, small, bract-like leaves. Leaves medium green on upper surface, much paler beneath; basal and lower cauline with lamina up to 30 × 18 cm, oblong-elliptical to broadly elliptical in outline, more or less acute at apex, flat, pinnatisect, the segments numerous, oblong or oblong-lanceolate, obtuse or acute at apex, dentate to lobed and with weak marginal spines up to 2 mm, semiamplexicaul and auriculate at base; all shortly hairy. Capitula 16–22 mm in diameter, solitary or in apical clusters of 2–3(–5), nodding; peduncles short, arachnoidhairy. Involucral bracts in numerous rows, 5–20 × 2–3 mm, pale green, darker towards apex, lanceolate, the spinose tips usually patent. Flowers all tubular, the corolla 14– 20(–22) mm, yellow, 5-lobed. Receptacle flat, pitted, with

dense bristles. Achenes 4–5 mm, obovoid; pappus 15– 20 mm, of several rows of plumose hairs. Flowers 7–9. 2n = 34. Introduced. Naturalised in a disused quarry in Nightingale Valley, Leigh Woods, Somerset since 1980. Native of wet grassland in the mountain regions of Europe from southcentral France and the Carpathians southwards to central Apennines and the Balkan peninsula and in the lowlands of the western margin of Russia. 6. C. heterophyllum (L.) Hill Melancholy Thistle Carduus heterophyllus L.; C. helenioides auct.; Cnicus heterophyllus (L.) Roth; Cynara diversifolia Stokes nom. illegit. Perennial herb with obliquely ascending stock and subterranean stolons. Stems (30–)40–100(–150) cm, pale green, sometimes tinted purplish, erect, ridged, simple or sparingly branched above, not winged, arachnoid-hairy, usually leafless towards the apex. Leaves flat, yellowish-green on upper surface, grey beneath; basal and lower cauline with lamina 20–40 × 4–8 cm, narrowly to broadly elliptical, lanceolate or oblong-lanceolate, acute at apex, usually spinulose-serrate, sometimes more or less pinnatifid with forwardly directed, oblong lobes, narrowed to a spinulose, winged petiole; upper cauline with lamina lanceolate or oblong-lanceolate, long drawn out to an acute apex, spinulose-serrate, rounded, sessile and amplexicaul at base; all more or less glabrous on upper surface, arachnoidtomentose beneath. Capitula solitary or 2–3 in a cluster, 30–50 mm in diameter; peduncles arachnoid-hairy. Involucral bracts in numerous rows, erect, 10–28(–32) × 3–4 mm, green, dark towards apex, lanceolate, weakly spinulose at apex, the outermost acute, the median obtuse with a scarious margin, the inner with a membranous appendage at apex. Flowers all tubular, the corolla (20–)25–30 mm, reddishpurple, rarely white, deeply divided into 5 lobes at apex. Receptacle flat, pitted, with dense bristles. Achenes 4–5 mm, pale brown, obovoid, smooth; pappus (18–)22–26(–32) mm, pure white, of several rows of plumose hairs. Flowers 7–8. Visited chiefly by bees. 2n = 34. Native. Hay meadows, roadsides, scrub, open woodland, streamsides and tall-herb communities on rock-ledges in hilly country. Locally common in Great Britain northwards from Derbyshire and Radnorshire and in Ireland northwards from Co. Fermanagh and Co. Leitrim. Northern Europe and mountains of central Europe; Siberia; introduced in Greenland. A member of the Eurosiberian Boreal-montane element. × palustre = C. × wankelii Reichard Carduus carolorum Howie & Jenner; Cirsium carolorum (Howie & Jenner) Nyman; Cnicus carolorum (Howie & Jenner) Hook. fil.; Cnicus wankelii (Reichard) F. Hanb. This hybrid has the stem branched above, leaves grey-felted beneath, irregularly serrate, lobed or pinnatifid, shortly decurrent and weakly spinous and capitula solitary or clustered. It occurs in scattered localities in Scotland and is probably more common than is recorded. It is widespread in Continental Europe.

6. Cirsium 7. C. oleraceum (L.) Scop. Cabbage Thistle Cnicus oleraceus L. Perennial herb with a rhizome. Stem (20–)50–150(–170) cm, pale yellowish-green, sometimes tinted purplish, markedly ridged, sparsely greyish arachnoid-hairy in the upper part, simple or sparingly branched, leafy. Leaves with lamina 5–30 × 3–15 cm, dull green on upper surface, much paler and greyish-green beneath, flat, broadly ovate to broadly elliptical in outline, acute at apex, slightly to deeply lobed, the terminal and lateral segments lanceolate-oblong or ovate-oblong, acute with a yellow-spined tip at apex, dentate, the teeth with yellow-spined tips and the area between the teeth ciliate with small, ascending, yellowish spines, the interlobe area near the midrib weakly ciliate-spinulose, the upper surface glabrous or with minute pricklets making the surface rough, the under surface glabrous, the lower with short, sparsely spinulose, winged petioles, the upper sessile and auriculate-amplexicaul. Capitula 20–35 mm in diameter, solitary or in clusters of 2–6; peduncles short, ridged and arachnoid-hairy. Involucral bracts in numerous rows, 15–23(−26) × 1.8–2.0 mm, pale yellowish-green, linear-lanceolate, long-attenuate to an acute apex, the outer with a short spine, glabrous. Flowers (14–)18–25 mm, all tubular, pale yellow, rarely reddish, deeply divided into 5, narrowly linear lobes. Receptacle convex, deeply pitted, densely bristly. Achenes 4.0–5.5 mm, pale grey, obovoid, angled; pappus (12–)15–21 mm, whitish, of several rows of plumose hairs. Flowers 7–9. The usually bisexual capitula are freely visited by bees and butterflies. 2n = 34. Introduced. Naturalised since at least 1912 in marshes by the River Tay in Perthshire and since 1978 by a stream in Lancashire. Native of a large part of Europe where it grows in damp meadows and woods but absent from most of the Mediterranean region and the north; central Russia and Siberia. 8. C. acaule Scop. Dwarf Thistle Carduus acaulos L.; Cnicus acaulos (L.) Willd.; C. acaulon (L.) Scop., non C. acaule Scop. Perennial herb with a horizontal, branched, sympodial rhizome from which several, stout, white roots are produced each year. Stem 2–3(–30) cm, with joined hairs, simple or branched. Leaves in a compact rosette of 12–20, spirally arranged, the lamina 10–15(–20) × 2–3 cm, dark green on upper surface, paler beneath, oblong-lanceolate in outline, spinose-acute at apex, pinnatifid, with 8–10 segments on each side, the upper segments with 3–5 lobes each bearing a long, stiff, pale yellow terminal spine and smaller marginal spines, the lowest segments reduced to spines, the lobes on the lower side of each segment curved upwards and overlapping the midrib, nearly glabrous on the upper surface, with sparse, long white hairs on the veins beneath; petiole pale green and purplish at base. Capitula 1–3(–8), 12–20 mm in diameter, ovoid, usually sessile on the rosette. Involucral bracts in numerous rows, 6–30 × 2.0–2.5 mm, green with a narrow, membranous margin or sometimes purplish, the outer ovate, with a minute apical spine, the inner oblanceolate, and obtuse at apex, fringed with short hairs. Flowers 23–35 mm, all tubular, bright purplish-red, rarely pale pink or white, divided into 5 narrow segments at apex,

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1 longer than the other 4. Receptacle flat, covered with long hairs. Achenes 3–5 mm, pale to dark brown, ovate or conical, smooth; pappus 21–27 mm, white, of many rows of plumose hairs, united at the base. Flowers 7–9. Entomophilous, bees being the most important agents. Gynodioecious. 2n = 34. Dwarf plants transplanted into shade or moist soil usually produce a tall, often leafy stem forma caulescens Rchb., but some remain dwarf. Bisexual capitula are usually larger than the all-female ones. Individuals with white or pale pink flowers are rare. Native. Closely grazed pastures especially on chalk or limestone up to 380 m in Derbyshire. Widespread in the outcrops of chalk and Jurassic limestone in south-eastern England, and more sparsely on chalk, oolite and magnesium limestone in Yorkshire, on the Carboniferous limestone in Derbyshire, the Welsh borderland, south Wales and the Mendips and in a few localities on Devonian limestone and other rocks in Devonshire and Cornwall. Spain, north Italy, and the north-west Balkan peninsula to south Scandinavia and Estonia. A member of the European Temperate element. Our plant is subsp. acaule which occurs throughout the range of the species except southern Spain. Closely related taxa, including C. esculentum (Stev.) C. A. Mey. and C. rhizocephalum C. A. Mey., occur from eastern Europe to central Asia and may be only subspecifically distinct. × arvense = C. × boulayi E. G. Camus C. × babingtonii Rouy; Carduus acaulos var. dubius auct.; C. gibsonii H. C. Wats. nom. nud. This hybrid has a creeping rootstock, branched stems 20–60 cm and leaves resembling those of C. arvense. The indumentum contains both jointed and arachnoid hairs. The capitula are numerous on long peduncles with strongly spinose involucral bracts. It has occurred for certain only in Essex, but other records exist for England and Wales. It is recorded also for central Europe and Sweden. Named after Abb´e Jean Nicolas Boulay (1837–1905). × dissectum = C. × woodwardii (H. C. Wats.) Nyman Carduus woodwardii H. C. Watson; Cnicus woodwardii (H. C. Watson) Hook. fil. This hybrid has a creeping rootstock which bears unthickened roots and branched stems. The leaves are deeply pinnatifid, less spinose than in C. acaulon and arachnoid-hairy beneath. The upper part of the stem bears arachnoid and jointed hairs. The capitula are solitary and terminal. The plants are functionally female, but set some seed, the resulting offspring showing no segregation. Known only from near Swindon in Wiltshire where it has persisted for nearly a century. Named after Samuel Pickworth Woodward (1821–65). × palustre = C. × kirschlegeri Sch. Bip. This hybrid has erect stems to 40 cm and the upper part bears arachnoid and jointed hairs. The leaves are pinnatifid, some forming decurrent wings. The few capitula have shortly spinous, slightly arachnoid-hairy involucral bracts. It has been found in a few localities in southern England. Occurs also in widely scattered localities in Europe. Named after Fr´ed´eric Kirschleger (1804–69).

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× tuberosum = C. × medium All. C. × zizianum W. D. J. Koch; C. fraserianum Druce nom. nud. The non-creeping rootstock of this hybrid bears more or less thickened roots and branched stems 20–60 cm which have arachnoid and jointed hairs in their upper parts. The leaves are narrower in outline with broader, closer lobes than in C. tuberosum and the capitula are more elongated. F1 hybrids are more or less intermediate between the parents, but backcrossing produces plants that grade into both. It is known from several places in southern England and Wales particularly on the Wiltshire Downs where it may occur in the absence of C. tuberosum. It is also known from central Europe. × vulgare = C. × sabaudum M. Loehr This hybrid grows up to 30 cm and has strongly spinose margins of the leaves, normally a few spinules on the upper surface of the lobes and a shortly decurrent wing on the stem. The stem and leaves bear jointed and arachnoid hairs. Capitula are about 5 with appressed involucral bracts with squamose spinose tips. Some achenes are apparently fertile. It has occurred in a few localities in England south of Lincolnshire. It is also known for central Europe and Sweden. 9. C. palustre (L.) Scop. Marsh Thistle Carduus palustris L.; Cnicus palustris (L.) Willd.; Polycantha palustris (L.) Hill; Cynara palustris (L.) Stokes Biennial herb with a short, erect, premorse stock and a tap-root. Stem 30–150 cm, erect, furrowed, pale yellowishgreen, sometimes suffused brownish-purple, narrowly but continuously spiny-winged, with arachnoid and jointed hairs, unbranched or with long ascending and arching branches, leafy. Leaves yellowish-green on upper surface, sometimes tinted brownish-purple, paler beneath; basal flat in a rosette, the lamina 10–20 × 5–10 cm, oblanceolate or oblong-lanceolate in outline, acute at apex, pinnatifid, the segments triangular-ovate to narrowly lanceolate or oblonglanceolate, acute at apex, undulate-dentate and spreading with unequal, yellow spines at the ends of segments and teeth, with a short winged petiole; lower cauline often longer, then becoming gradually smaller upwards, and more or less sessile and long decurrent; all with jointed hairs on upper surface and arachnoid-hairy beneath. Capitula numerous, 10–20 mm in diameter, sessile, in a crowded cluster at the end of the stem or at the ends of branches. Involucral bracts in numerous rows, 3–13 × 1.5–2.0 mm, pale yellowish-brown, often flushed purplish-red towards the apex, lanceolate to oblong-lanceolate, narrowed to a spine at apex, slightly arachnoid-hairy, the inner with a subrotund to lanceolate, membranous, ciliate appendage. Flowers all tubular, corolla 10–15 mm, purple, rarely white, 5-lobed. Receptacle flat, pitted, with dense, long bristles. Achenes 3–4 mm, dirty white, of several rows of plumose hairs. Flowers 7–9. Visited by many bees, flies and Lepidoptera, the nectar being more readily accessible than in the other species of Cirsium. 2n = 34. On the whole plants from Great Britain and Ireland are much more spiny than those of Continental Europe and

should perhaps be recognised as a distinct subspecies. They can be divided into two ecotypes. (i) Var. palustre Stem up to 150 cm, with long ascending-arching branches and clusters of capitula at the end of each branch. (ii) Var. ferox Druce Stem up to 50 cm, even more spiny than var. palustre, with capitula in a tight cluster at the end of the stem and not branched. Native. Marshes, hedgerows, woodland rides and clearings and moorland pasture up to 760 m. Common throughout Great Britain and Ireland. Europe to 68◦ N in Scandinavia; western Asia; North Africa; widely naturalised in North America and New Zealand. A member of the Eurosiberian Boreo-temperate element. Var. ferox is the plant of upland pasture in the north and west. Plants in all other habitats are var. palustre. White-flowered variants are found in both varieties, sometimes as whole populations, and are said to be more frequent in upland areas. × tuberosum = C. × semidecurrens Richt. This hybrid has more or less thickened roots, the stems are about 60 cm and unbranched and the cauline leaves are pinnatipartite and decurrent. The 6–15 capitula are borne on felted peduncles, the involucral bracts appressed and spine-tipped. It is sterile. Recorded with C. tuberosum at Nash Point in Glamorganshire; formerly occurred at Great Ridge in Wiltshire and recently recorded at another Wiltshire site in the same area. It is also recorded from France, Germany, Switzerland and Italy. × vulgare = C. × subspinuligerum Peterm. Intermediate in capitulum size and shape and colour of involucral bracts. Only known from a roadside near Inverliever in Argyll in 1974 and on a roadside at Arthog in Merionethshire in 1986. It is also known from central Europe. Section 3. Cephalonoplos DC. Section Breea (Less.) W. D. J. Koch; Breea Less. Leaves without spine-like hairs on upper surface. Flowers unisexual. Corolla limb 5-partite almost to the base. Mature pappus much longer than the corolla. 10. C. arvense (L.) Scop. Creeping Thistle Serratula arvensis L.; Carduus arvensis (L.) Hill; Ixine arvensis (L.) Hill; Cnicus arvensis (L.) Roth; Cynara repens Stokes nom. illegit.; Breea dioica Less.; Cephalonoplos arvensis (L.) Fourr. Perennial, sometimes more or less dioecious herb, initially with a slender tap-root, later producing far-creeping, whitish, lateral roots which bear numerous, adventitious non-flowering and flowering shoots. Stems 20–120(–150) cm, pale yellowish-green through various degrees of brownish-purple tinting to suffused deep brownish-purple, erect, channelled and ridged, unwinged but often with some pricklets in the lower part, glabrous or slightly arachnoidhairy above, leafy, simple or variously branched. Leaves dull medium yellowish-green with a pale midrib on upper surface, paler beneath or rarely whitish; basal not in a

7. Onopordon compact rosette and usually dead at anthesis, the lamina 10–20 × 2–5 cm, lanceolate, oblong-lanceolate or oblong in outline, acute at apex, usually more or less pinnatifid, the segments broadly to narrowly triangular-ovate, pointing in various directions making the leaf more or less 3dimensional, rounded to acute at apex and spinose-dentate with weak to stout spines 1–10 mm, the interlobe area near the midrib with narrow to broad wings, rarely shallowly or scarcely lobed and flat and weakly spinose-dentate, narrowed at base to a short, winged petiole; cauline gradually decreasing in size upwards, similar to basal but sessile, semiamplexicaul and not or slightly decurrent; all more or less glabrous, arachnoid-hairy or densely tomentose beneath. Capitula 15–25 mm in diameter, solitary or in terminal clusters of 1–5, together forming an irregular condensed to open corymb; peduncles short, yellowish-green or purplish, more or less arachnoid-hairy. Involucral bracts in many rows, 4–17 × 1.2–2.0 mm, pale green, often suffused purplish especially near the apex, paler near the margins, the outer ovate and obtuse at apex with a short, spreading spine, the inner lanceolate and narrowed to an erect, acute apex, all appressed and glabrous or slightly arachnoid-hairy. Flowers (10–)13–18 mm, all tubular, usually dull purple, sometimes whitish, deeply divided into 5 linear lobes, strongly honeyscented. Receptacle flat, deeply pitted, with dense, long bristles. Achenes 3–4 mm, dark brown, oblong, smooth; pappus (15–)20–30 mm, pale brownish, of many rows of plumose hairs united at the base. Flowers 7–9. Male capitula have abortive ovaries, but sometimes ripen a few achenes; in the female capitula the anthers are abortive. The strong scent attracts a great variety of insects. 2n = 34. (i) Var. maritimum Fr. Stems usually solitary, up to 50 cm. Leaves green beneath, deeply lobed, the lamina 3-dimensional, the undivided area near the midrib up to 10 mm, strongly spinose. Capitula few, in a condensed, unbranched inflorescence. (ii) Var. arvense Stems often more than 1, up to 120(–150) cm. Leaves green beneath, deeply obtuse-lobed, the lamina 3-dimensional, the undivided area near the midrib up to 15 mm, strongly spinose. Capitula usually numerous in an open, branched inflorescence. 2n = 34. (iii) Var. horridum Wimm. & Grab. Stem often more than 1, up to 120(–150) cm. Leaves green beneath, deeply, narrowly and acute-lobed, the lobes strongly spine-tipped, the lamina 3-dimensional, the undivided area near the midrib up to 10 mm. Capitula numerous in an open, branched inflorescence. (iv) Var. integrifolium Wimm. & Grab. Serratula setosa Willd.; Cirsium setosum (Willd.) M. Bieb.; C. arvense var. setosum (Willd.) C. A. Mey.; Carduus arvensis subsp. setosus (Willd.) Hook. fil.; Cnicus arvensis subsp. setosus (Willd.) Hook. fil.; Cirsium arvense var. mite Wimm & Grab.; Carduus arvensis var. latifolius Bab.; Cirsium arvense subsp. setosum (Willd.) Iljin Stems often more than 1, up to 120(–150) cm. Leaves green, but sometimes arachnoid-hairy beneath, subentire to undulate-lobed and flat, the lobes obtuse, the undivided

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area by the midrib up to 35 mm, weakly spinose-margined. Capitula numerous in an open, branched inflorescence. (v) Var. vestitum Wimm. & Grab. Serratula incana S. G. Gmel.; Cirsium incanum (S. G. Gmel.) Fisch. ex M. Bieb.; Cirsium arvense var. incanum (S. G. Gmel.) Ledeb.; Cirsium arvense subsp. incanum (S. G. Gmel.) Iljin Stems often more than 1, up to 120(–150) cm. Leaves white or grey tomentose beneath, subentire to shallowly undulate-lobed and flat, the lobes obtuse, the undivided area by the midrib up to 35 mm, weakly spinose-margined. Capitula numerous, usually in a branched inflorescence. 2n = 34. Native. Grassy, cultivated and waste places, waysides, hedgerows, dunes and shingle. Common throughout Great Britain and Ireland. Europe to 68◦ 50 N in Scandinavia; Asia; North Africa; introduced in North America and Australasia. A member of the Eurasian Temperate element which has now attained Circumpolar Temperate distribution. Var. maritimum is the plant of dunes and shingle by the sea. It is probably common on the coasts of western Europe. Var. arvense is the inland weed of arable, grassy and waste places. When it was abundant in cornfields, before herbicides almost eradicated it there, it formed small vegetative clones recognisable from one another by the amount of brownish-purple colouring, amount of spines and number and size of capitula. It is now most common in open waste places and is not so abundant in grassland. Var. horridum occurs in scattered localities, apparently most often near the sea. It occurs also in Continental Europe. Var. integrifolium and var. vestitum are apparently introduced casuals but widely represented in British herbaria. Plants of var. integrifolium in fens may be native or they may be a slightly different plant for which the name var. latifolium is available. They often do not appear to flower or have only a few capitula. Babington himself finally put these two varieties together and we have found difficulty in distinguishing them. Both var. integrifolium and var. vestitum seem to be native of south and south-east Europe and western Asia and may be good subspecies. × palustre = C. × celakovskianum Knaf C. × mixtum Druce nom. nud. This hybrid has the stems winged below with decurrent, sinuate-lobed leaves and more or less naked above, the capitula are small and clustered or shortly stalked, the corolla-tube is longer than the corolla-limb which is 5-lobed for three-quarters of its length. Native. Occurs where C. palustre grows on dry ground or where C. arvense grows at the edge of marshy places. There are very scattered records in Great Britain and Ireland where it is uncommon. It is also recorded for central Europe. Named after Ladislav Joseph Celakovsky (1834–1902). Galactites tomentosa Moench occurs as a bird-seed casual. 7. Onopordon L. Biennial herbs. Stems erect, with spiny wings. Leaves alternate, dentate to shallowly lobed, arachnoid-hairy, strongly spiny. Capitula solitary or in terminal clusters of 2–3.

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Involucral bracts in several rows, simple, linear or linearlanceolate, strongly spine-tipped. Flowers all tubular and bisexual. Corolla tubular, usually pale purple, rarely white, with 5 linear lobes. Anthers with terminal, subulate appendages and short basal tails; filaments papillose-hairy. Style with long branches, which are dorsally hairy and with a ring of hairs below the branches. Receptacle flat, pitted, without bristles but the pits fringed with teeth. Achenes subtetragonal, glabrous; pappus of many rows of denticulate hairs, united at base, deciduous. Close to Carduus, but lacks receptacular bristles. About 50 species in Europe, the Mediterranean region and western Asia. O. macranthum Schousboe has been recorded as a wool casual. 1. Leaves densely arachnoid-hairy on both surfaces 1. acanthium 1. Leaves nearly glabrous on upper surface, sparsely 2. nervosum arachnoid-hairy beneath

1. O. acanthium L. Cotton Thistle O. vulgare Gray nom. illegit. Biennial herb with a stout tap-root. Stems 45–300 cm, greyish-green, stiffly erect, with 4 wings, the wings up to 60 mm wide and undulate-spinose, the spines up to 15 mm, arachnoid-hairy, simple to much branched, the branches rigid and ascending, leafy. Leaves with lamina 4–45 × 1–30 cm, greyish-green with a pale midrib on both surfaces, ovate-oblong, elliptic-oblong, ovate or lanceolate, spinose-acute at apex, divided halfway to midrib into 2–8 pairs of lobes, the lobes with large mammiform spinosetipped teeth, sessile at base and decurrent to the stem wings, arachnoid-hairy on both surfaces. Capitula 10–80 mm in diameter, hemispherical, solitary or in clusters of 2–3 at the ends of stems or branches; peduncles winged, armed and clothed like stem. Involucral bracts in several rows, 8–20 × 1.0–1.5 mm, dull medium green with a yellowish or brown tip, linear or linear-lanceolate, with a yellowish spine up to 5 mm at apex, arachnoid-hairy, the outer spreading, the inner erect. Flowers 22–25 mm, all tubular, pale purple, rarely white, with 5 linear lobes. Receptacle deeply pitted, the pits with toothed membranous margins, without scales or hairs. Achenes 4–5 mm, marbled greyish-black, compressed, subtetragonal, glabrous; pappus 7–9 mm, pale reddish, of denticulate hairs united into a ring at base, deciduous. Flowers 6–9. Visited chiefly by bees. 2n = 34. Introduced or possibly native. Fields, waste ground, heaths and rough places, especially on sandy soils. Locally frequent in England especially the south-east and perhaps native in East Anglia; scattered records in Wales, Scotland and the Channel Islands. Known since at least the sixteenth century. Europe northwards to southern Scandinavia and central Russia; western Asia; introduced in North America and Australasia. A member of the Eurosiberian Temperate element. Our plant is subsp. acanthium which occurs throughout the range of the species. 2. O. nervosum Boiss. Reticulate Thistle Biennial herb with a stout tap-root. Stems up to 270 cm, yellowish-green, stiffly erect, with 4 wings up to 20 mm

wide with an undulate margin, with densely reticulate veins and spines up to 10 mm, with rather dense, short hairs, leafy. Leaves with lamina up to 50 × 20 cm, gradually decreasing in size upwards, green with whitish veins on upper surface, greyish-green beneath, oblong-lanceolate, acute at apex, pinnatifid, with 6–8 pairs of lobes, the lobes triangular, spinose-acute at apex and with 2–4 spines at the sides, sessile, nearly glabrous on upper surface, sparsely arachnoidhairy beneath, densely reticulate-veined. Capitula 30–50 mm in diameter, conical-ovoid, solitary or in clusters of 2–6 at the ends of branches; peduncles winged, armed and clothed like the stem. Involucral bracts in several rows, pale green, 8–30 × 4–6 mm, ovate or ovate-lanceolate, acute at apex, with a rigid spine up to 4 mm, glabrous or nearly so. Flowers 32–35 mm, all tubular, pink, with 5 linear lobes. Receptacle with deep pits with dentate margins, glabrous. Achenes 4–5 mm, greyish-brown, obovoid; pappus 8–10 mm, of numerous hairs united into a ring at the base. Flowers 6–9. 2n = 34. Introduced. Garden escape naturalised in a few places in southern England in old sand and chalk pits. Native of south and central Spain and south and central Portugal. 8. Cynara L. Perennial herbs. Stems erect, arachnoid-hairy. Leaves alternate, pinnately lobed, arachnoid-hairy beneath, spiny. Capitula large, solitary and terminal or in small corymbs. Involucral bracts in numerous rows, appendage separated from lower part by a constriction, at least the outer with an apical spine. Flowers all tubular, bisexual. Corolla blue, lilac or whitish, deeply 5-lobed. Anthers spurred and tailed; filaments hairy. Style with long branches, which are dorsally hairy and with a ring of hairs below the branches. Receptacle flat and fleshy, with rigid, white bristles. Achenes oblong, 4-angled, glabrous; pappus of many rows of plumose hairs. About 10 species in the Mediterranean region and western Asia. Phillips, R. & Rix, M. (1993). Vegetables. London. 1. Leaves and involucral bracts strongly spiny 1(i). cardunculus var. cardunculus 1. Leaves and involucral bracts spineless 1(ii). cardunculus var. sativa

1. C. cardunculus L. C. horrida Aiton Perennial herb. Stems up to 2 m, pale yellowish green, erect, robust, conspicuously sulcate, thinly arachnoid-hairy. Leaves numerous, extending up the stems almost to the capitulum, 30–40 × 10–15 cm, green on upper surface, greyish-green beneath, oblong-lanceolate in outline, acute at apex, pinnately lobed, the lobes up to 7 × 1.5 cm, narrowly oblong and caudate-acuminate at apex, the margins with deltoid, spine-tipped lobes, the rhachis and leaf-base with digitate spine-clusters, or without spines, sparsely hairy becoming glabrous on the upper surface, thinly arachnoid-hairy beneath. Capitula 80–110 mm in diameter, solitary and terminal or in sparsely branched corymbs. Involucral bracts in numerous series, the outer about 10 × 10 mm, with a broad, flattish base, abruptly constricted above into a rigid, suberect terminal spine

10. Serratula 20–30 mm, the inner with a broad, ovate, acute, mucronate or very shortly spinose, purplish, reflexed apical appendage 10–13 × 5–7 mm. Flowers all tubular, slightly exceeding the involucre, the tube 30–40 mm, slender, glabrous, the limb about 14 × 2 mm, the 5 lobes about 12.0 × 0.6 mm, linear and erect. Receptacle flat, pitted, the walls densely rigidhairy with long, white bristles. Achenes 6–8 × 3–4 mm, brown-spotted, oblong, 4-angled, glabrous; pappus 25– 40 mm, of copious, slightly brown, free or almost free, rigid plumose hairs. Flowers 8–9. (i) Var. cardunculus Cardoon Leaves and involucral bracts strongly spiny. 2n = 34. (ii) Var. sativa Moris Globe Artichoke C. scolymus L.; C. cardunculus var. scolymus (L.) Fiori Leaves and involucral bracts spineless. 2n = 34. Introduced. A long-persistent garden escape, recorded from 1947 to 1978 at Bawdsey in Suffolk and cultivated as a vegetable in southern England. Probably native of the Mediterranean region, where it has been long grown as a vegetable and was certainly known to the ancient Greeks and Romans. The fleshy leaf-bases of var. cardunculus are tied up and blanched to be eaten raw or cooked, like celery. The dried flowers can be used as a substitute for rennet and in parts of France are used in the making of a soft, curd cheese. The immature flowers of var. sativa are eaten and later the fleshy bases of the involucral bracts and the thickened receptacle. The young flowers stalks and leaves are also edible. 9. Silybum Adans. nom. conserv. Mariana Hill nom. rejic. Biennial herbs. Stems erect, glabrous or slightly arachnoidhairy, not spiny. Leaves alternate, shallowly to deeply lobed, white-veined or variegated, glabrous or nearly so, strongly spiny. Capitula solitary at the ends of branches. Involucral bracts in many rows, the outer and median with spine-tipped lateral lobes or teeth and a strong apical spine. Flowers all tubular and bisexual. Corolla tubular, usually purplish, rarely white, 5-lobed. Anthers shortly tailed; filaments connate into a papillose tube. Style with long linear branches which are dorsally hairy and with a ring of hairs below the branches. Receptacle flat, with dense rigid hairs. Achenes obovoid-oblong, compressed, glabrous; pappus of many rows of rigid, scabridulous hairs united proximally. Two species in the Mediterranean region and eastwards to Afghanistan. 1. S. marianum (L.) Gaertn. Milk Thistle Carduus marianus L.; Mariana mariana (L.) Hill nom. illegit.; Mariana lactea Hill; Carduus lactifolius Stokes nom. illegit.; S. mariae Gray nom. illegit. Biennial herb with a stout tap-root. Stems 25–250 cm, erect, shortly branched above, conspicuously sulcate, unwinged, glabrous or slightly arachnoid-hairy, not spiny. Leaves pale green, white-veined or variegated; basal with lamina 9– 50 × 5–25 cm, oblanceolate in outline, shallowly to deeply sinuate-lobed, with yellowish-white spines up to 8 mm along the margins, petiolate; cauline diminishing in size upwards, sessile, shortly decurrent, with distinct, spinose

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basal auricles; all glabrous. Capitula 25–50 mm in diameter, solitary at the end of branches, erect or drooping; peduncles long, sometimes with a few small leaf-like bracts. Involucral bracts in many rows, 8–15 × 6–10 mm, broadly oblong, the outer and middle pectinate-spinose with a stout apical spine 25–70 mm. Flowers all tubular, the corolla usually purplish, rarely white, 5-lobed. Receptacle flat, fleshy, with dense, rigid hairs. Achenes 6–8 × 2.5–4.0 mm, shiny black with grey spots, obovoid-oblong, compressed, glabrous; pappus of rigid, scabridulous or remotely plumed hairs up to 15 mm, united proximally. Flowers 6–8. Visited by various bees. 2n = 34. Introduced since at least the seventeenth century. Frequent casual, sometimes naturalised in waste places and rough ground especially near the coast. Scattered throughout Great Britain and Ireland north to Ross-shire. South Europe from Spain to Russia; North Africa; Caucasus; Near East; introduced in central Europe, North and South America and Australasia. So named because the marks on the leaves were supposed to have resulted from the milk of the Virgin Mary falling on them. 10. Serratula L. Perennial gynodioecious or more or less dioecious herbs. Stems erect, sparsely hairy, not spiny. Leaves alternate, serrate to pinnately lobed, not spiny. Capitula in a rather lax panicle or subsessile in a compact cluster. Involucral bracts in many rows, simple, not spiny. Flowers bisexual or the marginal female. Corolla tubular, usually purple, rarely white, deeply 5-lobed. Anthers without or with short tails; filaments papillose-hairy. Styles with long, linear branches, which are dorsally hairy and with a ring of hairs below the branches. Receptacle flat, with dense chaffy scales. Achenes oblong, slightly compressed, glabrous; pappus of many rows of free, simple hairs, the outermost much shorter than the inner. Differs from Saussurea only in the anthers having short or no tails and the pappus hairs not plumose. About 70 species in Europe, North Africa and central Asia. 1. Stem up to 100 cm; capitula few to numerous, usually in an open inflorescence, sometimes in a condensed one 1(i). tinctoria var. tinctoria 1. Stem 9–12(–20) cm; capitula 1–3, on short peduncles 1(ii). tinctoria var. reducta

1. S. tinctoria L. Saw-wort Carduus tinctorius (L.) Scop.; S. inermis Gilib. nom. illegit. Perennial, glabrous, gynodioecious or more or less dioecious herb with a short, stout, more or less erect stock. Stem 4–100 cm, pale yellowish-green, often tinted purplish especially near the base, more or less erect, slender, wiry, grooved, with some arachnoid hair or nearly glabrous, simple or branched above. Leaves with lamina 3–25 × 1–5 cm, yellowish-green on upper surface, paler beneath, lanceolate to ovate-lanceolate or oblong-lanceolate in outline, obtuse or acute at apex, entire to lyrate-pinnatifid or almost pinnate, the lobes narrowly lanceolate or oblong-lanceolate and acute or obtuse at apex, the leaf or the lobes serrulatespinulose, the teeth with fine bristle tips, glabrous or with a

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few, scattered hairs, the basal and lower cauline short to long petiolate, the upper sessile. Capitula 12–15 mm in diameter, the female are larger than male, solitary, loosely corymbose or crowded; peduncles 10–40 mm, straight, sparsely to densely mealy or minutely glandular-hairy. Involucral bracts in many rows, 3–15 × 1–2 mm, pale green, flushed in the upper part with reddish-purple, with minute glandular hairs and more or less arachnoid-hairy; outer ovate, acute at apex, with a gradual transition to the linear-oblong, acute inner ones. Flowers all tubular, equalling the involucral bracts, the corolla usually purple, rarely white, deeply 5lobed. Receptacle flat; scales numerous and chaffy. Achenes 5–6 mm, pale brown, oblong, slightly compressed, glabrous; pappus 5–6 mm, yellowish, of many rows of stiff, rough, deciduous simple eglandular hairs, all free at the base, the outermost shortest. Flowers 7–9. Visited by flies and bees. 2n = 22. A very variable species, especially in continental Europe. Var. reducta grows in distinct populations and is a recognisable dwarf ecotype which retains its characters in cultivation. Most of our plants are tall and have an open inflorescence, but some plants, which have been referred to var. monticola (Boreau) Syme (S. monticola Boreau), have a congested one and are said to occur in more upland localities. It is not known if they occur as individual plants or in distinct populations. (i) Var. tinctoria Stem up to 100 cm. Cauline leaves numerous. Capitula few to numerous, usually in an open inflorescence, sometimes in a condensed one. (ii) Var. reducta Rouy Stem 9–12(–20) cm. Cauline leaves few. Capitula 1–3, on short peduncles. Native. Wood margins, clearings and rides and open, unimproved grassland on basic soils over chalk and limestone; also on cliff-tops and rocky streamsides. Local in Britain north to south-west Scotland and decreasing in many areas; formerly in Co. Wexford in Ireland. Central Europe from northern Spain, central Italy and the north Balkan peninsula northwards to southern Scandinavia, Estonia and central Russia; Siberia; Algeria. A member of the European Temperate element. Var. tinctoria is the common British plant. Var. reducta occurs on cliff-tops in short grassland in the Isle of Wight, Cornwall, Pembrokeshire and Caernarvonshire and should be looked for elsewhere in such habitats. Intermediates sometimes occur when there is an inland population close by. It was originally described from France. In Continental Europe the amount of variation in the species is much greater. Amberboa moschata (L.) DC. has been recorded as a casual garden escape and Mantisalca salmantica (L.) Briq. & Cavill. as a grain and bird-seed casual. 11. Acroptilon Cass. Perennial herbs with rhizomes. Stem erect, very leafy, not spiny. Leaves alternate, entire or remotely dentate, not spiny. Capitula solitary on branches. Involucral bracts in many rows, the outer with a broadly ovate to lunulate appendage,

the inner with an oblong appendage, not spiny. Flowers bisexual. Corolla tubular, pink, rarely white, deeply 5lobed. Anthers tailed; filaments papillose-hairy. Style with long, linear branches which are dorsally hairy and with a ring of hairs below the branches. Receptacle flat, with numerous bristles. Achenes obovoid; pappus of free, barbed hairs, soon falling. A single species in eastern Europe and south-west and central Asia and introduced in North America and Australia. 1. A. repens (L.) DC. Russian Knapweed Centaurea repens L.; Centaurea picris Pall. ex Willd.; A. picris (Pall. ex Willd.) C. A. Mey.; Serratula picris (Willd.) M. Bieb. Perennial herb with rhizomes. Stems numerous, 40–70 cm, greyish-green, ridged, erect, more or less arachnoid-hairy, with numerous, erect-ascending branches, leafy. Leaves with lamina 1–8 × 0.2–1.5 cm, greyish-green on upper surface, only slightly paler beneath; lower linear or linearlanceolate, more or less acute at apex, with widely spaced teeth, the teeth spreading, ascending or cusped and usually more or less acute, more or less sessile; upper mostly linear, acute and entire, sometimes with 1–2 teeth; all minutely hairy. Capitula 20–25 mm in diameter, numerous, solitary at the ends of branches; peduncles arachnoid-hairy, rough with small projections. Involucral bracts in many rows, 6– 20 × 3–4 mm, green with a brown margin, the outer subrotund with a broadly ovate to lunulate appendage, the inner ovate to lanceolate with an oblong appendage, arachnoidhairy, not spiny but with an entire or lacerate margin. Flowers 20–22 mm, all tubular and bisexual, pink or rarely white, deeply 5-lobed. Receptacle flat, with numerous whitish bristles. Achenes 3–4 mm, obovoid; pappus 6–8 mm, of free, barbed hairs, soon falling. Flowers 7–9. 2n = 26. Introduced. Naturalised on a railway bank near Hereford station, Herefordshire since 1959. Native of east Europe and south-west Asia. 12. Centaurea L. Hippophaestum Gray nom. illegit. Annual to perennial herbs. Stems erect to spreading, not spiny. Leaves alternate, simple and entire to pinnately divided, not spiny. Capitula solitary or in groups of 2–3 at the ends of branches. Involucral bracts in many rows, with a distinct, scarious, toothed or spiny apical appendage. Flowers in the centre bisexual, the outer often sterile, often longer than the inner. Corolla tubular, purple to pink or blue, white or yellow, deeply 5-lobed. Anthers tailed; filaments papillose-hairy. Style with long, linear branches, which are dorsally hairy and with a ring of hairs below the branches. Receptacle more or less flat, with dense bristles. Achenes more or less compressed; pappus absent, or of many rows of simple to toothed, free hairs, sometimes also with some scales. About 500 species, widespread in Eurasia, North and East Africa and particularly numerous in the Mediterranean region and the near East, with a few species in South America and widely introduced elsewhere. C. dealbata Willd. has been recorded as a casual garden escape.

12. Centaurea Many attempts have been made to divide this large genus into many smaller ones, but none of the arrangements has become common practice. The subgenera here recognised show where these divisions would take place. Within these subgenera hybridisation is widespread and causes great difficulty in interpretation of taxa. Britton, C. E. (1923). Centaurea scabiosa L.: varieties and a hybrid. Rep. Bot. Soc. Exch. Cl. Brit. Isles 6: 767–773. Britton, C. E. (1921). British forms of Centaurea jacea. Rep. Bot. Soc. Exch. Cl. Brit. Isles 6: 163–173. Britton, C. E. (1922). British Centaureas of the nigra group. Rep. Bot. Soc. Exch. Cl. Brit. Isles 6: 406–417. Britton, C. E. (1927). Centaurea pratensis. Rep. Bot. Soc. Exch. Cl. Brit. Isles 8: 149–152. Elkington, T. T. & Middlefell, L. C. (1972). Population variation within Centaurea nigra L. in the Sheffield region. Watsonia 9: 109–116. Grime, J. P. (1988). Comparative plant ecology. London. [C. scabiosa.] Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 13: 220–221 [C. cyanus.] Marsden-Jones, E. M. & Turrill, W. B. (1954). British Knapweeds. London. Ockendon, D. J., Walters, S. M. & Whiffen, T. P. (1969). Variation within Centaurea nigra L. Proc. B.S.B.I. 7: 549–552. Palmer, R. C. (1995). Centaurea jacea not yet extinct in Britain? B.S.B.I. News 69: 75. Valentine, D. H. (1979). Presidential Address, 1978: Experimental work on the British flora. Watsonia 12: 201–207. fide C. scabiosa. 1. Apical area of appendages to involucral bracts with more 2. than one spine 1. Apical area of appendages to involucral bracts without 8. spines or with only one spine 3. 2. Flowers pink or purple, rarely white 5. 2. Flowers yellow 6. calcitrapa 3. Leaves not decurrent at base 4. 3. Leaves decurrent at base 4. Appendanges of involucral bracts with 5 palmately 7. aspera arranged spines 4. Appendages of involucral bracts with more than 11. diluta 5 curved fimbriae 5. hyalolepis 5. Leaves not decurrent at base 6. 5. Leaves decurrent at base 10. eriophora 6. Capitula more than 15 mm in diameter 7. 6. Capitula less than 15 mm in diameter 7. At least some spines of apical appendages of involucral bracts more than 10 mm; corolla without sessile glands; pappus about twice as long as body of inner achene 8. solstitialis 7. Spines of apical appendage of involucral bracts all less than 10 mm; corolla with sessile glands; pappus about as 9. melitensis long as body of achene 8. Appendages of involucral bracts scarious, variously toothed or fimbriate, with 1–few terminal spines 11. diluta 8. Whole of appendage of involucral bracts surrounded by 9. fimbriae or toothed lobes 9. Appendages of outer involucral bracts more than 10 mm 12. macrocephala wide; flowers yellow

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9. Appendages of outer involucral bracts less than 10 mm 10. wide; flowers pink or blue to purple, rarely white 10. Appendage gradually blended (decurrent) with and not clearly differentiated from the lower part of the 11. involucral bract 10. Appendage clearly distinguished from the lower part of 17. the involucral bract by a waist 11. Easily uprooted annuals; basal leaves absent when 12. flowering 11. Deeply rooted perennials; basal leaves or non-flowering 13. shoots present 4. diffusa 12. Capitula 8–11 mm in diameter; flowers pink 12. Capitula 15–30 mm in diameter; flowers blue, pinkish, 17. cyanus mauve or white 14. 13. Leaves more or less entire or crenate-dentate 15. 13. Leaves deeply lobed 14. Leaves entire, strongly decurrent at base; outer flowers 16. montana blue 14. Leaves entire or crenate-dentate, not decurrent; flowers 1(iii). scabiosa var. succisiifolia reddish-purple 3. paniculata 15. Capitula 15–20 mm in diameter 16. 15. Capitula 20–70 mm in diameter 16. Plant up to 20 cm; capitula solitary or few on short 1(i). scabiosa var. nana peduncles 16. Plant up to 120 cm; capitula numerous on long branches 1(ii). scabiosa var. scabiosa 2. cineraria 17. Leaves densely felted at least beneath 18. 17. Leaves not densely felted 18. Appendage of outer involucral bracts denticulate to 13. jacea pectinate-lacerate but not fimbriate 19. 18. Appendage of outer involucral bracts fimbriate 20. 19. Involucre 9–14 mm in diameter 23. 19. Involucre 15–20 mm in diameter 21. 20. Outer flowers scarcely longer than inner 22. 20. Outer flowers much longer than inner 21. Plant up to 30 cm; capitula few on short peduncles 14(a,i). debeauxii subsp. nemoralis var. minima 21. Plant up to 80 cm; capitula numerous on long branches 14(a,ii). debeauxii subsp. nemoralis var. nemoralis 22. Plant up to 30 cm; capitula few on short peduncles 14(b,i). debeauxii subsp. thuillieri var. pumila 22. Plant up to 80 cm; capitula numerous on long branches 14(b,ii). debeauxii subsp. thuillieri var. thuillieri 23. Outer flowers scarcely longer than inner 15(a). nigra subsp. nigra 23. Outer flowers much longer than inner 15(b). nigra subsp. rivularis

Subgenus 1. Lopholoma (Cass.) Dobrocz. Sagmen Hill; Colymbada Hill.; Lopholoma Cass. Perennial herbs. Stems erect, branched above the middle. Lower leaves pinnatisect. Capitula comparatively large. Involucral bracts with appendages decurrent and fimbriate. Pappus present. 1. C. scabiosa L. Greater Knapweed Jacea scabiosa (L.) Lam.; Lopholoma scabiosa (L.) Cass.; Sagmen scabiosa (L.) Hill; Sagmen jacoidea Hill nom. illegit.; Phrygia major Gray nom. illegit.

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Perennial herb with a stout, woody, oblique, branching stock, enclosed about in fibrous scales. Stems 10–120 cm, erect, pale green, striate, slightly angled, with numerous, pale simple eglandular hairs especially below, usually branched above the middle, leafy. Leaves with lamina 3– 25 × 2–10 cm, gradually decreasing in size upwards, dull medium green on upper surface, paler beneath, oblanceolate, obovate or elliptical in outline, obtuse or subacute at apex, usually 1–2-pinnatisect, rarely simple, the lobes linear-lanceolate, obtuse or acute at apex and toothed or deeply divided again, the lower petiolate, the upper sessile, with numerous, short, subrigid, pale simple eglandular hairs on both surfaces and the margins. Capitula 30–70 mm in diameter, solitary at the end of branches, ovoid-globose; peduncles long, striate, angled, glabrous or with a few, very short simple eglandular hairs. Involucral bracts in many rows, 9–11 × 3–4 mm, ovate, glabrous or with arachnoid hairs; appendages 1–2(–3) × 1–2 mm, not covering the green bracts, triangular-ovate, brown or black, decurrent, with pale brown fimbriae, inconspicuous and paler on inner bracts. Flowers all tubular, usually purple, rarely white, the outer 25–30 mm, deeply divided into 5 linear spreading lobes and spreading, the inner 14–16 mm, divided into 5 erect lobes. Styles purple. Receptacle flat, pitted, with numerous to dense bristles. Achenes 4.0–5.5 mm, greyish or brownish, oblanceolate, truncate at apex, shortly hairy; pappus 4–5 mm, greyish or brownish-white, of many rows of stiff simple hairs. Flowers 6–9. Freely visited by various bees and flies. 2n = 20. (i) Var. nana P. D. Sell Plant up to 20 cm. Leaves deeply pinnatisect. Capitula few, on short peduncles. (ii) Var. scabiosa Leaves 1- to 2-pinnatisect, but very variable. Capitula on long peduncles. (iii) Var. succisiifolia E. S. Marshall ex C. E. Britton Leaves simple, subentire or crenate-dentate. Capitula on long peduncles. Native. Dry grassland, hedgebanks, roadsides and clifftops especially on calcareous soils, up to 320 m in Derbyshire. Locally common in Great Britain and Ireland north to northern England and central Ireland, very local in mainland Scotland. Europe northwards to 70◦ N in Scandinavia, Finland and Karelia; Caucasus; naturalised in North America and New Zealand. A member of the Eurosiberian Temperate element. Var. scabiosa is the common plant. Var. succisiifolia is recorded from Coalbackie and Melness on the east and west coasts of Tongue Bay in Sutherland and on cliffs in Glamorganshire. It should be looked for elsewhere on coasts. It reproduces itself from seed, but intermediates have been found with var. scabiosa. Var. nana occurs on cliffs by the sea on the Gower Peninsula, Glamorganshire and at Bettyhill in Sutherland and should also be looked for elsewhere. Subgenus 2. Acrolophus (Cass.) Dobrocz. Acosta Adans.; Acrolophus Cass.; Centaurea Section Acrolophys (Cass.) DC.

Annual to perennial herbs. Stems erect and much branched. Leaves pinnatisect with narrow segments. Capitula comparatively small. Involucral bracts with appendages hardly decurrent and fimbriate. Pappus present or absent. Section 1. Pannophyllum Hayek Perennial herb. Stems herbaceous, the branches not spiny. Leaves not rigid or spiny. Capitula ovoid to campanulate. Involucral bracts with lower fimbriae of appendages usually free. Flowers purple, glandular. 2. C. cineraria L. Ragwort Knapweed Jacea cineraria (L.) Delarbre Perennial herb. Stems 50–75 cm, pale green, erect, ribbed, with dense silvery indumentum, branched above, leafy. Leaves greyish-green on upper surface, silvery beneath; basal and lower cauline with lamina 8–20 × 3–6 cm, ovate in outline, deeply 1- to 2-pinnatisect, the segments linear to oblong, petiolate; upper cauline smaller, 1(–2)-pinnatisect, sessile; all densely felted beneath, sparsely so on upper surface. Capitula 13–16 mm in diameter, ovoid to campanulate, solitary at the end of branches; peduncles grey-felted. Involucral bracts in many rows, the outer and middle ovate to lanceolate, 3(–5)-veined, with sparse, flattened, arachnoid hairs, the appendages erect, fimbriate, not covering the bracts, membranous, narrowed at junction with bract and hardly decurrent, the fimbriae 7–10 on each side, pale to dark brown and 1 mm long. Flowers all tubular, the outer slightly longer than the inner, the corolla purple, glandular, deeply 5-lobed. Receptacle flat, with dense, white bristles. Achenes 4–5 mm, oblong, hairy; pappus about 0.5 mm. Flowers 7–9. 2n = 18. Introduced. Established on sea-cliffs in south-west England; also a bird-seed casual. Native of Italy and Sicily. Section 2. Paniculatae (Hayek) Dostal Stems herbaceous, the branches not spiny. Leaves not rigid or spiny. Capitula ovoid. Involucral bracts with lower fimbriae of decurrent appendages free. Flowers purple, not glandular. 3. C. paniculata L. Jersey Knapweed Acrolophus paniculatus (L) Cass.; Jacea paniculata (L.) Lam. Said to be a biennial herb but has a strong, woody root and a branched stock. Stems 10–80 cm, pale green often suffused purplish, ridged, erect, arachnoid-hairy, much branched nearly to the base, the branches ascending, leafy. Leaves with lamina 0.5–8.0 × 0.1–3.0 cm, greyish with woolly and arachnoid hairs, the lower oblong or elliptic-oblong in outline, pinnatisect with linear, more or less acute, entire segments, shortly petiolate, the upper linear, acute and entire or trilobed with the lobes linear and sessile. Capitula 15– 20 mm in diameter, ovoid, sessile at the ends of branches. Involucral bracts in several rows, 2–10 × 1–2 mm, yellowish with brown appendages, the outer ovate, the inner oblong, shortly arachnoid-hairy, the appendages shortly triangular, said to be acuminate but not spiny at apex but on herbarium sheets appearing to have a short spine, with fimbriae 0.7–1.0 mm, and decurrent. Flowers all tubular, 10–15 mm, purple, the outer slightly larger and sterile, the

12. Centaurea inner bisexual, deeply 5-lobed. Receptacle flat, with dense, white bristles. Achenes 1.8–2.5 mm, ovoid, more or less compressed, glabrous; pappus 0.7–1.0 mm, of scale-like bristles. Flowers 7–9. 2n = 18. Introduced. Naturalised on a hillside at St Ouen’s Bay, Jersey from 1851 to 1981. In Great Britain a rare casual. Native of southern Europe from Portugal to Italy. Our plant seems to be subsp. paniculata which occurs in south France and south and east Spain and north-west Italy. Section 3. Cylindracea (Hayek) Dost´al Stem herbaceous, the branches not spiny. Leaves not rigid or spiny. Capitula cylindrical. Involucral bracts with lower fimbriae of appendages free and with an apical spine. Flowers pink, not glandular. 4. C. diffusa Lam. Small-flowered Star-thistle C. parviflora Sibth. & Sm., non Lam. Annual or biennial herb with a tap-root. Stems 10–50(–60) cm, pale yellowish-green, sometimes slightly suffused brownish-purple, erect, ridged, glabrous and rough or with very short, curled hairs, divaricately much branched, leafy. Leaves dull green on upper surface, paler beneath; lower with lamina 3–5 × 1.0–1.5 cm, oblong-oblanceolate in outline, acute at apex, 2-pinnatisect, the segments linear and acute, shortly petiolate; upper cauline gradually smaller, and less divided, the uppermost linear, dentate, entire and obtuse at apex; all with very short, rigid hairs and gland-dotted. Capitula numerous, 8–11 mm in diameter, cylindrical, solitary at the ends of branches; peduncles ridged, glabrous or nearly so. Involucral bracts in numerous rows, 2–10 × 2– 3 mm, uniform pale yellowish-brown, broad ovate, the appendage with an apical spine 2–3(–5) mm and longpectinate fimbriae on the margins and decurrent at base, glabrous. Flowers all tubular, the corolla pink, deeply 5lobed. Receptacle flat, with long bristles. Achenes about 2.5 mm, obovate; pappus absent. Flowers 7–8. 2n =18. Introduced. Bird-seed and grain casual. Native of southeast Europe, northwards to north-central Ukraine, widely naturalised especially in central Europe. Subgenus 3. Calcitrapa (Heist. ex Fabr.) Hayek Calcitrapa Heist. ex Fabr. Annual or biennial herbs. Stems ascending to erect, much divaricately branched. Leaves pinnatisect, not decurrent. Capitula fairly large. Involucral bracts with appendages not decurrent at base, palmately or pinnately spiny at apex, the apical spine much longer than the others, not fimbriate. Pappus present or absent. 5. C. hyalolepis Boiss. Eastern Star-thistle C. pallescens subsp. hyalolepis (Boiss.) Holmboe; C. pallescens var. hyalolepis (Boiss.) Boiss. Annual or biennial herb. Stems 20–60 cm, pale green, erect, channelled or angled, glabrous or with few, white simple eglandular hairs and glandular hairs in the upper part, usually much branched, the branches spreading, leafy. Leaves greyish-green on upper surface, paler beneath; basal with lamina 4–15 × 2.4–7.0 cm, oblong in outline, acute at apex, coarsely lacerate-pinnatisect or 2-pinnatisect, the segments narrow to rather wide with irregular serrations which ter-

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minate in a minute, hard spine or apiculus, the petiole up to 7 cm, flat and winged; cauline becoming small upwards, oblanceolate, the uppermost almost subtending the capitulum, coarsely pinnatisect, with spinulose-serrulate margins, sessile, not decurrent; all sparsely scabrid-hispidulous on both surfaces. Capitula 6–14 mm in diameter, solitary at the ends of spreading branches. Involucral bracts in several rows, 3–10 × 2–3 mm, pale green with a conspicuous hyaline margin, broadly ovate, the outer terminating in a spinose appendage, the spines usually 10–30 mm, stout, rigid and yellowish and usually with 2–3, short spines on either side of the base, the innermost bracts with a broad, rounded or emarginate, denticulate, decurrent, membranous, hyaline or brownish appendage. Flowers all tubular, 14–16 mm, yellow, glandular-hairy, deeply 5-lobed. Receptacle flat, with dense white bristles. Achenes 2.0–2.5 mm, brown, oblong, glabrous; pappus up to 3 mm, of whitish, unequal bristles, the innermost row much shorter than the others. Flowers 7–9. 2n = 20. Introduced. Grain, wool and bird-seed casual. Few localities in England. Native of south-east Greece, Anatolia, Cyprus, western Syria and southern Iran. 6. C. calcitrapa L. Red Star-thistle Calcitrapa hippophaestum Gaertn.; Calcitrapa stellaris Hill; Hippophaestum vulgare Gray nom. illegit. Biennial herb with a stout tap-root and an erect, branched stock. Stem 15–100 cm, yellowish-green, ascending to erect, grooved, slightly arachnoid-hairy and glandular in upper part, leafy, branched. Leaves greyish-green becoming yellowish-green, gradually decreasing in size upwards; basal and lower with lamina 5–13 × 2–5 cm, lanceolate or ovate in outline, acute at apex, pinnatifid, with distinct lanceolate, acute, entire or remotely serrate lobes and petiolate but withered at anthesis; median pinnatifid with linear-lanceolate lobes, sessile, not decurrent; upper linear or lanceolate, undulate or slightly toothed, sometimes somewhat hastate, sessile; all greyish-lanate when young, later rather short glandular-hairy. Capitula 25–30 mm in diameter, sessile at the ends of branches. Involucral bracts in several rows, 6–8 × 4–5 mm, coriaceous, indistinctly veined, green with a scarious margin, ovate, the appendage with a pale yellow, patent apical spine 10–18 mm which is channelled above and strongly thickened at the base, and with 2–6, pale yellow, ascending basal spines 3–5 mm. Flowers all tubular and similar, 14–16 mm, pale reddish-purple, 5lobed, glandular. Receptacle flat, with dense, white bristles. Achenes 3–7 mm, whitish or with brown mottling, ovoid or oblong, truncate at apex, glabrous; pappus absent, but puberulous at apex of achenes. Flowers 7–9. Visited by bees and flies. 2n = 20. Introduced (although sometimes considered native). Waste and rough ground and waysides on sandy and gravelly ground and on chalk; formerly also a weed of arable fields. Naturalised in a few scattered localities in southern England, especially on chalk downs near the coast in Sussex and decreasing; a casual from wool, bird-seed and other sources elsewhere. South and south-central Europe north to Switzerland and Czechoslovakia; North Africa and Canary Islands; western Asia; widely naturalised beyond its native range but in these sites it is tending to decrease.

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Subgenus 4. Seridia (Juss.) Czerep. Seridia Juss. Perennial herbs. Stems erect or ascending, much branched. Lower leaves pinnatisect; upper decurrent. Capitula small. Involucral bracts with appendages not decurrent, palmately spiny at apex. Pappus present. 7. C. aspera L. Rough Star-thistle C. heterophylla Willd.; Seridia aspera (L.) Sweet; C. isnardii L.; Polyacantha procumbens Gray nom. illegit. Perennial herb with a slender stock and thick tap-root. Stems 20–60 cm, pale yellowish-green, often suffused brownishpurple, erect or ascending, markedly striate or ridged, with few to numerous, short simple eglandular hairs or the bases of hairs or nearly glabrous below, becoming slightly arachnoid-hairy above, with many, slender, spreading branches, leafy. Leaves greyish-green on upper surface, paler beneath; basal numerous, but usually dying before anthesis, the lamina 5–10 × 1.0–2.5 cm, narrowly oblongelliptical or oblong-oblanceolate in outline, obtuse at apex, pinnatisect, the lobes triangular and obtuse at apex, gradually narrowed to a winged petiole; cauline 2–8 × 0.2–3.0 cm, either linear and entire or oblanceolate in outline and pinnatifid like the basal, narrowed at base, sessile and sometimes semiamplexicaul, decurrent; all with numerous, very short or short, pale, stiff hairs on both surfaces and the margins. Capitula 2.5–3.5 mm in diameter, solitary at the ends of branches and subsessile. Involucral bracts in several rows, 3.5–10 × 2.5–3.5 mm, yellowish, leathery, ovate, with spreading or reflexed appendages which have 5, palmately arranged spines up to 3 mm, glabrous or slightly arachnoidhairy. Flowers all tubular, usually pale reddish-purple, rarely white, 5-lobed, the outer 15–17 mm, sterile, a little larger than the bisexual inner. Receptacle flat, with numerous, whitish bristles. Achenes 3–5 mm, greyish-white, with reddish stripes, obovate, more or less compressed; pappus 1–2 mm, of reddish hairs. Flowers 7–9. 2n = 22. Introduced. Naturalised on maritime dunes and in waste places. Known in Guernsey since at least 1788 and Jersey since at least 1839. It is casual or more or less naturalised in a few places in southern England and Wales. Native in south and west France, Spain, Portugal, Corsica, Sardinia and Italy. Our plant is subsp. aspera which occurs throughout the range of the species. Subgenus 5. Solstitiaria (Hill) Dobrocz. Solstitiaria Hill; Centaurea section Mesocentron Hayek Annual to perennial herbs. Stems ascending or erect, branched. Leaves usually decurrent at base, the lower lobed to pinnatifid, the upper entire to dentate. Capitula medium to large. Involucral bracts with appendages not decurrent and palmately spiny at apex, the apical spine usually much longer than the others. Pappus present. 8. C. solstitialis L. Yellow Star-thistle Seridia solstitialis (L.) Sweet; Solstitiaria solstitialis (L.) Hill; Solstitiaria flava Hill nom. illegit.; Leucantha cyanifolia Gray nom. illegit.

Annual or rarely biennial herb. Stems (10–)20–80 cm, pale green, ascending or erect, stiff, striate, with broad, continuous, wavy wings and arachnoid tomentum, branched above, leafy. Leaves greyish-green on upper surface, paler beneath; basal and lower cauline with lamina 6–15 × 1–6 cm, deeply lyrate-pinnatifid, with distant, narrow, entire or toothed lateral lobes and a larger, triangular, acute, denticulate terminal lobe, with a winged petiole; middle and upper cauline lanceolate, 1–6 × 0.2–0.8 cm, acute at apex, more or less entire or sinuate-dentate, sessile and decurrent into the wings of the stem; all with arachnoid hairs on both surfaces. Capitula (5–)7–12 mm in diameter, ovoid-globose, solitary at the ends of branches; peduncles arachnoid-hairy. Involucral bracts in many rows, pale green, ovate, the outer tipped with a slender yellow spine 5–40 mm and bordered basally on either side with 1–2 much shorter lateral spines, the inner elongate, with a rounded or emarginate, hyaline apical appendage. Flowers all tubular, 12–13 mm, yellow, with 5 lobes at apex. Receptacle flat, with long simple eglandular hairs. Achenes dimorphic, 2.0–2.5 mm, oblong-fusiform, the marginal blackish and without a pappus, the inner brown, with a pappus of white, scabridulous bristles up to 4.5 mm. Flowers 7–9. Visited chiefly by bees. 2n = 16. Introduced. Formerly naturalised in arable fields especially Lucerne, now a casual of waste ground mostly from wool and bird-seed. Scattered records, mainly in south and central Great Britain. Native of south and south-east Europe and western Asia; introduced elsewhere in Europe and in New Zealand. 9. C. melitensis L. Maltese Star-thistle Seridia melitensis (L.) Sweet; Solstitiaria melitensis (L.) Hill Annual or biennial herb with a tap-root and fibrous sideroot. Stems (10–)25–80 cm, pale green, erect, ribbed, winged, with sparse to dense arachnoid hairs, branched above, leafy. Leaves greyish-green on upper surface, paler beneath; lower cauline with lamina 5–8 × 0.5–1.5 cm, spathulate to oblanceolate, obtuse at apex, dentate to lyrate-pinnatifid, decurrent at base onto the stems; upper cauline gradually becoming smaller, linear-spathulate to linear-lanceolate, decurrent as lower; all with both short, curved and arachnoid hairs. Capitula (6–)8–12 mm in diameter, broadly ovoid to globose, solitary or in groups of 2–3; peduncles short or absent. Involucral bracts 4– 12 × 3–5 mm, pale yellowish-brown, broadly elliptical or broadly ovate, glabrous or very shortly hairy, veinless, the appendages short, not decurrent, with a patent apical spine 5–8 mm and 1–3 remote, short lateral spines on each side. Flowers all tubular, the outer patent, the corolla yellow, 5lobed and glandular-papillose. Receptacle flat, with long bristles. Achenes about 2.5 mm, obovoid; pappus as long as achene body. Flowers 7–8. 2n = 22, 24, 26. Introduced. Grain, wool, linseed, bird-seed and esparto casual. Native of the Mediterranean region; introduced in Australia and North America. 10. C. eriophora L. Woolly Star-thistle Annual herb with fibrous roots. Stems 20–70 cm, pale green, erect, markedly striate, winged, rather sparsely arachnoidhairy, branching at first from below the terminal capitulum,

12. Centaurea later from lower nodes, leafy. Leaves medium green on upper surface, paler beneath; lower with lamina oblanceolate, obtuse to subacute at apex, incise-dentate to pinnatifid, narrowed at base and decurrent on the stem; upper linear, mucronate at apex, entire or dentate, all with short, rigid hairs on both surfaces and particularly the margins and slightly arachnoid-hairy. Capitula 30–50 mm in diameter, the first opening sessile or nearly so, the remainder solitary at the ends of branches; peduncles long, sparsely arachnoidhairy. Involucral bracts in many rows, pale yellowish-green, ovate-elliptical to lanceolate, the outer with an appendage of a terminal brown spine 15–26 mm, and three lateral spines on each side up to 5 mm, densely arachnoid-hairy. Flowers all tubular, 16–18 mm, yellow, with 5 very narrow lobes at apex. Receptacle flat, with long, silky simple eglandular hairs. Achenes 4.0–4.5 mm, blackish, shortly oblong or ovoid, compressed, hairy; pappus 1.5–2.0 mm, of numerous hairs, crimson at their base. Flowers 7–8. 2n = 24. Introduced. A bird-seed casual. In a few scattered localities. Native of Spain and Portugal. 11. C. diluta Aiton Lesser Star-thistle Perennial herb often behaving as an annual. Stems 20–100 cm, pale green, sometimes tinted purplish, markedly striate, glabrous, much branched, sometimes from the base, leafy. Leaves dull medium green on upper surface, paler beneath; the lamina 2–10 × 0.5–2.5 cm, oblong, lanceolate, lanceolate-oblong, lyrate or linear, mostly obtuse at apex, the lower undulate-dentate, or slightly lobed, the upper more or less entire, more or less amplexicaul with rounded auricles, glabrous on the surfaces, but glandular-punctulate, very shortly ascending ciliate on the margins. Capitula 35–50 mm in diameter, numerous, solitary at the end of branches; peduncles slightly swollen below capitula, striate, rough with small projections. Involucral bracts in several rows, 5–14 × 1.5–5.0 mm, pale yellowish-green, brownish on the margin, oblong to very broadly ovate, emarginate at apex with a rigid, filiform spine in the notch, a membranous, fimbriate-lacerate margin and shortly decurrent at base, glabrous. Flowers all tubular, the outer up to 25 mm, sterile, rose, purple or violet and patent, the inner shorter, bisexual and erect, all with 5 long linear lobes at apex. Receptacle flat, with dense, long bristles. Achenes 2.5–3.0 mm, ovoid, glabrous or hairy on veins; pappus 4.5–5.0 mm, whitish. Flowers 7–10. 2n = 20. Introduced. Casual on waste ground and tips, mainly from bird-seed. Scattered localities mainly in south and central England, but north to Inverness-shire. Native of southwest Spain and north-west Africa.

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Boiss.; Grossheimia macrocephala (Muss. Puschk. ex Willd.) Sosn. & Takht. Perennial herb with a stout root. Stems 50–100 cm, pale yellowish-green, markedly striate, erect with long, stout, pale simple eglandular hairs, usually simple, leafy. Leaves medium yellowish-green on upper surface, paler beneath; lamina 10–20 × 5–6 cm, lanceolate or oblong-lanceolate, gradually narrowed to an acute apex, entire, narrowed at base, the lower petiolate, the upper sessile and shortly decurrent, more or less glabrous on the surfaces with short, pale simple eglandular hairs on the margin and midrib beneath. Capitula 50–70 mm in diameter, hemispherical, solitary at the end of the stem, enveloped by the upper leaves; peduncles swollen below the capitulum, with numerous, long, wavy, pale simple eglandular hairs. Involucral bracts in several rows, 25–30 × 15–20 mm, ovate and green below, with a very large, pale brown scarious appendage which is denticulate to lacerate and ciliate and completely concealing the basal part of the bracts. Flowers all more or less similar, 30–50 mm, all tubular, yellow, divided into 5 linear lobes at apex. Receptacle flat, densely long bristly. Achenes 6.0–7.5 mm, obovate; pappus 7–8 mm, whitish, of scabrid hairs. Flowers 6–8. 2n = 18. Introduced. A garden escape persisting in an abandoned orchard at Belstead and possibly bird-sown in a farmyard at Middleton, both in Suffolk. Native of south-west Asia.

Subgenus 6. Chartolepis (Cass.) Spach Chartolepis Cass. Perennial herb. Stems erect, usually unbranched. Leaves decurrent at base, entire. Capitula large. Involucral bracts with appendage not decurrent and denticulate to lacerate. Pappus present.

Subgenus Jacea (Mill.) Hayek Jacea Mill.; Phrygia (Pers.) Gray Perennial herbs. Stems ascending to erect, branched above. Leaves subentire to pinnatisect. Capitula small to rather large. Involucral bracts with appendages not decurrent, entire to fimbriate. Pappus absent or short. All the taxa in this subgenus seem to be replacing ecologically or geographically and fertile intermediates are common where two taxa grow together. Our native species are C. debeauxii and C. nigra, which hybridise freely when they grow adjacent. This is further complicated by the introduction of C. jacea, mainly in the area of C. debeauxii, but it is felt that only the kind of plant described as C. × monctonii is referable with certainty to C. debeauxii × jacea, and that other plants which have been referred to that hybrid are either part of C. debeauxii or its hybrid with C. nigra. The situation is further complicated by ecological races, particularly in coastal areas. The best method of trying to understand the group would seem to be to first find uniform populations of all the taxa in the field and then try to interpret the hybrid swarms. One of the most important things to do is base the account of C. nigra on a population in the north of England where C. debeauxii is mostly absent. This immediately shows the hybrid nature of much of what is thought to be C. nigra in the south. To work only from herbarium sheets is not helpful. This species and subspecies in this account are based on J. Dost´al’s account in Flora Europaea (1976). The account by E. M. Marsden-Jones & W. B. Turrill (1954) contains much data, but it is difficult to interpret.

12. C. macrocephala Muss. Puschk. ex Willd. Large-headed Star-thistle Phaeopappus macrocephalus (Muss. Puschk. ex Willd.)

13. C. jacea L. Brown Knapweed Jacea communis Delarbre; Jacea nigra Gray, non (L.) Hill; Jacea pratensis Lam.

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Perennial herb with an oblique branching stock. Stems 15– 60 cm, pale yellowish-green, erect or ascending, grooved, glabrous or slightly arachnoid-hairy, with a few long, slender branches, leafy. Leaves medium yellowish-green on upper surface, paler beneath; basal and lower cauline with lamina 1.5–15 × 0.5–3.0 cm, ovate to broadly lanceolate, more or less acute at apex, entire, dentate or pinnately lobed, gradually narrowed into a stalk-like base; upper cauline lanceolate or oblong-lanceolate, acute at apex, entire or dentate, sessile; all scabrid or shortly and roughly hairy on the margin and beneath. Capitula 20–40 mm in diameter, solitary at the ends of stems and branches sometimes forming a corymb; peduncles thickened below the capitula. Involucral bracts in several rows, 5–18 × 1.5–3.0 mm, pale brown, darker in centre, ovate to oblong; appendages usually covering the bracts, but distinct from them and not decurrent, the outer rotund or very broadly ovate, denticulate to pectinate-lacerate, the innermost entire. Flowers all tubular, reddish-purple, rarely white, the outer usually much larger than the inner, deeply 5-lobed. Receptacle flat, pitted, with numerous long bristles. Achenes about 3 mm, pale greyish to blackish-brown, narrowly obovoid; pappus absent or very short. Flowers 8–9. 2n = 44. Introduced. Grassy places and waste land. That this species has occurred in the past in southern England is beyond question, but whether it still does so seems to be questionable. It occurs throughout most of temperate Continental Europe. 14. C. debeauxii Gren. & Godr. Chalk Knapweed Perennial herb with a stout, branching, oblique stock. Stems 1–several, 10–80 cm, pale green, erect or ascending, channelled, tough, rigid, with numerous to dense, curled, pale simple eglandular hairs, branched, leafy. Leaves dull, medium green, paler beneath; basal with lamina 4–19 × 1– 5 cm, acute at apex, entire, sinuate-dentate or slightly pinnatifid, attenuate at base and petiolate; cauline rather rapidly becoming smaller up the stem, oblong, linear or linear-lanceolate, entire or lobed at base, sessile; all with numerous, rather soft, simple eglandular hairs. Capitula numerous, solitary at the ends of branches, involucre 9–14 mm in diameter, ovoid-globose; peduncles greyish-green, not much swollen below the capitulum, shortly and stiffly hairy. Involucral bracts in several rows, 7–12 × 1.5–2.0 mm, green, lanceolate, with a brown or brownish-black, non-decurrent, pectinately divided appendages, the fimbriae longer than the undivided part and not completely concealing the lower portion of the bract. Flowers all tubular, reddish-purple, all of them of more or less similar length or the outer much longer than the inner, deeply 5-lobed. Receptacle flat, pitted, with dense white bristles. Achenes about 3 mm, pale brown, oblong, compressed, with more or less numerous, short simple eglandular hairs; pappus absent, or of short, dirty white, subrigid simple eglandular hairs. Flowers 6–9. 2n = 44. (a) Subsp. nemoralis (Jord.) Dost´al C. nemoralis Jord. Outer flowers more or less the same length as the inner. (i) Var. minima (C. E. Britton) P. D. Sell C. nemoralis var. minima C. E. Britton

Plant up to 30 cm. Capitula few on short peduncles. (ii) Var. nemoralis (Jord.) P. D. Sell Plant up to 80 cm. Capitula numerous on long branches. (b) Subsp. thuillieri Dost´al C. pratensis Thuill., non Salisb. Outer flowers much longer than the inner. Named after Jean Louis Thuillier (1757–1822). (iii) Var. pumila P. D. Sell Plant up to 30 cm. Capitula few, on short peduncles. (iv) Var. thuillieri (Dost´al) P. D. Sell Plant up to 80 cm. Capitula numerous on long branches. Native. Especially common on light calcareous grassland, heaths, roadsides and cliff-tops. Mostly south of a line between the Humber and North Wales with a few localities further north. The most widespread plant is subsp. nemoralis var. nemoralis. This is also widespread in Western Europe. Var. minima occurs in southern coastal areas. Subsp. thuillieri seems to be confined to southern England and is widespread in Continental Europe. Its var. pumila occurs on southern and western coasts. The exact distribution of all these taxa is not understood and much more recording needs to be done. The species is a member of the Suboceanic European element. It is naturalised outside its native range. Named after Jean Odon Debeaux (1826–1910). × jacea = C. × moncktonii C. E. Britton This hybrid can only be recognised with certainty by the very irregular laciniations of the appendages of the outer involucral bracts. 2n = 44. In scattered localities in southern England, but is probably over-recorded. Named after Horace Wollaston Monckton (1857–1931). × nigra This hybrid is frequent where the two parents grow in adjacent populations in the area of distribution of C. debeauxii. They are intermediate in capitulum, shape and division of the appendages of the involucral bracts and the way in which the appendages cover the lower part of the bract. They are particularly common where chalk grassland is adjacent to fenland. 15. C. nigra L. Common Knapweed Jacea nigra (L.) Hill; Jacea vulgaris Bernh.; Phrygia nigra (L.) Gray Perennial herb with a stout, branching, oblique stock. Stems 1–several, 15–60(–90) cm, pale green, erect, channelled, tough, rigid, with numerous to dense, curled, pale simple eglandular hairs, leafy, branched above. Leaves dull, medium green on upper surface, paler beneath, basal with lamina 4–19 × 1–5 cm, elliptical, oblong or lanceolate, entire, sinuate-dentate or slightly pinnatifid, more or less acute at apex, attenuate at base, the petioles up to 150 mm, pale green or brownish-purple suffused and with pale, unequal simple eglandular hairs; cauline gradually decreasing in size upwards, oblong, linear or linear-lanceolate, acute at apex, entire, dentate or lobed, sessile; all with numerous, short, pale, curly simple eglandular hairs. Capitula

12. Centaurea numerous, solitary at the ends of branches, involucre 15–20 mm in diameter, ovoid-globose; peduncles greyishgreen, very shortly and stiffly hairy, more or less swollen below the capitulum. Involucral bracts in several rows, 7– 12 × 1.5–2.0 mm, green, lanceolate, with a brown or blackish, non-decurrent, more or less deeply pectinately divided appendage which more or less conceals the lower portion of the bract. Flowers 15–30 mm, all tubular, the outer sometimes much larger than the inner, reddish-purple, deeply divided at apex into 5 linear segments. Receptacle more or less flat, with dense, white, linear bristles. Achenes about 3 mm, pale brown, oblong, compressed, with more or less numerous, short simple eglandular hairs; pappus of short, dirty white, subrigid simple eglandular hairs. Flowers 6–9. Freely visited by a great variety of insects. (a) Subsp. nigra C. obscura Jord. Outer flowers more or less the same length as the inner. 2n = 22. (b) Subsp. rivularis (Brot.) Cout. C. rivularis Brot. Outer flowers much longer than inner. 2n = 22. Native. Grassland, waysides, cliffs, fens and heaths. Common in Wales, northern England and southern Scotland and extending more locally to northernmost Scotland and the islands. Its distribution in Ireland is not known, but it appears to be of scattered occurrence. In southern England it appears to be mainly in wet places and absent from the light calcareous soils. It occurs in Europe eastwards to Sweden and central Italy, and is naturalised in New Zealand. It is a member of the Suboceanic European element. Subsp. nigra occurs throughout most of the range of the species both in Great Britain, Ireland and Europe. Subsp. rivularis probably occurs most in western coastal areas, but its distribution is not really known. It was originally described from Portugal and it is not known if it is exactly the same plant as ours. Subgenus 8. Cyanus (Mill.) Hayek Cyanus Mill. Annual or perennial herbs. Stems ascending or erect, branched or not. Leaves entire to lyrate-pinnatifid. Capitula medium-sized. Involucral bracts with appendages decurrent and dentate to fimbriate. Pappus present. 16. C. montana L. Perennial Cornflower Cyanus montanus (L.) Hill; Jacea alata Lam. Perennial herb, often stoloniferous and with a stout, creeping rhizome, slightly foetid. Stems 40–80 cm, pale yellowish-green, erect or flopping, with long, pale, wavy simple eglandular, arachnoid and glandular hairs, simple or rarely branched above, leafy. Leaves 5–40 × 1–8 cm, dull medium green with a pale midrib on upper surface, paler with a prominent midrib beneath, soft, patent or ascending, oblong, elliptic-oblong, lanceolate or linear, more or less acute at apex, entire or the lower rarely remotely dentate to lobed, long attenuate at base, with long, pale arachnoid hairs at least some of them glandular, the lowest with short, winged petioles, the median and upper sessile and long-decurrent as a wing down the stem. Capitula 1–3,

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ovoid, terminal and usually solitary at the apex of the stem and branches, 70–100 mm in diameter; peduncles long, slightly thickened below the capitulum, arachnoidhairy. Involucral bracts in several rows, 6–15 × 4–6 mm, the outer smaller than the inner, medium green with a narrow, pale border and a brownish-black margin and appendage, ovate, with a fimbriate margin and appendage, the fimbriae half as long as the dark margin, glabrous. Flowers all tubular, honey-scented, the outer 40–50 mm, sterile, blue and divided at the apex into 5, linear, spreading segments, the inner 20–25 mm, bisexual, purple, plants with white and pink flowers occur in cultivation, divided into 5 narrowly linear, erect segments. Receptacle flat, pitted, with numerous bristles. Achenes 5–7 mm, brown, oblanceolate, compressed; pappus about 1.5 mm, of pale simple eglandular hairs in several rows. Flowers 5–6 and sometimes again in the autumn. Visited by bees. 2n = 24, 40, 44. Introduced. Much grown in gardens and naturalised in grassy and waste places. Throughout most of Great Britain, especially the centre and north. Native of the mountains of Europe, from the Ardennes and the Carpathians to the Pyrenees, central Italy and central Yugoslavia; widely cultivated for ornament. 17. C. cyanus L. Cornflower Cyanus segetum Baumg.; Cyanus cyanus (L.) Hill nom. illegit.; Jacea segetum Lam.; Cyanus vulgaris Delarb. Annual or overwintering herb with fibrous roots. Stems 20– 90 cm, pale green, erect, grooved, wiry, arachnoid-hairy, usually with many, ascending, slender branches, leafy. Leaves medium green on upper surface, paler beneath; lower with lamina 10–20 × 0.2–4.0 cm, lanceolate or oblanceolate in outline, acute at apex, entire or remotely dentate to lyrate-pinnatifid with narrow, distant lobes, petiolate; upper smaller, linear-lanceolate and sessile; all with arachnoid hairs on both surfaces making them look greyish when young. Capitula 15–30 mm in diameter, solitary at the end of main stem and branches, ovoid-globose; peduncles long, arachnoid-hairy. Involucral bracts in many rows, 4–15 × 1.5–2.5 mm, green in lower part, with a broad pale margin, ovate, arachnoid-hairy, with narrow, decurrent appendages cut halfway into long, spreading, narrowly triangular teeth, those of the outer bracts usually silvery-white, of the inner bracts brown with white-edged teeth. Flowers all tubular, the wild form with the outer dark blue and the inner reddish-purple, with garden forms ranging from white to mauve or pinkish; outer 15–30 mm, deeply divided into 5, linear, spreading lobes, the inner 12–15 mm, divided into 5, erect lobes. Receptacle flat, with numerous bristles. Achenes 3–4 mm, silvery-grey, narrowly ellipsoid, shortly hairy; pappus 3–4 mm, greyish, of many rows of stiff simple hairs. Flowers 6–8. Freely visited by insects, especially bees and flies. 2n = 24. Native or an ancient introduction. Formerly widespread and a pestilential weed in cornfields and occasionally reappearing from buried seed but now mostly a bird-seed alien or garden escape on tips and in waste places or deliberately sown in wild flower sites. Formerly common throughout Great Britain but now much rarer. Probably native of

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southern Europe and the Near East, but widely introduced elsewhere as a cornfield weed. 13. Cnicus L. nom. conserv. Carbenia Adans. Annual herbs. Stems erect, arachnoid-villous, not spiny. Leaves alternate, pinnately divided, with prominent white veins beneath, minutely spinose-dentate. Capitula solitary, surrounded by the upper leaves. Involucral bracts in several rows, narrowly oblong, the outer with a simple spine at apex, the inner longer with a pectinate, spine-like apical appendage. Flowers in centre bisexual, the outer small and sterile. Corolla tubular, yellow, deeply 5-lobed. Anthers tailed; filaments papillose-hairy. Style with slender, linear branches which are dorsally hairy and with a ring of hairs below the branches. Receptacle flat, with numerous setaceous scales. Achenes cylindrical-fusiform, sparsely hairy, crowned by a pale, dentate corona; pappus of 2 rows of rigid hairs, the outer long and minutely scabridulous, the inner much shorter and ciliate. Contains a single species widely distributed in southern Europe and the Mediterranean region; western Asia. 1. C. benedictus L. Blessed Thistle Centaurea benedicta (L.) L.; Carbenia benedicta (L.) Arcang. Sprawling or tufted annual herb. Stems up to 25 cm or absent, reddish-purple, usually branched only at base, thinly arachnoid-hairy to conspicuously villous. Leaves alternate, often forming basal rosettes, the lamina 3–15 × 0.8–4.0 cm, bright green on upper surface, paler with prominent white veins beneath, narrowly oblong in outline, acute at apex, coarsely sinuate-pinnatifid, with acute, erose-denticulate lobes, the marginal teeth normally ending in a short spine, thinly villous, the basal with purplish petioles up to 8 cm, the cauline normally sessile. Capitula 15–30 mm in diameter, usually solitary, terminal, subsessile or shortly pedunculate, surrounded by the upper leaves. Involucral bracts in several rows, 8–30 × 6–12 mm, greenish, narrowly oblong, the outer with a simple, purple spine at apex, the inner with a conspicuous, purple, spreading pectinately spinulose appendage, arachnoid-hairy. Flowers pale straw-coloured, the marginal 15–16 mm, slender, sterile and divided almost to the base into 3(–5), linear lobes, the inner fertile flowers divided into 5, linear lobes. Receptacle flat, with numerous, setaceous scales. Achenes 7–8 × 2.0–2.5 mm, greyish, shining, cylindrical-fusiform, prominently ribbed, sparsely hairy, with a pale, toothed apical corona and a very conspicuous lateral hilum; pappus of bristles, the outer 7–8 mm, spreading, pale and sparingly scabridulous, the inner about 2.5 mm, erect and conspicuously ciliate. Flowers 6–8. 2n = 22. Introduced. A bird-seed casual; formerly a wool alien. Scattered records in Britain in waste places. Native of southern Europe and the Mediterranean region, eastwards to Iran and Afghanistan. 14. Carthamus L. Annual herbs. Stems erect, sparsely hairy to glandular hairy and arachnoid hairy. Leaves alternate, simple to pinnately

divided, entire to spinose-dentate. Capitula solitary on stems and branches forming a loose corymb. Involucral bracts in many rows, the outer leaf-like, the inner usually spine-tipped. Flowers bisexual. Corolla tubular, yellow to orange, rarely whitish, deeply 5-lobed. Anthers tailed; filaments papillose-hairy. Style with long, linear branches which are dorsally hairy and with a ring of hairs below the branches. Receptacle convex, with dense, rigid hairs. Achenes turbinate to obpyramidal, 4-angled, glabrous; pappus absent or of linear, pointed scales. About 15 species, mainly in the Mediterranean region, but extending to Afghanistan and India. ¨ Hanelt, P. (1963). Monographische Ubersicht der Gattung Carthamus L. (Compositae). Feddes Rep. 67: 41–180, plus plates. 1. Leaves entire to spinose-serrate; flowers reddish-orange; 1. tinctorius pappus absent 1. Leaves pinnatifid; flowers yellow, rarely white; pappus of 2. scales 2. Plant more or less densely arachnoid-hairy; outer involucral bracts 1.5 times as long as inner; corolla and 2(a). lanatus subsp. lanatus anthers yellow 2. Plant sparsely arachnoid-hairy; outer involucral bracts twice as long as inner; corolla pale yellow, rarely whitish; anthers white with violet lines 2(b). lanatus subsp. baeticus

Section 1. Carthamus Flowers deep reddish-orange. Pappus absent. 1. C. tinctorius L. Safflower Annual to biennial herb with a tap-root and fibrous sideroots. Stem 10–60(–150) cm, very pale green, erect, slender to robust, striate, glabrous or nearly so, leafy, branched above. Leaves yellowish-green on upper surface, paler beneath with only the midrib prominent; basal with lamina 3–9 × 1–2 cm, ovate-oblong or oblong, obtuse or acute at apex, entire or spinose-serrate, narrowed into a short petiole; cauline gradually decreasing in size upwards, oblong, ovateoblong, oblong-lanceolate or lanceolate, obtuse to acute at apex, entire or spinose-serrate, sessile and amplexicaul; all glabrous. Inflorescence a lax corymb; peduncles striate, glabrous. Capitula 2–5 cm in diameter, subglobose. Involucral bracts in many rows, the outer 15–25 × 5–7 mm, green, ovate to ovate-lanceolate, long-spinose-acute at apex, with long pale spines on the margin, spreading, glabrous and veins prominent, the inner similar, but with even longer and narrower apex. Flowers all tubular, 15–25 mm, deep reddish-orange, with 5 linear lobes at apex. Receptacle convex, with long, dense, rigid hairs. Achenes 7–8 mm, pale brown, obpyramidal, 4-angled; pappus absent. Flowers 7–9. 2n = 24. Introduced. Bird-seed casual on tips and waste and rough ground. Fairly frequent in southern Great Britain and scarcer further north. Of uncertain origin, having been cultivated in the Mediterranean region, Africa and Asia since ancient times, the flowers providing a yellow or reddish dye, which was used as a cheap substitute for the true Saffron. The large achenes are edible and included in food for cage birds, poultry and cattle.

15. Scolymus Section 2. Atractylis Rchb. Kentrophyllum Necker ex A. DC. Flowers yellow. Pappus of scales. 2. C. lanatus L. Downy Safflower Kentrophyllum lanatum (L.) DC. Annual to biennial herb with a tap-root and fibrous sideroots. Stem 15–60(–100) cm, very pale green, slender to robust, erect, markedly striate, more or less arachnoidhairy and sometimes also with short glandular hairs, leafy, branched. Leaves rather greyish-green on upper surface, paler with prominent veins beneath, basal with lamina 5– 15 × 1.5–6.0 cm, oblong in outline, acute at apex with a terminal spine, deeply pinnatisect into 5–8 or more pairs of narrow, spine-tipped serrate or lobulate segments, with a short, ill-defined petiole; cauline gradually decreasing in size upwards, oblong, ovate-lanceolate or ovate, tapering from a broad, sessile, amplexicaul base to an acute, spine-tipped apex, with 3–4 pairs of rigid, spreading or forward-pointing, spine-tipped lobes; all with short, unequal glandular and arachnoid hairs. Inflorescence a lax corymb; peduncles rigid, with arachnoid and glandular hairs. Capitula 20–50 mm in diameter, ovoid. Involucral bracts in many rows, the outer foliaceous, 25–40 × 3–5 mm, green, with a pale yellowish base, linear-lanceolate, spinose-acute at apex, spinose-ciliate on the margin, nearly glabrous to glandular and arachnoid-hairy and erect, spreading or recurved, the inner 15–25 × 3–8 mm, scarious, pale yellowish-green or purplish, ovate-oblong, narrowing to an entire or irregularly fissured, spine-tipped apex and arachnoid-hairy. Flowers all tubular, 20–30 mm, yellow, rarely whitish, with 5, linear lobes at apex. Receptacle convex, with dense, rigid hairs. Achenes 4.5–6.0 mm, pale brown, turbinate, obscurely angled, crenulate at apex; pappus 6–12 mm, whitish, of linear, acute or obtuse, ciliate scales, the innermost row shorter and oblong. Flowers 7–9. A complex of allopolyploid taxa here treated as subspecies as morphological intermediates seem to occur. (a) Subsp. lanatus Plant more or less densely arachnoid-hairy. Outer involucral bracts about 1.5 times as long as inner. Corolla and anthers yellow. 2n = 44. Allotetraploid, perhaps with C. dentatus (Forssk.) Vahl as one parent. (b) Subsp. baeticus (Boiss. & Reut.) Nyman C. creticus L.; Kentrophyllum baeticum Boiss. & Reut.; C. lanatus subsp. creticus (L.) Holmboe Plant sparsely arachnoid-hairy. Outer involucral bracts twice as long as inner. Corolla pale yellow, rarely whitish; anthers white with violet lines. 2n = 64. An allohexaploid derived from C. lanatus subsp. lanatus × C. leucocaulos Sibth. & Sm. Introduced. Casual on tips and waste and rough ground, mostly a wool alien. Fairly frequent in the south of Great Britain and scarce further north. Southern Europe, Mediterranean region, south-west Asia and Canary Islands. Both subspecies seem to occur in Great Britain. Subsp. lanatus is native throughout most of the range of the species.

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Subsp. baeticus is native of the Mediterranean islands, south Greece and south Spain. Carduncellus caeruleus (L.) C. Presl has occurred as a casual on refuse tips. Tribe 2. Cichorieae Lam. & DC. Tribe Lactuceae Cass.; Tribe Scolymeae Kostel.; Tribe Catanancheae D. Don; Tribe Lapsaneae Kostel.; Tribe Hypochaerideae D. Don; Tribe Leontodontineae (Sch. Bip.) W. D. J. Koch; Subtribe Leontodoninae Sch. Bip.; Tribe Picrideae Sch. Bip.; Tribe Tragopogoneae Sch. Bip.; Tribe Taraxaceae D. Don; Tribe Crepideae Lindl.; Tribe Hieracieae D. Don; Tribe Chondrilleae W. D. J. Koch; Tribe Hyoserideae Kostel. Plants producing white latex, rarely spiny. Capitula with the flowers all bisexual and ligulate, the ligules usually yellow and 5-lobed. Anthers with apical appendage elongate and obtuse, tailed; filaments smooth; pollen mostly echinolophate or sometimes echinate. Style slender, mostly with long, slender branches, evenly hairy, with long and short hairs on the shaft and branches. Contains 11 subtribes, 98 genera and more than 1,550 species (plus a large number of apomicts), worldwide, but mainly in the northern hemisphere. 15. Scolymus L. Annual to perennial herbs with latex, without rhizomes or stolons. Stems erect, leafy, solitary, with spinose-dentate wings. Leaves alternate, pinnately divided, spiny. Capitula in a narrow subspicate to subcorymbose panicle. Involucral bracts in several rows, lanceolate to ovate-lanceolate. Flowers bisexual. Corolla ligulate, yellow, 5-lobed at apex. Anthers with elongate appendage, tailed; filaments smooth. Style with long, slender branches, evenly hairy on shaft and branches. Receptacle conical, with ovate, slightly winged, deciduous scales. Achenes broadly obovate or oblongobovate, dorsally compressed, not beaked, tightly enclosed in and adnate to the receptacular scales; pappus absent or of a few, rigid hairs. Three species widely distributed in the Mediterranean region. 1. Leaves and wings of stem with strongly thickened, white margins and veins; corolla with dark brown hairs in the 1. maculatus lower half; pappus absent 1. Leaves and wings of stem without thickened, white margin and veins; corolla with white hairs in the lower 2. hispanicus half; pappus of 2–4, barbed, rigid hairs

1. S. maculatus L. Spotted Golden Thistle Annual herb with a tap-root. Stems 15–100 cm, whitish, stout, erect, conspicuously, irregularly and continuously winged, the wings with deltoid, denticulate, spine-tipped lobes, glabrous, sparingly branched above, leafy. Leaves greyish-green, with much thickened, white margins and veins; basal with lamina 10–20 × 2–5 cm, oblanceolate in outline, spine-tipped, pinnatifid, spinose-dentate, attenuate at base to an indistinct petiole; cauline 2–7 × 2–5 cm, oblong-deltoid, oblong-lanceolate or ovate, spinosetipped, very thick and rigid, sinuate-pinnatifid and coarsely

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spinose, the uppermost regularly pectinate-spiny, sessile. Capitula 30–40 mm in diameter, in a subcorymbose panicle, sessile; all glabrous. Involucral bracts in several rows, dull medium green, the outer 12–30 × 3–5 mm, ovatelanceolate, coarsely and rigidly spinose and spreading or loosely erect, the inner 4–15 × 3–4 mm, ovate-lanceolate, lanceolate or oblong, spine-tipped or mucronate at apex and with narrow or membranous margins. Flowers 14–16 mm, all ligulate, the ligules yellow, with dark brown hairs in the lower half, minutely 5-lobed at apex. Receptacle conical, with pale, folded, adnate scales almost totally enveloping the achenes and winged all round. Achenes 4–5 × about 3 mm, oblong-obovate, dorsally compressed; pappus absent. Flowers 6–8. 2n = 20. Introduced. A wool and bird-seed casual. Native of southern Europe and the Mediterranean region. 2. S. hispanicus L. Golden Thistle Biennial to perennial herb with a tap-root. Stems (10–)20– 80(–100) cm, pale, with narrow, interrupted wings which have very unequal, sharply spinose lobes, arachnoid-hairy, usually branched, leafy. Leaves yellowish-green with margins and veins not conspicuously thickened or whitish; basal with lamina 1.2–20.0 × 2.5–4.0 cm, narrowly oblong, spine-tipped, pinnatifid, with broadly deltoid, spine-tipped and spinose-denticulate lobes; cauline slowly diminishing in size upwards, rigidly spinose, sessile and amplexicaul; all scabridulous-hairy or glabrous on both surfaces. Capitula 30–40 mm in diameter, in a subspicate inflorescence and usually solitary and sessile in the axil of upper cauline leaves. Involucral bracts in several rows, dull medium green, the outer leafy, 15–40 × 5–10 mm, ovate, spinose and suberect, the inner 8–20 × 3–7 mm, lanceolate, acuminate at apex, spine-tipped and with a very narrow, shortly ciliate, membranous margin. Flowers 20–25 mm, all ligulate, the ligules golden yellow, minutely 5-lobed at apex, with short, white hairs in the lower half. Receptacle conical, with pale, folded, adnate scales almost totally enveloping the achenes and forming membranous-margined, laciniate, lateral wings. Achenes 8–9 × 7–8 mm, broadly obovate, dorsally compressed; pappus up to 6 mm of 2–4, pale, barbed, rigid hairs. Flowers 6–8. 2n = 20. Introduced. Grain, bird-seed and shoddy alien, casual in waste places. In very scattered localities in southern England and Wales. Native of southern Europe and the Mediterranean region. 16. Cichorium L. Perennial herbs with latex, without rhizomes or stolons. Stems leafy, erect, solitary. Leaves alternate, dentate or pinnately divided. Capitula numerous, terminal and axillary in a spike-like inflorescence. Involucral bracts in 2 rows, the inner twice as long as the outer. Flowers bisexual. Corolla ligulate, usually blue or pink, rarely white, 5-lobed at apex. Anthers tailed, apical appendage elongate; filaments smooth. Style with long, slender branches, both shaft and branches evenly hairy. Receptacle more or less flat, without scales. Achenes obovoid or turbinate, more or less angled, not beaked; pappus of 1–2 rows of short, obtuse scales.

About 6 species in Europe, the Mediterranean region and south to Ethiopia. Hult´en, E. (1971). The circumpolar plants. II. Dicotyledons. Kungl. Svenska Vet.-Akad. Handl. ser. 4, 13: 280–281. [C. intybus.] Philips, R. & Rix, M. (1993). Vegetables. London.

The wild Chicory, C. intybus subsp. silvestre, once known is easy to recognise. The cultivated variants of Chicory and Endive, however, need to be examined more closely. There is much confusion between the use of the vernacular names Chicory and Endive; particularly in France some forms of C. intybus are called Endive and some forms of C. endivia Chicor´ee. 1. Outer involucral bracts 4–5 mm wide, ovate-elliptical; 2. endivia pappus 0.4–0.5 mm 1. Outer involucral bracts 1–3 mm wide, linear-lanceolate; 2. pappus about 0.2 mm 2. Basal and lower cauline leaves pinnatisect 1(a). intybus subsp. silvestre 2. Basal and lower cauline leaves sinuate-denticulate to 3. repand-dentate 3. Basal and lower cauline leaves lanceolate to oblong-lanceolate, sinuate-denticulate 1(b). intybus subsp. intybus 3. Basal and lower cauline leaves broadly ovate to reniform, crisped like curly lettuce, cordate at base 1(c). intybus subsp. foliosum

1. C. intybus L. Chicory C. sylvestre Lam. nom. illegit.; C. perenne Stokes nom. illegit. Perennial herb with a pale, slender to thick, woody taproot and fibrous side-roots and a short, vertical stock. Stems 30–200 cm, pale yellowish-green, often suffused brownishpurple, stiffly erect, tough, slightly angled, grooved, sometimes flexuous, sometimes hollow, with numerous, pale, short to long, spreading, sometimes subrigid simple eglandular hairs at least below and sometimes with glandular hairs above, branched, leafy. Leaves very variable, dull yellowish-green or medium green, rarely dark green, sometimes tinted pinkish-red or brownish purple on upper surface, paler beneath, sometimes with paler veins; basal in a rosette, with lamina 6–30 × 3–20 cm, oblong or oblonglanceolate in outline and pinnatisect, oblong to oblongoblanceolate and sinuous-denticulate or broadly ovate or reniform and lettuce-like; cauline gradually decreasing in size upwards, similar to basal, more or less amplexicaul; all with very short to medium, subrigid, pale simple eglandular hairs on both surfaces and the margins or only below particularly on the veins. Capitula 35–40 mm in diameter, few to numerous, terminal or in groups of 2–3 in the axils of upper leaves in a spike-like inflorescence; peduncles somewhat thickened, glabrous or with a few simple eglandular hairs. Involucral bracts in 2 rows, 7–15 × 1–3 mm, pale to medium yellowish-green with red or reddish-purple tips, linear-lanceolate to lanceolate or oblong-lanceolate, obtuse to acute and sometimes aristate at the apex, lax and sometimes keeled near the apex, with more or less reddish glandular hairs sometimes confined to the median line and very short, white simple eglandular hairs near the apex. Flowers

16. Cichorium 18–25 mm, all ligulate, the ligules pale to deep mauve to blue, rarely pink or white, with 5 short lobes at apex. Receptacle flat, pitted, the pits with raised, toothed margins, without scales. Achenes 1–3 mm, pale brown, often with darker mottling, obovoid, irregularly angular; pappus of 1–2 rows of scales about 0.2 mm, one-tenth to one-eighth as long as achene, the scales obtuse-fimbriate. Flowers 6–10. Flowers open in early morning and close soon after midday and are visited chiefly by bees and hoverflies. 2n = 18. (a) Subsp. silvestre (Bisch.) Janch. C. intybus taxon silvestre Bisch. Plants up to 100 cm, sometimes branched from base. Basal leaves with lamina oblong or oblong-lanceolate in outline, obtuse at apex, pinnatisect, the terminal lobe oblong, obtuse at apex and sinuate or shallowly lobed, the lateral lobes lanceolate or oblong, obtuse at apex, denticulate to dentate or shallowly lobed and retrorse, the area between the lobes narrowly winged and irregularly toothed, widened at base and semiamplexicaul; lower cauline similar to basal, the upper narrowly triangular-lanceolate, gradually narrowed to an obtuse apex, toothed or lobed only near the amplexicaul base; all more or less hairy on both surfaces. Inner involucral bracts 10–12 mm, linear-lanceolate. (b) Subsp. intybus C. intybus subsp. sativum (DC.) Janch.; non (Bisch.) Janch.; C. intybus subsp. sativum Janch.; C. intybus convar. radicosum (Alef.) Holub; C. intybus var. sativum DC.; C. intybus convar. radicosum Alef.; C. intybus taxon sativum Bisch. nom. illegit. Plant up to 150 cm, branched mainly in upper part. Basal leaves with lamina oblanceolate to oblong-oblanceolate, acute at apex, sinuate-denticulate or dentate; lower cauline similar to basal, the upper ovate-oblong, lanceolate-oblong or lanceolate, acute at apex, more or less amplexicaul with rounded auricles; all more or less hairy on both surfaces. Inner involucral bracts 13–15 mm, oblong-lanceolate. (c) Subsp. foliosum (Hegi) Janch. C. intybus var. foliosum Hegi Plant up to 200 cm, much branched in upper part. Basal leaves with lamina lettuce-like, broadly ovate to reniform, rounded at apex, repand-dentate, cordate at base; cauline gradually decreasing in size upwards, similar to basal, the lower shortly petiolate, the upper more or less amplexicaul; all more or less glabrous above with pale simple hairs beneath, particularly on the veins. Inner involucral bracts 10–12 mm, lanceolate. Native or introduced. Waysides, old tracks, rough grassland, waste places and field margins, particularly on calcareous soils. Frequent in central and south Britain with scattered localities elsewhere in Great Britain and Ireland. Europe northwards to central Scandinavia, Finland and central Russia; western Asia; North Africa; introduced in eastern Asia, North and South America, South Africa and New Zealand. It is a member of the Eurosiberian Southerntemperate element. The three subspecies have not been studied throughout our area and herbarium sheets are mostly inadequate. The above account is of our personal experience in East Anglia. Subsp. silvestre is believed to be native

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and is mostly found on ancient trackways on the boulder clay. Subsp. intybus is the plant grown as a crop and occurs as a relic of cultivation. Subsp. foliosum is grown in market gardens and may escape. All three subspecies occur in wild flower seed, and in one locality at Melbourn in Cambridgeshire they have all been introduced, so plants should always be looked at carefully where there have been introductions. In our area when subsp. intybus is grown as a crop it is for the seed. Large-rooted forms of it have the root chopped, roasted and ground and mixed with coffee to give it a slight bitterness which some people find refreshing. The leaves of subsp. foliosum are used as a salad and the blanched leaves and young shoots are also cooked as a vegetable. 2. C. endivia L. Endive Annual herb with a slender tap-root and fibrous side-roots. Stems (5–)10–60(–100) cm, pale yellowish-green, erect, channelled, glabrous or with simple eglandular hairs below, usually much branched, with spreading branches, leafy. Leaves very variable, dull yellowish or medium green, sometimes tinted purplish, paler beneath; basal with lamina 15–30 × 2–6 cm, usually oblong or oblanceolate in outline, more or less obtuse at apex, subentire, pinnatifid, lyrate-runcinate or variously crisped and curled, narrowed at base; cauline numerous and fairly close together, becoming steadily smaller upwards, similar to basal but sometimes broader and more obovate, rounded or acute at apex, entire, toothed or pinnatifid, with an amplexicaul base often prominently auriculate, the auricles often exceeding the stem diameter; all generally more or less glabrous but sometimes crispate hairy. Capitula 15–40 mm in diameter, terminal or in dense lateral clusters on the flowering branches, forming a spike-like inflorescence; peduncles often conspicuously swollen, glabrous or with a few simple eglandular hairs. Involucral bracts in 2 rows, the outer 8–12 × 4–5 mm, with a large, pale, indurated median zone surrounded by a broad, green, ciliate margin, the cilia often remote, erect, ovate-elliptical and acute or shortly cristate and sometimes recurved at apex, the inner 10.0– 12.5 × 2–3 mm, loosely erect, oblong, obtuse or subacute at apex, glabrous or the margins and midrib long-ciliate. Flowers 10–14 mm, all ligulate, the ligules blue, with 5, sharp lobes at apex. Receptacle flat, somewhat pitted, without scales. Achenes 2–3 mm, brownish with darker mottling, obovoid or turbinate, 5-angled, prominently ribbed, densely and minutely verrucose-papillose; pappus whitish, of a single row of oblong or ovate scales about 0.4–1.5 mm, one-sixth to half as long as achene. Flowers 6–10. 2n = 18. Introduced. A wool and bird-seed alien and a relic of cultivation found in a few waste and rough places. Southern Europe; cultivated elsewhere. Our plant is the cultivated subsp. endivia. The wild form, subsp. divaricatum (Schousb.) Bonnier & Layens (C. divaricatum Schousb.) is a shorter plant with much narrower runcinate-pinnatifid, hairy leaves and divaricate branches which is a native of the Mediterranean region. The cultivated Endive produces a dense rosette of usually glabrous leaves, which in the popular variety var. crispum Hegi are much divided and curled

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and are sometimes blanched to make them more palatable as a salad plant. The types with non-curly leaves are called Escarole and Scarole. 17. Catananche L. Perennial herbs with latex, without rhizomes or stolons. Stems erect, solitary or few. Leaves alternate, linear, entire or with few, narrow teeth. Capitula 1–5(–numerous) on very long peduncles. Involucral bracts in several rows, ovatecuspidate, scarious, with a darker midrib. Flowers bisexual. Corolla ligulate, mauve-blue, with 5 lobes at apex. Anthers tailed, apical appendage elongate; filaments smooth. Style with slender branches, evenly hairy on both shaft and branches. Receptacle flat, with long filiform scales. Achenes conical, obscurely 5-angled; pappus of one row of 5–7, ovate, long-aristate scales. Five species in southern Europe, North Africa, Turkey and the Middle East. 1. C. caerulea L. Blue Cupidone Perennial herb. Stems solitary or few, 20–90 cm, dull medium green, ridged, erect, with numerous to dense, short, appressed simple eglandular hairs, sparingly leafy, much branched. Leaves dull medium green on upper surface, hardly paler and with a prominent midrib beneath, basal numerous, the lamina 2–30 × 0.2–0.7 cm, linear, obtuse to acute at apex, entire or with up to 4, linear, usually forwardly directed teeth, sessile; cauline similar but few and remote; all with dense, short, appressed simple eglandular hairs on both surfaces and the margins. Capitula 25–50 mm in diameter solitary at the end of very long peduncles; peduncles medium green, with dense, short, appressed simple eglandular hairs. Involucral bracts in several rows, 10–25 × 3.5– 5.0 mm, greyish-scarious except for the dark midrib, ovate or oblong-ovate, acute at apex with an excurrent midrib making them cuspidate, glabrous. Flowers 25–28 mm, all ligulate, the ligules mauve-blue, deeply 5-lobed at apex. Receptacle flat, with long, scarious, filiform scales. Achenes 5–6 mm, oblong, obscurely 5-angled, 5- to 10-ribbed, appressed-hairy; pappus scarious, of ovate, long-aristate scales. Flowers 9–10. 2n =18. Introduced. A garden escape, which persisted on a road verge at Priddy in Somerset for some years after its discovery in 1988. Native of the western Mediterranean region. 18. Arnoseris Gaertn. Annual herbs with latex. Stems numerous, leafless, but with small, scale-like bracts. Leaves all basal, more or less dentate. Capitula 1–3 per stem, on swollen peduncles. Involucral bracts usually in 2 rows, the outer small and incomplete. Flowers bisexual. Corolla ligulate, yellow, sometimes turning greenish when dry, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with short branches, both shaft and branches shortly hairy. Receptacle flat, without scales. Achenes obconical, angled, slightly compressed, not beaked; pappus a very short, membranous border. A single species in Europe and western Asia.

1. A. minima (L.) Schweigg. & K¨orte Lamb’s Succory Hyoseris minima L.; A. pusilla Gaertn. nom. illegit.; Lapsana pusilla Willd. nom. illegit. Annual herb with a slender tap-root. Stems numerous, 5– 32 cm, pale green, often tinted reddish, erect, faintly striate, glabrous or with a very occasional simple eglandular hair, simple or with long branches above in the axils of minute bracts. Leaves all basal, few to numerous, the lamina 1–9 × 0.3–2.0 cm, yellowish-green, sometimes tinted purplish on upper surface, paler beneath, spathulate or oblanceolate, obtuse to acute at apex, with patent to ascending, narrowly mammiform or aquiline-mammiform teeth, narrowed at base to a short, winged petiole, scabridulous or with short, stubby hairs especially on the margin. Capitula solitary at the end of stem or branches, 7–11 mm in diameter, peduncles long, markedly thickened beneath the capitula, curving upwards and finally overtopping the main stem, glabrous or with a very occasional simple eglandular hair. Involucral bracts in 2 rows, glabrous or minutely chaffyhairy, green with a prominent pale keel, connivent in fruit; outer 1.5–3.0 × about 0.5 mm, linear or linear-lanceolate, acute at apex; inner 5–6 × 1.0–1.5 mm, lanceolate, longacuminate to a narrow drawn-out apex. Flowers all ligulate, 8–9 mm; ligules yellow, sometimes turning greenish when dry, with 5 lobes at apex. Receptacle flat, without scales. Achenes 1.5–2.2 mm, dark with paler angles and ribs, obovoid, slightly flattened, with 5 strong and 5 weaker intermediate ribs, transversely wrinkled between the ribs; pappus a very short, membranous border, sometimes toothed above the ribs. Flowers 6–8. Sparingly visited by flies. 2n = 18. Probably native. A plant of the poorest and most acidic sandy arable fields especially over Lower Greensand. Formerly very local in eastern England from Dorset and Kent to Yorkshire and in eastern Scotland; extinct since 1971. Western Europe from Spain to Portugal, Corsica, northern Italy, Hungary, Romania and southern Russia north to South Sweden, north Poland and central Russia; introduced in North America, Australia and New Zealand. A member of the European Temperate element. 19. Lapsana L. Annual to perennial herbs with latex, without rhizomes or stolons. Stems usually solitary, leafy, branched. Leaves alternate, entire to pinnately lobed. Capitula small and numerous in an open panicle. Involucral bracts in 2 rows, the outer few and scale-like. Flowers bisexual. Corolla ligulate, yellow, with 5 lobes at apex. Anthers tailed, apical appendage elongate; filaments smooth. Style with slender branches, the shaft and branches evenly hairy. Receptacle flat, without scales. Achenes slightly flattened, about 20ribbed, not beaked; pappus absent. About 4 species in Europe and Asia. Burtt, B. L. (1950). Lapsana intermedia in Britain. Watsonia 1: 234–237. Grime, J. P. et al. (1988). Comparative plant ecology. London. Sell, P. D. (1981). Lapsana intermedia Bieb. or Lapsana communis L. subsp. intermedia (Bieb.) Hayek. Watsonia 13: 299–302.

20. Hedypnois 1. Capitula 10–20 mm in diameter; involucral bracts 5–7(–8) mm; ligules about 1.5 times the inner involucral bracts 1(a). communis subsp. communis 1. Capitula 20–30 mm in diameter; involucral bracts 7–10 mm; ligules up to 2.5 times the inner involucral bracts 1(b). communis subsp. intermedia

1. L. communis L. Nipplewort Annual to perennial herbs with fibrous roots. Stems usually solitary, 10–125(–150) cm, erect, leafy, pale yellowishgreen, sometimes tinted purplish, striate, often flexuous, with few to numerous, pale, wavy simple eglandular hairs especially below. Leaves dull medium to yellowish-green on upper surface, pale beneath with prominent veins; basal often numerous but dying before anthesis, the lamina 1– 20 × 1–10 cm, oblong-obovate in outline, lyrate-pinnatifid, obtuse at apex, the terminal lobe large, ovate or triangularovate, undulate and sinuate-dentate and more or less cordate at the base, the lateral lobes several and much smaller, gradually decreasing in size downwards, very unequal, patent or recurved, ovate, oblong or pinnatifid, obtuse to rounded, dentate and sessile or narrowed at base to a winged petiolule, the area between the lobes with a narrow, irregular wing, with a narrowly winged petiole; cauline gradually decreasing in size, the lower usually similar to basal, the upper ovate, obtuse to acute at apex, petiolate or sessile, the uppermost often lanceolate, denticulate or entire and sessile; all with few to numerous simple eglandular hairs on the upper surface and numerous, unequal ones below especially on the veins. Capitula 3–100(–380), 10–30 mm in diameter, in an open, more or less corymbose panicle; peduncles usually glabrous, rarely with glandular hairs. Involucral bracts in 2 rows; outer 0.5–1.2 × 0.5–1.0 mm, pale green, triangularovate, obtuse at apex, glabrous; inner 5–10 × 1.2–1.5 mm, pale yellowish-green, often reddish tipped, linear-oblong, more or less obtuse at apex, keeled, usually glabrous, rarely with glandular hairs. Flowers all ligulate, bisexual; ligules a little longer than to 2.5 times as long as the involucral bracts, yellow, with 5 lobes at apex. Receptacle flat, without scales. Achenes 3.5–5.0 mm, the outer longer than the inner, linear-oblanceolate, slightly flattened, about 20-ribbed, not beaked; pappus absent. Flowers 6–9. (a) Subsp. communis Annual herb. Lateral segments of leaves not as wide as terminal ones. Capitula 10–20 mm in diameter. Involucral bracts 5–7(–8) mm. Ligules about 1.5 times the inner involucral bracts. 2n = 14. (b) Subsp. intermedia (M. Bieb.) Hayek L. intermedia M. Bieb. Annual to perennial herb. Lateral segments of leaves often about as wide as the terminal ones. Capitula 25–30 mm in diameter. Involucral bracts 7–10 mm. Ligules up to 2.5 times as long as the inner involucral bracts. 2n = 14. Native. Subsp. communis occurs in cultivated and waste land, waysides, hedgerows, wood margins and clearings, walls and on cliffs. Common throughout Great Britain and Ireland. Throughout Europe north to 70◦ N, but perhaps absent from the south-east; introduced in North America

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and Australasia. Subsp. intermedia is introduced and has been naturalised on a chalk bank in Bedfordshire since 1945, Frosterley quarries in Co. Durham, in limestone grassland near Cilcain in Flintshire since about 1970, Great Ormes Head in Caernarvonshire since 1977, and in at least two localities in Middlesex, one since 1982. Native of southeast Europe and south-west Asia. Other subspecies occur in south-east Europe, south-west Asia and North Africa. The species is a member of the European Temperate element. 20. Hedypnois Mill. Annual herbs with latex, without rhizomes or stolons. Stems usually many, leafy, branched. Leaves alternate, entire, dentate or lobed. Capitula solitary or many, at the ends of branches; peduncles usually swollen. Involucral bracts in 2 rows, the outer very small. Flowers bisexual. Corolla ligulate, yellow, the outer sometimes with a greenish stripe on the outer face, with 3–5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long, slender branches, both shaft and branches with short hairs. Receptacle flat, without scales. Achenes more or less cylindrical, the outer usually partly enclosed in the involucral bracts, not beaked; pappus of outer achenes a corona, that of inner achenes of narrow, long-aristate scales. One very variable species, sometimes split into 2 or 3 species, widely distributed in the Mediterranean region and western Asia. 1. H. cretica (L.) Dum. Cours. Scaly Hawkbit Hyoseris cretica L.; Hyoseris hedypnois L.; Hyoseris rhagadioloides L.; Hedypnois rhagadioloides (L.) F. W. Schmidt; H. monspeliensis Willd.; H. polymorpha DC.; H. tubiformis Ten.; Rhagodiolus creticus (L.) All. Annual herb. Stems often many, 3–45(−60) cm, more or less hairy, branched. Leaves 0.5–18(−25) × 0.2–2.5(−3.5) cm, greyish-green on upper surface, paler and sometimes tinged purple beneath; basal elliptical, oblanceolate or obovate, obtuse at apex, entire to deeply dentate or lobed, the teeth or lobes diminishing in size downwards, attenuate and decurrent at base which is sessile or with a short, winged petiole; cauline similar to basal, or linear-lanceolate, lanceolate or oblong, decreasing in size upwards and usually sessile, but sometimes shortly petiolate; all with stiff, 2- to 3-furcate bristles on both surfaces, sometimes densely so on the upper surface or sometimes only along the veins and margins of the lower surface. Capitula (6–)8–10(−13) mm in diameter, subcylindrical at anthesis, suborbicular or hemispherical in fruit, solitary at the ends of stems or branches; peduncles usually swollen and hollow just below the capitulum, striate and often densely, minutely hispid, with forked bristles much shorter than those on the leaves. Involucral bracts in 2 rows, green; outer few, 2–5 mm, linear-lanceolate or subulate, with an apical tuft of minute hairs and with or without longer, white bristles on the surface and margins; inner 6– 8 × 1.0–1.5 mm, narrowly lanceolate or linear-lanceolate, glabrous or variably hispid on the surface and margins with bristles of various lengths or sometimes hispid only on the upper surface. Flowers all ligulate, the ligules 5–7 × 1.5– 2.5 mm, dull golden yellow, the outer surface sometimes with a greenish stripe, truncate with 3–5 lobes at the apex.

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Receptacle flat, without scales. Achenes erect and incurved, more or less cylindrical, with minute, rigid hairs, the outer 5.0–7.5 mm, each enveloped by an accrescent, indurated, boat-shaped involucral bract and with a coroniform pappus of erose scales about 1 mm, the inner naked, less curved, 5–7 mm with 5, subulate pappus scales extended into a distinct, scabrid awn 4–6 mm, each scale usually with many, short, hair-like teeth near the base. Flowers 6–8. 2n = 8, 12, 13, 14, 16. Very variable in size, habit, hairiness, dissection of leaf and thickening of peduncle. Many variants have been named, but the characters occur in various combinations, even in the same locality, and there seems to be no variants of any geographical or ecological significance. Introduced. Bird-seed, wool and esparto casual in waste places. Scattered records in England. Native of southern Europe, the Mediterranean region and western Asia; introduced in Australia and North America. 21. Rhagadiolus Juss. nom. conserv. Annual herbs with latex, without rhizomes or stolons. Stems usually solitary, branched, leafy. Leaves alternate, dentate to pinnately divided. Capitula few to numerous, stellate in fruit. Involucral bracts in 2 rows, the outer very short, the inner accrescent and patent in fruit and enclosing the outer achenes. Flowers bisexual. Corolla ligulate, yellow, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long branches, both shaft and branches with very short hair. Receptacle flat, without scales. Achenes subulate to narrowly cylindrical, the inner soon caducous, the outer enclosed in the involucral bracts, spreading star-like and long-persistent; pappus absent. Two species in the Mediterranean region, southern Europe, North Africa, Turkey and the Middle East to Iran. Miekle, R. D. (1979). Rhagadiolus: a misunderstood genus. Taxon 28: 133–141.

1. R. stellatus (L.) Gaertn. Star Hawkbit Lapsana stellata L.; R. intermedius Ten.; R. lyratus Lam.; R. stellatus var. hebelaenus DC.; R. stellatus var. leiocarpus DC.; R. stellatus var. indivisus Viv.; R. stellatus var. leiolaenus Boiss.; R. stellatus var. eriocarpus Faure & Maire Annual herb with tap-root and fibrous side-roots. Stems (10–)20–50(–70) cm, pale yellowish-green, erect, with few to dense, stiff, pale simple eglandular hairs in the lower part, glabrous above, simple or with wide, spreading branches, sparingly leafy. Leaves yellowish-green on upper surface, paler beneath; basal numerous, forming a rosette, the lamina 2–24 × 0.5–5.0 cm, oblanceolate to narrowly obovate, obtuse to acute at apex, subentire, repand-dentate or deeply and closely pinnatifid, with numerous, often toothed lobes, diminishing downwards to a short, indistinct petiole; cauline oblong, elliptical or narrowly lanceolate and remotely toothed or entire; all stiffly hairy on both surfaces or nearly glabrous. Inflorescence a lax panicle, often with divaricately spreading branches. Capitula 4–5 mm in diameter, stellate in fruit. Involucral bracts in 2 rows, the outer

about 1.0–0.8 mm, greenish with erose, membranous margins and deltoid-ovate with an apiculate apex, the inner 8–9 × about 1.0 mm, greenish, strap-shaped and acute at apex, and incurved with rigid hairs dorsally throughout their length, at first strictly erect but later accrescent, thickened and enveloping the fruit. Flowers 7–8 mm, all ligulate, the ligules yellow and 5-lobed at apex. Receptacle flat, without scales. Achenes of 2 kinds, the outer regularly 8, 15–18 mm, pale brown, subulate, distinctly incurved, patent and smooth, enclosed in the outer involucral bracts and long persistent, the inner (1–)3, rarely wanting, 9–10 mm, pale brown narrowly cylindrical, glabrous or hairy on the dorsal surface, soon caducous; pappus absent. Flowers 7–9. 2n = 10. Introduced. A bird-seed, grain and wool casual. A few scattered records in Great Britain. Native of southern Europe and the Mediterranean region, North Africa and south-west Asia. R. edulis Gaertn., which may occur, is distinguished by its lyrate-pinnatisect basal leaves with a large terminal lobe and small, remote, lateral ones, the 5–6 narrow, glabrous inner involucral bracts and the straight to slightly recurved fruits 10–15 mm. 22. Hypochaeris L. Achyrophorus Adans. nom. illegit.; Hypochoeris orthogr. var.; Porcellites Cass. Annual to perennial herbs with latex, without rhizomes or stolons. Stems solitary to few, usually leafless but with scalelike bracts. Leaves all basal, dentate to pinnately divided, with few to numerous simple hairs. Capitula solitary to few. Involucral bracts in several rows. Flowers bisexual. Corolla ligulate, yellow, the outer sometimes with a greenish or reddish stripe on the outer face, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long, slender branches, both shaft and branches with very short hairs. Receptacle flat, with numerous, scarious scales. Achenes more or less cylindrical, at least the inner usually beaked; pappus of 1–2 rows of dirty to pale brown, scabrid or plumose hairs. About 60 species, chiefly in South America; about 10 species in Europe, the Mediterranean region and western Asia. Chater, A. O. (1983). Those Latin names. B.S.B.I. News 33: 28. Grime, J. P. et al. (1988). Comparative plant ecology. London. [H. radicata.] Parker, J. S. (1975). Aneuploidy and isolation in two Hypochaeris species. Chromosoma 52: 89–101. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [H. glabra.] Turkington, R. & Aarssen, L. W. (1983). Hypochaeris radicata L. (Achyrophorus radicatus (L.) Scop. in Biological flora of the British Isles. Jour. Ecol. 71: 999–1022. Wells, T. C. E. (1976). Hypochoeris maculata L. in Biological flora of the British Isles. Jour. Ecol. 64: 757–774. 1. Leaves usually spotted, blotched or streaked with brownish-purple; flowers 36–50 mm, lemon yellow; 3. maculata pappus hairs in 1 row, all plumose 1. Leaves rarely with purplish markings; flowers 4–27 mm, deep yellow; pappus hairs in 2 rows, the outer usually 2. simple, the inner plumose

22. Hypochaeris 2. Capitula 5–15 mm in diameter, opening only in full sunlight in the morning and then not very widely; flowers 1. glabra 3–15(−21) mm 2. Capitula 20–50 mm in diameter, opening only in the 3. morning but widely; flowers 14–27 mm 3. Stems 30–60(–100) cm, usually more or less erect; leaves 8–25 cm, often ascending; inner involucral bracts 2(a). radicata subsp. radicata 16–25 mm 3. Stems (1–)7–30 cm, prostrate, decumbent or ascending; leaves 1–12 cm, usually prostrate; inner involucral bracts 2(b). radicata subsp. ericetorum 10–17 mm

1. H. glabra L. Smooth Cat’s-ear H. hispida Roth; H. dimorpha Brot.; H. ascendens Brot. Annual herb with a slender tap-root. Stems 1–several, 5– 40 cm, medium green to greyish-green, erect, ascending or decumbent, grooved, glabrous, simple or branched, with scales. Leaves pale to medium green on upper surface, paler beneath with an even paler midrib; basal in a rosette, the lamina 1–20 × 0.5–3.0 cm, oblanceolate or oblongoblanceolate, obtuse at apex, shallowly sinuate-dentate to pinnatifid, the lobes triangular to oblong and acute to obtuse, narrowed gradually to a stalk-like base; glabrous or with few to numerous, short, pale simple eglandular hairs;cauline absent, but with 1–few scales 1.5–3.5 mm, green, suffused purplish at tip, elongate-triangular, acute at apex. Capitula usually solitary, sometimes few, 5–15 mm in diameter, opening only in full sunlight in the morning and then not very widely; peduncles slightly swollen beneath the capitula, with a few scales. Involucral bracts in several rows, 3–15(–21) × 1.0–2.5 mm, medium yellowish-green, with a narrow scarious margin, the upper part often suffused dark brownish-purple, oblong, asymmetrically obtuse at apex, glabrous except for the very short hairy apex, or with a line of short, stiff hairs down the centre. Flowers 4–15(–21) mm, all ligulate, the inner shorter, the ligules golden yellow, with a purplish-brown stripe on the outer face of the outer, but sometimes only near the apex, with 5 lobes at apex. Styles yellow. Receptacle flat, pitted; scales 4–15(–21) mm, narrowly lanceolate, toothed at apex. Achenes reddish-brown, oblanceolate or cylindrical, muricate, body of the outer 2.5–5.0 mm, usually unbeaked but sometimes unbeaked achenes absent, the body of inner 4.5–5.5 mm, longbeaked or unbeaked; pappus of 2 rows of dirty white hairs, the outer row 3–6 mm and scabrid or sparsely plumose, the inner 9.5–15.0 mm, plumose and not dilated at base. Flowers 6–10. Visited by bees and also self-compatible. 2n = 10. Variation occurs in the achenes, but the variants do not show ecological or geographical isolation. Capitula with inner achenes only with a beak are var. glabra; these with all achenes with a beak are var. loiseleuriana Godr.; those with no achenes with a beak are var. erostris Boiss. Native. Disturbed, usually nutrient poor, light soils in acidic sandy, heathy or gravelly places. Very local and decreasing in England and Wales, the Channel Islands and the coasts of Fifeshire, Morayshire and Invernessshire in Scotland. It is easily missed because it is often small and does not open very widely and then only in the morning in bright sunlight. In East Anglia it often grows

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with H. radicata subsp. ericetorum. Europe northwards to southern Scandinavia; North Africa; Middle East; naturalised in North America, Australia, New Zealand and elsewhere. A member of the European Southern-temperate element. × radicata = H. × intermedia Richt. This hybrid is a short-lived perennial with many branched stems and looks nearer to H. radicata subsp. ericetorum and opens in dull weather like that species. It is of low fertility and most capitula have no viable achenes. When good achenes are produced the plants can backcross with H. radicata. Flowers 5–10. 2n = 9. Recorded for Surrey, Suffolk, Norfolk and Merionethshire. We have seen H. glabra growing with H. radicata subsp. ericetorum many times in East Anglia, but not with subsp. radicata. The hybrid is also recorded for Germany, Switzerland and Sweden. 2. H. radicata L. Cat’s-ear Achyrophorus radicatus (L.) Scop.; Porcellites radicata (L.) Cass.; H. infesta Salisb. nom. illegit. Perennial herb with a short, erect, branching, premorse stock and a strong, fleshy root. Stems 1–numerous, (1–) 7–60(–100) cm, dull, medium green, prostrate, decumbent, ascending or erect, slender to robust, striate, usually forking, glabrous, usually not leafy or with 1–2 small leaves below, with numerous small scales. Leaves yellowish-green, paler beneath, usually with numerous, pale, rather stiff simple eglandular hairs on both surfaces; basal in a rosette, 0.5–25.0 × 0.5–6.0 cm, erect or prostrate, oblong, oblonglanceolate, oblanceolate or oblong-oblanceolate, rounded at apex, sinuate-dentate to sinuate-pinnatifid, the lobes ovate or oblong, obtuse at apex and occasionally obscurely dentate, long-attenuate at base to a winged petiole; cauline usually absent, sometimes 1–2, small ones near the base, but stem with numerous but widely spaced scales 1.0– 2.5 × about 1.0 mm, ovate, pale in middle with a dark green margin and blackish at the obtuse to acute apex. Capitula 20–50 mm in diameter, solitary, or more usually 2–several, solitary at the end of branches; opening in morning, closing in afternoon; peduncles swollen beneath the capitulum, glabrous, with scales. Involucral bracts in several rows, 10– 25 × 1.0–2.5 mm, medium to dark green with scarious margins and blackish-purple apices, oblong, obtuse and shortly white hairy at apex, otherwise glabrous or with a line of short, rigid, tooth-like hairs down the midrib. Flowers 14– 27 mm, all ligulate, the ligules bright yellow, with 5 lobes at apex, the outer often with a dirty yellowish-green stripe on the outer face. Styles orange. Receptacle flat, pitted; scales 5–20 mm, long-linear, long-acute at apex, scarious. Achene body 3–7 mm, orange-brown, cylindrical, ribbed, strongly muricate, at least the inner narrowed into a long beak; pappus dirty white, of 2 rows of hairs, the outer 3–6 mm and scabrid or sparsely plumose, the inner 9.3–12.5 mm and plumose, not dilated at base. Flowers 5–11. Visited freely by many kinds of insects, especially bees, and not automatically self-pollinated. The following two subspecies are easily identified when they grow in colonies but individual plants may be difficult

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to identify. When subsp. radicata is cut early in the year, especially on roadsides, the resulting secondary growth may look like subsp. ericetorum, but it usually has larger capitula. (a) Subsp. radicata Stems 30–60(–100 cm), usually more or less erect. Leaves 8–25 × 1–6 cm, often ascending. Inner involucral bracts 16–25 mm. 2n = 8. (b) Subsp. ericetorum Van Soest H. radicata var. salina (Gren.) Rouy; H. salina Gren. Stems (1–)7–30 cm, prostrate, decumbent or ascending. Leaves 1–12 × 0.5–3.0 cm, usually prostrate. Inner involucral bracts 10–17 mm. 2n = 8. Native. Meadows, pastures, waysides, waste ground, heaths, churchyards, moors, cliff-tops, dunes and shingle. Throughout Great Britain and Ireland. Europe northwards to about 63◦ N in Scandinavia and to northern Russia; Asia Minor; North Africa; naturalised in North America, Australia, New Zealand and elsewhere. A member of the European Southern-temperate element. Subsp. ericetorum is the plant of shingle, dunes and sandy heaths (see note under Leontodon saxatilis). It also occurs elsewhere on the coasts of western Europe. Subsp. radicata is the plant of other localities, particularly non-sandy areas. Variable intermediates occur where the two subspecies grow adjacent or are divided by intermediate habitat. 3. H. maculata L. Spotted Cat’s-ear Achyrophorus maculatus (L.) Scop.; Porcellites maculata (L.) Cass. Long-lived perennial herb with a stout, cylindrical, branched, blackish stock and several main roots. Stems 1–several, 20–75(−90) cm, yellowish-green, erect, striate, with pale, stiff, few to numerous, short to medium simple eglandular hairs, simple or branched, sometimes with small leaves, sometimes with small bracts. Leaves medium yellowish-green, usually spotted, blotched or streaked with brownish-purple and with a pale or reddish midrib on upper surface, paler beneath; basal in a rosette, the lamina 4– 15(–30)× 2–5 cm, elliptical, ovate, obovate or obovateoblong, subacute at apex, subentire to more or less sinuatedentate, with shallow, mammiform teeth, narrowed at the base to a short, winged petiole; cauline absent, or 1–2 small ones similar to basal; all with few to numerous, pale, stiff simple eglandular hairs on both surfaces and margins. Capitula 30–65 mm in diameter, campanulate, solitary at the end of the stem or 2–4, solitary at the ends of branches; peduncles sometimes slightly enlarged just below the capitulum, with few or no simple eglandular hairs. Involucral bracts in several rows, 12–25 × 2.0–2.5 mm, blackishgreen, the outer lanceolate, the inner linear-lanceolate, acute at apex, with pale, stiff simple eglandular hairs and often a tomentose margin. Flowers 36–50 mm, all ligulate, the ligules lemon yellow, shallowly 5-lobed at apex. Receptacle flat, pitted, with colourless, lanceolate, acute scales almost as long as the flowers. Achene body 5–7 mm, brown, lanceolate, transversely ridged, slightly muricate, all with a rather long, narrow beak; pappus 6–11 mm, whitish, of

1 row of plumose hairs not dilated at the base. Flowers 5–8. Hymenoptera and Diptera are the most frequent visitors, but there is a high proportion of infertile achenes. Plants also reproduce by axillary buds on the stock. 2n = 10. Plants on coastal cliffs tend to be smaller in all their parts and often have solitary heads, while those inland are often branched, but there are no clear-cut differences. Native. Grassy or open ground, mostly on calcareous or sandy soils, maritime cliffs on calcareous or serpentinederived soils, and on acid, granite cliffs in Jersey. Very local and decreasing in Great Britain north to Westmorland and in Jersey. Europe from the Pyrenees, southern France, northern Italy, north Balkan peninsula and southern Russia, northwards to about 66◦ N in Scandinavia and to Karelia and central Russia extending eastwards to central Asia. A member of the Eurosiberian Temperate element with a continental distribution. There is great variation in Continental Europe where colonies of very large plants are sometimes found. 23. Scorzoneroides Moench Oporinia D. Don nom. illegit.; Leontodon subgenus Oporinia A. R. Clapham Perennial herbs with latex, without rhizomes or stolons. Stems solitary to numerous, usually branched, thickened below the capitulum, leafless but with small, scale-like bracts. Leaves all basal, nearly entire to dentate or pinnately divided, glabrous or with few to numerous simple hairs. Capitula 1–7(–numerous), erect when young. Involucral bracts in several rows, merging into the bracts of the peduncles or stems. Flowers bisexual. Corolla ligulate, yellow, the outer with a reddish stripe on the outer face, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style branches long and slender, with both shaft and branches with very short hair. Receptacle flat, hairy but without scales. Achenes cylindrical; pappus of 1 row of plumose hairs. About 9 species in Europe, North Africa and western Asia. This species is reluctantly removed from the genus Leontodon because it seems to be much more nearly related to Hypochaeris. With Leontodon it has only the absence of receptacular scales in common, with Hypochaeris it has the characters of the branched stem, the capitula gradually narrowed into the stem, bracts on the stem and peduncles and simple, not forked, eglandular hairs in common. It has been made a Subgenus Oporinia of Leontodon, but it would be more sensibly made a subgenus of Hypochaeris. However, as it will not hybridise with either species of Leontodon (fide R. A. Finch pers. comm.) or with species of Hypochaeris (fide J. S. Parker pers. comm.), it is better regarded as a distinct genus. See also the relationship of Leontodon sensu stricto to Picris. Grime, J. P. et al. (1988). Comparative plant ecology. London. [Leontodon autumnalis L.] Holub, J. (1977). New names in Phanerogamae 5. Folia Geobot. Phytotax. 12: 293–311. 1. Leaves entire to dentate 1. Leaves deeply divided almost to midrib

2. 7.

23. Scorzoneroides 3. 2. Involucral bracts glabrous or with sparse hair 4. 2. Involucral bracts densely hairy 3. Stems 5–20 cm; leaves entire, denticulate or narrowly toothed; involucral bracts usually glabrous 1(i). autumnalis var. salina 3. Stems 8–45 cm; leaves usually undulate-dentate; involucral bracts often with some hairs 1(vi). autumnalis var. autumnalis 4. At least some of the leaves with the undivided portion in 5. the centre 10 mm or more wide 6. 4. Undivided portion in centre of leaf not more than 8 mm 5. Leaves subentire, undulate-dentate or with short, broad, downturned segments; involucral bracts 10–12 mm 1(x). autumnalis var. dentata 5. Leaves with broad, long, triangular teeth; involucral 1(xi). autumnalis var. latifolia bracts 8–12 mm 6. Dwarf plant of coasts (2–10 cm) in north Scotland; 1(iii). autumnalis var. nigrolanata leaves with stiff hairs 6. Taller plant of mountains (10–20 cm); leaves glabrous 1(iv). autumnalis var. alpina or nearly so 7. Plant with basal rosette of prostrate to ascending leaves and prostrate, decumbent or ascending stems 1(ii). autumnalis var. simplex 8. 7. Plant with erect stems and usually erect leaves. 9. 8. Involucral bracts glabrous or relatively sparsely hairy 10. 8. Involucral bracts densely hairy 9. Leaves with short lobes 1(vi). autumnalis var. autumnalis 9. Leaves with long narrow lobes 1(vii). autumnalis var. pinnatifida 10. Fairly short mountain or northern plant ((7–30(–40) cm); usually with few capitula (1–3(−5)); hairs on 1(v). autumnalis var. pratensis involucre blackish 10. Tall lowland plants (10–60 cm); usually with numerous capitula (3 or more); hairs on involucre pale, fuscous or 11. yellowish 11. Involucral bracts with pale or fuscous hair 1(viii). autumnalis var. coronopifolia 11. Involucral bracts with yellowish hair 1(ix). autumnalis var. cinerascens

1. S. autumnalis (L.) Moench Autumn Hawkbit Leontodon autumnalis L.; Hedypnois autumnalis (L.) Huds.; Apargia autumnalis (L.) Hoffm.; Virea autumnalis (L.) Gray; Oporinia autumnalis (L.) D. Don; Picris autumnalis (L.) All.; Scorzonera autumnalis (L.) Lam.; Virea pyreniaca Gray nom. illegit. Perennial herb with a premorse, oblique, usually branched stock and tap-root with fibrous side-roots, each branch of the stock terminating in a rosette of leaves. Stems single to numerous, 2–60 cm, yellowish-green, sometimes tinted brownish-purple, prostrate to ascending or erect, markedly ridged and channelled, often flexuous, glabrous or with sparse simple eglandular hairs, leafless, but with narrow bracts, simple or branched. Leaves few to numerous, all basal, the lamina 15–20 × 0.3–3.0 cm, dull medium green on upper surface with a pale midrib, paler beneath, sometimes flushed reddish-purple especially on the midrib, rarely fleshy, narrowly to broadly oblanceolate, obtuse to acute at apex, entire to remotely sinuate-dentate or deeply pinnatisect, the segments narrowly lanceolate to linear,

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obtuse or acute at apex, patent or recurved and usually very irregular, gradually narrowed at base to short or medium, winged petioles, sometimes bearing an occasional tooth, glabrous or with few to numerous, sometimes rather stiff simple eglandular hairs. Capitula 1– numerous on each stem, solitary at the ends of branches, 12–40 mm in diameter; peduncles like the stem, with several small, lanceolate, often colourless bracts particularly just below the capitulum. Involucral bracts in several rows, 6–15 × 0.8–1.5 mm, dull green with dark tips, linearlanceolate, gradually narrowed to an acute apex, usually with some arachnoid hairs, without or with few to dense simple eglandular hairs and minute, whitish, soft hairs. Flowers 4–16 mm, all ligulate, the ligules deep yellow, rarely lemon yellow, the outer usually with a reddish or purplish stripe on the outer face, shortly 5-lobed at apex. Stigmas discoloured. Receptacle slightly convex, pitted, without scales. Achenes 3.5–8.0 mm, reddish-brown to dark brown, cylindrical, slightly narrowed above, transversly muricate; pappus 3–8 mm, very pale brown or grey, of 1 row of plumose hairs. Flowers 6–11. Freely visited by a great variety of insects and automatically self-pollinated. 2n = 12, 24. Extremely variable in habit, height, division of leaves, branching of stem and indumentum. More than one of the following eleven varieties is not usually found in one colony and most of them can be regarded as ecotypes. A division into two subspecies, as has been previous accepted, does not seem satisfactory. The nomenclature of the varieties is unsatisfactory, but is the best that can be done without bringing together all the types. (i) Var. salina (Aspegren) P. D. Sell Leontodon autumnalis var. salinus (Aspegren) Lange Stems slender, few, 5–20 cm, erect. Leaves narrow, entire, denticulate or narrowly toothed, often fleshy. Capitula 1–2 per stem, 15–35 mm in diameter. Involucral bracts 8–12 mm, glabrous. This variety may often meet the next where salt-marsh runs into shingle. (ii) Var. simplex (Duby) P. D. Sell Leontodon autumnalis var. simplex Duby; Oporinia autumnalis var. minima auct.; Leontodon autumnalis var. minimus auct. Stems slender, few to numerous, 2–25(–30) cm, prostrate, decumbent or ascending. Leaves in a spreading rosette often nearly flat on the ground to ascending, deeply divided almost to the midrib with rather narrow lateral lobes. Capitula 1–3 per stem, 20–30 mm in diameter. Involucral bracts 6–10 mm, with usually few to numerous, rather dark simple eglandular hairs. (iii) Var. nigrolanata (Fr.) P. D. Sell Leontodon autumnalis var. nigrolanatus Fr. Stems slender, few, 2–10 cm, ascending. Leaves few in a rosette, dentate or undulate-dentate with rather small teeth, with rather stiff hairs. Capitula 1–2 per stem, 20–30 mm in diameter. Involucral bracts 8–14 mm, with dense, long, greyish to blackish simple eglandular hairs which extend on to the upper stem.

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(iv) Var. alpina (Gaudin) P. D. Sell Leontodon autumnalis var. alpina (Gaudin) Gren. & Godr.; Apargia autumnalis var. alpina Gaudin; Leontodon autumnalis subsp. borealis Ball; Oporinia autumnalis var. alpina (Gaudin) DC. Stems slender, few, 10–20 cm, erect. Leaves in a rosette, usually ascending or erect, subentire to undulate-dentate, the teeth often curved down, glabrous or nearly so. Capitula usually solitary, rarely 2 per stem, 20–30 mm in diameter. Involucral bracts 8–12 mm, with numerous to dense, brownish to blackish, long simple eglandular hairs which extend on to the upper stem. (v) Var. pratensis (Hornem.) P. D. Sell Apargia pratensis Hornem.; Hieracium taraxaci L.; Leontondon autumnalis var. taraxaci (L.) Hartm.; Leontodon pratensis (Hornem.) Rchb.; Oporinia pratensis (Hornem.) Less.; Leontodon autumnalis var. pratensis (Hornem.) W. D. J. Koch; Oporinia autumnalis var. sordida Bab.; Hedypnois taraxaci (L.) Vill.; Apargia taraxaci (L.) Willd. Stems slender, few, 7–30(–40) cm, erect or ascending. Leaves usually erect or ascending, deeply divided to the midrib with narrow lateral lobes. Capitula 1–3(–5) per stem, 25–40 mm in diameter. Involucral bracts 9–15 mm, with dense, long, blackish simple eglandular hairs which extend on to the upper part of the stem. (vi) Var. autumnalis Leontodon autumnalis var. vulgaris Neilr. nom. illegit. Stems slender, few, 8–45 cm, flat to ground, erect or ascending. Leaves erect or ascending, of medium width, subentire to shortly and broadly toothed, rarely more deeply toothed. Capitula 1–numerous per stem, 18–30 mm in diameter. Involucral bracts 8–12 mm, with few, sometimes fairly numerous, short, pale simple eglandular hairs. (vii) Var. pinnatifida (Schur) P. D. Sell Leontodon autumnalis var. pinnatifidus Schur Stems slender, often very numerous, 30–45 cm, usually erect. Leaves usually erect, divided almost to the midrib with linear lateral segments. Capitula 3–numerous per stem, 15– 30 mm in diameter. Involucral bracts 8–11 mm, with few to fairly numerous, short to medium, pale simple eglandular hairs. (viii) Var. coronopifolia (Lange) P. D. Sell Leontodon autumnalis var. coronopifolius Lange Stems slender, often very numerous, 25–60 cm, usually erect. Leaves usually erect, divided almost to the midrib, with very long, narrow linear segments. Capitula 3– numerous per stem, 15–30 mm in diameter. Involucral bracts 8–12 mm, with numerous to dense, short to medium, pale or fuscous simple eglandular hairs. (ix) Var. cinerascens (Briq.) P. D. Sell Leontodon autumnalis var. cinerascens Briq. Stems robust, often several, 20–30 cm, erect. Leaves usually erect, divided almost to the midrib with linear lateral segments. Capitula 3–5 per stem, 25–30 mm in diameter. Involucral bracts 10–15 mm, with dense, long, yellowish simple eglandular hairs.

(x) Var. dentata (Holuby ex Schur) P. D. Sell Leontodon autumnalis var. dentatus Holuby ex Schur Stems robust, several, erect. Leaves usually erect, subentire or runcinate-dentate with short, broad teeth. Capitula 3–numerous per stem, 20–40 mm in diameter. Involucral bracts 10–12 mm, with dense, long, pale to slightly dusky simple eglandular hairs. (xi) Var. latifolia (Schur) P. D. Sell Leontodon autumnalis var. latifolius Schur Stems 25–60 cm, robust, several, erect. Leaves erect, wide in the undivided part, with widely spaced, long, spreading or ascending triangular teeth or lobes. Capitula 3–numerous per stem, 20–30 mm in diameter. Involucral bracts 9– 11 mm, with numerous pale to dusky simple eglandular hairs. Native. Meadows, pastures, waysides, waste places, screes, dunes, shingle, salt-marsh, heaths, moors and cliff ledges. Abundant throughout Great Britain and Ireland. Throughout Europe, though rarer in the Mediterranean region and absent from some of the islands, north and west Asia and north-west Africa; introduced in North America, Greenland and probably elsewhere. A member of the European Boreo-temperate element. Var. salina is the plant of salt-marshes. It also occurs in western and northern Europe. Var. simplex is the plant of heaths, moors, shingle and dunes by the sea and cliff-tops by the sea (see note under Leontodon saxatilis). Var. nigrolanata occurs by the coast in northern Scotland and the islands. It is also by the coast elsewhere in north-west Europe. Var. alpina is a plant of mountain cliff ledges in Scotland and Wales. It is widespread in the mountains of Continental Europe. Var. pratensis occurs in the mountains of Wales, northern England and Scotland coming down to the coast in the north. It is widespread in Continental Europe. Var. autumnalis, var. pinnatifida and var. coronopifolia are the common plants of roadsides and waste and grassy places. Where grass has been cut short they can form a solid mat of yellow, but in many cases may be secondary growth when the leaves can be narrower than normal. Occasionally they form mixed colonies. They all occur widely in Continental Europe. Var. cinerascens was collected at the Red Head in Forfar in 1905 and has recently been found in quantity at Histon and Impington in Cambridgeshire. It is a striking plant and may have been introduced. It was described from the Alps. Var. dentata has been seen from Pembrokeshire, Monmouthshire, Cardiganshire, Caernarvonshire and Co. Cork. In the field its general appearance is that of Hypochaeris radicata subsp. ericetorum with which it grows on the Cardiganshire coast. Var. latifolia is a plant of roadsides, seeming to grow in occasional clumps and not large sheets. It is widely scattered in Continental Europe. 24. Leontodon L. Thrincia Roth Perennial herbs with latex, without rhizomes or stolons. Stems solitary to numerous, usually simple, leafless but with small, scale-like bracts, or rarely with a small leaf-like bract. Leaves all basal, dentate to pinnately divided,

24. Leontodon usually with forked hairs, rarely glabrous. Capitula usually solitary. Involucral bracts in several rows, not merging with the bracts of the stem. Flowers bisexual. Corolla ligulate, yellow, the outer with an orange, reddish or greyishviolet stripe on the outer face, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches with very short hair. Receptacle flat, hairy, but without scales. Achenes more or less cylindrical, at least the inner beaked; pappus of 2 rows of plumose hairs. About 36 species throughout Europe, North Africa and south-west Asia to Iran. Over its total range it is impossible to distinguish Leontodon from Picris. The best character is the presence of anchor-shaped hairs in Picris and only forked hairs in Leontodon, but in some species it is difficult to distinguish its genus using even these characters. In our area Picris always has large cauline leaves and Leontodon has none. L. saxatilis has been separated as a separate genus Thrincia, because of the pappus of scales on the outer flowers, but frequent hybrids between it and L. hispidus suggest it is not worth even putting it in a different section. Finch, R. A. (1967). Natural chromosome variation in Leontodon. Heredity (London) 22: 359–386. Grime, J. P. et al. (1988). Comparative plant ecology. London. [L. hispidus.] 2. 1. All achenes with a pappus of hairs 1. Outer row of achenes with a pappus of a cup of scales, 3. inner achenes with a pappus of hairs 2. Leaves and involucral bracts clothed with rigid hairs 1. hispidus subsp. hispidus 2. Leaves and involucral bracts glabrous or nearly so 1. hispidus subsp. danubialis 3. Stems up to 12 cm; leaves up to 8 × 1 cm, forming a flat rosette; capitula 10–20 mm in diameter; involucral bracts 2(iii). saxatilis var. arenarius glabrous or hairy 3. Stems up to 30(–45) cm; leaves up to 20 × 3 cm, ascending or erect; capitula 15–30 mm in diameter; 4. involucral bracts hairy 4. Leaves more or less hairy; involucral bracts hairy but not 2(i). saxatilis var. saxatilis densely so (widespread) 4. Leaves at base and involucral bracts densely hairy 2(ii). saxatilis var. pristis (Channel Islands)

1. L. hispidus L. Rough Hawkbit Apargia hispida (L.) Hoffm.; Hedynois hispida (L.) Huds.; Virea hispida (L.) Gray Perennial herb with vertical or oblique, usually branched, truncate stock, each branch terminating in a rosette of leaves. Stems 1–numerous, (5–)10–60 cm, yellowish-green, erect, striate, simple, with numerous, white, subrigid simple and 2- to 3-fid eglandular hairs, without bracts. Leaves all basal, the lamina 3–35 × 0.3–4.0 cm, yellowish-green, the lower surface hardly different from the upper, with a pale midrib, oblanceolate to linear-oblanceolate, more or less acute to obtuse at apex, sinuate-dentate to pinnatifid, with broad to narrow mammiform teeth or triangular lobes, long attenuate at base to a winged, hairy petiole on

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which the teeth sometimes descend, usually with few to numerous, white, subrigid simple or 2- to 3-fid eglandular hairs, rarely glabrous or almost so. Capitula 25–40 mm in diameter, solitary, drooping in bud; peduncle clothed like the stem. Involucral bracts in several rows, imbricate, or the outer somewhat spreading, 5–15 × 1.0–1.5 mm, linearlanceolate, obtuse to acute at the densely, very shortly white hairy apex, with numerous to dense, whitish, subrigid simple or 2- to 3-fid eglandular hairs or rarely glabrous. Flowers 11–20 mm, all ligulate, the ligules golden yellow, the outermost orange, reddish or rarely greyish-violet on the outer face, shallowly 5-lobed at apex. Styles yellow. Receptacle flat, pitted, without scales. Achenes 5–8 mm, pale brown, fusiform, narrowed above but not beaked, with muricate longitudinal ribs and numerous transverse ridges; pappus 9–11 mm, dirty white, of 2 rows of hairs, the inner plumose, the outer simple and scabrid. Flowers 6–9. Freely visited by a variety of insects, especially bees and flies. 2n = 14. Native. Meadows, pastures, grassy slopes, waysides and other grassy places, mostly on calcareous soils, up to 600 m in northern England. Common in England, Wales and much of southern Scotland, but local further north and absent from the Isle of Man and the Channel Islands; locally frequent in Ireland. Europe from Portugal and central Spain, southern Italy and Greece northwards to Norway, Sweden and Karelia; Turkey; Caucasus; northern Iran. A member of the European Temperate element. Our plant is mostly subsp. hispidus which occurs through much of the range of the species and has the leaves and involucral bracts with rigid hairs. If you travel across central Europe, somewhere about Vienna all the plants have glabrous leaves and the involucral bracts are nearly glabrous, and this situation continues across the north Balkan peninsula and into central Russia. Such plants have been referred to subsp. danubialis (Jacq.) Simonk. (L. hastilis L.; L. danubialis Jacq.; L. hastilis var. glabratus W. D. J. Koch; L. hispidus var. glabratus (W. D. J. Koch) Bisch.) Similar glabrous plants are rarely found in Great Britain and it is not known if they are introductions or local variants of subsp. hispidus. × saxatilis = L. × vegetus Finch & P. D. Sell This vigorous hybrid can usually be recognised by its having many outer achenes with a pappus of scales or scales and a few hairs, and the leaf shape and indumentum and involucre size and indumentum intermediate between the accompanying parents. It has low pollen and achene fertility. Backcrosses probably occur. 2n = 11. Probably widespread and should be looked for wherever the parents grow adjacent. Not known outside Great Britain. 2. L. saxatilis Lam. Lesser Hawkbit Hyoseris taraxacoides Vill.; L. taraxacoides (Vill.) M`erat; L. hirtus auct.; Thrincia hirta Roth; Hedypnois hirta auct.; Thrincia hispida auct.; Thrincia leysseri Wallr. nom. illegit.; Apargia hirta auct.; Thrincia taraxacoides (Vill.) Gaudin; L. leysseri Beck; Thrincia nudicaulis auct.; L. nudicaulis auct.; Colobium taraxacoides (Vill.) Holub Perennial, rarely biennial herb with a short, erect, premorse stock and rather fleshy roots. Stems few to numerous,

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Var. nigrolanata (Fr.) P. D. Sell (black hairy)

Var. salina (Aspegren) P. D. Sell (glabrous)

Var. alpina (Gaudin) P. D. Sell (black hairy)

Var. simplex (Duby) P. D. Sell (hairy)

Var. pratensis (Hornem.) P. D. Sell (black hairy) Var. autumnalis (pale hairs)

Var. pinnatifida (Schur) P. D. Sell (rather few pale hairs)

Leaves of varieties of Scorzoneroides autumnalis (L.) Moench Leaf shape should be used in conjunction with hairiness of involucral bracts indicated in brackets

24. Leontodon

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Var. cinerascens (Briq.) P. D. Sell (dense yellow hairs)

Var. coronopifolia (Lange) P. D. Sell (numerous pale hairs)

Var. dentata (Holuby ex Schur) P. D. Sell (dense pale hairs) Scorzoneroides autumnalis (L.) Moench

Var. latifolia (Schur) P. D. Sell (numerous pale hairs)

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2.5–30(–45) cm, pale yellowish-green, decumbent, ascending or erect, striate, simple, glabrous or with few to numerous, short or medium, white, forked eglandular hairs, without bracts. Leaves numerous, all basal, the lamina 2– 20(–25) × 0.5–3.0 cm, yellowish-green on upper surface, slightly paler beneath, prostrate to erect, oblanceolate, linear-oblanceolate or linear-oblong, obtuse-mucronate to more or less acute at apex, entire or remotely sinuate-dentate to runcinate-pinnatifid, gradually narrowed at base to a winged petiole, nearly glabrous or with few to numerous, white, forked eglandular hairs on both surfaces and the margins. Capitula 10–30 mm in diameter, solitary, drooping in bud; peduncles clothed like the stem. Involucral bracts in several, imbricate rows, 5–11 × 1.0–1.5 mm, olive green, the inner with paler margins, linear-lanceolate, obtuse at apex, glabrous or with few to numerous, short to medium, white, forked eglandular hairs, particularly along the midrib. Flowers 10–14 mm, all ligulate, the ligules golden yellow, the outermost with a greyish-violet stripe on outer face, shallowly 5-lobed at apex. Receptacle flat, pitted, without scales. Achenes of 2 kinds, the outer pale brown, more or less cylindrical, curved, faintly transversely wrinkled, with longitudinal ribs, surmounted by a cup of small, scarious scales and not beaked, the inner about 5 mm, chestnut, attenuate above and more or less shortly beaked, straight, with strongly muricate, longitudinal ribs and a brownish-white pappus of plumose hairs. Flowers 6–9. Visited by many kinds of bees and syrphids. 2n = 8. (i) Var. saxatilis Stems to 30(–45) cm, with few, medium hairs or none. Leaves up to 20 × 3 cm, ascending or erect, usually hairy. Capitula 15–30 mm in diameter. Involucral bracts up to 11 mm, usually hairy. (ii) Var. pristis (Druce) P. D. Sell L. nudicaulis var. pristis Druce Stems up to 15 cm, with numerous long, hairs. Leaves up to 8 × 2 cm, ascending, with numerous, long, stiff hairs, dense and shaggy near the base. Capitula 15–20 mm in diameter. Involucral bracts up to 9 mm, densely hairy. (iii) Var. arenarius (Duby) P. D. Sell Thrincia hirta var. arenaria (Duby) DC.; L. hastilis var. arenarius Duby; Thrincia laevis Lag.; Thrincia psilocalyx Rchb.; L. psilocalyx (Rchb.) M`erat; Thrincia hirta var. psilocalyx (Rchb.) DC.; Thrincia nudicalyx Lag. Stems up to 12 cm, glabrous or slightly hairy. Leaves up to 8 × 1 cm, forming a flat rosette, more or less hairy. Capitula 10–20 mm in diameter. Involucral bracts 3–8 mm, glabrous or hairy. The pappus of the outer achenes is the only character that will always separate this species from L. hispidus. Var. pristis strongly resembles L. hispidus. Small plants of L. hispidus found on dry chalk grassland and autumnal flowering plants of L. hispidus strongly resemble the general appearance of L. saxatilis. L. saxatilis var. arenarius, Scorzoneroides autumnalis var. simplex and Hypochaeris radicata subsp. ericetorum grow intermixed on maritime shingle and fixed dunes and can cause untold trouble to those not familiar with them. To make matters more difficult

the Leontodon and Hypochaeris often flower very early and have a second flowering when Scorzoneroides flowers later in the year. Native. Dry grassland, especially on base-rich soils and maritime shingle and dunes. Common in Great Britain north to central Scotland and frequent in Ireland, the Channel Islands and the Isle of Man. Europe, northwards to Scotland, Denmark, Gotland and central Russia. A member of the Suboceanic Southern-temperate element. All our plants are referable to subsp. saxatilis. Var. arenarius is the plant of shingle, fixed dunes and open grassland. Var. saxatilis occurs in more closed grassland. Var. pristis is known only from Guernsey and Jersey in the Channel Islands. 25. Picris L. Biennial to perennial herbs with latex, without rhizomes or stolons. Stems usually solitary, usually branched, leafy. Leaves alternate, undulate-dentate with numerous stiff hairs at least some of which have hooked, anchor-like apices, the upper more or less amplexicaul. Capitula few to numerous. Involucral bracts in several rows, all lanceolate to narrowly elliptical. Flowers bisexual. Corolla ligulate, yellow, the outer often with a reddish stripe on the outer face, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches with very short hairs. Receptacle flat, without scales. Achenes fusiform, rather flattened, the inner more shortly beaked; pappus in 2 rows, the outer of rough simple hairs, the inner of plumose hairs. About 40 species in Europe, Asia, North and Tropical Africa and Australia. Beadell, A. (1936). Picris spinulosa Bertol. Rep. Bot. Soc. Exch. Cl. Brit. Isles 11: 178–179. Britton, C. E. (1936). Picris spinulosa Bertol. Jour. Bot. (London) 74: 354–355. 1. Capitula with very short curled hairs and occasionally a longer, straight hair on the median line of the involucral bracts, all on short peduncles and some sessile or nearly 1(a). hieracioides subsp. spinulosa so 1. Capitula with numerous longer, straight hairs as well as very short curly hairs on the involucral bracts; on long or 2. short peduncles or some sessile 2. Involucral bracts dull green; capitula on long branches, none of the capitula more or less sessile 1(b). hieracioides subsp. hieracioides 2. Involucral bracts blackish; capitula on long or short 3. branches, some often more or less sessile 3. Capitula on long branches; involucral bracts 10–15 × 2.0–2.5 mm, narrowly elliptical to 1(d). hieracioides subsp. grandiflora oblong-elliptical 3. Capitula in a small cluster on long branches often forming a subumbellate inflorescence with some capitula subsessile; involucral bracts 8–13 × 1.0–1.5 mm, 4. lanceolate or oblong-lanceolate 4. At least some leaves more than 1.5 cm wide 1(c,i). hieracioides subsp. villarsii var. villarsii 5. 4. Leaves rarely exceeding 1.0 cm wide 5. Stem with long branches often from the base 1(c,ii). hieracioides subsp. villarsii var. leteae

25. Picris 5. Stem without branches or with very short branches above 1(c,iii). hieracioides subsp. villarsii var. arenaria

1. P. hieracioides L. Hawkweed Oxtongue Hedypnois hieracioides (L.) Huds.; Crepis hieracioides (L.) Lam. Biennial or perennial herb with a thick stock and long, fleshy roots. Stems 15–100(–145) cm, pale green, sometimes suffused brownish-purple, erect, rigid, furrowed, with few to numerous, short to medium, simple, forked and anchor-like eglandular hairs, often with spreading or ascending branches sometimes from the very base, leafy. Leaves yellowish-green on upper surface, paler beneath, sometimes tinted purplish; basal and lower cauline with lamina 6–20 × 1–5 cm, lanceolate, ovate, narrowly elliptical or oblong, obtuse to acute at apex, entire to sinuatedentate with mammiform or sharply mammiform teeth, narrowed at base into a short petiole; median and upper cauline similar but smaller, sessile and more or less amplexicaul; all with more or less numerous, short to medium, simple, forked and anchor-like, rigid hairs on both surfaces and the margins. Capitula few to numerous, 20–40 mm in diameter, terminal on the main stem and branches, some often sessile; peduncles short to long, bracteate and somewhat thickened distally, with short, curled hairs and numerous to dense, short to long, rigid simple, forked and anchor-shaped hairs. Involucral bracts in several rows, 9–15 × 1.5–2.5 mm, green to blackish, linear-lanceolate to narrowly elliptical, obtuse to acute at apex, glabrous or with numerous very short curled hairs and few to numerous, pale to dark simple, forked and anchor-like hairs. Flowers all ligulate, the ligules bright deep yellow and 5-lobed at apex, the outer often with a reddish stripe on the outer face. Receptacle flat, pitted, without scales. Achenes 3–6 mm, reddish-brown, fusiform, somewhat flattened, slightly curved, with fine, interrupted, transverse wrinkles, weakly ribbed, shortly beaked; pappus cream-coloured, in 2 rows, the outer of simple hairs, the inner of plumose hairs. Flowers 7–9. Freely visited by flies and bees, but may be apomictic. 2n = 10. Very variable in hairiness, mode of branching and size of capitulum. The following four subspecies may not cover all the variation. They are more easily recognised in the field than in the herbarium. (a) Subsp. spinulosa (Bertol. ex Guss.) Arcangeli P. spinulosa Bertol. ex Guss.; P. stricta Jord.; P. setulosa Guss. ex Ces., Pass. & Gibelli; P. hieracioides subsp. setulosa (Guss. ex Ces., Pass. & Gibelli) Arcangeli; P. hieracioides var. incana Druce; P. hieracioides var. stricta (Jord.) Druce; P. hieracioides var. umbellata auct. Capitula crowded at apex of stems on very short peduncles, the lateral capitula often solitary and sessile or nearly so. Involucral bracts 9–11 × 1.0–1.5 mm, dull green, linearlanceolate or lanceolate, with very short, curled hairs and sometimes an occasional longer, straight hair on the median line. (b) Subsp. hieracioides P. hieracioides var. gracilis auct. Capitula without clustering in an open corymb on long peduncles. Involucral bracts 11–13 × 1.0–1.5 mm, dull

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green, lanceolate or oblong-lanceolate, with very short curled hairs and numerous medium to long hairs mainly down the median line. (c) Subsp. villarsii (Jord.) Nyman P. sonchoides Vest; P. crepoides Saut.; P. villarsii Jord.; P. hieracioides subsp. crepoides (Saut.) Nyman; P. pyrenaica auct.; P. spinulosa auct.; P. hieracioides subsp. sonchoides (Vest) Thell. Capitula in small clusters on long or short branches, often subumbellate in upper part and some capitula subsessile. Involucral bracts 10–13 × 1.0–1.5(–2.0) mm, blackish, lanceolate to oblong-lanceolate, with very short, curled hairs and few to numerous, short to long, pale hairs. (i) Var. villarsii (Jord.) P. D. Sell Stem much branched. At least some leaves more than 1.5 cm wide. Named after Dominique Villars (1745–1814). (ii) Var. leteae (Ny´ar.) P. D. Sell P. sonchoides var. leteae Ny´ar. Stem with numerous long branches often from the base. Leaves up to 1.0(–1.5) cm wide. (iii) Var. arenaria (Ny´ar.) P. D. Sell P. sonchoides var. arenaria Ny´ar. Stem unbranched or with short branches in the upper part. Leaves up to 1.0(–1.5) cm wide. (d) Subsp. grandiflora (Ten.) Arcangeli P. paleacea Vest; P. grandiflora Ten.; P. auriculata Sch. Bip.; P. tatrae Borb´as; P. hieracioides subsp. paleacea (Vest) Domin & Podp.; P. hieracioides subsp. tatrae Domin & Podp.; P. hieracioides subsp. auriculata (Sch. Bip.) Hayek Capitula solitary or in clusters at the end of long branches, the inflorescence open. Involucral bracts 10–15 × 2.0–2.5 mm, blackish, narrowly elliptical to oblong-elliptical, with very short, curled hairs and short to long pale and duskybased hairs. Native. Grassland, especially on calcareous slopes, waysides, railway banks and rough ground. Locally common in the lowlands of England and Wales and sparingly north to Cumberland and Northumberland, introduced in south Scotland; and on railway banks in Ireland. Europe northwards to Denmark, southern Sweden and Karelia; Asia east to Kamchatka and Japan; introduced in North America, Australia and New Zealand. A member of the Eurasian Temperate element. The common native plant is subsp. hieracioides. It occurs also throughout much of the European range of the species. Subsp. spinulosa is the plant of Jersey. It is not known if any other subspecies occur there. It is the subspecies of south and south-central Europe which may be at its northernmost limit in Jersey. A plant collected by G. C. Druce at Princes Risborough in 1904 and called P. gracilis Jord. appears to be it, but its status is doubtful. Subsp. villarsii was first found in Kent in 1935 and still persists in the area. It was initially confused with P. spinulosa. It occurs also in a sandpit between Mepal and Chatteris and in great abundance on the old railway and in other places at Histon in Cambridgeshire where all three varieties occur. A specimen from Southwick in Sussex also appears to be

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it. A plant of west and central Europe it is almost certainly introduced in Great Britain. The varieties are retained as they appear to be very distinct, but one experiment suggests they may not be. Subsp. grandiflora occurs at Milton and along the old railway to Histon in Cambridgeshire where it could have been introduced on the tracks of army tanks. On the Gog Magog hills near Cambridge it was probably introduced with wild flower seed, as it could also have been at Litlington chalk pit in that county. There are old specimens from Tenby in Glamorganshire and Buxton in Derbyshire which also appear to be it. It is native of central and southeast Europe. In the field it is a robust plant often over one metre high with large black heads and is easily recognised. Any populations of this species not obviously native need careful checking. 26. Helminthotheca Zinn Helminthia Juss. Annual or biennial herbs with latex, without rhizomes or stolons. Stems usually solitary, usually branched, leafy. Leaves dentate to pinnately lobed, with numerous stiff hairs at least some of which have hooked, anchor-like apexes (glochidiate), more or less amplexicaul. Capitula few to numerous. Involucral bracts of 2 kinds, the outer ovate, acuminate at apex and cordate or rounded at base, the inner lanceolate and awned at apex. Flowers bisexual. Corolla ligulate, yellow, the outer often with a reddish stripe on the outer face, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches with very short hairs. Receptacle flat, pitted, without scales. Achenes of 2 kinds, the inner more or less straight, the outer curved, all with a slender beak; pappus of 2 rows of plumose hairs. Four species in Europe and the Mediterranean region east to Iran; some species introduced as weeds elsewhere. Lack, H. W. (1975). A note on Helminthotheca Zinn (Compositae). Taxon 24: 111–112. 1. Plant spreading or prostrate; outer involucral bracts 1(i). echioides var. mollis rounded at base 2. 1. Plant erect; outer involucral bracts cordate at base 1(iv). echioides var. incisa 2. Leaves incised to halfway 3. 2. Leaves dentate but not incised 3. Leaves narrowly elliptical, oblong-elliptical or 1(ii). echioides var. pratensis oblong-lanceolate 1(iii). echioides var. echioides 3. Leaves broadly elliptical

1. H. echioides (L.) Holub Bristly Oxtongue Picris echioides L.; Helminthia echioides (L.) Gaertn.; Picris ruderalis Salisb. nom. illegit. Annual or biennial herb with numerous, stout, pale brown roots. Stem 15–90 cm, pale yellowish-green often completely suffused brownish-purple, markedly striate, usually erect sometimes spreading or prostrate, with numerous to dense, rigid, very unequal, short to long, pale to brownishpurple, often bulbous-based anchor-shaped hairs throughout or the longer hairs absent, usually branched above. Leaves dull yellowish-green on upper surface, slightly paler beneath, basal more or less in a rosette, numerous, the lamina 3.5–25 × 1.5–10 cm, elliptical, oblong-oblanceolate or

oblanceolate, more or less acute or sometimes obtuse at apex, entire to sinuate-dentate, gradually narrowed at base to a winged, hairy petiole up to 19 cm; lower cauline similar to basal but with semiamplexicaul petioles; middle and upper cauline gradually smaller upwards, ellipticaloblong, oblong, lanceolate or ovate, more or less acute at apex, more or less sinuate-dentate, sessile, semiamplexicaul with rounded auricles and sometimes decurrent; veins impressed above and prominent beneath; all with numerous to dense, medium to long, rigid, pale simple eglandular hairs and shorter, rigid anchor-shaped hairs making them very rough on both surfaces and the margins, the longer hairs sometimes absent. Capitula few to numerous, 20–37 mm in diameter, rather crowded on short lateral stalks in an irregular corymb; peduncles pale green, with numerous, very unequal, short to long, anchor-shaped hairs. Involucral bracts of 2 kinds, the outer 3–5, 15–20 × 7–10 mm, ovate, acuminate at apex, cordate or rounded at base, the inner 12– 15 × 2–4 mm, linear, obtuse at apex, with a spiny-pectinate awn from just below the apex, all yellowish-green the inner often suffused brownish-purple at the apex and pectinateciliate, the hairs rigid very unequal and sometimes anchorshaped, the longer hairs sometimes absent. Flowers 20–22 mm, all ligulate, the ligules golden yellow, rarely pale yellow, the outermost sometimes more or less reddish-purple beneath, with 5 lobes at apex. Styles greenish. Receptacle flat, pitted, without scales. Achenes 5–7 mm (with beak), the central reddish-brown, oblong or obovoid, more or less straight and glabrous, the marginal whitish, curved, and downy on the ventral side, all transversely wrinkled and with a slender beak about as long as the body; pappus 6–8 mm, pure white, of 2 rows of plumose hairs, deciduous with the end of the beak. Flowers 5–10. Visited by bees, but said to be apomictic. 2n = 10. Very variable in height, branching, leaf dissection and amount of purplish suffusion. (i) Var. mollis (Duby) P. D. Sell Picris humifusa Willd.; Helminthia humifusa (Willd.) Trevis; Helminthia echioides var. mollis Duby; Helminthia echioides subsp. humifusa (Willd.) Arcangeli Plant spreading or prostrate with few longer rigid hairs. Leaves medium to large, dentate but not incised. Outer involucral bracts rounded at base. (ii) Var. pratensis (Chevall.) P. D. Sell Helminthia pratensis Chevall.; Helminthia echioides var. pratensis (Chevall.) DC. Plant erect with longer rigid hairs as well as short ones. Leaves narrowly ellipical, oblong-elliptical or oblonglanceolate, dentate, but not incised. Outer involucral bracts cordate at base. (iii) Var. echioides Plant erect, with longer rigid hairs as well as shorter ones over much of the plant. Leaves broadly elliptical, large, dentate but not incised. Outer involucral bracts cordate at base. (iv) Var. incisa P. D. Sell Plant erect with longer rigid hairs as well as short ones. Leaves large and incised halfway to the midrib into

27. Scorzonera broad, dentate lobes. Outer involucral bracts cordate at base. Possibly introduced. Waysides, hedgebanks, field margins, and waste and rough places and coastal cliffs. Frequent in south and east England, scattered elsewhere in England and in the Channel Islands; rare in Scotland; very local in south and east Ireland. Native in the Mediterranean region, Canary Islands and south-west Asia; introduced elsewhere in Europe and North America. A member of the European Southern-temperate element. Var. echioides is the common form. There is a large population of var. mollis at Portland Bill, Dorset, and it should be looked for elsewhere on the south coast where it may be native. Occasional plants have been seen inland. It seems to be a plant of Mediterranean coasts. Var. pratensis and var. incisa occur in Cambridgeshire and var. pratensis occurs in a few other places in southern England, but their general distribution is unknown. Var. pratensis was described from France. 27. Scorzonera L. Perennial herbs, with latex, without rhizomes or stolons. Stems solitary or few, leafy, simple or with 1–2 branches. Leaves alternate, entire. Capitula solitary at the ends of stems or branches. Involucral bracts in several rows. Flowers bisexual. Corolla ligulate, yellow, the outermost usually flushed purplish, brownish or reddish on outer face, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches with very short hairs. Receptacle flat, without scales. Achenes cylindrical, but angled by ribs, slightly flattened, not beaked; pappus of several rows of plumose hairs. About 175 species throughout Europe and Asia, but mainly in south-west and central Asia, rarely in North Africa. Druce, G. C. (1929). Scorzonera humilis L. Jour. Bot. (London) 67: 26–27. Phillips, R. & Rix, M. (1993). Vegetables. London. [S. hispanica.] Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [S. humilis.] Woodman, J. P. (1997). Scorzonera humilis – new to Wales. B.S.B.I. News 75: 70–71. 1. Capitula 20–35 mm in diameter; involucral bracts 3.0–4.5 mm wide; at least the outer achenes with 1. humilis crenate-tuberculate ribs 1. Capitula 40–60 mm in diameter; involucral bracts 4–7 mm wide; at least the outer achenes with peg-like 2. hispanica tubercles on the ribs

1. S. humilis L. Viper’s Grass Perennial herb with a stout, vertical, black rootstock. Stems solitary or few, 5–50(–120) cm, yellowish-green, erect or ascending, more or less arachnoid-hairy at first, becoming almost glabrous, usually simple, leafy. Leaves with lamina 5–30 × 0.3–3.5 cm, medium yellowish-green on upper surface, paler beneath, flat, linear to lanceolate-elliptical or ovate-elliptical, long-acute to long-acuminate at apex, entire, gradually narrowed at base into a half-sheathing, usually distinct petiole, the upper cauline usually much

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narrower than the lower cauline and basal but more abruptly widened into a semiamplexicaul base, all at first greywoolly becoming nearly glabrous. Capitula usually solitary, 20–35 mm in diameter, rounded at base. Involucral bracts in several rows, 7–30 × 3.0–4.5 mm, green, with scarious tips and margins, woolly at base, the outer ovate, the inner oblong-lanceolate, all obtuse at apex. Flowers 30– 60 mm, all ligulate, the ligules pale to medium yellow, rarely whitish, often reddish or brownish on the outer face of the outer, shallowly 5-lobed at apex. Styles yellow. Receptacle flat, pitted, without scales. Achenes 7–11 mm, the inner straw-coloured, the outer purplish-brown, all cylindrical but angled with minutely crenate-tuberculate ribs; pappus 10– 15 mm, dirty white, of several rows of plumose hairs. Flowers 5–7 but actual flowering period very short. Visited by various bees and other insects as well as being automatically self-pollinated. 2n = 14. The flowering period of S. humilis is short and when not in flower it could be passed over at a casual glance for Plantago lanceolata. Native. Marshy lowland meadows of unimproved grassland and fen-meadow. Now confined to one locality in Dorset where it was once more common and a recently discovered site of five, species-rich meadows in Glamorganshire where there are thousands of plants; formerly in another site in Dorset and one site in Warwickshire. Europe from Portugal, central Spain, southern France, northern Italy, the north Balkan peninsula and southern Russia northwards to Denmark, southern Sweden, Karelia and central Russia; Caucasus. A member of the European Temperate element. 2. S. hispanica L. Black Salsify Perennial herb with a stout, vertical, black rootstock. Stems solitary or few, 25–150 cm, pale or bluish-green, sometimes flecked brownish, erect, hollow, smooth, glabrous or sparsely arachnoid-hairy, often branched, leafy. Leaves with lamina 7–40 × 0.3–6.0 cm, pale or bluish-green on upper surface with a pale midrib, paler beneath, gradually decreasing in size upwards, linear to ovate-elliptical or oblongoblanceolate, narrowed to a long-acuminate, tail-like apex, entire, sometimes undulate, long-attenuate at the base, the lower sometimes to a winged petiole, all glabrous. Capitula few to numerous, 40–60 mm in diameter, rounded at base, solitary at the end of long peduncles or branches, often with a group at the apex of the stem. Involucral bracts in several rows, 10–30(–40 in fruit) × 4–7 mm, medium yellowishgreen with a scarious margin, the outer ovate and acute at the very shortly hairy apex, the inner oblong-lanceolate and acute at the shortly hairy apex. Flowers 25–60 mm, all ligulate, the ligules bright medium yellow, sometimes purplish on outer face, shallowly 5-lobed at apex. Styles lemon yellow. Receptacle flat, pitted, without scales. Achenes 10–15(–20) mm, greenish-white to brownish, cylindrical, angled, at least the outer with prominent, peg-like tubercles on the ribs; pappus 10–15(–20) mm, dirty white, of several rows of plumose hairs. Flowers 6–7. 2n = 14. Introduced. Grown on a small scale as a vegetable and sometimes existing briefly as an escape on roadsides and tips. Native of southern Europe and western Asia. It was

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first cultivated as a vegetable in Italy in the early sixteenth century and came to England via Spain and France possibly as early as 1560. The roots are boiled, skinned and eaten and the young shoots and flower buds are cooked like asparagus. 28. Tragopogon L. Annual to perennial herbs with latex, without rhizomes and stolons. Stems usually solitary, leafy, simple or branched. Leaves alternate, narrow, entire and sheathing. Capitula solitary or few. Involucral bracts in 1 row. Flowers bisexual. Corolla ligulate, yellow, mauve, pink or purple with 5 lobes at apex. Anthers tailed, with elongate apical appendage, filaments smooth. Style with long, slender branches, with both shaft and branches with very short hairs. Receptacle flat, without scales. Achenes fusiform, beaked; pappus of plumose or plumose and simple hairs. About 110 species in Europe, the Mediterranean region and western Asia. Phillips, R. & Rix, M. (1993). Vegetables. London. [T. porrifolius.] 1. 1. Flowers yellow 4. 1. Flowers mauve, pink or purple 2. Flowers about half as long as involucral bracts 1(a). pratensis subsp. minor 2. Flowers nearly as long as to longer than involucral bracts 3. 3. Ligules pale yellow; beak about as long as body of achene 1(b). pratensis subsp. pratensis 3. Ligules golden yellow; beak usually shorter than body of 1(c). pratensis subsp. orientalis achene 3. hybridus 4. Marginal achenes with pappus of simple hairs 5. 4. Marginal achenes with pappus of some plumose hairs 5. Stems 30–125 cm; ligules about as long as involucral bracts; body of achene abruptly narrowed into the beak 2(a). porrifolius subsp. porrifolius 5. Stems 20–50 cm; ligules about half as long as the involucral bracts; body of achene gradually narrowed into 2(b). porrifolius subsp. australis the beak

1. T. pratensis L. Goat’s-beard Annual to perennial herb with a long, pale tap-root surmounted by the remains of old leaves. Stem 18–70 cm, pale green to slightly glaucous, sometimes tinted purplish at base, simple or branched above, leafy, glabrous. Leaves 8–35 × 0.8–1.7 cm, becoming gradually smaller upwards, glaucous-green with a pale midrib on upper surface, only slightly paler beneath, erect, long linear, gradually tapered to a long, thin, acute apex, minutely serrulate, long-sheathing at the semiamplexicaul base, all glabrous. Capitula 1–5, solitary at the end of the main stem and branches; peduncles slightly swollen below the capitulum, glabrous. Involucral bracts in 1 row, 25–31 × 3.5–4.0 mm, elongating up to 40 × 9 mm in fruit, slightly glaucous-green, the outer with a purplish, pale or whitish margin, the inner with a dark margin, linear-lanceolate, long-attenuate to an acute apex, glabrous or with some woolly hairs at their base. Flowers from half as long as to exceeding the involucral bracts, all ligulate, the ligules pale to golden yellow, the outer sometimes with a treacle-coloured stripe on the

outer face, with 5 lobes at apex. Styles yellow. Receptacle flat, without scales. Achene body 9–25 mm, yellowish, fusiform, those of the centre usually smooth, those of the outer ones scaly muricate on the ribs and sometimes also tuberculate between the ribs, but sometimes all smooth or all more or less muricate, with a long slender beak usually about equalling the body; pappus 19–21 mm, whitish, of 1 row of plumose hairs, the hairs interwoven. Flowers 6–7. Visited by various insects and finally also self-pollinated. The capitula close about noon, hence one of its popular names, Jack-go-to-bed-at-noon. The following three subspecies have been called species but intermediates occur. (a) Subsp. minor (Mill.) Wahlenb. T. minor Mill. Involucral bracts with purplish margins. Flowers bright yellow, about half as long as the involucral bracts. Anthers brownish above. Achenes of outer flowers scaly-muricate on the ridges, tuberculate between, beak about as long as body. 2n = 12. (b) Subsp. pratensis Involucral bracts with pale margins. Flowers pale yellow, almost or quite equalling the involucral bracts. Anthers yellow below and dark violet at tips. Achenes of outer flowers smooth or slightly scaly-muricate, beak about as long as body. 2n = 12. ˇ (c) Subsp. orientalis (L.) Celak. T. orientalis L. Involucral bracts with white margins. Flowers golden yellow, as long as or exceeding the involucral bracts. Achenes of outer flowers with cartilaginous scales on ribs, beak shorter than body. 2n = 12. Native. Grassy, open, rough and cultivated ground, dunes and waysides. Common in much of Great Britain north to central Scotland, scattered records in coastal localities elsewhere in Scotland, in Ireland and in Alderney. Most of Europe northwards to 64◦ N; Caucasus; Armenia, Iran; central Asia. A member of the Eurosiberian Temperate element; widely naturalised outside its native range. The common native plant is subsp. minor with a westerly distribution in Continental Europe. Subsp. pratensis is recorded from scattered localities in central and southern Britain and may be introduced. It also occurs in western and central Europe. Subsp. orientalis is a rare casual which is native of centraleast Europe and western and central Asia. 2. T. porrifolius L. Salsify Annual or biennial herb with a white, branched, irregularly cylindrical tap-root. Stems 20–125 cm, glaucous, some areas very pale, sometimes suffused brownish-purple, hollow, erect, striate, glabrous, with long branches in upper half, very leafy. Leaves all cauline, numerous, the lamina 10–40 × 1–2 cm, glaucous with a pale midrib on upper surface, hardly paler beneath, long-linear, gradually narrowed to an acute apex, entire, widened below to a sheathing base, channelled above, with a very prominent midrib beneath, glabrous, ascending-erect. Capitula solitary at the ends of stem or branches, 40–60 mm in diameter; peduncles

29. Aetheorhiza conspicuously swollen just below the capitulum, glabrous. Involucral bracts usually 8 in 1 row, 35–60 × 2–5 mm, yellowish-green, linear-lanceolate, acute at apex, glabrous. Flowers from half as long to as long as the involucre, all ligulate, the ligules purple, minutely 5-lobed at apex. Styles purple. Receptacle flat, without scales. Achene body 3–12 mm, pale brown, narrowly fusiform, 10-ribbed with the ribs muricate-tuberculate, abruptly narrowed into the beak; beak up to 27 mm; pappus shorter than body, whitish or pale brown, of 2 series of rigid, plumose hairs, the outer series thicker than the inner. Flowers 6–8. Visited by various insects. (a) Subsp. porrifolius T. porrifolius subsp. sativus (Gaterau) Br.-Bl.; T. eriosperma Ten. Stems 30–125 cm. Ligules about as long as the involucral bracts, lilac to reddish-purple. Achenes with the body abruptly narrowed into the beak. 2n = 12. (b) Subsp. australis (Jord.) Nyman T. australis Jord.; T. coelesyriacus Boiss.; T. longirostris Bisch. ex Sch. Bip.; T. porrifolius subsp. longirostris (Bisch. ex Sch. Bip.) Holmboe; T. sinuatus Av´e-Lall. Stems 20–50 cm. Ligules about half as long as involucral bracts. Achenes with the body gradually narrowed into the beak. 2n = 12. Introduced. Casual or rarely naturalised in waste or rough ground and waysides. Scattered records in Great Britain and Ireland, mainly in southern England and the Channel Islands. Both subspecies occur, but it is not known how they are distributed. Subsp. porrifolius is perhaps native in the central and east Mediterranean region. It is widely cultivated for its roots which are boiled or baked or used in a soup. Its tender young leaves are sometimes used as a salad and its young shoots and flower buds can be cooked like asparagus. It is sometimes called Oyster-plant because its roots are said to taste like oysters. Subsp. australis is widespread in the Mediterranean region and the Atlantic Islands. × pratensis = T. × mirabilis Rouy This hybrid has the ligules with a yellow base but suffused with purple distally to a variable extent. It has a low level of fertility. 2n = 12. Introduced. Roadsides and waste places. A few records in central and south England. Also in western Europe. 3. T. hybridus L. Slender Salsify Geropogon glaber L.; Geropogon hirsutus L.; T. hirsutus (L.) Kotschy; T. glaber (L.) Hoffm.; Geropogon hybridus (L.) Sch. Bip.; Tragopogon geropogon Rouy Annual herb with a tap-root. Stems 10–50(–80) cm, pale green, channelled, glabrous, leafy, branched or unbranched. Leaves numerous, sometimes crowded towards the base, the lamina up to 18 × 0.8 cm, greyish-green on upper surface, paler beneath, long-linear, acuminate at apex, entire, sheathing at base, all glabrous. Capitula solitary, terminal; peduncle glabrous, often swollen at apex. Involucral bracts in 1 row, 7–8, 28–32 × 2.5–3.0 mm, linear-subulate, finely acuminate at apex, glabrous. Flowers 12–14 mm, about half as long as involucral bracts, all ligulate, pale mauve

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or pink, shallowly 5-lobed at apex. Receptacle flat, with filiform scales up to 20 mm. Achene body 20–25 × about 1.5 mm, pale brown, very narrowly fusiform or subcylindrical, of 2 kinds; outer almost smooth, gradually tapering into a scabridulous beak about 25 mm, the pappus consisting of 5, unequal, scabridulous, erecto-patent awns, the shortest 2.0–2.5 mm, the longer up to 9 mm, with minute atrophied awns sometimes present between the developed ones; inner obscurely ribbed with 10 minutely scabridulous ribs with the beak a little shorter than those of the outer, the pappus consisting of about 20, thinly plumose, unequal bristles, the stouter about 15–18 mm, the more slender about 12 mm. Flowers 6–8. 2n = 14. Introduced. A bird-seed, spice and grain casual in gardens, parks, tips and waste ground. Scattered records in southern Great Britain. Native of the Mediterranean region and western Asia. 29. Aetheorhiza Cass. Perennial herbs with latex and a long, thin rhizome bearing tubers and leafy stolons. Stems usually several, simple or forked. Leaves usually all basal, but sometimes low on stem, entire or dentate. Capitula usually solitary, rarely several. Involucral bracts in several rows. Flowers bisexual. Corolla ligulate, yellow, sometimes with a reddish-purple or greenish stripe on the outer face with 5 lobes at apex. Anthers tailed, with elongate apical appendage; filaments smooth. Style with long, slender branches, both shaft and branches with long hairs. Receptacle flat, pitted, without scales. Achenes narrowly fusiform, not flattened, attenuate above, swollen at base, with 4 deep grooves, not beaked; pappus of several rows of pure white, simple hairs. A single species in the Mediterranean region and coastal areas of south-west Europe. 1. A. bulbosa (L.) Cass. Tuberous Hawk’s-beard Leontodon bulbosus L.; Crepis bulbosus (L.) Tausch; Intybus bulbosus (L.) Fr.; Prenanthes bulbosus (L.) DC. Perennial herb with long, thin rhizomes bearing large, whitish, subglobose tubers and leafy stolons. Stems usually several, 7–55 cm, simple or forked, erect, slender, striate, glabrous. Leaves glaucous, basal numerous, the lamina 1– 25 × 0.4–3.5 cm, elliptical to obovate, acute or obtuse at apex, entire to sinuate-dentate, gradually attenuate at base, the petiole up to 50 mm and winged; cauline usually absent, but sometimes 1–2 near the base of the stem and similar to the basal; all usually glabrous, rarely with a few hairs. Capitula 1(–8), 20–25 mm in diameter; peduncles up to 11 cm, slender, with unequal, black, clavate glandular hairs, few below but often condensed just under the capitulum. Involucral bracts in several rows, but often weakly 2-rowed, (8–) 13–15 × 1.8–2.1 mm, blackish-green, with a pale margin, lanceolate to oblong, abruptly narrowed to the obtuse apex, with black, clavate glandular hairs at the base and minute white hairs at the apex. Flowers all ligulate, 17–20 mm; ligules yellow, sometimes with a reddish-purple or greenish stripe on the outer face, with 5 lobes at apex. Receptacle flat, pitted, without scales. Achenes 3.0–4.5 mm, pale brown, narrowly fusiform, more or less attenuate at apex and with a hollow, swollen base, with 4 deep grooves; pappus

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7–8 mm, white, of several rows of pure white simple hairs. Flowers 6–8. 2n = 32. Introduced. Long known as an irreducible weed in a few gardens. Very local in eastern Ireland and has occurred as a casual elsewhere. Native of the Mediterranean region and the Atlantic coast of Portugal, Spain and France. Our plant is subsp. bulbosa, which occurs throughout the range of the species except the Aegean region. 30. Sonchus L. Annual to perennial herbs with latex, sometimes with rhizomes. Stems usually solitary, leafy, usually branched above. Leaves alternate, denticulate to pinnately divided, the cauline more or less amplexicaul, often more or less prickly. Capitula few to very numerous. Involucral bracts in several rows. Flowers bisexual. Corolla ligulate; the ligules yellow with 5 lobes at apex. Anthers tailed, with elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches with very short hairs. Receptacle flat, pitted, without scales. Achenes usually elliptical or oblanceolate, more or less compressed, not beaked; pappus of 2 rows of 2 kinds of hairs, deciduous, rough solitary hairs and more or less persistent, softer hairs in fascicles. About 60 species throughout the Old World but especially in the Mediterranean region and Africa. Barber, H. N. (1941). Spontaneous hybrids between Sonchus asper and S. oleraceous. Ann. Bot. new ser. 5: 375–377. Lewin, R. A. (1948). Sonchus L. in Biological flora of the British Isles. Jour. Ecol. 36: 203–223. Lousley, J. E. (1968). A glabrous perennial Sonchus in Britain. Proc. B.S.B.I. 7: 151–157. Pegtel, D. M. (?1976). On the ecology of two varieties of Sonchus arvensis L. Groningen. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough [S. palustris.] 1. Perennial herbs with thick, more or less erect 2. underground portion or rhizomatous 1. Annual or biennial herb with a main tap-root and fibrous 6. lateral roots 2. Stems arising from a short, more or less erect, tuberous stock; cauline leaves with pointed auricles; achenes with 1. palustris cream ribs and pale brown between them 2. Plant with a strong rhizome; cauline leaves with rounded 3. auricles; achenes dark brown 3. Upper parts of peduncles and involucral bracts without 2(b). arvensis subsp. uliginosus glandular hairs 3. Upper parts of peduncles and involucral bracts with 4. glandular hairs 4. Leaves broadly oblong, entire, denticulate or with very shallow lobes 2(a,iii). arvensis subsp. arvensis var. integrifolius 4. Leaves broadly oblong and pinnately lobed, if entire or 5. shallowly lobed then linear or narrowly oblong 5. Leaves linear or narrowly oblong in outline, subentire or the lobes including the terminal narrow, the laterals up to 2(a,i). arvensis subsp. arvensis var. maritimus 6 pairs 5. Leaves elliptical or oblanceolate in outline, the lobes short and broad, the terminal wide, the laterals up to 4 pairs 2(a,ii). arvensis subsp. arvensis var. arvensis

6. Auricles of cauline leaves pointed; achenes transversely 7. rugose between the ribs 6. Auricles of cauline leaves rounded; achenes smooth 8. between the ribs 7. Stem 10–50 cm; leaves often tinted reddish-purple; capitula few in a dense cluster 3(i). oleraceus var. litoralis 7. Stem up to 150 cm; leaves occasionally tinted reddish-purple; capitula usually numerous in an open 3(ii). oleraceus var. oleraceus inflorescence 8. Leaves coriaceous and very spiny; achenes with dense, minute, recurved spicules on the ribs 4(b). asper subsp. glaucescens 8. Leaves only sometimes coriaceous; achenes with few or 9. no spicules on the ribs 9. Leaves simple, bidentate, often panduriform 4(a,iii). asper subsp. asper var. integrifolius 10. 9. Leaves much pinnately divided 10. Stems up to 50 cm; capitula few in a dense cluster 4(a,i). asper subsp. asper var. sabulosus 10. Stems up to 120(–200) cm; capitula numerous in an 4(a,ii). asper subsp. asper var. asper open inflorescence

1. S. palustris L. Marsh Sow-thistle Perennial herb with a short, more or less erect, tuberous stock. Stem 90–300 cm, pale yellowish-green, sometimes striated or tinted purplish-brown, angled and channelled, hollow, with the large central cavity which is square in cross section, erect, glabrous except for the inflorescence, leafy, branched in the upper sixth to one-quarter, with a sticky, yellowish, foetid latex. Leaves numerous, the lower usually dead at the time of flowering, the lamina 4–28 × 0.5–9.0 cm, yellowish-green with a pale yellow midrib on upper surface, bluish-green beneath with a prominent pale yellowish midrib; lower and median with lamina broadly ovate or ovate-oblong in outline, acute at apex, pinnately divided, the terminal segment long, narrowly linear-lanceolate, acute at apex and with its whole margin with minute denticulations, the 1–5 lateral segments linear or linear-lanceolate, acute at apex, entire or with an occasional large denticulation and patent or recurved; the inter segment area broadly winged and with irregular, pointed auricles at the sessile, semiamplexicaul base; upper leaves linear-lanceolate, with a long-drawn-out, acute apex, the margin minutely denticulate, with acute auricles at the semiamplexicaul base; all leaves glabrous. Capitula numerous, 10–40 mm in diameter, in corymbs at the end of branches, the whole forming a large corymbose panicle; branches and peduncles with dark and yellowish, unequal glandular hairs which become more dense upwards and with some arachnoid hairs. Involucral bracts in several rows, 9–15 × 1–2 mm, yellowish-green to medium green, the inner with paler margins, linearlanceolate, acute at apex, with dense dark and yellowish, unequal glandular hairs. Flowers 10–14 mm, all ligulate, pale yellow, the inner darker than the outer, with 5 short lobes at apex. Receptacle flat, pitted, without scales. Achenes 3.5–4.0 mm, pale creamy-brown, ribbed, the 6–10 ribs minutely muricate and cream, the 4 strongest making the slightly compressed body more or less tetragonal, the intercostal area pale brown; pappus 5.0–6.5 mm, creamy-white, of 2 rows of simple eglandular hairs. Flowers 7–9. 2n = 18.

30. Sonchus Native. Marshes, fens and streamsides, characteristically in Phragmites australis. Local in coastal areas of southeast England from Kent to Norfolk, formerly north to Lincolnshire and west to Leicestershire and Oxfordshire. A population in Hampshire was not discovered until 1959, but appears to be native. Along the River Thames it has declined, but holds its own in Broadland and eastern Suffolk. In the Cambridgeshire Fenland it became extinct due to drainage, but in the last 30 years it has spread from stock introduced into Woodwalton Fen, Huntingdonshire. Central Europe from Spain, Corsica, Balkans and southern Russia northwards to the Netherlands, Denmark, southern Sweden and central Russia; Caucasus; northeastern Anatolia. A member of the Eurosiberian Temperate element. 2. S. arvensis L. Perennial Sow-thistle Perennial herb with far-creeping rhizomes, bearing adventitious buds. Stems 30–150 cm, pale yellowish-green, erect or ascending, furrowed, hollow with the small central cavity elliptical in cross-section, glabrous or glandular-hairy above, leafy, branched in upper half. Leaves 5–40 × 0.5– 8.0 cm, dull medium to dark green with a pale midrib on upper surface, paler and slightly glaucous beneath; basal with lamina oblong, linear, elliptical or oblanceolate in outline, more or less acute at apex, runcinate-pinnatifid, the terminal lobe broadly or narrowly triangular-ovate and irregularly undulate-spinulose, the lateral lobes irregular, broadly or narrowly triangular-ovate, more or less acute at apex and undulate-dentate-spinulose, area between the lobes broad and entire to spinulose-denticulate or spinulosedentate, narrowed to a broadly winged petiole; cauline gradually decreasing in size, the lower and median similar to basal but sessile and amplexicaul with rounded, spinulose-dentate, appressed auricles, the upper with lamina lanceolate to linear-lanceolate, gradually narrowed to an acute apex, nearly entire to spinulose-dentate, sessile and amplexicaul with rounded, spinulose-dentate auricles; all glabrous. Capitula 30–50 mm in diameter, campanulate, in a loose corymb; peduncles long or short, yellowishgreen, glabrous or with yellowish or dark glandular hairs. Involucral bracts in several rows, 10–20 × 1.5–2.5 mm, yellowish-green with a narrow, scarious margin, linearlanceolate, obtuse at apex, glabrous or with yellowish or blackish glandular hairs. Flowers 15–20 mm, all ligulate, the ligules golden yellow, shallowly 5-lobed at apex. Receptacle slightly convex, pitted, the pit margins finely toothed, without scales. Achenes 2.5–3.5 × 1.0–1.5 mm, yellow turning dark brown, elliptical, compressed, with 4–5 ribs on each face, rugulose between the ribs; pappus 10–14 mm, white, of 2 equal rows of simple eglandular hairs. Flowers 7–10. Visited by many kinds of insects, especially bees. There seems to be no clear correlation between chromosome number and subspecific differentiation, but usually they are different. (a) Subsp. arvensis S. arvensis var. glandulosus Coss. & Germ.; S. vulgaris Rouy nom. illegit.

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Leaves darker green. Upper parts of peduncles and involucre and sometimes branches and upper stem with glandular hairs. Involucral bracts 14–20 mm. 2n = 54. (i) Var. maritimus G. F. W. Meyer S. vulgaris var. marinus Rouy; S. arvensis var. angustifolius auct. Leaves linear or narrowly oblong in outline, the lobes subentire or including the terminal narrow, the laterals up to 6 pairs. (ii) Var. arvensis Leaves elliptical or oblanceolate in outline, the lobes short and broad, the terminal wide, the lateral up to 4 pairs. (iii) Var. integrifolius Bisch. Plant up to 1 m. Leaves fleshy, narrowly to broadly oblong, entire, denticulate or with faint signs of broad lobes. (b) Subsp. uliginosus (M. Bieb.) Nyman S. uliginosus M. Bieb.; S. arvensis forma uliginosus (M. Bieb.) Javorka; S. arvensis subsp. glabrescens (Wimm. & Grab.) Guenth.; S. arvensis var. laevipes W. D. J. Koch Leaves paler green. Upper parts of peduncles and involucre glabrous. Involucral bracts 10–15 mm. 2n = 36. Native. Formerly an abundant and pernicious weed of arable land where it is now almost eradicated. Common in waste land, waysides, dunes and shingle by the sea, ditches and river banks. Throughout the lowlands of Great Britain and Ireland. Throughout Europe to 70◦ 33 N in Scandinavia, western Asia; widely introduced in Asia, the Americas and Africa. A member of the Eurosiberian Temperate element, but now through naturalisation with a Circumpolar Temperate distribution. Subsp. arvensis is the common subspecies, the distribution and ecology of subsp. uliginosus is unknown. Var. maritimus is the plant of coastal dunes and shingle. It occurs elsewhere on the coast of western Europe. Var. arvensis is the plant of cultivated and waste places. Var. integrifolius is known only from the banks of the River Alde in Suffolk among Phragmites with Rumex crispus subsp. uliginosus and Sonchus palustris. 3. S. oleraceus L. Smooth Sow-thistle S. piquetii Druce Annual or overwintering herb with a long, slender taproot and numerous, pale brown, fibrous roots. Stem 10– 150 cm, more or less 5-angled, pale yellowish-green, often tinted brownish-purple, sometimes deep purplish flushed, erect, stout, hollow except at the nodes, glabrous, usually branched. Leaves very variable, dull medium green to slightly bluish and sometimes flushed reddish-purple on upper surface, paler and rather glaucous beneath; basal few to numerous, the lamina 8–30 × 2–13 cm, oblanceolate to obovate in outline, obtuse or rounded at apex, undivided to runcinate-pinnatifid, the terminal lobe usually wider than the uppermost laterals, more or less ovate and repandspinulose-dentate or sometimes trilobed, the laterals of 1–3 pairs, more or less ovate, recurved, more or less acute at apex and repand-dentate, the interlobe area winged, with winged petioles up to 10 cm; lower cauline similar, but semiamplexicaul and with rounded, sharply denticulate,

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acute auricles; upper cauline gradually becoming smaller, similar to lower but sessile with the lower lateral lobes merging into the auricles; all glabrous or slightly arachnoid-hairy when young. Capitula few to numerous, 15–35 mm in diameter, in an irregular cymose umbel; peduncles up to 80 mm, yellowish-green, glabrous or rarely with glandular hairs, rarely white tomentose at apex just under the capitulum. Involucral bracts in several rows, 10–15 × 1.5–2.0 mm, dull yellowish-green, linear-lanceolate, obtuse and with short, tufted hairs at apex, arachnoid-hairy in bud, then glabrous except for apex, or rarely glandular-hairy. Flowers all ligulate, 12–17 mm, the ligules yellow, the outer sometimes much paler or white and sometimes purple-tinged or lilac below, very shallowly 5-lobed at apex. Styles dark. Receptacle flat, pitted, without scales. Achenes 2.5–3.7 mm, yellow turning brown, oblanceolate, compressed, never winged, longitudinally 3-ribbed on each face and transversely rugose between ribs; pappus 5–8 mm, white, of 2 rows of simple eglandular hairs, the outer thickened at the base. Flowers 5–9. Visited by bees, hoverflies and other insects. 2n = 32. Variable in size, division of leaf, colour of ligules and hairiness of peduncles and involucral bracts. Believed to be an allopolyploid derived from S. asper (2n = 18) and S. tenerrimus L. (2n = 14). Some plants sometimes have the white tomentum at the apex of the peduncle as in S. tenerrimus, but not the other characters. The two following varieties seem to be distinct ecotypes, the remaining variation seems to show no geographical or ecological preference. (i) Var. litoralis P. D. Sell Stems 10–50 cm. Leaves often tinted reddish-purple. Capitula few, in a dense cluster. (ii) Var. oleraceus S. oleraceus var. albescens Neuman; S. oleraceus var. glandulosus auct.; S. oleraceus var. integrifolius Wallr.; S. oleraceus var. triangularis Wallr.; S. oleraceus var. lacerus Wallr.; S. oleraceus var. purpurascens Druce Stem up to 150 cm. Leaves only occasionally tinted reddishpurple. Capitula usually numerous and the inflorescence more open. Native. Coastal shingle, dunes and cliffs, and waste and cultivated land and waysides inland. Abundant throughout the lowland areas of Great Britain and Ireland. Throughout Europe to 66◦ 13 N in Scandinavia; western Asia; North Africa and the Canary Islands; widely introduced elsewhere as a weed of cultivation. A member of the European Southern-temperate element, but now with a Circumpolar Southern-temperate distribution. Var. litoralis is the plant of the coast. Var. oleraceus is the inland weed. 4. S. asper (L.) Hill Prickly Sow-thistle S. oleraceus var. asper L.; S. oleraceus subsp. asper (L.) Hook. fil. Annual or overwintering herb with a slender tap-root and fibrous side-roots. Stems 10–120(–200) cm, pale, shiny yellowish-green, often tinted reddish-purple, sometimes heavily suffused reddish-purple, thick, hollow, glabrous, leafy, usually branched above. Leaves numerous, sometimes with a basal rosette, getting gradually smaller upwards or the median largest, the lamina 4–21 × 2–9 cm, shining

yellowish- to dark green on upper surface, paler and glaucous beneath, sometimes tinted purplish, oblanceolate, obovate or elliptical in outline, entire to pinnately divided, the terminal segment triangular-ovate and large in the lower leaves and smaller in the upper, the lateral segments linear to ovate, gradually decreasing in size downwards, acute at apex, all segments undulate and crisply spinose-dentate, the lower with a spinose-winged petiole, the upper sessile, more or less amplexicaul and with spinose, rounded auricles; all glabrous. Capitula few to numerous, 15–25 mm in diameter, in corymbose cymes at the ends of branches; peduncles pale green, curved, glabrous or with a cluster of clavate glandular hairs just below the capitulum. Involucral bracts in several rows, 14–17 × 1.6–1.8 mm, dull olive green, linearlanceolate, obtuse at the very shortly hairy apex and with scattered spicules down the midline, otherwise glabrous. Flowers 15–16 mm, all ligulate, the ligules golden yellow, the outer with a red stripe on the outer face, shallowly 5lobed at apex. Receptacle slightly concave, pitted, with hairs or scales. Achenes 2–3 mm, pale brown, oblanceolate, compressed, longitudinally 3-ribbed on each face and smooth between them, the margin and ribs sometimes with minute, recurved spinules; pappus 10–12 mm, white, of 2 rows of simple eglandular hairs, the outer thickened at the base. Flowers 6–9. Visited by bees and hoverflies. There is great variation in the leaves from being entire, flat and denticulate to deeply pinnately divided, 3-dimensional and spinose. Some plants have a cluster of glandular hairs just beneath the capitulum, but most are glabrous. The only certain way of distinguishing S. asper from S. oleraceus is by the rugosity of its achenes. When the leaves are dark shining green and prickly it is usually safe to identify the plant as S. asper, but they can be much less prickly and pale, though still shiny green. S. oleraceus has dull, pale green leaves. (a) Subsp. asper Annual herb. Leaves mostly cauline, not coriaceous. Achenes with sparse spicules on margins and ribs. Pollen grains 35–42 µm 2n = 18. (i) Var. sabulosus P. D. Sell Stem up to 50 cm. Leaves usually much divided and very prickly. Capitula few in a dense cluster. (ii) Var. asper Stems up to 120(–200) cm. Leaves variable, but pinnately divided. Capitula numerous in an open inflorescence. (iii) Var. integrifolius Lej. S. asper var. laevis Wallr.; S. asper var. inermis Bisch. Stems usually above 100 cm. Leaves simple but bidentate, often panduriform. Capitula numerous in an open inflorescence. (b) Subsp. glaucescens (Jord.) Ball S. glaucescens Jord.; S. nymanii Tineo & Guss.; S. asper subsp. nymanii (Tineo & Guss.) Hegi Biennial herb. Leaves often forming a rosette, coriaceous, glaucous beneath and very spiny. Capitula forming an irregular umbel at the end of stems and branches. Achenes with dense recurved, minute spicules on the ribs and margins.

31. Lactuca Native. Shingle, sand and cliffs by the sea and waste and cultivated land and waysides inland. Abundant throughout the lowlands of Great Britain and Ireland. Europe northwards to 66◦ N in Scandinavia; western Asia; North Africa; and widely introduced elsewhere as a weed of cultivated land. A member of the European Southern-temperate element; now with a Circumpolar Southern-temperate distribution. Our common plant is subsp. asper var. asper, while var. sabulosus is the plant of shingle and sand on the coast. Var. integrifolius is a strikingly distinct plant, but has only been recorded in a few places. Nothing is known of its ecology. It occurs widely in Continental Europe. Subsp. glaucescens occurs and has been seen in quantity at Histon, Cambridgeshire, but its general habitat, distribution and status are unknown. It is a plant of south, west and central Europe. × oleraceus This hybrid, which is difficult to recognise, has the auricles of the upper leaves rounded and dentate as in S. asper, with a single elongate tooth directed downwards or backwards. The achenes are small, white and abortive with wings and lacking transverse rugosity. 2n = 25. Apparently rare, being recorded only for Cambridgeshire and Shropshire. 31. Lactuca L. Annual to perennial herbs with latex, sometimes with rhizomes. Stem usually solitary, leafy, branched. Leaves alternate, entire to pinnately divided, often prickly. Capitula few to numerous, not usually open after 11 a.m. Involucral bracts in several rows. Flowers bisexual. Corolla ligulate, yellow or bluish, with 5 lobes at apex. Anthers tailed, with elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches hairy. Receptacle flat, without scales. Achenes elliptical or obovate, compressed, beaked; pappus of 2 rows of simple hairs. About 95 species, more or less cosmopolitan but chiefly in drier temperate and subtropical regions. L. viminea (L.) J. & C. Presl has been recorded as a rare casual. Pre Second World War records of L. serriola and L. virosa are unreliable unless supported by herbarium specimens, which need careful checking (cf. Oswald, 2000). Carter, R. N. & Prince, S. D. (1982). A history of the taxonomic treatment of unlobed-leaved prickly lettuce, Lactuca serriola L., in Britain. Watsonia 14: 59–62. Fer´akova, V. (1977). The Genus Lactuca L. in Europe. Univerzita Komensk´eho. Leick, E. & Steubring, L. (1957). Lactuca tatarica (L.) C. A. Meyer als Wanderpflanze und Insel-Endemit. Feddes Rep. 59: 179–189. Oswald, P. H. (2000). Historical records of Lactuca serriola L. and L. virosa L. in Britain, with special reference to Cambridgeshire (v.c. 29). Watsonia 23: 149–159. Phillips, R. & Rix, M. (1993). Vegetables. London. Prince, S. D. & Carter, R. N. (1977). Prickly Lettuce (Lactuca serriola L.) in Britain. Watsonia 11: 331–338. Prince, S. D. & Hare, A. D. R. (1981). Lactuca saligna and Pulicaria vulgaris in Britain in Synge, H. (Edit.) The biological aspects of rare plant conservation, pp. 379–388. Chichester. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [L. saligna.]

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1. Perennial with rhizomes; ligules blue; achenes with a beak less than half as long as the body and the same 5. tatarica colour 1. Annual to biennial without rhizomes; ligules yellow; achenes with a beak more than half as long as the body 2. and much paler in colour 2. Midrib on lower side of leaves with numerous, strong, 3. rigid hairs 2. Midrib on lower side of leaves glabrous or with a very 6. occasional rigid hair 3. Stems and midribs of leaves pale cream, rarely with a touch of red; latex not strong-smelling; diameter of open capitula 8–10 mm; body of ripe achenes (2.8–)3–4(–4.2) 4. mm, olive-grey 3. Stems and midribs of leaves usually suffused brownish-purple; latex smelling strongly of opium; diameter of open capitula (14–)17–20 mm; body of ripe 5. achenes (4–)4.2–4.8(–5.2) mm, purplish to blackish 4. Cauline leaves runcinate-pinnatifid and spinulose-toothed 1(1). serriola forma serriola 4. Cauline leaves unlobed but spinulose toothed to undulate 1(2). serriola forma integrifolia 5. Cauline leaves unlobed, irregularly sinuate-dentate 3(1). virosa forma virosa 5. At least the upper cauline leaves pinnatisect with wide 3(2). virosa forma lactucarii lobes 6. Middle and upper cauline leaves well separated, linear with a sagittate-amplexicaul base; inflorescence very 4. saligna narrow 6. Cauline leaves dense, broadly elliptical, ovate or 7. subrotund, amplexicaul; inflorescence broad 7. Basal leaves crisped and toothed forming a loose, open 2(c). sativa subsp. crispa centre 8. 7. Basal leaves not crisped and toothed 8. Basal leaves broadly rounded, ovate, entire and spreading, forming a flat rosette with a globular centre 2(a). sativa subsp. sativa 9. 8. Basal leaves long and erect 9. Basal leaves forming a tight subcylindrical centre 2(b). sativa subsp. romana 9. Basal leaves loose with a stout succulent stem 2(d). sativa subsp. asparagina

Section 1. Lactuca Inflorescence a dense panicle of many capitula. Achenes elliptical to obovate, with a pale beak at least as long as the body. 1. L. serriola L. Prickly Lettuce L. scariola L.; L. officinarum Crantz; L. silvestris Garsult; L. sylvestris Lam.; L. schimperi Jord. Overwintering or spring-germinating annual herb or rarely biennial, with latex but smelling of garden lettuce when bruised, with a pale, slender tap-root and fibrous sideroots. Stems (15–)30–210 cm, pale cream or whitish, sometimes faintly reddish-tinged, stiffly erect, smooth, glabrous or with numerous, yellowish, very rigid, unequal, patent, swollen-based simple eglandular hairs in the lower part, leafy, usually branched above and sometimes with leafy

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branches in the axils of the lower cauline leaves. Leaves rigid, medium or sometimes slightly bluish-green on upper surface, slightly paler beneath, with a very pale midrib; basal overwintering in a rosette, the lamina oblong-obovate, rounded at apex, irregularly spinulose-toothed or pinnatifid, the cauline gradually decreasing in size, 4–15 × 1–4 cm, obovate-oblong, oblong, oblong-lanceolate or narrowly elliptical, unlobed and spinulose-toothed to sinuate-lobed or runcinate-pinnatifid with an acuminate terminal lobe and a few distant pairs of narrowish, acute lateral lobes which are back-curved distally, the upper less lobed and the uppermost hastate or sagittate; when fully exposed to the sun all are held vertically in a north–south plane; all with short to medium, rigid, pale simple eglandular hairs on the margin and the midrib. Capitula very numerous, 8–10 mm in diameter, narrowly ovoid, of (7–)10–30(–50) flowers, closely spaced along the distal halves of the ascending panicle branches, in an elongated, irregular, often flat-topped, pyramidal panicle, with the lower branches leafy; peduncles pale, tinted pale brownish, glabrous; bracts small, ovate, obtuse at apex, sagittate, with rounded, spreading auricles. Involucral bracts in several rows, 4–14 × 1.2–1.5 mm, yellowish-green with reddish tips and sometimes suffused reddish, linear-lanceolate, obtuse at apex, glabrous except for a very short tuft of white hairs at the apex. Flowers 10–16 mm, all ligulate, closing before midday, the ligules pale yellow, often mauve-tinged, with 5 shallow lobes at apex. Receptacle flat, pitted, without scales. Achene body (2.8–)3–4(–4.2) mm, olive-grey and mottled, broadly elliptical, compressed, 5- to 7-ribbed on each face, narrowly bordered, with many very short, rigid simple eglandular bristles near the apex; beak white and equalling or longer than the length of the body; pappus 3–7 mm, of 2 equal rows of soft, white simple eglandular hairs. Flowers 7–9. Little visited by insects and automatically self-pollinated. 2n = 18. (1) Forma serriola L. scariola var. vulgaris Bisch.; L. scariola var. typica Rouy Cauline leaves lobed, runcinate-pinnatifid and spinulosedentate. (2) Forma integrifolia (Gray) S. D. Prince & R. N. Carter L. virosa var. integrifolia Gray; L. scariola var. integrata Gren. & Godr.; L. scariola var. integrifolia (Gray) Bogenhard; L. dubia Jord.; L. integrata (Gren. & Godr.) A. Nelson; L. serriola var. dubia (Jord.) Rouy Cauline leaves unlobed, but spinulose-dentate. Probably native. Waste and rough ground, field margins and waysides. Frequent in England south of line between the Rivers Humber and Severn and much increased in the last 50 years especially in the south-east and East Anglia; very scattered elsewhere in England, Wales, Ireland and the Channel Islands. South and central Europe, but introduced further north, west Asia and North Africa, introduced in North America, Argentina and South Africa and elsewhere. A member of the Eurosiberian Southern-temperate element. Forma integrifolia is our common plant and forma serriola occurs here and there but does not seem to show any ecological preference.

2. L. sativa L. Garden Lettuce Annual or biennial herb with a slender, pale tap-root. Stems 30–120 cm, pale whitish to yellowish- to glaucous-green, thick and fleshy with much latex when allowed to flower, erect, solid, ridged and slightly angled, glabrous, branched in upper half, very leafy. Leaves yellowish-green with a pale midrib on upper surface, paler beneath or suffused purple and soft or crisp; basal in a dense rosette, the lamina 5–20 × 4–20 cm, broadly to narrowly ovate, elliptical, obovate or oblong-obovate, rarely lanceolate, roundedobtuse at apex, undulate and entire to runcinate-pinnatifid or crispate-dentate, cuneate or rounded to a very short petiole; cauline numerous and dense, patent, gradually decreasing in size upwards, broadly elliptical to ovate or subrotund, rounded or obtuse at apex, entire or denticulate and usually undulate, or crispate-dentate, cuneate, rounded or amplexicaul at the sessile base, the auricles slightly decurrent on the stem; all glabrous and smooth. Capitula numerous (up to 400) and dense, 10–15 mm in diameter, of 7– 15(–35) flowers, in a corymbose panicle; peduncles and branches erect or erecto-patent, yellowish-green, glabrous, with numerous, small, broadly to narrowly ovate, obtuse, amplexicaul bracts. Involucral bracts in 3–4 rows, more or less appressed, 2.5–12.0 × 1.0–1.7 mm, pale yellowishgreen, the inner with a very narrow paler margin, lanceolate to ovate, obtuse at apex, glabrous but minutely granulate. Flowers 12–14 mm, all ligulate, the ligules medium yellow, sometimes violet-streaked and with 5 shallow lobes at apex. Receptacle flat, pitted, without scales. Achene body 3–4(–5) mm; brownish-grey, narrowly obovate, compressed, with 5–7 ribs on each face, often finely muricate at apex; beak white, half as long as to as long as body; pappus 3–4 mm, of 2 rows of white simple hairs. Flowers 7–9. Visited by flies, but also automatically self-pollinated. 2n = 18. There are many cultivars, most of which can be fitted into the following subspecies, which are based on the young rosettes. The type specimen is a flowering plant and it is not known which is the nominate subspecies. For this account the most common form of northern Europe is taken as the nominate subspecies. (a) Subsp. sativa Cabbage Lettuce L. sativa var. capitata L.; L. sativa subsp. capitata (L.) Sch¨ubl. & G. Martens Stem short when young. Basal leaves broadly rounded, ovate, entire and spreading, forming a flat rosette with a globular centre. (b) Subsp. romana Sch¨ubl. & G. Martens Cos Lettuce L. sativa var. longifolia Lam. Stem short when young. Basal leaves long and erect, forming an elongated, upright rosette with a subcylindrical centre. (c) Subsp. crispa (L.) Sch¨ubl. & G. Martens Curled Lettuce L. sativa var. crispa L. Stem short when young. Basal leaves very broad, crispatedentate and forming a loose, open centre.

31. Lactuca (d) Subsp. asparagina Janch. Celtuce L. sativa var. angustana Irish ex Bremer Stem stout and succulent when young, conspicuously marked with horizontal leaf scars. Basal leaves lanceolate, alternate and ascending, not forming a centre. Introduced. Many varieties have been long cultivated in gardens as salad plants. Some 6,300 hectares are grown commercially countrywide; Kent and the north-west are the most significant areas, but lettuces are also grown in the Thames Valley, Vale of Evesham and Bedfordshire. It occurs as an escape from cultivation on tips, waste land and abandoned arable land; also as a bird-seed casual in scattered localities in Great Britain, mainly in the south. Lettuces were grown by the Romans, but are thought to have been first cultivated by the Egyptians in about 4500 BC, although they may have been first produced for the edible oil of their seeds. The Romans probably introduced it to Great Britain, but it is not next mentioned until Gerard’s Herball of 1597, when he lists eight varieties. It probably arose by selection from L. serriola, and those early varieties were still bitter and had to be blanched. The bitterness is in the milky latex which still occurs in the stem of cultivated varieties when they bolt. The bitter latex is soporific and has often been used as a substitute for opium or laudanum. Soft Cabbage Lettuces were the older form of northern Europe. The Cos Lettuce was so-called because it was said to be common on the Greek island of Kos. The heavy, crisp Iceberg Lettuces have been introduced from America but have not become as popular here as they are there. Celtuce or Asparagus Lettuce was developed in China and introduced to France in 1885. As well as being eaten as a salad, like all the other lettuces, the swollen stem is peeled, thinly sliced and stir-fried. About 95 per cent of the lettuce is water, but it contains vitamin A. 3. L. virosa L. Great Lettuce L. ambigua Schrad.; Wiestia virosa (L.) Sch. Bip.; L. flavida Jord.; L. serratifolia Sennen Overwintering annual or biennial herb with a branched tap-root, fibrous side-roots and very sticky, opium-smelling latex. Stems 60–250 cm, pale yellowish-green, but usually suffused throughout with brownish-purple, erect, flexuous, with numerous, bulbous-based, rigid, yellow to orange simple eglandular hairs below, leafy in lower two-thirds, much branched. Leaves rigid, dull medium or slightly bluishgreen, often suffused brownish-purple round the margins and on the midrib of the upper surface, paler and more bluish beneath; basal and lower cauline with lamina of 15–23 × 5–8 cm, obovate or obovate-oblong in outline, rounded at apex, irregularly sinuate-dentate, the teeth ending in a small prickle, narrowed at base to a winged petiole; median and upper gradually decreasing in size, the lamina oblong or oblong-ovate in outline, irregularly undulatedentate, or pinnatisect with the lobes oblong or ovate, obtuse with a spinose tip and irregularly dentate with spinose teeth, sessile and cordate-amplexicaul with appressed auricles; all with rigid, prickly, yellowish simple eglandular hairs on the veins and sometimes the surface. Capitula very numerous, (14–)17–20 mm in diameter, of about 15 flowers, in

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an elongated, characteristically rhomboid, almost leafless panicle; peduncles deep blackish-purple, glabrous; bracts amplexicaul, with appressed, more or less rounded auricles. Involucral bracts in several rows, 4–16 × about 1.0 mm, pale green, much suffused and spotted dark brownishpurple, linear-lanceolate, obtuse at apex, glabrous or with minute hairs. Flowers 13–14 mm, all ligulate, closing before midday, the ligules pale greenish-yellow, 5-lobed at apex. Receptacle flat, pitted, without scales. Achene body (4–)4.2–4.8(–5.2) mm, blackish-purple or maroon when young, blackish when ripe, narrowly elliptical, compressed, with a narrow, wing-like border, rugose but not spiculate at apex, 5-ribbed on each face, glabrous; beak white and about equalling the length of the body; pappus 5–8 mm, of 2 equal rows of soft, white simple eglandular hairs. Flowers 6–9 (a fortnight earlier than L. serriola). 2n = 18. (1) Forma virosa Cauline leaves unlobed, irregularly undulate-dentate. (2) Forma lactucarii (Lamotte) P. D. Sell L. lactucarii Lamotte; L. scariola var. lactucarii (Lamotte) Rouy; L. scariola var. altissima Lecoq & Lamotte nom. nud. Cauline leaves pinnatisect with wide lobes. Probably native. Occasional on coastal dunes, marshes and cliff ledges and inland cliffs, recently becoming more frequent on disturbed roadsides, tracks, streamside banks, walls, gravel pits and quarries. Mainly south of a line between the Rivers Severn and Tees, with scattered records elsewhere in Great Britain. South and central Europe from Portugal to Turkey and northwards to Scotland, Belgium, Austria and Romania. A member of the Suboceanic Southern-temperate element. Forma virosa is the common form. Forma lactucarii is of scattered occurrence. 4. L. saligna L. Least Lettuce L. minima Gray nom. illegit.; L. angustifolia Gilib.; L. salicifolia Salisb.; L. caucasica K. Koch; L. cyanea K. Koch; L. adulteriana Gren. & Godr.; L. saligna var. runcinata Gren. & Godr.; L. saligna var. cracoviensis auct. Annual, rarely biennial herb with a fusiform tap-root and fibrous side-roots. Stem 30–100 cm, whitish, erect, glabrous or with bristly simple eglandular hairs in the lower part, with long, slender, ascending branches, the lowest often arising near the base of the stem, leafy. Leaves more or less glaucous with a conspicuous, broad white midrib on upper surface and paler beneath; basal withered at time of flowering, the lamina 5–15(–22) × 2–7 cm, oblong, acute at apex, sinuate-pinnatifid or runcinate-pinnatifid, with narrow, distant, acute, entire or denticulate lateral lobes and a long, slender, acute terminal lobe, narrowed below into a petiolate-like base; cauline with lamina linear-lanceolate, acute at apex, entire, with a sagittate-amplexicaul base or pinnatifid with a few, narrow, distant lobes, commonly held vertically and all more or less in one plane; all glabrous. Capitula 2–4 mm in diameter, narrowly cylindrical with 6– 15 flowers, borne singly or in small clusters in the axils of sagittate bracts on the long branches of the narrow, striate panicle; peduncles very short, glabrous. Involucral bracts in

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several rows, 5–15 × 1.0–1.3 mm, greenish, with a narrow, white margin, linear-lanceolate, obtuse at apex, glabrous. Flowers 10–12 mm, all ligulate, closing before midday, the ligules pale yellow, often reddish on the outer face of the outer, becoming deep blue when dry, with 5 lobes at apex. Receptacle flat, pitted, without scales. Achene body 5–8 mm, pale brown or greyish, elliptical, compressed, with 7– 8 ribs, with a narrow margin, finely muricate at apex; beak white, 1.5 to 3.0 times as long as the body; pappus 2.5–3.0 mm, of 2 equal rows of soft, white simple hairs. Flowers 7–9. 2n = 18. Native. Salt-marshes, shingle, waste places, and walls by the sea. Very local at Fobbing in Essex, and Rye Harbour, Sussex; formerly in southern England from Cornwall to Norfolk, sometimes on waste and grassy places inland. Much of Europe northwards to southern England, central Germany and south-central Russia; introduced in Australia. A member of the European Southern-temperate element. Section 2. Mulgedium (Cass.) C. B. Clarke Mulgedium Cass. Inflorescence with ascending branches and few capitula. Achenes with a short beak the same colour as the body. 5. L. tatarica (L.) C. A. Mey. Blue Lettuce Sonchus tataricus L.; Mulgedium tataricum (L.) DC.; Mulgedium salicifolium K. Koch; L. kochiana Beauverd; L. salicifolia (K. Koch) Grossh., non Salisb.; Agathysus tataricus (L.) D. Don; L. sonchifolia DC.; Crepis charbonnelii L´evl.; Cicerbita tatarica (L.) Beaverd ex Sosn.; Lagedium tataricum (L.) Soj´ak Perennial herb with a vertical stock and rhizomes. Stems 30–80(–100) cm, very pale green, erect, glabrous, branched above, leafy. Leaves bluish-green on upper surface with a pale midrib, paler beneath; basal with lamina 11–19 × 1.5–5.5 cm, elliptic-lanceolate, acute at apex, unlobed to runcinate-pinnatifid, with a long terminal lobe and 4–10, smaller, recurved lateral lobes, attenuate to a winged, petiole-like base, the median smaller, with fewer lobes or only denticulate and less attenuate at the sessile base, the upper much reduced, entire, sessile and semiamplexicaul; all glabrous. Capitula 10–12 mm in diameter, of 16–23 flowers, few to numerous in a more or less corymbose panicle; peduncles glabrous. Involucral bracts in 3–4 rows, 3.0– 16.5 × 1.5–2.0 mm, pale green, with a pale margin, often purple tinged, the outer ovate, the inner lanceolate to linearlanceolate, obtuse at apex, glabrous. Flowers 15–17 mm, all ligulate, the ligules blue, with 5 lobes at apex. Receptacle flat, pitted, without scales. Achene body 5.5–6.5 mm, yellowish, brownish or blackish, narrowly obovate, 5–7 ribbed, slightly compressed, with a stout, concolorous beak 1.0– 1.5 mm; pappus 8–9 mm, of numerous, whitish bristles. Flowers 7–8. 2n = 16, 18. Sometimes placed in a separate genus, Mulgedium Cass. Introduced. Naturalised in rough and waste ground, mostly by the sea. Scattered records in England, Wales, Isle of Man, Guernsey and Co. Galway since 1886. Native of Turkey, Balkans, southern Russia, Caucasus, Iran, Afghanistan, central Asia and north and west China.

32. Cicerbita Wallr. Perennial herbs with latex and often with rhizomes. Stems usually solitary, branched, leafy. Leaves alternate, pinnately lobed, the cauline semiamplexicaul. Capitula few to numerous. Involucral bracts in several rows. Flowers bisexual. Corolla ligulate, blue, lilac or violet, with 5 lobes at apex. Anthers tailed, with elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches hairy. Receptacle flat, without scales. Achenes linear-oblong to elliptical, compressed, ribbed, not beaked; pappus of 2 rows of simple hairs, the outer shorter than the inner. Differs from Sonchus only in its blue to violet flowers and unequal pappus hairs. Includes about 35 species, in the mountains of Europe, south-west and central Asia, China and 1 species in Libya. Akeroyd, A. J. [J. R.], Jury, S. L. & Rumsey, F. J. (1983). Cicerbita macrophylla (Willd.) Wallr. B.S.B.I. News 33: 14–15. Beauverd, G. (1910). Contribution a` l’´etude des Compos´ees le genre Cicerbita. Bull. Soc. Bot. Gen`eve ser. 2, 2: 99–145. Marren, P. R., Payne, A. F. & Randall, R. E. (1986). The past and present status of Cicerbita alpina (L.) Wallr. in Britain. Watsonia 16: 131–142. Sell, P. D. (1986). The genus Cicerbita Wallr. in the British Isles. Watsonia 16: 121–129. Wegm¨uller, S. (1994). Cicerbita plumieri (L.) Kirschl. eine cytologische arealkundliche und o¨ kologische Studie. Bot. Jahrb. 116: 189–219. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [C. alpina.] 1. Plant glabrous or with very occasional simple eglandular 3. plumieri hairs 1. Peduncles and/or upper part of stem with few to 2. numerous glandular or simple eglandular hairs 2. Stems and sometimes peduncles with rather rigid simple 4. bourgaei eglandular hairs, without glandular hairs 2. At least the peduncles and usually also the upper part of 3. the stem and involucral bracts with glandular hairs 3. Lower leaves glabrous, with a triangular terminal lobe and a few pairs of small lateral lobes; capitula in an elongated panicle; achenes linear or linear-oblong 1. alpina 3. Lower leaves hairy on the veins beneath, with a cordate terminal lobe and usually only a single pair of lateral lobes; panicles wider, more or less corymbose; achenes 2. macrophylla subsp. uralensis narrowly elliptical

1. C. alpina (L.) Wallr. Alpine Blue Sow-thistle Sonchus alpinus L.; Mulgedium alpinum (L.) Less.; Sonchus caeruleus Sm. nom. illegit.; Lactuca alpina (L.) A. Gray Perennial herb with a cylindrical rootstock and fibrous roots. Stem 50–250 cm, simple or branched, stout, hollow, erect, furrowed, with bristly hair below and usually with dense, reddish glandular hairs in the upper part. Leaves with lamina 8–25 × 2–12 cm, dark green on upper surface, glaucous beneath; lowest lyrate or runcinate-pinnatifid, with a large, broadly triangular, acuminate, sharply denticulate terminal lobe and a few pairs of much smaller, triangular,

32. Cicerbita acute, denticulate lateral ones, narrowed at the base into a broad, winged petiole; upper smaller and less divided, with a winged petiole widened into a cordate-amplexicaul base; uppermost with lamina more or less lanceolate; all glabrous. Capitula in an elongated panicle, 20–25 mm in diameter; peduncles with dense, reddish glandular hairs. Involucral bracts in several rows, 10–15 × 1.0–1.5 mm, purplish-green, linear, obtuse at apex, usually with numerous, reddish glandular hairs. Flowers 15–18 mm, all ligulate, the ligules pale blue to violet, linear, with 5 lobes at apex. Receptacle flat, naked. Achenes 4.5–5.0 mm, linear or linear-oblong, compressed, not beaked, with 5 strong ribs and several weaker ones; pappus about 7 mm, of 2 rows of simple hairs, the outer shorter. Flowers 7–9. Pollinated by bees and butterflies, but in most years does not appear to set good seed. 2n = 18. Native. Moist acidic mountain rock ledges in remote gullies and cliffs between 530 and 1,090 m. Extremely local in Glen Callater, Caenlochan, Glen Canness and Glen Clova in Angus and Lochnager in Aberdeenshire. Continental Europe from Fennoscandia southwards to the Pyrenees, northern Apennines and Bulgaria. It reaches over 2,100 m in Switzerland. A member of the European Boreal-montane element. 2. C. macrophylla (Willd.) Wallr. Common Blue Sow-thistle Sonchus canadensis Froel., non L.; Sonchus macrophyllus Willd.; Mulgedium macrophyllum (Willd.) DC.; Lactuca macrophylla (Willd.) A. Gray Perennial herb with pale brown, fibrous roots and long, whitish rhizomes which send up flowering stems at regular intervals. Stems 60–200 cm, pale green, rather slender, slightly angular, glabrous below with long, slender glandular hairs in the upper part. Leaves up to 12, all cauline with up to 3 sometimes close together near the base, becoming gradually smaller up the stem, rather dull yellowishgreen; lowest with lamina 6–20(–40) × 5–17(–21) cm, ovate, rounded-obtuse to subacute at apex, sinuateglandular-denticulate, more or less cordate at base, the petiole 10–30 cm with a denticulate, irregular wing for most or all of its length, sometimes tinted purplish at base and with pale simple eglandular hairs; medium and lower 16–30 × 8–12 cm, lyrate, the terminal lobe 12–16 × 8–12 cm, ovate, shortly acute or acuminate at apex, sinuateglandular-denticulate, often cordate at base, the midrib between the lateral and terminal lobes broadly winged with occasional small teeth, the basal lobes forming a subrotund outline with a cordate-amplexicaul base; upper with lamina 6–14 × 1.5–6.0 cm, lanceolate to narrowly ovate, longacuminate at apex, sinuate-glandular-denticulate, roundedamplexicaul at base; glabrous or nearly so on the upper surface, slightly glaucous with prominent venation on the lower surface and with numerous simple eglandular hairs on the veins. Capitula in a compact group near the apex with several long branches from the upper leaf axils, 40–50 mm in diameter; peduncles 3–25 mm, slender, with numerous, yellowish glandular hairs. Involucral bracts in several rows, 4.5–14.0 × 1.5–2.0 mm, pale yellowish-green

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with a very narrow pale margin and slightly purplish at apex, linear to linear-lanceolate, more or less obtuse at apex, with numerous yellowish or slightly reddish glandular hairs. Flowers 15–23 mm, all ligulate, the ligules pale lilac with slightly darker veins, shallowly and irregularly 5-lobed at apex. Receptacle shallowly pitted, flat, naked. Achenes about 5 mm, elliptical, compressed, narrowly winged, with 3 ribs on each face, glabrous; pappus of 2 rows of simple hairs, the outer shorter. Flowers 6–8. Visited by bees and flies. Introduced. Well naturalised on rough and waste ground and roadsides, but probably mostly spread by pieces of rhizome rather than seed. Frequent throughout much of Great Britain and in scattered localities in Ireland. Native of the Urals. All our plants belong to subsp. uralensis (Rouy) P. D. Sell (Sonchus hispidus Ledeb., non C. hispida (M. Bieb.) Beauverd; Mulgedium hispidum Korsh.; Mulgedium uralensis Rouy; C. uralensis (Rouy) Beauverd; C. gmelinii Beauverd). 3. C. plumieri (L.) Kirschl. Hairless Blue Sow-thistle Sonchus plumieri L.; Mulgedium plumieri (L.) DC.; Lactuca plumieri (L.) Gren. & Godr. Perennial herb with a rootstock. Stem 60–130 cm, erect, stout, furrowed, glabrous or with a very occasional simple eglandular hair. Leaves 5–60 × 2–17 cm, green on upper surface, slightly glaucous and paler beneath; basal few, the lamina lyrate-pinnatifid, with a large, triangular, entire or remotely denticulate terminal lobe and several pairs of more or less ovate, patent or slightly recurved lateral lobes which are shorter than, but at least as wide as the terminal lobe, the lobes more or less undulate with mammiform teeth, the midrib between the lobes and the petiole with a broad wing; cauline similar but gradually becoming smaller upwards, sessile and semiamplexicaul; all glabrous or with a very occasional simple eglandular hair. Capitula in a wide, more or less corymbose panicle. Involucral bracts in several rows, 10–17 × 2–3 mm, yellowish-green, linear or linearlanceolate, more or less obtuse at apex. Flowers 17–20 mm, all ligulate, the ligules blue, with 5 lobes at apex. Receptacle flat, naked. Achenes 5.5–6.5 mm, linear to narrowly elliptical, conspicuously narrowed at apex, compressed, pappus of 2 rows of simple hairs, the outer shorter. Flowers 6–9. 2n = 16. Introduced. Garden escape naturalised in waste and grassy places. In a few scattered localities in England and Scotland. Known at Tighnabruich in East Inverness since 1950 and at Bothwell in Lanarkshire since 1957. Native of the Pyrenees and mountains of France and west-central Europe and south-west Bulgaria. Named after Charles Plumier (1646–1704). 4. C. bourgaei (Boiss.) Beauverd Pontic Blue Sow-thistle Mulgedium bourgaei Boiss.; Lactuca bourgaei (Boiss.) Irish & N. Taylor Perennial herb with stout roots and many stems from 1 base. Stem up to 3 m, pale green, solid, erect, slightly ridged, with scattered, pale simple eglandular hairs. Leaves numerous, all cauline, the lower soon dying, gradually

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decreasing in size up the stem, yellowish-green, the lamina 8–26 × 2–7 cm, thick, obovate or oblanceolate in outline, acute at apex, the lower with a large, rhombic, minutely undulate-denticulate terminal lobe and 1–2 pairs of rounded-denticulate lateral ones, the uppermost often undivided but dentate, all narrowed to a semiamplexicaul base, glabrous above and with numerous, stiff, white simple eglandular hairs beneath particularly on the prominent veins. Capitula up to 100, in an elongate, partly leafy panicle, 20–40 mm in diameter; peduncles glabrous or with simple eglandular hairs. Involucral bracts in several rows, 1–12 × 1.7–2.0 mm, pale green, suffused purple, ovatelanceolate to linear, obtuse at apex, glabrous. Flowers 12–19 mm, all ligulate, the ligules pale lilac with darker veins and a nearly white base, 5-lobed at apex. Receptacle flat, naked. Achenes 4.0–4.5 mm, pale brown, many-ribbed, compressed; pappus 7.0–7.5 mm, of 2 rows of simple hairs, the outer shorter. Flowers 6–9. Introduced. Grown in gardens from which it escapes on to roadsides and waste places. In scattered localities in England, Scotland and the Isle of Man. Native of Georgia and north-eastern Turkey. Named after Eug`ene Bourgeau (1815–77). 33. Mycelis Cass. Perennial herbs with latex, without rhizomes or stolons. Stems usually solitary, leafy, branched. Leaves pinnately lobed, the cauline more or less amplexicaul. Capitula few to numerous. Involucral bracts in 2 distinct, but unequal rows. Flowers bisexual. Corolla ligulate, yellow with 5 lobes at apex. Anthers tailed, with elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches hairy. Receptacle flat, without scales. Achenes oblanceolate, compressed, beaked; pappus of 2 unequal rows of white simple hairs. One species in Europe, south-west Asia and north-west Africa. Clabby, G. & Osborne, B. A. (1999). Mycelis muralis (L.) Dumort. in Biological flora of the British Isles. Jour. Ecol. 87: 156–172. Grime, J. P. et al. (1988). Comparative plant ecology. London.

1. M. muralis (L.) Dumort. Wall Lettuce Prenanthes muralis L.; Lactuca muralis (L.) Gaertn.; Cicerbita muralis (L.) Wallr.; Chondrilla muralis (L.) Lam. Perennial herb with a short, premorse stock. Stem 2–100 cm, solitary, erect, pale yellowish-green often suffused purplish, striate, glabrous, leafy, much branched above. Leaves yellowish-green, often suffused purplish; lower with lamina 5–22 × 3.5–9.0 cm, lyrate-pinnatifid, obtuse to acute at apex, the terminal lobe often hastately 3-lobed, triangulardentate and cordate-based, the lateral lobes gradually getting smaller towards the base, patent, hastate or ovate, dentate and parallel-sided at base, the area between the lobes winged and often dentate, with narrowly winged, often dentate petioles up to 70 mm; upper gradually becoming smaller, similar to lower but sessile and semiamplexicaul, the uppermost sometimes with lamina linear and entire;

all glabrous or nearly so. Capitula numerous, in a large, open panicle, cylindrical, 12–15 mm in diameter. Involucral bracts in 2 rows, pale green but usually suffused purplish, glabrous; inner 7–10 × 0.5–1.0 mm, linear to linearlanceolate, obtuse and often slightly frilled at apex; outer 0.5–2.5 × 0.5–0.7 mm, triangular-lanceolate, more or less obtuse at apex. Flowers 12–14 mm, all ligulate, the ligules yellow, with 5 lobes at apex. Receptacle flat, naked. Achenes 3.5–5.0 mm, blackish-brown, oblanceolate, compressed, many-ribbed, with a yellow beak 0.6–1.3 mm; pappus of an inner ring of scabrid hairs 5–6 mm, and an outer ring of thick obtuse hairs 0.1–0.2 mm. Flowers 7–9. Visited by flies and bees. 2n = 18. Native. On walls and rocks, on limestone pavement, in woods particularly of beech on chalk, in cultivated and waste places and in hedgerows, usually on nutrient-rich soils. Locally common in England and Wales, scattered in Scotland, Ireland and the Isle of Man. In the Burren it was not recorded until 1939, but is now frequent. Europe northwards to about 68.5◦ N in Norway; rare in north-west Africa; Turkey; Caucasus. A member of the European Temperate element. 34. Taraxacum Wigg. nom. conserv. By A. A. Dudman, A. J. Richards and P. D. Sell Perennial herbs with tap-roots. Stems few to many, hollow, simple, leafless. Leaves all basal, entire to laciniate-dentate or pinnately lobed. Capitula solitary. Involucral bracts in 2 rows, the inner erect and more or less oblong-lanceolate, the outer shorter, usually wider, often with paler margins, both rows sometimes with a small lump (callosed) or small appendage (corniculate) just below the apex. Flowers bisexual, all ligulate. Corolla yellow, often with a darker reddish, violet or brownish stripe on the outer face, with 5 lobes at apex. Anthers tailed, apical appendage elongate; filaments smooth. Style with slender branches, evenly hairy on both shaft and branches. Receptacle more or less flat, pitted, without scales. Achenes fusiform to oblanceolate, often with spiniform projections near the apex, usually with a slender beak and a more or less clearly demarcated swollen region (cone) between the beak and the body; pappus of many rows of simple, scabrid, rough, usually whitish hairs. There are probably at least a thousand species, mainly in Europe and Asia, North Africa and North America, and some have become cosmopolitan weeds. Dandelions are usually considered to be weedy plants, and the greater number of species, particularly of the Section Ruderalia, are to be found in pastures, roadside verges (especially recently disturbed ones) and urban wastelands. On recently sown motorway verges many may prove to be rare introductions from Scandinavia and the Netherlands which do not persist very long. This sort of species is often difficult to define as native or introduction, and the suggestions made in this account of the species are based on the experience of the authors of the B.S.B.I. Handbook. In native habitats dandelions occur on heaths, dunes, shingle, old grassland, cliff ledges and woodland rides. One of the best localities for native species is an old sunken lane, particularly in the west of Great Britain, where

34. Taraxacum species of the Sections Naevosa, Celtica, Hamata and Erythrosperma may be found and introduced Ruderalia rarely occur. Seeds are dispersed on the wheels of vehicles, on the feet of humans and animals, in animal feed and on clothing. They are also wind-dispersed, but we do not know for what distance. They will swiftly colonise any open or disturbed ground, but are poor competitors. Their deep tap-roots can stand almost any damage and fresh plants will propagate from buried root-cuttings. History Until the start of the twentieth century only a few aggregate species of Taraxacum were recognised in western Europe. Modern Taraxacum taxonomy, in which narrowly defined microspecies were recognised, started with the work of the Scandinavian botanists G. A. H. Dahlstedt (1856–1934) and C. C. Raunkiaer (1860–1938), who were followed by H. Lindberg (1871–1963), A. Palmgren (1880–1960), G. G. Marklund (1892–1964), M. M. W. Brenner (1843–1930), M. P. Christiansen (1889–1975), G. E. Haglund (1900–55), A. N. Railonsala (1902–82), C. I. Schlin (1912–20), J. L. van Soest (1898–1983) and Hans Øllgaard (b. 1943). The first work carried out on the British and Irish species was when G. C. Druce sent specimens to the ageing Dahlstedt for identification. Much of this early work was of poor quality, and between 1930 and 1964 little further work was carried out. The first serious study of our dandelions was carried out by John Richards, which resulted in his monograph in 1972. In his early years Richards was helped by the veteran van Soest and by Carl-Frederick Lundeval, but he largely worked alone. Further stimulus was added when Chris Haworth and Hans Øllgaard collaborated with Richards, with the result that the 1980s were a fruitful decade for the study of British and Irish dandelions. On the untimely death of Haworth in 1990, his friend Andrew Dudman took over and has produced with Richards the B.S.B.I. Handbook, Dandelions of Great Britain and Ireland. It is based on a large herbarium they have brought together and a database of some 30,000 records. Dahlstedt’s early naming has been revised and brought up to date by Haworth and Richards (1990). The account for this flora is largely based on the Handbook, the manuscript of which the authors kindly lent us at an early stage, and the specimens they have determined. Gaps in the descriptions have been filled in by reference to the original publications all of which have been examined. Original statements in the Handbook, however, are not altered even if they disagree with the original description. Andrew Dudman had agreed to go through the manuscript and bring the distributions up to date, but his untimely death has prevented this. John Richards has seen the final manuscript. Morphology Most dandelions are long-lived perennials with a simple or branched tap-root, but in some circumstances behave as

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biennials. They are referred to as small, medium or large to give an approximation of size. The leaves are in a basal rosette and are usually numerous. They are prostrate, spreading or erect. In outline they are usually broader above the middle but elongated and are thus often oblong-oblanceolate. Sometimes they are almost oblong and in the smaller species almost linear. In the large species they can become almost oblong-obovate. Sometimes the plant is heterophyllous, with the outer leaves different from the inner. Their colour is important. The general green colour given is that of the upper surface; the lower surface is usually paler. The midrib on the upper surface is often tinted purplish, but it is more important to know if it is at all purple, pinkish, pale green or even whitish on the lower surface. Sometimes the leaves are spotted or they are blotched with brownish-purple in the winged area of the midrib between the lobes. The lamina is more or less deeply pinnately divided with a terminal lobe and up to 10 pairs of lateral lobes. The lamina is flat, or the wings of the midrib between the lobes and the teeth of the lobes, spreading or erect, giving the leaf a three-dimensional appearance in nature, when it is referred to as crisped. The lamina is glabrous and smooth or with simple eglandular hairs which make it rough. The lobes are triangular, deltoid, hastate or hamate. Triangular lobes form two sides of an isosceles triangle, so that the apex is patent to the centre of the base of the lobe. Deltoid lobes have the margin towards the proximal part of the leaf shorter than that of the distal. Hastate lobes are triangular with the basal corners spreading. Hamate lobes have the margin towards the distal part of the leaf convex and the proximal one concave. The lobes may be acute or obtuse and sometimes have the apex and/or basal corners elongated into a narrow process, and they may be entire to deeply dentate. The margins of the lobes may be straight, curved or S-shaped (sigmoid). The lateral lobes may be spreading, recurved or, rarely, upturned. Sometimes the shape of the interlobe space is important. The winged area of the midrib between the lobes may be entire or dentate. The petioles are short to long, unwinged or with narrow to broad wings. It is important whether they are pale green to whitish or pink to purplish on the lower surface. The upper and lower surfaces of the leaf are defined according to how the leaf is fixed at the base, regardless of whether it is prostrate, spreading or erect. Dandelion leaves differ according to age, time of the year, different soils, amount of moisture or amount of shade and grazing, so that it is only possible to identify them with certainty for a short time of the year, in late bud and early flower. The following points should be considered. (1). Juvenile and shaded leaves are less complex in shape and essentially undivided. (2). Trodden and drought-stressed plants have leaves more complex in shape. (3). During the season, successive leaves develop an increasing complexity of shape, the outer early leaves usually being the basic phenotype. (4). Some species are heterophyllous, the inner leaves developing large terminal lobes quite different from the small terminal lobes of the early outer ones. (5). Sometimes only some of the leaves have the distinctive shape and toothing of the species. (6). In general, the colour, markings and texture of a leaf are

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inner involucral bracts

distal margin

reflexed outer involucral bracts

flowering stem (scape) proximal margin midrib dentate interlobe area

Deltoid

Triangular

Shapes of leaf lobes Taraxacum

34. Taraxacum

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lobes at apex inner involucral bracts

pappus

ligule

style outer appressed involucral bracts

pale scarious margin

beak

cone

pappus

spiniform projection cone

seed head

ovary

achene body

flower inner involucral bracts

inner involucral bracts ligules achene

outer involucral bracts

spreading outer involucral bracts

reflexed outer involucral bracts

capitulum in bud capitulum in fruit capitulum in flower Taraxacum

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more reliable guides to identification than leaf shape; however, anthocyanin pigments develop more strongly in good light and on early outer leaves than in shade or on later inner leaves. These pigments may also fade with age, and, in one case at least, that of T. pseudolarssonii, they appear to be water-soluble. (7). Petiole undersides which appear green in pressed specimens may be whitish when fresh, and pink or purplish ones may be whitish at the very base. Petiole colour may be modified by habit. (8). Hairiness is best judged on young leaves, as they tend to become glabrous with age. Flower stems are true scapes in the dandelions, but are not so called in this account, because as in other genera of the Asteraceae there is a gradual transition to leafy stems. They are pale green to suffused brownish-purple and are arachnoid hairy (best seen in young plants) to glabrous. Capitulum size can be judged in two ways, by its diameter when it is open and by the length of the inner involucral bracts. In a few species the capitulum does not open. Outer involucral bracts provide important characters. Size is given by measurements. They can be reflexed, recurved, spreading, erect or erect-appressed, a character best judged when they are in late bud before the capitulum opens. The shape and apex are important. In the Sections Erythrosperma and Obliqua there is a small appendage on the outer face near the apex, in which case the bract is referred to as corniculate. In this account the outer face and the inner face are determined by how the bract is fixed at the base; thus they always refer to the same face whichever direction the bract is pointing. Colour of the bracts is important. The outer face is usually darker than the inner and the inner is more likely to be pinkish-tinted or pruinose. The pale margin is usually more obvious on the outer face. Inner involucral bracts are not usually used by dandelion taxonomists, but their length gives an indication of the size of the capitulum and the corniculation on them can be clearly seen in bud in the species of the Sections Erythrosperma and Obliqua. Ligule colour and the colour of the stripe on the outer face are important characters. The apical lobes are sometimes red or black. Style colour, which usually refers to the stigmatic arms as well, whether yellow or discoloured, is important. In some species it is yellow both when fresh and when dried, although poor drying or disease can discolour the styles. Those that are dark when fresh usually dry black. Achenes can be divided into the body, the cone, the beak and the pappus. Size of the body is consistent in each species, but varies considerably in respect to its position in the capitulum, the inner tending to be smaller than the outer. The colour of the body is important, but it changes with maturity and fades easily. Little attention has been given to the shape of the body, but an examination of H. Lindberg (1935) shows how this varies in different species. At the apex of the body are spiniform projections, which vary in number and size. Between the body and the beak is a short projection from the body called the cone, which varies in size according to the species. It is usually the same

colour as the body, but sometimes paler. The beak is long and connects the body to the pappus; it is paler than the body. The pappus is whitish and varies in size according to the species. Collecting Dandelion identification is best carried out by comparing specimens with a reference herbarium. Specimens are best collected by cutting with a knife below the rosette. The root left in the ground will regenerate, so that conservation rarely presents a problem. However, rare native species, particularly those of the Sections Palustria and Taraxacum, should not be collected. The best material is collected early in the season, preferably just as it is coming into flower. Wellgrown, typical specimens should be selected, and diseased, grazed, shaded, gross, stressed or juvenile plants ignored. Specimens should be put in the press immediately and dried as quickly as possible so as to retain the colour. If the most advanced capitulum is put in a pot of water, mature achenes can be obtained as well. The collecting of good dandelion specimens is time-consuming and hard work. Cultivation If there is any doubt about two plants being different they should be grown side by side in the same conditions. In a similar way plants collected out of season can be cultivated. The Czech botanists Jan Kirschner and Jan Stepanek have found that the best way to grow them is in tomato boxes, the root restriction so brought about seeming to provide the nearest state to the early-flowering forms that it is recommended to collect. The species Although the sections in Taraxacum seem to be more interesting ecologically and geographically than they do in Hieracium, they are not the equivalent of sexual species. The microspecies are the most important entity within the genus and they should be regarded as the equivalent of sexual species in other genera. Virtually all our species are apogamous, producing seed asexually, the offspring being an exact copy of the mother. Sexual reproduction of dandelions in Great Britain and Ireland, if it occurs at all, is rare. As in Hieracium, when the dandelions went south ahead of the last glaciation they may have been mainly if not entirely sexual, species which had formerly been geographically isolated and hybridising coming together. An even greater mixture may have taken place when they moved north again. While they were still sexual some geographical and ecological segregation may have taken place. At what stage apomixis appeared we do not know, but it is almost certain that some species were still sexual while others were apomictic. Thus, further crossing between apomictic and sexual species furthered new apomictic taxa. More than in Hieracium, with the coming of Man many species became weedy and probably spread rapidly into new areas. There would have been a rapid increase in the numbers of plants, with a greater chance of producing new apomicts by mutation. Asexual species fall prey to the mutants which accumulate within

34. Taraxacum them. They cannot shed these mutants by the sexual processes of recombination or segregation (Richards, 1986). This is best illustrated by pollen-less mutants. Approximately 16 per cent of the British and Irish dandelion species lack pollen. Although the remainder have pollen, they are mostly asexual and thus the pollen is not beneficial unless they encounter the rare sexual species. Pollenproducing species cannot become pollen-less unless the correct mutations arise. Recent evidence suggests that dandelion species may not be as invariable as was once thought. They do of course accumulate mutants, which provide some variability. This process has now been detected in dandelions at the molecular level (King & Schaal, 1990). Also, although their genes are not recombined at meiosis, they have unusually high levels of chromosome breakage and reunion in dividing cells (Richards, 1989). This phenomenon may enable disadvantageous mutants in apomictic dandelions to move potentially into a position in the genome where they are silenced and can do no harm. All diploid Taraxaca (2n = 16) are sexual and almost all polyploids apomictic. Most polyploids are triploids (2n = 24). Chromosome gains and losses in the polyploids are accumulated. Although triploids usually have 24 chromosomes, 25, 26 and occasionally 23 are sometimes found, some of which are partially sexual. Every number between 32 and 39 has been recorded in T. ceratolobum (Richards, 1973). Borgvall, T. (1960). The Taraxacum Flora of Bohusl¨an. Acta Horti Gotob. 23: 1–40. Christiansen, M. P. (1936). Nye Taraxacum-arter af Gruppen Vulgaria. Dansk Bot. Ark. 9(2): 1–32. Christiansen, M. P. (1942). Taraxacum L. in C. Raunkiaer, Dansk Ekskursions – Flora. Ed. 6, by K. Wiinsted. 321. Christiansen, M. P. (1942). The Taraxacum-flora of Iceland in Rosenvinger, J. L. A. K. et al. (but (3) by J. Gr¨ontved, O. Paulsen & T. Søvensen.) (Edits.) The Botany of Iceland 3(3): 229–343, plus plates 1–44. Christiansen, M. P. (1971). Nye Taraxacum-arter i Danmark. Bot. Tidsskr. 66: 76–97. Dahlstedt, H. (1905). Om Scandinavska Taraxacum former. Bot. Not. 1905: 145–172. Dahlstedt, H. (4 Nov., 1906). Einige wildwachsende Taraxaca aus dem Botanischen Garten zu Upsala in Sernander, R. et al. (Edits.) Botaniska Studier till¨agnade F. R. Kjellman, pp. 164–183. Uppsala. Dahlstedt, H. (13 Nov., 1907). Taraxacum Weber in Warming, E. et al. (Edits.) Botany of the Faer¨oes, vol. 3, p. 840. Copenhagen & Christiania. Dahlstedt, H. (1909). Nya skandinaviska Taraxacum-arter. Bot. Not. 1909: 167–179. ¨ Dahlstedt, H. (1910). Ostsvenska Taraxaca. Ark. Bot. (Stockh.) 9(10): 1–74. ¨ Dahlstedt, H. (1911). Nya Ostsvenska Taraxaca. Ark. Bot. (Stockh.) 10(6): 1–36. Dahlstedt, H. (1911). V¨astsvenska Taraxaca. Ark. Bot. (Stockh.) 10(11): 1–74. Dahlstedt, H. (1912). Nordsvenska Taraxaca. Ark. Bot. (Stockh.) 12(2): 1–122. Dahlstedt, H. (1913). Taraxaca Scandinavica exsiccata 3: 14. Stockholm. Dahlstedt, H. (1920). Taraxacum anglicum Dahlst. Plant notes for 1919. Rep. Bot. Soc. Exch. Cl. Brit. Isles 5: 567.

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Dahlstedt, H. (1921). De svenska arterna av slaktet Taraxacum. I. Erythrosperma, II Obliqua. Acta Fl. Sueciae 1: 1– 160. Dahlstedt, H. (1923). Some new English species of Taraxacum. Rep. Bot. Soc. Exch. Cl. Brit. Isles 6: 773–780. Dahlstedt, H. (1925). Taraxaca fr˚an vastra Norge. Bergen Mus. Årbok. ¨ Dahlstedt, H. (1925). Om Olands Taraxacum-flora. Ark. Bot. (Stockh.) 19(18): 1–19. Dahlstedt, H. (1926). In Johnston, H. H., Scottish Taraxaca. Trans. & Proc. Bot. Soc. Edinb. 29: 300–305. Dahlstedt, H. (1927). In Johnston, H. H., Scottish Taraxaca. Trans. & Proc. Bot. Soc. Edinb. 29: 416–423. Dahlstedt, H. (1927). Plant notes for 1926. Rep. Bot. Soc. Exch. Cl. Brit. Isles 8: 30–32. Dahlstedt, H. (1929). Plant notes for 1928. Rep. Bot. Soc. Exch. Cl. Brit. Isles 8: 619–629. ¨ Dahlstedt, H. (1929). Uber einige orientalische Taraxacum-Arten. Acta Hort. Berg. 9: 1–36, plus plates. Dahlstedt, H. (1930). Plant notes for 1929. Rep. Bot. Soc. Exch. Cl. Brit. Isles 9: 26–30. Dahlstedt, H. (1930). De svenska arterna av slaktet Taraxacum. Kungl. Svenska Vet.-Akad. Handl. ser. 3, 9(2): 1–97. Dahlstedt, H. (1932). Plant notes for 1931. Rep. Bot. Soc. Exch. Cl. Brit. Isles 9: 562–566. Dahlstedt, H. (1935). Nya skandinaviska Taraxaca. Bot. Not. 1935: 295–316. Dudman, A. A. & Richards, A. J. (1994). Seven new species of Taraxacum Wigg. (Asteraceae), native to the British Isles. Watsonia 20: 119–132. Dudman, A. A. & Richards, A. J. (1997). Dandelions of Great Britain and Ireland. London. Florstr¨om, B. (1914). Studier o¨ fver Taraxacum-floran i Satakunta. Acta Soc. Fauna Fl. Fenn. 39(4): 1–125. Hagendijk, A., Soest, J. L. van & Zevenbergen, H. A. (1972). Neue Taraxacumarten der Niederlande. Acta Bot. Neerl. 21(5): 491– 511. Hagendijk, A., Soest, J. L. van & Zevenbergen, H. A. (1973). Neue Taraxacumarten der Niederlande II. Acta Bot. Neerl. 22(6): 616– 636. Hagendijk, A., Soest, J. L. van & Zevenbergen, H. A. (1974). Neue Taraxacumarten der Niederlande III. Acta Bot. Neerl. 23(4): 439–459. Hagendijk, A., Soest, J. L. van & Zevenbergen, H. A. (1976). Neue Taraxacumarten der Niederlande IV. Acta Bot. Neerl. 25(1): 81– 105. Hagendijk, A., Soest, J. L. van & Zevenbergen, H. A. (1978). Neue Taraxacumarten der Niederlande V. Acta Bot. Neerl. 27(5–6): 307–331. Haglund, G. E. (1934). N˚agra nya eller i Sverige nyfunna Taraxacum-arter. Bot. Not. 1934: 1–42. Haglund, G. E. (1935). Taraxaca fr˚an s¨odra och mellersta Sverige samt Danmark. Bot. Not. 1935: 96–130. Haglund, G. E. (1935). Some Taraxacum-species from Ireland and Wales cultivated in the Botanical Garden of Lund. Bot. Not. 1935: 429–438. Haglund, G. E. (1936). N˚agra Taraxaca huvudsakligen fr˚an v¨astra Sverige. Acta Horti Gotob. 11: 19–41. Haglund, G. E. (1937). On some Taraxacum species of the Group Vulgaria Dahlst. Bot. Not. 1937: 56–62. Haglund, G. E. (1938). Bidrag till k¨annedomen om Skandinaviens Taraxacum-flora II. Bot. Not. 1938: 499–508. Haglund, G. E. (1942). Taraxacum in Holmgren, B. Blekinges Flora, pp. 322–346. Karlshamn. Haglund, G. E. (1943). N˚agra nya Taraxaca fr˚an Sk˚ane och Danmark. Bot. Not. 1943: 232–242.

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¨ Haglund, G. E. (1946). Zur Taraxacum – Flora der Insel Oland. Bot. Not. 1946: 335–363. ¨ Haglund, G. E. (1947). Uber die Taraxacum-flora der Insel R¨ugen. Svensk Bot. Tidskr. 41: 81–103. Haglund, G. E. & Lillieroth, C. G. (1941). Beitr¨age zur Taraxacum der Inselgruppe Lofoten. Nytt. Mag. Naturvidensk. 82: 83– 99. Haglund, G. E. & Morander, R. (1937). Till Hj¨almarlandskapens Taraxacum-flora. Svensk Bot. Tidskr. 31: 339–353. Haworth, C. C. (1990). Six native species of Taraxacum new to the British Isles. Watsonia 18: 131–138. Haworth, C. C. (1997). Taraxacum in Halliday, G. A flora of Cumbria. Haworth, C. C. & Richards, A. J. (1990). The lectotypification and revision of Dahlstedt’s species of Taraxacum Weber based on British or Irish plants. Watsonia 18: 125–130. Hughes, J. & Richards, A. J. (1988). The genetic structure of populations of sexual and asexual Taraxacum (dandelions). Heredity (London) 60: 161–171. Hughes, J. & Richards, A. J. (1989). Isozymes, and the status of Taraxacum (Asteraceae) agamospecies. Bot. Jour. Linn. Soc. 99: 365–376. ˇ ep´anek, J. (1987). Again on the sections in TaraxKirschner, J. & Stˇ acum (Cichoriaceae) (Studies in Taraxacum 6). Taxon 36: 608– 617. ˇ ep´anek, J. (1995). Souˇcasn´y stav taxonomick´eho Kirschner, J. & Stˇ ˇ v´yzkumu rodu Taraxacum v Cesk´ e a Slovensk´e republice. Zpr. ˇ Bot. Spaleˇc. Praha 29: 1–9. (Gives other papers on TaraxCes. acum by the same authors.) ˇ ep´anek, J. (1996). Modes of speciation and evoKirschner, J. & Stˇ lution of sections in Taraxacum. Folia Geobot. Phytotax. 31: 415–426. ˇ ep´anek, J. (1997). A nomenclatural checkKirschner, J. & Stˇ list of supraspecific names in Taraxacum. Taxon 46: 87–98. ˇ ep´anek, J. (1998). A monograph of Taraxacum Kirschner, J. & Stˇ sect. Pruhonice. Lange, Th. (1938). J¨amtlands K¨arlv¨axtflora. Acta Bot. Fenn. 21: 1–204. (H. Dahlstedt species ex T. A. Lange, the remainder ex G. Haglund.) Lambinon, J. & Soest, J. L. van (1962). Deux Taraxacum nouveaux de Belgique. Lejeunea nov. ser. 8: 1–2. Lindberg, H. (1908). Taraxacum-former fr˚an s¨odra och mellersta Finland. Acta Soc. Fauna Fl. Fenn. 29(9): 1–48. Lindberg, H. (1909). Nytt bidrag till k¨annedomen af Taraxacumformerna i s¨odra och mellersta Finland. Meddeland. Soc. Fauna Fl. Fenn. 35: 13–31. Lindberg, H. (1910). Finska Taraxacum-former. Meddeland. Soc. Fauna Fl. Fenn. 36: 5. Lindberg, H. (1935). Die fr¨uchte der Taraxacum-arten Finnlands. Acta Bot. Fenn. 17: 1–22, tafel 1–38. Lindberg, H. & Marklund, G. (1911). Acta Soc. Fauna Fl. Fenn. 34(7): 5. Marklund, G. (1925). Nya Taraxaca. Acta Soc. Fauna Fl. Fenn. 55(5): 1–25, plus plates. Marklund, G. (1938). Die Taraxacum-Flora Estlands. Acta Bot. Fenn. 23: 1–150. Marklund, G. (1940). Die Taraxacum-Flora Nylands. Acta Bot. Fenn. 26: 1–187. Mogie, M. & Richards, A. J. (1983). Satellited chromosomes, systematics and phylogeny in Taraxacum (Asteraceae). Pl. Syst. Evol. 141: 219–229. Øllgaard, H. (1972). Om nogle danske maelkebøtter. Bot. Tidsskr. 67(1–2): 139–145. Øllgaard, H. (1978). New species of Taraxacum from Denmark. Bot. Not. 131(4): 497–521.

Øllgaard, H. (1983). Hamata, a new section of Taraxacum (Asteraceae). Pl. Syst. Evol. 141: 199–217. Palmer, R. C. & Scott, W. (1995). A forgotten Dandelion. Watsonia 20: 279–281. Palmgren, A. (1910). Bidrag till k¨annedom om a˚ lands Vegetation och Flora. I Taraxaca. Acta Soc. Fauna Fl. Fenn. 34(1): 1–53 and plates 1–12. Palmgren, A. (1910). Acta Soc. Fauna Fl. Fenn. 34(4): 1–16. Puolanne, M. E. (1932–1933). Helsingin ja sen l¨ahiseudun kasvisto. 1. Taraxacum-lajit. Memoranda Soc. Fauna Fl. Fenn. 8: 136– 181. Railonsala, A. (1942). Ann. Soc. Zool. Bot. Fenn. ‘Vanamo’ 16(5): 11. Railonsala, A. (1957). Taraxaca nova I. Arch. Soc. Zool. Bot. Fenn. ‘Vanamo’ 11: 148–171. Railonsala, A. (1967). Taraxaca nova VI. Ann. Bot. Fenn. 4: 102– 115. Raunkiaer, C. (1903). Kimdannelse uden befr¨ugtning hos Maelkebotte (Taraxacum). Bot. Tidsskr. 25(2): 109–139. Raunkiaer, C. (Jan.–June, 1906). Dansk Ekskursions-Flora. Ed. 2. pp. 254–258. København & Kristiania. Raunkiaer, C. (1922). Dansk Ekskursions-Flora. Ed. 4. Ostenfeld, C. H. & Raunkiaer, C. (Edits.). (Taraxacum, pp. 303–307 by C. Raunkiaer.) Raunkiaer, C. (1934). Dansk Ekskursions-Flora. Ed. 5. Wiinstedt, K. & Jessen, K. (Edits.). København & Kristiania. (Taraxacum, pp. 302–318, by M. P. Christiansen & K. Wiinstedt, but new species by individual authors.) Richards, A. J. (1972). The Taraxacum Flora of the British Isles. Watsonia 9 suppl.: 1–141. Richards, A. J. (1973). The origin of Taraxacum agamospecies. Bot. Jour. Linn. Soc. 66: 189–211. Richards, A. J. (1981). New species of Taraxacum from the British Isles. Watsonia 13: 185–193. Richards, A. J. (1985). Sectional nomenclature in Taraxacum (Asteraceae). Taxon 34(4): 633–644. Richards, A. J. (1989). A comparison of within-plant karyological heterogeneity in nucleolar organiser chromosome. Pl. Syst. Evol. 163: 177–185. Richards, A. J. (1992). The Taraxacum flora of the Sierra de Guadarrama and its surroundings (Spain). Anales Jard. Bot. Madrid 50: 201–208. Richards, A. J. (1996). Genetic variability in obligate apomicts of the Genus Taraxacum. Folia Geobot. Phytotax. 31: 405– 414. Richards, A. J. & Haworth, C. C. (1984). Further new species of Taraxacum from the British Isles. Watsonia 15: 85–94 and plates 1–3. Sahlin, C. I. (1983). Taraxacum cenabense Sahlin, a new AngloFrench species. Watsonia 14: 281–282. Sahlin, C. I. & Soest, J. L. van (1970). Taraxacum atactum, spec. nov. Acta Bot. Neerl. 19: 197–200. S˚altin, H. (1971). Sieben neue nordische Taraxaca. Mem. Soc. Fauna Fl. Fenn. 47: 38–59 and plates 1–9. Soest, J. L. van (1956). New Taraxaca from the Netherlands. Acta Bot. Neerl. 5: 94–101. Soest, J. L. van (1961). Quelques nouvelles esp`eces de Taraxacum, native d’Europe. Acta Bot. Neerl. 10: 280–306. Soest, J. L. van (1975). Quelques esp`eces nouvelles de Taraxacum, de France. Acta Bot. Neerl. 24: 139–154. Soest, J. L. van, Hagendijk, A. & Zevenbergen, H. (1968). Taraxacum kernianum, spec. nov. Gorteria 4: 123–126.

The following key has been constructed from the key to the sections by Stace (1997), the key to the groups of species by Dudman and Richards (1997) and from

34. Taraxacum information taken from the descriptions in the latter work. To use the key good living material is needed, or good pressed specimens plus detailed notes of characters, particularly colour, made when the plant was fresh. Identification should always be confirmed by checking with the detailed descriptions in the text. If the user is experienced and can recognise the sections, Section Erythrosperma starts at break 4, Section Obliqua at break 3, Section Palustria at break 42, Section Taraxacum at break 46, Section Spectabilia at break 52, Section Naevosa at break 55, Section Hamata at break 68, Section Celtica at break 95 and Section Ruderalia at break 138. 1. Plants delicate; usually with strongly dissected leaves more or less to the midrib; outer involucral bracts rarely more than 7 mm; capitula rarely more than 30 mm in 2. diameter 1. Plants usually medium-sized to robust; rarely with such deeply dissected leaves; outer involucral bracts usually more than 7 mm; capitula usually more than 30 mm in 41. diameter 2. Leaves often with 6 or more pairs of lateral lobes; achenes greyish-brown, with a pyramidal cone less than 3. 0.5 mm (Section Obliqua) 2. Leaves rarely with more than 6 pairs of lateral lobes; achenes usually violet-purple or reddish- to yellowish-brown, with a cylindrical cone about 1.0 mm 4. (Section Erythrosperma) 3. Leaves pale green, the lateral lobes obtuse at apex and scarcely dentate; capitulum 10–15 mm in diameter, 32. obliquum usually closed 3. Leaves medium to dark green, the lateral lobes more acute at apex and more dentate; capitulum 25–30 mm in 33. platyglossum diameter, usually flat and open 5. 4. Petioles narrow and deep purple 12. 4. Petioles not as above 8. rubicundum 5. Flowering stem completely glabrous 5. Flowering stem arachnoid-hairy, at least just below the 6. capitulum when young 11. dunense 6. Achenes dark violet 7. 6. Achenes not dark violet 8. 7. Achenes pale brown 9. 7. Achenes not pale brown 30. tortilobum 8. Capitulum less than 25 mm in diameter 8. Capitulum more than 25 mm in diameter 31. pseudolacistophyllum 10. 9. Outer involucral bracts spreading 11. 9. Outer involucral bracts erect 1. lacistophyllum 10. Achenes chestnut to purple 16. pseudoproximum 10. Achenes greyish-brown 11. Ligules with a reddish-purple stripe on outer face; 7. commixtum achenes dark red 11. Ligules with a greyish-violet stripe on outer face; 18. oxoniense achenes cinnamon 13. 12. Outer involucral bracts strongly recurved 17. 12. Outer involucral bracts spreading to erect 14. 13. Achenes chestnut, dark brown or puce 15. 13. Achenes pale

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14. Styles discoloured; achenes dark brown or puce 14. proximum 17. disseminatum 14. Styles blackish; achenes chestnut 24. glauciniforme 15. Styles yellow; pollen absent 16. 15. Styles discoloured; pollen present 16. Lateral leaf-lobes dentate, often on both margins; 23. retzii achenes cinnamon 16. Lateral leaf-lobes dentate only on distal margin of 25. wallonicum proximal lobes; achenes straw-coloured 18. 17. Pollen absent 30. 17. Pollen present 19. 18. Achenes chestnut to dark red 23. 18. Achenes brown or puce 2. inopinatum 19. Leaves spotted 20. 19. Leaves unspotted 20. Ligules inrolled, striped reddish-orange on outer face 5. argutum 21. 20. Ligules flat, striped otherwise 21. Outer involucral bracts with well-marked pale margins; ligules with purple-violet stripe on outer face 6. arenastrum 21. Outer involucral bracts with poorly marked pale 22. margins; ligules with purple stripe on outer face 22. Leaves with purple petioles; flowering stems arachnoid10. parnassicum hairy at apex; ligule teeth yellow 22. Leaves with green petioles; flowering stems glabrous at 9. cenabense apex; ligule teeth purple 14. proximum 23. Achenes dark brown or puce 24. 23. Achenes cinnamon or pale brown 25. 24. Styles yellow 27. 24. Styles discoloured 25. Capitulum 30–40 mm in diameter; achene cone about 29. tanylepis 0.5 mm 25. Capitulum 20–30 mm in diameter; achene cone 26. 0.7–1.0 mm 26. Leaves bright green, lateral lobes if present not dentate but dentate on proximal interlobes near the midrib; petiole winged; ligules with purple teeth 15. proximiforme 26. Leaves dull green, lateral lobes dentate; petiole 20. fulvum unwinged below; ligules with reddish teeth 27. Outer involucral bracts with a conspicuous pale margin; 26. degelii ligule teeth yellow 27. Outer involucral bracts without or with only an 28. inconspicuous pale margin; ligule teeth dark 28. Leaves bright green; ligule teeth purple 15. proximiforme 29. 28. Leaves dull green; ligule teeth reddish 29. Outer involucral bracts erect to spreading giving a 19. fulviforme star-shaped involucre 20. fulvum 29. Outer involucral bracts spreading to recurved 31. 30. Achenes chestnut to red 35. 30. Achenes brown or dark purple 13. gotlandicum 31. Styles yellow; ligule teeth dark 32. 31. Styles usually discoloured; ligule teeth yellow 33. 32. Ligule stripe purplish-violet 34. 32. Ligule stripe greyish-purple

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33. Outer involucral bracts glaucous and usually suffused purple on inner face, faintly pale margined, corniculate; 3. brachyglossum achene cone 0.4–0.7 mm 33. Outer involucral bracts green with a well-marked pale margin, scarcely corniculate; achene cone 0.7–0.8 6. arenastrum mm 34. Only the proximal lateral lobes dentate; outer involucral bracts somewhat corniculate; ligules rather pale 1. lacistophyllum yellow 34. All lateral lobes dentate on distal margin, sometimes on both margins; outer involucral bracts not corniculate; 4. scanicum ligules rather dark yellow 12. haworthianum 35. Styles yellow; achenes dark violet 36. 35. Styles usually discoloured; achenes pale 22. falcatum 36. Ligule lobes yellow 37. 36. Ligule lobes dark 37. Outer involucral bracts ascending or erect; styles 27. acutum discoloured; ligules with dark violet stripe 37. Outer involucral bracts spreading or recurved; styles yellowish; ligules with silver-grey or greyish-violet 38. stripe 28. placidum 38. Ligules with a silver-grey stripe 39. 38. Ligules with a greyish-violet stripe 16. pseudoproximum 39. Achenes greyish-brown 40. 39. Achenes straw-coloured or cinnamon 40. Outer involucral bracts about 8 × 2 mm; achene body 15. proximiforme about 3 mm 40. Outer involucral bracts 5–6 × 2–3 mm; achene body 21. scoticum 3.2–3.5 mm 41. Leaves very narrow, usually scarcely lobed; outer row of involucral bracts appressed, ovate, with a broad scarious 42. margin (Section Palustria) 41. Leaves usually distinctly lobed and broader; outer row of involucral bracts appressed to recurved, linear to narrowly ovate, with narrow to very narrow scarious 46. margin or without a scarious margin 43. 42. Pollen absent 45. 42. Pollen present 43. Outer involucral bracts with a broad, pale green margin; 34. palustre achene body 3.5–4.0 mm 43. Outer involucral bracts with a distinctly whitish margin; 44. achene body 4.5–5.5 mm 44. Outer involucral bracts 5.5–6.5 mm, broadest at base; achene with cone 0.8–1.0 mm, the beak 6.5–7.5 mm 35. webbii 44. Outer involucral bracts 6.5–7.5 mm, widest just below the middle; achenes with cone 0.7–0.9(–1.3) mm, the 36. amarellum beak 7.0–8.5 mm 45. Leaves oblanceolate-spathulate, usually with regularly spaced teeth; flowering stem arachnoid-hairy when 37. ciliare young 45. Leaves with shallow lateral lobes; flowering stem 38. anglicum glabrous 46. Leaves simple and bright green, with green petioles; rare plants of a few mountain cliffs in Scotland 47. (Section Taraxacum) 46. Lowland plants or, if on mountain cliffs with leaves usually dark green or blotched or spotted with purple 52. and petiole usually purple

47. 47. 48. 48. 49. 49. 50. 50. 51. 51. 52.

52.

53. 53. 54. 54. 55.

48. Styles yellow 49. Styles discoloured Achene body about 4.0 mm, cone 0.7 mm 42. ceratolobum Achene body about 3.2 mm, cone 0.3 mm 45. craspedotum 43. clovense Achene body 4.8–5.3 mm 50. Achene body not exceeding 3.8 mm 47. pycnostictum Leaves bluish with punctate spots 51. Leaves pure green, without spots Leaves unlobed, regularly though sparsely 44. xiphoideum deltoid-dentate 46. cymbifolium Leaves with 3–4 pairs of lateral lobes Outer row of involucral bracts erect to appressed; ligules usually with red stripes on outer face; pollen usually absent; achenes including cone but excluding beak more than 4.5 mm 53. and nearly cylindrical (Section Spectabilia) Outer row of involucral bracts rarely appressed; ligule stripe rarely red; pollen present or absent; achene body 55. less than 4 mm, narrowly turbinate Capitulum 45–50 mm in diameter; pollen present 40. geirhildae 54. Capitulum 30–40 mm in diameter; pollen absent Achene body 3.8–4.0 mm, almost lacking spiniform 39. faeroense projections at apex Achene body 4.8–5.0 mm, with short spiniform 41. serpenticola projections at apex Leaves with large dark spots covering more than 10 56. per cent of the surface (Section Naevosa)

55. Leaves unspotted or with spots covering less than 10 per 67. cent of the surface 56. Pollen absent or confined to the outer flowers of the 57. capitulum and very sparse 60. 56. Pollen present in all flowers of the capitulum 58. 57. Leaves with scattered small spots 59. 57. Leaves with many larger, often confluent spots 51. euryphyllum 58. Petiole broadly winged 53. maculosum 58. Petiole more or less unwinged 55. subnaevosum 59. Outer involucral bracts recurved 59. Outer involucral bracts suberect to spreading 59. richardsianum 61. 60. Leaves with scattered spots or none 64. 60. Leaves with many larger, often confluent spots 56. cornubiense 61. Achene body 3.6–3.8 mm 62. 61. Achene body 3.0–3.5 mm 57. drucei 62. Leaves mid-green, glabrous 63. 62. Leaves greyish-green, hairy 52. hirsutissimum 63. Outer involucral bracts recurved 63. Outer involucral bracts numerous and curved upwards to 58. stictophyllum give a rounded involucre 65. 64. Capitulum more than 50 mm in diameter 66. 64. Capitulum less than 50 mm in diameter 65. Outer involucral bracts 3–5 mm wide; achenes 48. naevosum straw-coloured 65. Outer involucral bracts 2.0–3.5 mm wide; achenes 50. rubellum brick-red 49. naevosiforme 66. Lateral leaf-lobes 4–7 pairs, broad

34. Taraxacum 54. pseudolarssonii 66. Lateral leaf-lobes 3–5 pairs, narrow 67. Leaves more or less flat and relatively simply lobed, upper side of petiole and midrib usually minutely striped red or purple; outer involucral bracts rarely more than 10 mm, usually patent to erect and dark green on outer 68. face 67. Leaves often complexly lobed and folded in 3 dimensions, the petiole and midrib green or solid red or purple; outer row of involucral bracts usually more than 10 mm, often recurved, not very dark on outer face 139. (Section Ruderalia) 68. Lateral leaf-lobes broad-based, with convex front and concave rear edge, commonly 4 pairs; outer row of involucral bracts usually arched to various degrees, often subobtuse at apex, pruinose on inner face, dark 69. olive green on outer face (Section Hamata) 68. Lateral leaf-lobes rarely as above, often 5–6 pairs; outer row of involucral bracts erect to recurved, all more or less to the same degree, often acute at apex 96. (Section Celtica) 69. Outer involucral bracts with a narrow but more or less 70. clearly demarcated pale margin 69. Outer involucral bracts without a pale margin or pale 73. margin indistinct 71. 70. Outer involucral bracts with a white margin 70. Outer involucral bracts with a pale margin, but not white 75. 95. hamatulum 71. Achene body 3.8–4.0 mm 72. 71. Achene body 3.0–3.5 mm 106. pruinatum 72. Ligule stripe greyish-violet 111. lamprophyllum 72. Ligule stripe dark brown 73. Capitulum 30–40 mm in diameter; achene body 99. quadrans 3.8–4.0 mm 73. Capitulum 45–55 mm in diameter; achene body about 74. 3.5 mm 74. Outer involucral bracts strongly pruinose on outer face 100. pseudohamatum 74. Outer involucral bracts not pruinose on outer face 102. prionum 75. Buds very dark, pruinose, inky black at apex; outer 94. hamatum bracts forming a stellate involucre 76. 75. Involucre and bud not so 77. 76. Terminal leaf-lobe more or less obtuse 79. 76. Terminal leaf-lobe more or less acute or acuminate 103. boekmanii 77. Terminal leaf-lobe entire 78. 77. Terminal leaf-lobe typically with one or more teeth 78. Leaves dark greyish-green, dentate, the petiole unwinged at least below; achene body 3.2–3.5 mm 104. atactum 78. Leaves dirty dark green, more dentate and more gross, the petioles distinctly winged; achene body 3.8–4.0 mm 105. sahlinianum 80. 79. Terminal lobe of leaf toothed 83. 79. Terminal lobe of leaf more or less entire 80. Terminal lobe of leaf obtuse or subacute without an 101. fusciflorum elongated apex 81. 80. Terminal lobe of leaf with an elongated apex 81. Ligule striped dark brown on outer face 111. lamprophyllum 82. 81. Ligules striped violet or greyish-violet on outer face

129

82. Lateral lobes of leaf regular, usually recurved and sometimes acutely so, the distal margin convex 97. marklundii 82. Lateral lobes of leaf irregular, spreading or recurved, the 110. kernianum distal margin straight to sinuate 84. 83. Terminal lobe of leaf with elongated apex 91. 83. Terminal lobe of leaf without elongated apex 85. 84. Midrib green or weakly pink or purple 86. 84. Midrib strongly coloured purple 85. Lateral lobes of leaf with distal margin slightly dentate proximally; outer involucral bracts pruinose, pale green 96. subhamatum on inner face 85. Lateral lobes of leaf recurved with the distal margin strongly dentate throughout; outer involucral bracts 97. marklundii suffused purple 86. Interlobe area near the midrib heavily blotched; maximum width of outer involucral bracts more than 87. 3.5 mm 86. Interlobe area near the midrib blotched or unblotched; maximum width of outer involucral bracts less than 88. 3.5 mm 87. Outer involucral bracts pale green on inner face; ligules 102. prionum with greyish-violet stripe on outer face 87. Outer involucral bracts suffused purplish on inner face; ligules with dark brown stripe on outer face 111. lamprophyllum 88. Ligules with dark brown stripe on outer face, those of inner flowers of capitulum with purple 109. lancidens teeth 89. 88. Ligules with greyish-violet stripe on outer face 89. Outer involucral bracts 11–13 × 3.5–4.5 mm 102. prionum 90. 89. Outer involucral bracts 10–11 × 2–3 mm 90. Terminal lobe of leaf triangular-sagittate with a more or less elongated apex; lateral lobes patent or recurved, narrowly triangular and acute at 107. hamatiforme apex 90. Terminal lobe of leaf markedly narrowly sagittate with an elongated apex; lateral lobes patent or recurved, sometimes markedly so, triangular, long, very narrow, 108. spiculatum tapering and acute at apex 91. Midrib bright purple to apex, usually with red 103. boekmanii side-veining 92. 91. Not as above 92. Ligule with brown stripe on outer face without yellow 101. fusciflorum margin at apex 92. Ligule with yellow margin at apex of stripe on outer 93. face 93. Inner involucral bracts not equally wide and usually 98. hamiferum irregularly connate 94. 93. Inner involucral bracts more or less equally wide 94. Maximum width of outer involucral bracts more than 100. pseudohamatum 3.5 mm 94. Maximum width of outer involucral bracts less than 95. 3.5 mm 95. Outer involucral bracts blackish-green and heavily 99. quadrans pruinose on outer face 95. Outer involucral bracts very dark, shiny, brownish-green 107. hamatiforme on outer face 97. 96. Pollen absent

130 96. 97. 97. 98. 98. 99. 99. 100. 100. 101. 101. 102.

102. 103. 103. 104. 104. 105. 105. 106. 106. 107. 107. 108. 108. 109. 109. 110. 110. 111. 111. 112.

112.

113.

1 4 8 . A S T E R AC E A E 109. Pollen present 98. Styles yellow 100. Styles discoloured or dark Outer involucral bracts spreading to erect, pruinose, 92. unguilobum with a clear white margin Outer involucral bracts spreading to recurved, pruinose 99. or not, more or less without a pale margin Ligules flat; styles exserted; achenes brown 88. ostenfeldii Ligules involute; styles inserted; achenes 89. breconense straw-coloured 101. Outer involucral bracts spreading to recurved Outer involucral bracts spreading to erect or 102. appressed 69. celticum Terminal lobe of leaf 3-lobed 72. inane Terminal lobe of leaf more or less obtuse Lateral lobes of leaves coarsely dentate; outer involucral bracts pruinose throughout; achenes 91. fulvicarpum cinnamon, oxidising to brown 103. Not as above 104. Outer involucral bracts with a clear pale margin Outer involucral bracts scarcely showing a pale 106. margin Outer involucral bracts dark glaucous-green; ligules 86. lancastriense short, with a grey stripe on outer face Outer involucral bracts green, pruinose or not; ligules 105. with a purple stripe on outer face Flowering stems purplish and arachnoid-hairy; achene 84. olgae body 2.6–3.0 mm Flowering stems glabrous and green; achene body 87. palustrisquamum 3.1–3.5 mm Outer involucral bracts not pruinose; ligules with a 80. landmarkii purple stripe on outer face Outer involucral bracts pruinose; ligules short, with a 107. puce or purple stripe on outer face Capitulum convex or closed, up to 20 mm in diameter 90. caledonicum 108. Capitulum open, 25–40 mm in diameter 81. nordstedtii Achene body 3.2–3.5 mm, olive-brown 83. berthae Achene body 3.6–3.9 mm, straw-coloured 93. luteum Ligules lacking a stripe 110. Ligules with a stripe on outer face Proximal lateral lobes of leaf filiform or acuminate at 111. apex Proximal lateral lobes of leaf acute to obtuse at apex 114. Ligules with purple stripe on outer face; achene body 112. straw-coloured Ligules with greyish-violet or brownish-purple stripe 113. on outer face; achene body olive-brown Leaves pale or yellowish-green, without spots, the petioles narrowly winged; flowering stems pale green, 66. oellgaardii becoming glabrous Leaves dull dark green the interlobes near the midrib sometimes with a slight dark blotch, the petiole unwinged; flowering stems dull green or purple, 68. porteri moderately arachnoid-hairy Leaves dull green, without spots; ligules with a greyish-violet stripe on outer face; styles darkly 67. duplidentifrons discoloured to blackish

113. Leaves dark, pure green, the interlobes near the midrib with dark blotches; ligules with a dark brownish-purple 77. haematicum stripe on outer face; styles discoloured 114. Leaves heavily and distinctly dark blotched on the 115. lower surface of the internodes 121. 114. Leaves unblotched or indistinctly blotched 116. 115. Terminal lobe of leaf obtuse to rounded at apex 118. 115. Terminal lobe of leaf acute to trilobate at apex 74. tamesense 116. Petiole unwinged 117. 116. Petiole winged 117. Ligules with a dark greyish-violet stripe on outer face 71. excellens 117. Ligules with a brownish-purple stripe on outer face 73. fulgidum 65. subbracteatum 118. Achene body straw-coloured 118. 119. 119. 120. 120. 121. 121. 122. 122. 123. 123. 124. 124. 125. 125. 126. 126. 127. 127. 128. 128. 129.

129.

130. 130. 131. 131. 132.

119. Achene body olive-brown or pale brown 69. celticum Petiole winged, sometimes broadly so 120. Petiole unwinged 70. hesperium Outer involucral bracts pruinose 77. haematicum Outer involucral bracts not pruinose 122. Midrib of leaf bright purple almost to apex Midrib of leaf not bright purple or, if so, not distally 130. 123. Lateral lobes of leaves 2–3(–4) pairs 125. Lateral lobes of leaves (3–)4 or more pairs 62. orcadense Capitulum 40–50 mm in diameter 124. Capitulum 20–35 mm in diameter 63. nietoi Outer involucral bracts erect 75. texelense Outer involucral bracts spreading Ligules short, with a puce, brown or purple stripe on 126. outer face Ligules longer, with a grey to purple stripe on outer 127. face Distal margin of lateral leaf-lobes concave; ligules with a brown or puce stripe on outer face 81. nordstedtii Distal margin of lateral leaf-lobes not concave; ligules 82. pseudonordstedtii with a purple stripe on outer face Outer involucral bracts blackish on outer surface; 64. britannicum styles blackish Outer involucral bracts dark green on outer surface; 128. styles not blackish Petiole unwinged; ligules with a purple stripe on outer 61. bracteatum face Petiole winged; ligules with a grey or greyish-violet 129. stripe on outer face Capitula 30–40 mm in diameter; ligules with a greyish-violet stripe on outer face and purple teeth at 65. subbracteatum apex Capitula 40–45 mm in diameter; ligules with a grey stripe on outer face and yellow teeth at apex 85. cambricum 131. Lateral lobes of leaf 2–3 pairs 133. Lateral lobes of leaf (3–)4 pairs 79. beeftinkii Petiole wine red 132. Petiole whitish or tinted purplish Outer involucral bracts 4–5 × 2.0–2.5 mm 76. hygrophilum

34. Taraxacum 132. 133. 133. 134. 134. 135. 135. 136.

136. 137. 137. 138. 138.

139. 139. 140. 140. 141. 141. 142.

142. 143. 143. 144. 144. 145. 145. 146.

146.

78. akteum Outer involucral bracts 7–10 × 2.5–3.5 mm 134. Outer involucral bracts pruinose 135. Outer involucral bracts green Ligules with greyish-violet stripe on outer face; styles 64. britannicum blackish Ligules with brown or puce stripe on outer face; styles 81. nordstedtii discoloured 136. Teeth of inner flowers blackish or purplish 137. Teeth of inner flowers yellow Ligules with greyish-violet stripe on outer face and teeth of inner flowers of capitulum blackish 67. duplidentifrons Ligules with dark greyish-violet stripe on outer face 69. celticum with purple teeth at apex Distal margin of lateral lobes not toothed or with a 60. gelertii single tooth Distal margin of lateral lobes with filiform or 138. triangular teeth Capitulum 30–40 mm in diameter; ligules with purple 66. oellgaardii stripe on outer face Capitulum 40–45 mm in diameter; ligule with a greyish-violet stripe on outer face and teeth of inner 67. duplidentifrons flowers of capitulum blackish 140. Pollen absent 143. Pollen present Outer involucral bracts more or less erect 174. exsertiforme 141. Outer involucral bracts recurved Outer involucral bracts 10–12 × 2–3 mm; styles 141. cherwellense discoloured Outer involucral bracts 12–16 × 3.5–4.5 mm; styles 142. yellow when fresh Interlobe area near the midrib of leaves unblotched; outer involucral bracts with a pale margin 130. subhuelphersianum Interlobe area near the midrib of leaves blotched; outer 148. speciosum involucral bracts without a pale margin When fresh underside of petioles white to green with 144. little or no trace of red When fresh at least the underside of the petioles of the 201. inner leaves pink to purple 145. Styles yellow or yellowish 148. Styles discoloured Ligules with olive-brown or greyish-brown stripe on 146. outer face Ligules with greyish-violet or greyish-purple stripe on 147. outer face Distal margin of lateral lobes of leaves often narrowing sharply from a very broad base and forming a conspicuous and characteristic hump; leaves crispate 142. porrigens Distal margin of lateral lobes of leaves straight to 147. chloroticum convex and dentate; leaves flat

147. Leaves flat with 5–6 pairs of lateral lobes and a fairly 146. undulatiflorum long petiole 147. Leaves somewhat crispate with 5–8 pairs of lateral 160. aequisectum lobes almost to the base 148. Upper surface of midrib darker than petioles and pink 149. or purplish

131

156. 148. Upper surface of midrib not darker than petiole 149. Leaves flat; terminal lobe large, subacute to obtuse; 157. ancistrolobum lateral lobes 2–3(–4) pairs 150. 149. Not as above 150. Some outer involucral bracts more than 5 mm in width; midrib discoloured, often bordered by black lines 158. sellandii 151. 150. Outer involucral bracts less than 5 mm wide 151. Outer involucral bracts more or less violet on inner 152. face 155. 151. Outer involucral bracts pale green on inner face 152. Outer involucral bracts leadon-violet on inner face 149. piceatum 153. 152. Outer involucral bracts clear violet on inner face 153. Leaves obviously blotched in the interlobe area near 150. cyanolepis the midrib; achene body olive-brown 153. Leaves not obviously blotched in the interlobe area 154. near the midrib; achene body straw-coloured 154. Outer involucral bracts slightly recurved with erect 151. curtifrons tips, without a pale margin 154. Outer involucral bracts more or less horizontally spreading, with a more or less pale margin 152. acutifrons 155. Lateral lobes of leaves 6–8 pairs; capitulum 35–40 mm 153. chrysophaenum in diameter 155. Lateral lobes of leaves 3–5 pairs; capitulum 40–50 mm 226. sublongisquameum in diameter 156. Plant small, with fleshy, greyish-green leaves, petiole 145. obtusilobum and midrib 157. 156. Not as above 158. 157. Petioles unwinged in basal half 169. 157. Petioles winged basally, sometimes narrowly so 158. Outer involucral bracts erect to appressed or erect to 159. spreading 161. 158. Outer involucral bracts spreading to recurved 159. Leaves flat; ligules with a brownish-purple stripe on 114. pannucium outer face; achene body about 2.8 mm 159. leaves more or less crispate; ligules with a greyish-violet stripe on outer face; achene body 160. 3.0–3.5 mm 160. Lateral lobes of leaves patent to slightly recurved; 115. subexpallidum outer involucral bracts spreading 160. Lateral lobes of leaves patent or forward pointing; outer involucral bracts erect to appressed 118. tenebricans 162. 161. Outer involucral bracts recurved 163. 161. Outer involucral bracts spreading to recurved 162. Ligules with an olive-brown stripe on outer face 140. stenacrum 162. Ligules with a greyish-violet stripe on outer face 146. undulatiflorum 163. Outer involucral bracts spreading, with a pale margin, 164. 4–5 mm wide 163. Outer involucral bracts often recurved, without a pale 165. margin, usually less than 4 mm wide. 164. Outer involucral bracts 9–11 mm; achene body 132. margettsii straw-coloured 164. Outer involucral bracts 11–15 mm; achene body olive159. altissimum brown

132

1 4 8 . A S T E R AC E A E

165. Outer involucral bracts pruinose or lead-coloured on 166. inner face 167. 165. Outer involucral bracts green on inner face 166. Leaves more or less flat, interlobes near the midrib 134. lepidum dentate or with small lobules 166. Leaves more or less crispate, interlobes near the midrib 155. intumescens often with a single large tooth 167. Ligules with an olive-brown stripe on outer face 140. stenacrum 168. 167. Ligules with a greyish-violet stripe on outer face 168. Interlobe area near the midrib of the leaves not 115. subexpallidum obviously blotched 168. Interlobe area near the midrib of the leaves with dark 156. angulare blotches 169. Petiole wings parallel and narrow; outer involucral 170. bracts recurved or reflexed but spreading at apex 169. Petiole wings not usually parallel; outer involucral 171. bracts not as above 170. Outer involucral bracts glaucous and pink-tipped on 120. alatum inner face 170. Outer involucral bracts pale green on inner face 122. densilobum 171. Outer involucral bracts usually less than 3 mm in 172. maximum width 171. Outer involucral bracts usually more than 3 mm in 179. maximum width 172. Outer involucral bracts suffused purple, narrow and 123. insigne spoke-like 173. 172. Not as above 173. Lateral lobes of leaves with patent, often with double 140. stenacrum linear processes 174. 173. Not as above 127. nigridentatum 174. Tips of ligules dark 175. 174. Tips of ligules orange-red or yellow 138. croceiflorum 175. Tips of ligules orange-red 176. 175. Tips of ligules yellow 176. Leaves with 5–7 pairs of lateral lobes; outer involucral 129. pallescens bracts spreading, 3.5–4.5 mm wide 176. Leaves with 3–5 pairs of lateral lobes; outer involucral bracts more or less recurved, 2.0–3.5 mm 177. wide 177. Outer involucral bracts pruinose on inner face 134. lepidum 178. 177. Outer involucral bracts not pruinose on inner face 178. Lateral leaf-lobes with an elongated tip; outer involucral bracts pale green lightly suffused violet on 135. expallidiforme inner face 178. Lateral leaf-lobes without an elongated tip; outer involucral bracts often suffused reddish-purple 136. subcyanolepis 180. 179. Inner face of outer involucral bracts whitish 179. Inner face of outer involucral bracts pale green or 183. darker 180. Terminal lobe of leaf abruptly acuminate at apex 144. leucopodum 181. 180. Not as above 181. Outer involucral bracts pale green on inner face 131. necessarium 182. 181. Outer involucral bracts greyish-white on inner face

182. Leaves greyish-green, with 3–7 pairs of lateral lobes; interlobes near the midrib sometimes blotched; petiole green and more or less broadly 121. horridifrons winged 182. Leaves rather pale green, with 3–5 pairs of lateral lobes; interlobe area near the midrib unblotched; 137. pallidipes petiole whitish and narrowly winged 184. 183. Leaves more or less strongly crispate 188. 183. Leaves not strongly crispate 185. 184. Outer involucral bracts spreading to recurved 186. 184. Outer involucral bracts erect to spreading 185. Lateral lobes of leaves narrowing from a broad base 116. corynodes 185. Lateral lobes of early (outer) leaves falcate and semilunar and those of later leaves broad, highly 193. lunare dentate 117. undulatum 186. Lateral lobes of leaves recurved 186. Lateral lobes of leaves spreading or pointing 187. forwards 187. Interlobe area near the midrib entire or dentate, more 113. macrolobum or less dark-margined 187. Interlobe area near the midrib usually bearing single 119. dilaceratum lobules, not dark-margined 188. Terminal lobes of outer leaves with a narrow elongated 189. apex 188. Terminal lobes of outer leaves sometimes acute but not 193. elongated at apex 189. Ligules with a brownish-purple stripe on outer face 114. pannucium 190. 189. Ligules with a greyish-violet stripe on outer face 190. Outer involucral bracts about 18 × 4.3 mm, reflexed 183. procerisquameum 191. 190. Outer involucral bracts 10–14 × 3.0–4.5 mm 191. Interlobe area near the midrib not obviously 115. subexpallidum dark-blotched or margined 191. Interlobe area near the midrib dark blotched or 192. margined 192. Interlobe area near the midrib dark margined; lateral lobes of leaves spreading or pointing forward; outer 113. macrolobum involucral bracts erect or spreading 192. Interlobe area of leaves near the midrib dark blotched; lateral lobes of leaves spreading or recurved; outer 124. laciniosum involucral bracts spreading to recurved 193. Lobes of leaves crowded and more or less overlapping 194. 195. 193. Lobes of leaves not crowded 128. laticordatum 194. Achene body 3.5–4.0 mm 131. necessarium 194. Achene body 2.5–3.0 mm 195. Plant very large, with the rather regular lateral lobes of 143. broddesonii leaves with large teeth 196. 195. Not as above 196. Outer involucral bracts 8–11 mm, spreading to suberect 197. 196. Outer involucral bracts 11–16 mm, spreading to 198. recurved 197. Outer involucral bracts dark green, suffused purplish 112. laeticolor on outer face; achene body 3.8–4.0 mm 197. Outer involucral bracts dark, shiny green on outer face; 133. sublaeticolor achene body 3.0–3.2 mm

34. Taraxacum 198. Interlobe area of the leaf near the midrib blotched 126. pannulatiforme 198. Interlobe area of the leaf near the midrib unblotched 199. 193. lunare 199. Outer involucral bracts recurved 200. 199. Outer involucral bracts more or less spreading 200. Lateral lobes of leaves spreading or pointing forwards 119. dilaceratum 125. atonolobum 200. Lateral lobes of leaves recurved 201. Under side of petioles of outer leaves green to white, 202. those of inner leaves pink 201. Under side of petioles of all leaves uniformly pink, red 229. or purple 203. 202. Styles yellow 204. 202. Styles discoloured 203. Interlobe area of leaves near the midrib not obviously blotched; ligules with an orange-brown stripe on outer 142. porrigens face 203. Interlobe area of leaves near the midrib somewhat blotched; ligules with a greyish-violet stripe on outer 171. edmondsonianum face 205. 204. Outer involucral bracts reflexed or recurved 208. 204. Outer involucral bracts recurved, spreading or erect 205. Flowering stems heavily arachnoid-hairy throughout 164. stereodes when young 206. 205. Flowering stem more or less glabrous 206. Petioles unwinged, or, if winged with petiole colour 180. lingulatum not clearly demarcated 206. Petioles pinkish or purplish, clearly demarcated from 207. the parallel-sided green wings 207. Interlobe area of leaves near the midrib unblotched, the 165. adiantifrons lateral lobes patent to recurved 207. Interlobe area of leaves near the midrib somewhat blotched, the lateral lobes patent 171. edmondsonianum 208. Petioles pink and clearly demarcated from the 209. parallel-sided green wings 208. Petioles pink but unwinged or with midrib indistinct 214. 201. diastematicum 209. Outer involucral bracts spreading 210. 209. Outer involucral bracts recurved 210. Leaves plicate, their lateral lobes 4–8 pairs, crowded 217. planum and recurved to falcate 211. 210. Leaves not so 211. Outer involucral bracts without a pale margin 185. remanentilobum 212. 211. Outer involucral bracts with a pale margin 170. latens 212. Ligule teeth reddish-purple 213. 212. Ligule teeth yellow 167. semiglobosum 213. Leaves obviously and densely hairy 202. tanyphyllum 213. Leaves more or less glabrous 214. Some leaves at least with the terminal lobe longer than 215. the pair of lateral lobes below it 214. Terminal lobe not larger than the pair of lateral lobes 221. below it, except sometimes on inner leaves 216. 215. Leaves flat 217. 215. Leaves more or less crispate 216. Leaves with 4–7 pairs of lateral lobes; outer involucral bracts spreading; ligules with greyish-violet stripe on 129. pallescens outer face

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216. Leaves with 3–4 pairs of lateral lobes; outer involucral bracts recurved; ligules with violet-brown stripe on 187. sagittipotens outer face 218. 217. Outer involucral bracts spreading 219. 217. Outer involucral bracts recurved 149. piceatum 218. Leaves dark green 190. ekmanii 218. Leaves pale lettuce green 219. Ligule with a violet-brown stripe on outer face 190. ekmanii 219. Ligule with a greyish-violet or greyish-purple stripe on 220. outer face 220. Capitulum about 50 mm in diameter; ligules with 181. macranthoides greyish-purple stripe on outer face 220. Capitulum 40–45 mm in diameter; ligules with 210. fagerstroemii greyish-violet stripe on outer face 221. Interlobe area of the leaves near the midrib dark 222. blotched 221. Interlobe area of the leaves near the midrib not dark 224. blotched 222. Leaves very pale green when fresh; capitulum 50–65 mm in diameter; achene body about 3.8–4.0 mm 225. maculatum 222. Leaves medium to dark green; capitulum 40–50 mm in 223. diameter; achene body 2.5–3.0 mm 223. Lateral lobes of leaves triangular to lingulate, acute to 126. pannulatiforme obtuse at apex 223. Lateral lobes of leaves deltoid, rather short, tapering to 209. huelphersianum a very acute apex 224. Some lateral lobes of the leaves with long linear elongations; open capitulum exceeding 50 mm in 192. aurosulum diameter 225. 224. Not as above 225. 225. 226. 226. 227.

227.

228. 228. 229.

229. 230. 230. 231. 231. 232.

217. planum Outer involucral bracts somewhat twisted 226. Outer involucral bracts not twisted 186. cordatum Lateral lobes of leaves never divided 227. Lateral lobes of leaves sometimes divided Outer involucral bracts more or less suffused purple; ligules with greyish-brown stripe on outer face; achene 139. lacerifolium body greyish-brown Outer involucral bracts pale green, sometimes pruinose on inner face; ligules with a greyish-violet stripe on 228. outer face; achene body straw-coloured Flowering stem heavily arachnoid-hairy throughout 164. stereodes 210. fagerstroemii Flowering stem glabrous Outer involucral bracts claw-shaped, pinkish or purplish and interlobe area near the midrib blotched 230. dark on young leaves 231. Not as above Plant heterophyllous; outer involucral bracts 2–3 mm 218. polyodon wide Plant not heterophyllous; outer involucral bracts 229. acutifidum 2.5–3.5 mm wide Terminal lobe of leaves large and rounded or 232. helmet-shaped on all or most leaves Terminal lobe of leaves large and rounded only on 234. inner leaves or not at all Outer involucral bracts 11–14 mm, pruinose, pale 195. aberrans green on inner face

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232. Outer involucral bracts 8–11 mm, often suffused purple 233. 233. Leaves often pale green with midrib clear reddish-purple; achene body bright golden brown 188. hexhamense 233. Leaves dull, dark green with a pinkish midrib; achene 198. cophocentrum body straw-coloured 235. 234. Styles yellow or yellowish 249. 234. Styles discoloured 235. Interlobe area of leaves near the midrib strongly dark 236. blotched 235. Interlobe area of leaves near the midrib not dark 243. blotched or scarcely so 237. 236. lateral leaf-lobes 5 or more pairs 240. 236. Lateral leaf-lobes 2 to 5 pairs 237. Outer involucral bracts suffused with purple 220. nitidum 237. Outer involucral bracts not suffused with purple 238. though sometimes pink-tipped 219. multicolorans 238. Outer involucral bracts spreading 239. 238. Outer involucral bracts recurved or reflexed 239. Leaves with 5–8 pairs of lateral lobes; outer involucral 160. aequisectum bracts 2.0–3.5 mm wide 239. Leaves with 4–6 pairs of lateral lobes; outer involucral 171. edmondsonianum bracts 4–5 mm wide 241. 240. Outer involucral bracts 4–5 mm wide 242. 240. Outer involucral bracts 3–4 mm wide 241. Leaves yellowish-green; flowering stems glabrous or nearly so; outer involucral bracts with a narrow, pale 171. edmondsonianum margin 241. Leaves dark to bluish-green; flowering stems thickly 224. scotiniforme arachnoid-hairy when young 242. Plant heterophyllous; petioles narrowly winged; 222. xanthostigma achene body straw-coloured 242. Plant not heterophyllous; petioles unwinged; achene 223. longisquameum body olive-brown 243. Terminal lobes of most leaves with an elongated apex 244. 243. Terminal lobes of the leaves lacking an elongated apex 246. 244. Leaves pale green and rather flaccid, with 4–8 pairs of 173. exsertum lateral lobes 244. Leaves medium to dark green, with 3–5 pairs of lateral 245. lobes 245. Outer involucral bracts 3.0–4.5 mm wide, pale green 166. retroflexum on inner face 245. Outer involucral bracts 2.0–3.5 mm wide, suffused 178. leptodon purple on inner face 247. 246. Outer involucral bracts with a pale margin 248. 246. Outer involucral bracts without a pale margin 247. Outer involucral bracts strongly reflexed, pale 171. edmondsonianum bluish-green on inner face 247. Outer involucral bracts spreading to recurved, suffused 189. amplum violet on inner face 248. Petioles pink; ligules with a greyish-violet stripe on 194. coartatum outer face 248. Petioles rose-purple; ligules with a brownish-purple 211. hepaticum stripe on outer face 249. Some outer involucral bracts not exceeding 3 mm in 250. maximum width 261. 249. Outer involucral bracts exceeding 3 mm in width

250. Petioles usually vivid purple; outer involucral bracts 251. recurved or hanging down 250. Petioles not vivid purple; outer involucral bracts erect 254. to recurved 252. 251. Lateral lobes of leaves 4–6 pairs 253. 251. Lateral lobes of leaves 6–10 pairs 252. Leaves hairy; petiole unwinged; outer involucral bracts 205. dahlstedtii recurved 252. Leaves glabrous; petioles narrowly winged; outer 207. pachylobum involucral bracts spreading 253. Leaves dark green, the midrib pinkish-red to the tip; 214. pectinatiforme petiole often poorly differentiated 253. Leaves medium green, the midrib green to faintly pink; 215. caloschistum petioles deep reddish-purple 254. Terminal lobe of leaf tripartite; petiole short and very 216. trilobatum broadly winged 255. 254. Not as above 256. 255. Outer involucral bracts twisted 257. 255. Outer involucral bracts not twisted 256. Lateral lobes of leaves 5–9 pairs; petiole often winged 168. aequilobum 256. Lateral lobes of leaves 4–7 pairs; petioles unwinged 169. latissimum 257. Ligules with a greyish-brown stripe on outer face and 127. nigridentatum dark teeth 257. Ligules with greyish-violet or greyish-purple stripe on 258. outer face and yellow teeth 259. 258. Outer involucral bracts spreading 260. 258. Outer involucral bracts recurved 259. Outer involucral bracts green and more or less pruinose 162. angustisquameum on inner face 259. Outer involucral bracts suffused with purple on inner 218. polyodon face 197. oblongatum 260. Outer involucral bracts 9–12 mm 229. acutifidum 260. Outer involucral bracts 11–14 mm 261. Petiole shining purple; outer involucral bracts broad 261. and spreading 261. Petiole rarely shining purple, but, if so, then the outer 264. involucral bracts not broad 262. Terminal lobe with a narrow, acuminate apex 213. pulchrifolium 263. 262. Terminal lobe with a much broader apex 263. Lateral lobes of leaves 3–5 pairs; terminal lobe tending 177. obtusifrons to be longer than pair of lobes below it 263. Lateral lobes of leaves 4–7 pairs; terminal lobe smaller 212. subundulatum than pair of lobes below it 264. Terminal leaf-lobe very acute or acuminate at apex or 265. elongated 264. Terminal leaf-lobe obtuse to acute at apex but not 271. ending in a fine point 265. Leaves flat, hairy at least when young; flowering stems 266. arachnoid-hairy at least when young 265. Leaves often more or less crispate; leaves and 267. flowering stems more or less glabrous 266. Outer involucral bracts recurved, 12–14 × 4.0–5.5 mm 172. acroglossum 266. Outer involucral bracts mostly spreading (to recurved), 175. exacutum 14 × 4.0–4.5 mm

34. Taraxacum 268. 267. Outer involucral bracts with a pale margin 269. 267. Outer involucral bracts without a pale margin 268. Lateral lobes of leaves 5–6 pairs; ligules with 170. latens reddish-purple teeth 268. Lateral lobes of leaves 3–5 pairs; ligules with yellow 196. pseudoretroflexum teeth 217. planum 269. Outer involucral bracts somewhat twisted 270. 269. Outer involucral bracts not twisted 161. interveniens 270. Leaves hairy 163. mimulum 270. Leaves glabrous 272. 271. Lateral lobes of leaves 2–3 pairs 274. 271. Lateral lobes of leaves usually 4 or more pairs 272. Leaves flat; outer involucral bracts not pruinose, 176. valens without a pale margin 272. Leaves crispate; outer involucral bracts pruinose, with 273. a pale margin 273. Outer involucral bracts 10–15 × 3.5–4.0 mm, 231. lucidum spreading to erect 273. Outer involucral bracts 14–16 × 4–6 mm, spreading to 232. sundbergii recurved 275. 274. Outer involucral bracts twisted 277. 274. Outer involucral bracts not twisted 275. Outer involucral bracts pale green on inner face 217. planum 276. 275. Outer involucral bracts pruinose on inner face 276. Leaves narrowly oblong-oblanceolate in outline, the 168. aequilobum interlobe area near the midrib dentate 276. Leaves broadly oblanceolate in outline, the interlobe area near the midrib often with one large tooth 169. latissimum 277. Petiole narrowly parallel-winged with a clearly 278. demarcated red midrib 280. 277. Not as above 278. Outer involucral bracts strongly reflexed, pale bluish-green on inner face, with a narrow pale margin 171. edmondsonianum 278. Outer involucral bracts spreading to recurved, brownish-purple to violet-coloured on inner face 279. without a pale margin 279. Capitulum 35–45 mm in diameter; outer involucral 200. dilatatum bracts brownish-purple on inner face 279. Capitulum 45–50 mm in diameter; outer involucral bracts violet-coloured on 228. subxanthostigma inner face 280. Interlobe area near the midrib beneath the leaves dark 281. blotched 280. Interlobe area near the midrib beneath the leaves 290. unblotched or weakly dark coloured 281. Lateral lobes of leaves strongly dentate, divided almost 221. incisum to midrib 281. Lateral lobes of leaves sometimes strongly dentate but 282. not divided almost to midrib 282. Leaves greyish-green with scattered punctate spots 230. melanthoides 283. 282. Not as above 283. Leaves smudged blackish on upper surface, lateral 199. pachymerum lobes more or less overlapping 284. 283. Not as above

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284. Distal margin of lateral lobes of leaves with many dark 127. nigridentatum teeth 285. 284. Not as above 286. 285. Midribs of leaves pink to purple 287. 285. Midribs of leaves green to slightly pinkish 286. Distal margin of lateral lobes of leaves strongly dentate 220. nitidum 286. Distal margin of lateral lobes of leaves entire 228. subxanthostigma 288. 287. Outer involucral bracts with inner face green 287. Outer involucral bracts with inner face suffused purple 289. 288. Outer involucral bracts spreading to recurved, 3–4 mm 154. tumentilobum wide, without a pale margin 288. Outer involucral bracts strongly reflexed, 4–5 mm wide, with a narrow pale margin 171. edmondsonianum 289. Leaves with 5–6 pairs of lateral lobes; outer involucral 218. polyodon bracts 11 × 2–3 mm, claw-like 289. Leaves with 3–5 pairs lateral lobes; outer involucral 227. fasciatum bracts 12–15 × 3.5–4.5 mm 290. Midribs golden brown on upper surface, especially on 291. centre of leaf on fresh specimens 292. 290. Midribs not golden brown on upper surface of leaf 291. Leaves with 5–8 pairs of crowded lateral lobes; outer involucral bracts 15–17 × 3.5–5.0 mm, green on outer 179. pannulatum face 291. Leaves with 3–6 pairs of lateral lobes; outer involucral bracts 9–13 × 2–4 mm, suffused with violet on outer 191. ochrochlorum face 292. Petioles very short, more or less obsolete, broadly 293. winged 292. Petioles not very short or, if short, then not broadly 294. winged 293. Leaves with 5–8 pairs of lateral lobes; outer involucral 179. pannulatum bracts 15–17 × 3.5–5.0 mm 293. Leaves with 3–5 pairs of lateral lobes; outer involucral 204. laciniosifrons bracts 10–15 × 3.0–3.5 mm 294. Leaves bright green, shiny, crispate, with lateral lobes 203. sinuatum pointing in all directions 295. 294. Leaves not as above 296. 295. Leaves with lateral lobes somewhat divided 297. 295. Leaves not as above 296. Flowering stem heavily arachnoid-hairy throughout 164. stereodes 296. Flowering stem more or less glabrous 210. fagerstroemii 297. Apex of ligules of inner flowers blackish 208. latisectum 298. 297. Apex of ligules of inner flowers reddish or yellow 298. Apex of ligules of inner flowers reddish 206. obliquilobum 299. 298. Apex of ligules of inner flowers yellow 299. Flowering stems arachnoid-hairy, at least when young 300. 302. 299. Flowering stems more or less glabrous 300. Involucre strongly pruinose in bud 185. remanentilobum 301. 300. Involucre green in bud 164. stereodes 301. Leaves crisped, lateral lobes 4–8 pairs 176. valens 301. Leaves flat; lateral lobes 3–5 pairs 303. 302. Outer involucral bracts less than 14 mm 307. 302. Outer involucral bracts more than 14 mm

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303. Outer involucral bracts pruinose on inner face 182. rhamphodes 304. 303. Outer involucral bracts not pruinose 304. Leaves with 3–5 pairs of lateral lobes; outer involucral 196. pseudoretroflexum bracts strongly reflexed 304. Leaves with 5–7 pairs of lateral lobes; outer involucral 305. bracts recurved 170. latens 305. Teeth of ligules reddish-purple 306. 305. Teeth of ligules yellow 184. vastisectum 306. Lateral lobes crowded in centre of leaf 306. Lateral lobes not crowded in centre of leaf 210. fagerstroemii 307. Outer involucral bracts with a pale margin 179. pannulatum 308. 307. Outer involucral bracts without a pale margin 308. Outer involucral bracts strongly reflexed 171. edmondsonianum 309. 308. Outer involucral bracts recurved 170. latens 309. Teeth of ligules reddish-purple 210. fagerstroemii 309. Teeth of ligules yellow

Section 1. Erythrosperma (H. Lindb.) Dahlst. Taraxacum taxon Erythrosperma H. Lindb.; Taraxacum subsection Erythrosperma (H. Lindb.) Schischk.; Taraxacum section Dissimilia Dahlst.; Taraxacum subsection Dissimilia (Dahlst.) R. Doll; Taraxacum section Fulva M. P. Christ.; Taraxacum subsection Proxima R. Doll. Small to medium-sized perennial herbs. Leaves variously coloured, deeply divided, usually almost to midrib, rarely with more than 6 pairs of lateral lobes. Capitulum rarely more than 30 mm in diameter. Outer involucral bracts 3– 8(–9) mm, spreading or erect, often more or less corniculate at apex. Inner involucral bracts corniculate at apex, most easily seen when in bud. Achenes with body reddish to yellowish-brown, narrow, with fine spiniform projections at apex; cone up to 1.3 mm, narrow. Usually obligate agamospermic, sometimes facultatively agamospermic or sexual. This section seems to be a natural one, and may have arisen spontaneously from the Asian and Mediterranean Section Erythrocarpa Hand.-Mazz. They are among the shortest-lived Taraxaca and in cultivation may flower within three months of germination, and they may be biennial in some localities. Dry places in rocky and sandy ground and short grassland, usually with a shallow soil and south-facing and unshaded. Throughout Great Britain and Ireland. Throughout Europe except for montane regions and the Arctic; Turkey, Iran and Afghanistan; introduced in North America and probably elsewhere. 1. T. lacistophyllum (Dahlst.) Raunk. Cut-leaved Dandelion T. erythrospermum subsp. lacistophyllum Dahlst. Small to medium-sized perennial herb with a stock-like rhizome. Leaves usually numerous, spreading; lamina 2– 8(–20) × 1–2(–3) cm, pale to dark but more often medium dull green on upper surface, slightly to deeply purplish on

both surfaces of the midrib, oblanceolate in outline, divided nearly or quite to the midrib; terminal lobe linear or narrowly triangular-subsagittate, acute at apex, usually entire; lateral lobes 3–6(–12), spreading to sloping down, often continuing as teeth onto the petiole, linear, often narrowly, sometimes narrowly triangular, subacute at apex, entire or with 1– few, narrow teeth, characteristically abruptly widening to form a triangular base; interlobe area near the midrib long and narrow, entire or with 1–few, linear teeth; glabrous or slightly arachnoid-hairy along the midrib; petiole 10–60 mm, usually brownish-purple at least above, unwinged. Flowering stems few to numerous, 5–15(–20) cm, erect, pale green, often with a flush of purplish-brown, rarely densely so, glabrous or sometimes with arachnoidhair just below the capitulum. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 5–7 × 1.5–2.0 mm, bluish-green on outer face, greyish-green, usually with a waxy covering, sometimes with a trace of purple on inner face, with a narrow but distinct pale margin, linearlanceolate, with an obtuse but jagged, somewhat corniculate apex, usually spreading, sometimes arcuate-recurved or erect. Inner involucral bracts 10–12 × 1.5–2.0 mm, dull medium green, with a narrow, usually indistinct paler margin and reddish near the apex, linear-lanceolate, with a jagged, obtuse, corniculate apex, erect. Ligules pale to deep lemon yellow with a sharply defined, dirty, greyish-purple stripe on the outer face and yellow apical lobes. Styles exserted, discoloured. Pollen little to copious. Achenes with body 2.5–3.0 mm, dark brownish-red to purple, oblanceolate, with numerous spiniform projections at the apex; cone 0.8–1.0 mm, dark brownish-red; beak 5–7 mm, very pale brown; pappus 4–7 mm, dirty white. Flowers 4–5. Obligate agamosperm. 2n = 24. Easily recognised by the abruptly expanded bases to the leaf-lobes, waxy outer involucral bracts, usually pollen rich stigmas and brownish-red achenes. Native. Dry places such as calcareous downs, sandy heaths, cliff-tops, walls, paths, waste ground and almost anywhere where the vegetation is not too dense. Widespread and locally common in southern England, becoming rarer in Scotland; coastal in Ireland; Channel Islands. Western Europe, Czechoslovakia, southern Scandinavia and the Baltic. 2. T. inopinatum C. C. Haw. Unexpected Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves usually numerous, spreading to erect; lamina 1.3–20.0 × 2–6 cm, medium to dark olive green, usually with scattered, faint, purple spots on upper surface, green on midrib, lanceolate or oblong-lanceolate in outline, divided almost to the midrib; terminal lobe medium-sized, more or less trilobate with a drawn-out, acute or subacute apex, entire; lateral lobes 4–6 pairs, sloping down, usually continuing as teeth on the petiole, triangular or deltoid, sometimes narrowing rather abruptly to the narrow apex which can be pointing in various directions, entire or with an occasional narrow tooth; interlobe area near the midrib short and narrow, usually with narrow teeth of various lengths; glabrous or with a few hairs; petioles up to one-quarter of the leaf length, usually green but occasionally slightly

34. Taraxacum dull purple, slender, unwinged. Flowering stems 15–20 cm, green or purple, ascending, slightly arachnoid-hairy below the capitulum. Capitulum 25–35 mm in diameter, opening. Outer involucral bracts 7–8 × 2–3 mm, pale glaucous and pruinose on the inner face, very dark glaucous and very pruinose on the outer face, and usually with a narrow, white margin, linear-lanceolate, acute at apex, scarcely corniculate, more or less erect. Inner involucral bracts 11– 12 × 1.2–1.5 mm, medium green, linear-lanceolate, slightly corniculate at apex. Ligules yellow, with a purple stripe on outer face. Styles greyish-yellow. Pollen absent. Achenes with body 3.2–3.4 mm, dark orange-brown or red, with fine and sharp spiniform projections in upper part; cone 0.6–0.9 mm, cylindrical; beak 7–8 mm, pappus 6–7 mm. Flowers 4–5. This species is unique among British species belonging to the section in having spotted leaves. Native. Bare limestone and sand and rocks especially in disused quarries. Western Britain from Cornwall to Cumberland. Endemic. 3. T. brachyglossum (Dahlst.) Raunk. Purple-bracted Dandelion T. erythrospermum subsp. brachyglossum Dahlst.; T. vachelliae Dahlst.; T. hispanicum auct. Small to medium-sized perennial herb with a stock-like rhizome. Leaves usually numerous, spreading; lamina 2–10 (–20) × 1–2(–3.5) cm, pale to dark, usually dull green, sometimes slightly glaucous on upper surface, often purplish on the midrib, oblong in outline, divided nearly or quite to the midrib; terminal lobe linear or narrowly triangular, usually entire; lateral lobes 4–6(–8) pairs, spreading to sloping down, usually continuing as teeth on to the petiole, usually linear-lanceolate, sometimes linear but not abruptly widening at the base, obtuse to acute at apex, entire or with an occasional tooth; interlobe area near the midrib more or less wide, usually with small, unequal teeth; glabrous or with scattered short hairs; petiole 10–60(–80) mm, usually purplish, sometimes intensely so, unwinged. Flowering stems few to numerous, 3–15(–20) cm, prostrate to erect, pale green, often suffused purplish especially towards the base, glabrous or with arachnoid hairs. Capitulum 20– 30 mm in diameter, opening. Outer involucral bracts (5–) 6–8 ×(1.5–)2.0–2.5 mm, dark glaucous-green, often suffused reddish or purplish and more or less waxy coated on both faces, lanceolate to ovate, obtuse but jagged at apex, corniculate, spreading or ascending, rarely reflexed. Inner involucral bracts 12–15 × 1.2–1.5 mm, medium green with a narrow, pale margin, not waxy and slightly purplish at apex, linear-lanceolate, corniculate at apex. Ligules very short, deep lemon yellow, with a sharply defined greyishpurple stripe on the outer face. Styles exserted, discoloured. Pollen little to copious. Achene with body 2.8–3.2 mm, reddish-brown, oblanceolate, with numerous very narrow, sharp spiniform projections at the apex; cone 0.4–0.7 mm, reddish-brown; beak 6–7 mm, very pale brown; pappus 6– 7 mm, whitish. Flowers 4–5. Sexual, facultative agamosperm or obligate agamosperm. 2n = 16, 17, 18, 20, 22, 24, 26.

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Often grows with T. lacistophyllum from which it is distinguished by its purplish outer involucral bracts and short ligules with a purple stripe. It sometimes grows in more acidic situations than that species. Native. Dry places on calcareous downs, sandy heaths, sand-dunes, walls and waste places. Widespread and locally frequent throughout Great Britain; coastal in Ireland; Channel Islands. Central Europe extending westwards to France and Great Britain and northwards to southern Scandinavia. 4. T. scanicum Dahlst. Sk˚ane Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous; lamina 7–18 × 2.5–6.0 cm, medium green on upper surface, brownish-purple on midrib, linear-oblanceolate to oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe narrowly triangular-hastate, the apex typically narrow and elongate, with a subobtuse tip, often with 2 pairs of spreading lobes at its base; lateral lobes 4–6 pairs, long and narrow with a broad base, acute at apex, finely dentate on both margins or only on the one towards the distal part of the leaf, spreading or sloping down; interlobe area near the midrib short to medium, narrow, often dentate; glabrous or nearly so; petiole fairly long, purplish, unwinged. Flowering stem 4–14 cm, pale green, often suffused purplish, somewhat arachnoid-hairy at apex. Capitulum 35–40 mm in diameter, opening. Outer involucral bracts 7–8 × 0.8–0.2 mm, dark green on both faces, with a pale margin, linear lanceolate, obtuse at apex, not corniculate. Inner involucral bracts 12– 15 × 1.2–1.5 mm, dark green, oblong-lanceolate, obtuse, purplish and corniculate at apex. Ligules rather dark yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, chestnut to dark brick red, with few, very narrow, sharp spinulose projections at apex; cone 0.8–1.0 mm; beak 8–9 mm; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. 2n = 25. Native. Grassy places in the Breckland of East Anglia and the Channel Islands; by a runway on sand-dunes and on a gravelly roadside in Shetland where it may be introduced. Widespread throughout much of Europe. 5. T. argutum Dahlst. Sharp-toothed Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves rather few, erect; lamina 3–10 × 1–2 cm, dark green and the interlobe area near the midrib sometimes blotched on the upper surface, green to faintly purplish on the midrib, oblong-oblanceolate in outline, divided almost to the midrib; terminal lobe triangular with a somewhat elongated apex, entire; lateral lobes 4–5 pairs, regular, deltoid, acute at apex, almost entire, spreading; interlobe area near the midrib short and narrow to fairly wide, with occasional subulate teeth; glabrous or nearly so; petiole short, white to green or dull purple, scarcely winged. Flowering stems 10–15 cm, pale green, arachnoid-hairy below the capitulum. Capitulum 15–20 mm in diameter, closed. Outer involucral bracts 3–6 × 1.5–2.0 mm, dark green, without or with an indistinct pale margin on outer face, ovate, obtuse at apex and scarcely corniculate, spreading. Inner

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involucral bracts 11–13 × 1.5–1.8 mm, dark green with a narrow pale, margin, oblong-lanceolate, obtuse at apex and corniculate. Ligules deep orange-yellow, involute, with a reddish-orange outer face. Styles exserted, yellow. Pollen absent or infrequent. Achenes with body 3.0–3.5 mm, deep red, with sharp, spiniform projections at apex; cone 0.5–0.8 mm, cylindrical; beak 5–7 mm; pappus 5–6 mm, whitish. Flowers 5–6. Seed production poor. Obligate agamosperm. 2n = 24. This is the only British or Irish species of the section with the capitulum habitually closed and the ligules involute. Native. Found especially in dry meadows in limestone areas and on sandy roadsides. Scattered over Great Britain where it is more common in the west with a single locality in the north of Ireland. Recently recorded from central Bohemia. 6. T. arenastrum A. J. Richards Sand Dandelion Small, delicate perennial herb with a stock-like rhizome surrounded by the remnants of old leaf bases. Leaves fairly numerous, more or less prostrate; lamina 5–8 × 1–3 cm, medium green on upper surface, green to faintly purplish on the midrib, divided two-thirds or more of the way to the midrib; terminal lobe triangular, with a narrow, attenuate apex or at least markedly mucronate, entire; lateral lobes 4– 5 pairs, often contracted from a broad base to a very narrow, downwards pointing, acute tip, scarcely dentate, strongly recurved; interlobe area near the midrib, short and thick, entire or rarely with a tooth; more or less glabrous; petiole short or up to one-quarter the length of the lamina, green or somewhat purple, unwinged. Flowering stems 5–8 cm, dull green or copper-coloured, arachnoid-hairy. Capitulum 20–25 mm in diameter, opening. Outer involucral bracts 6–9 × 2–3 mm, green, with well-marked white margins on outer face, lanceolate, obtuse at apex and scarcely corniculate, erect. Inner involucral bracts 9–14 × 1.0–1.5 mm, medium green, oblong-lanceolate, slightly corniculate at apex. Ligules pale yellow, with a violet stripe on the outer face. Styles exserted, yellow or somewhat discoloured. Pollen present or absent. Achenes with body 2.8–3.0 mm, chestnut or dark red, with short spiniform projections above, but otherwise smooth; cone 0.7–0.8 mm, abruptly demarcated, narrow; beak 6–7 mm; pappus 6–7 mm, white. Flowers 4–5. 2n = 24. Characterised by its chestnut or dark red achenes, erect outer involucral bracts with a white margin and yellow or somewhat discoloured styles. Native. Sand-dunes and chalk and limestone pasture, mainly by the sea. Rare to local in coastal localities throughout England, Wales and southern Scotland with a single locality in the west of Ireland. Holland and probably northern France. 7. T. commixtum G. E. Haglund Sociable Dandelion T. commutatum Dahlst., non Jord. Medium-sized perennial herb with a stock-like rhizome. Leaves rather few, erect; lamina 7–20 × 2.5–3.5 cm, dull green on upper surface, green on the midrib, elliptical in outline, divided to the midrib; terminal lobe triangular, the apex and basal corners elongated into long linear points,

entire; lateral lobes 4–6 pairs, with a triangular base and long linear apex the tip of which is often swollen or bifurcate, often, with narrow teeth, pointing forwards, spreading or sloping down; interlobe area near the midrib obvious and narrow, often with narrow teeth; glabrous or nearly so; petiole rather long, purple, narrow and unwinged. Flowering stem 15–25 cm, ascending to erect, rather stout, often purplish, arachnoid-hairy. Capitulum 25–40 mm in diameter, opening. Outer involucral bracts 8–10 × 1.5–5.0 mm, dark green on outer face, with a more or less pale margin, ovate, obtuse and sometimes corniculate at apex, erect to spreading. Inner involucral bracts 15–20 × 1.0– 1.5 mm, dark green, with a narrow, pale margin, obtuse and slightly corniculate at apex. Ligules deep yellow, with a reddish-purple stripe on outer surface. Styles exserted, yellow. Pollen present. Achenes with body 2.5–3.5 mm, dark red, with spiniform projections at apex; cone 0.8–1.0 mm; beak 6–7 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5. Easily recognised by its distinctive leaves, erect, dark green outer involucral bracts, reddish-purple ligule stripes, yellow styles and large achenes. Dutch material is said to differ from the Swedish type and the British plants may be like the Dutch. Native. Dunes and sandy heaths. East Anglia, a few localities in southern England, Cheshire, Flintshire and Northumberland. Scandinavia, France, Holland and Germany. 8. T. rubicundum (Dahlst.) Dahlst. Ruddy Dandelion T. erythrospermum subsp. rubicundum Dahlst.; T. rubicundum subsp. monspeliense Dahlst.; T. rubicundum subsp. pulvigerum H. Lindb.; T. monspeliense (Dahlst.) Dahlst. Small, delicate perennial herb with a stock-like rhizome, with persistent dark leaf-bases. Leaves fairly numerous, erect or spreading; lamina 2–8 × 0.7–1.7 cm, dull green on upper surface, reddish on the midrib, oblanceolate in outline, divided to the midrib; terminal lobe triangular, the apex and basal corners elongated into narrowly linear points; lateral lobes 4–7 pairs, triangular at base and elongated into linear apices, sometimes toothed, spreading or curved forwards; interlobe area near the midrib narrow, toothed with narrow, unequal teeth; glabrous or nearly so; petioles short to long, dark violet-purplish, thin, wiry, glabrous. Flowering stem 4–12 cm, ascending or erect, often purplish, glabrous. Capitulum 15–20 mm in diameter, opening. Outer involucral bracts 2.5–5.0 × 0.5–1.5 mm, dark glaucous-green, often suffused with purple on outer face and with a more or less pale border, similar on inner face, ovate or ovate-lanceolate, obtuse and conspicuously corniculate at apex, erect. Inner involucral bracts 10–15 × 1.0–1.2 mm, dark green with a pale margin, oblong-lanceolate, purplish, obtuse and corniculate at apex. Ligules pale yellow, with a dark violet stripe on outer face and blackish apical lobes. Styles exserted, discoloured. Pollen usually absent. Achenes with body 2.5–2.8 mm, dark violet, with long, narrow, very sharp, spiniform projections at apex, cone 0.8–1.0 mm; beak 6–8 mm, pale; pappus 4–5 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24.

34. Taraxacum Easily recognised by its completely glabrous flowering stem, ovate or ovate-lanceolate, erect outer involucral bracts and dark violet achenes. Native. Dry places, but mostly restricted to calcareous downland and locally on sandy heaths and dunes. Throughout Great Britain and in the Channel Islands, common in southern England, rarer northwards. Much of Europe, except the east. 9. T. cenabense Sahlin Cenabum Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, erect or spreading; lamina 9–12 × 1.5–2.2 cm, greyish-green on upper surface, green on the midrib, divided to the midrib; terminal lobe mediumsized, triangular or 3-lobed, sagittate at base, obtuse at apex; lateral lobes 3–6 pairs, falcate or in outer leaves broader and deltoid, more or less curved down, entire or with an occasional tooth, narrowed to an obtuse apex; interlobe area near the midrib rather short, with 1–2 teeth; glabrous or nearly so; petioles short to medium, pale green, narrow and almost unwinged. Flowering stems 5.0–13.5 cm, pale green, glabrous. Capitulum 20–25 mm in diameter, opening. Outer involucral bracts 6–8 × 1.8–2.0 mm, dark green on both faces, with a conspicuous white margin, lanceolate, with a recurved, corniculate apex, suberect. Inner involucral bracts 12–15 × 1.5–2.0 mm, dark green, linearlanceolate, corniculate at apex. Ligules golden yellow, with a dark violet stripe on the outer face, with a purple apex to the lobes of the inner. Styles greenish. Pollen absent. Achenes with body 4.1–4.5 mm, reddish, with short, spiniform projections; cone 0.7–0.9 mm; beak about 7 mm; pappus about 8 mm, whitish. Flowers 4–5. Intermediate between T. rubicundum and T. parnassicum, differing from the former by its larger, reddish achenes, pale petioles and darker flower colour, and from the latter by the glabrous scape, pale petioles and purple ligule teeth. Native. In Britain known only from Bignor Hill in Sussex. Described from between Satillieu and La Louvescon in Ard´eche and from Sully-sur-Loire in Loiret, France. 10. T. parnassicum Dahlst. Parnassus Dandelion T. silesiacum Dahlst. ex G. E. Haglund Small perennial herb with a stock-like rhizome. Leaves numerous, spreading or erect; lamina 3–10 × 0.5–3.0 cm, clear green on upper surface, green to faintly reddish on the midrib, regularly divided almost to the midrib; terminal lobe triangular, with a short, subacute tip, or the apex and basal corners elongated into linear tips, sometimes toothed; lateral lobes 4–5 pairs, crowded, narrowly triangular, subobtuse at apex, the margin towards the distal area of the leaf convex and sometimes dentate, usually slightly sloping down; interlobe area near midrib short and narrow, frequently with a single tooth; glabrous or nearly so; petioles short, purple, winged. Flowering stems 3–12(–20) cm, often purplish, arachnoid-hairy when young. Capitulum 20– 30 mm in diameter, opening. Outer involucral bracts 5– 6 × 1.5–2.0 mm, green on both faces, with an indistinct pale margin, triangular-lanceolate, obtuse and somewhat corniculate at apex, erect to spreading. Inner involucral bracts 10–13 × 1.5–2.0 mm, dark green, oblong-lanceolate, obtuse

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and somewhat corniculate at apex. Ligules yellow, with a purple stripe on the outer face and yellow apical lobes. Styles exserted, discoloured. Pollen more or less absent. Achenes with body 2.7–3.2 mm, dark red, with sparse spiniform projections at apex; cone 0.7–1.0 mm; beak 7–8 mm; pappus 6–9 mm, whitish. Flowers 4–5. Obligate agamosperm. 2n = 24. Native. Dry calcareous grassland. In scattered localities through England and Wales, especially in the west; in Galway in Ireland. Widespread in Europe. 11. T. dunense Soest Dune Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 4–15(–19) × 2–5 cm, dark green, often suffused purplish on the upper surface, green to purplish on midrib, oblong in outline, divided to midrib; terminal lobe linear, with linear side projections from the base, sometimes with a solitary tooth; lateral lobes 5–10 pairs, long and narrowly linear, but thinner in the middle than at base or apex, with occasional small teeth; interlobe area near midrib very narrow, with unequal, very narrow teeth; glabrous or nearly so; petioles fairly short, dark violet-purple, very narrow and unwinged. Flowering stems 5–15 cm, often suffused purplish, decumbent to erect, arachnoid-hairy. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 5–8 × 2.0–2.5 mm, dark green on both faces, with an obscure, pale, sometimes purplish, margin, ovate, obtuse, sometimes purplish and corniculate at apex, spreading. Inner involucral bracts 16–20 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse and corniculate at apex. Ligules deep yellow, with a purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achene with body 3.5–4.0 mm, dark violet to red, with long, thin spiniform projections at apex; cone 1.2–2.0 mm; beak 9–11 mm, pale; pappus 5–7 mm, whitish. Flowers 4–5. 2n = 24. A very distinct species with linear leaf-lobes, narrow midribs and large violet to red achenes with long cones. Early leaves can be entire or scarcely lobed and can be very misleading. Native. Sand-dunes. Coasts of southern England and Wales, north to Anglesey and Northumberland. Belgium, Holland and Sweden. 12. T. haworthianum

Dudman & A. J. Richards Haworth’s Dandelion T. laetum auct.; T. laetiforme auct.; T. discretum auct.; T. sublaetum auct. Small perennial herb with a stock-like rhizome. Leaves fairly numerous, spreading; lamina 5–10 × 0.5–3.0 cm, medium green on upper surface, green to faintly purplish on midrib, divided nearly to midrib; terminal lobe triangularsubsagittate, with an acute, elongated apex, entire; lateral lobes 6–8 pairs, regular, triangular, subacute at apex, sometimes sinuous on the margin towards the distal area of the leaf, entire, sloping down; interlobe area near midrib short and narrow, usually entire; glabrous or nearly so; petioles short to fairly long, green or purple at base, narrow and unwinged. Flowering stems 5–12 cm, purplish,

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somewhat arachnoid-hairy at apex. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 4–6 × 1.8–2 mm, green, with a white or rose, somewhat scarious margin on outer face, ovate, obtuse and markedly purplishcorniculate at apex, erect to appressed. Inner involucral bracts 12–15 × 1.0–1.5 mm, dark green with a narrow pale margin, obtuse, purplish and corniculate at apex. Ligules rather pale yellow, with a violet or grey stripe on the outer surface, the apical lobes purple. Styles dirty yellow. Pollen present. Achenes with body 2.8–3.0 mm, dark violet-purple, with sharp spiniform projections at apex; cone 0.8–1.0 mm; beak 6–7 mm, pale; pappus 5–7 mm, whitish. Flowers 4–5. 2n = 24. Recognised by the combination of violet achene bodies, dirty yellow styles and erect to appressed outer involucral bracts. It is difficult to separate from T. platyglossum without achenes. Native. Sand-dunes and dry grassland. Scattered throughout Great Britain and Ireland, but mostly coastal and commonest in the north and west. Endemic. Named after C. C. Haworth (1934–89). 13. T. gotlandicum (Dahlst.) Dahlst. Gotland Dandelion T. erythrospermum subsp. gotlandicum Dahlst. Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading or erect; lamina 3.5– 10 × 1.0–3.0 cm, medium green on upper surface, green on midrib, oblong in outline, divided almost to midrib; terminal lobe triangular, with obtuse, elongated, apical and basal points, entire; lateral lobes 4–6 pairs, narrowly triangular, mostly obtuse at apex, usually subentire, the margin towards the distal area of the leaf concave, mostly spreading or slightly sloping down; interlobe area near the midrib narrow to rather wide, with unequal narrow teeth; glabrous or almost so; petioles short to medium, usually green, winged or unwinged. Flowering stems many, 4–18 cm, decumbent to erect, pale green, arachnoid-hairy. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 6.5–7.0 × 2.0–2.5 mm, glaucous, suffused violet and with a pale margin on outer face, ovate, obtuse and slightly corniculate at apex, erect. Inner involucral bracts 15–16 × 2.5– 3.0 mm, dark green with a narrow pale margin, oblonglanceolate, abruptly narrowed to a linear, obtuse, corniculate apex. Ligules pale yellow, with a violet stripe on the outer face. Styles exserted, yellow. Pollen present. Achenes with body 3.0–3.5 mm, brick-red, with long, spiniform projections at apex; cone 0.8–1.0 mm; beak 5–6 mm, pale; pappus 5–6 mm, whitish. Flowers 5–6. 2n = 24. Native. Calcareous grassland and tracks. Forfarshire, Banffshire and Caithness in Scotland and possibly Co. Clare ¨ in Ireland, not seen recently. Gotland and Oland in Sweden, Estonia and Oslofjord in Norway. This species appears to have a relict distribution and fresh material may indicate that our plant is a distinct species. 14. T. proximum (Dahlst.) Raunk. Umber-fruited Dandelion T. erythrospermum subsp. proximum Dahlst. Small to medium-sized, heterophyllous perennial herb with a stock-like rhizome. Leaves few to numerous, spreading

or prostrate; lamina 5–15(–20) × 1.0–3.0 cm, dull green on upper surface, green to tinted purplish on midrib, oblong to oblong-oblanceolate in outline, divided almost to the midrib; terminal lobe triangular-subsagittate, with an acute, sometimes drawn out apex, entire; lateral lobes 5–8 pairs, spreading or sloping down, narrowly triangularlinear-lanceolate, acute at apex, regularly dentate; interlobe area near the midrib, short, narrow to fairly wide, dentate; glabrous or with an occasional hair; petioles rather short, dull purple, unwinged. Flowering stems 5–20 cm, green, sometimes slightly tinted purplish, erect, arachnoidhairy above when young. Capitulum 20–35 mm in diameter, opening. Outer involucral bracts 6–8 × 1.5–2.0 mm, spreading to recurved, green or sometimes rather strongly purplish-green on both faces, with a narrow pale border, pruinose especially on the inner face, ovate or ovate-lanceolate, acute at apex, slightly corniculate. Inner involucral bracts 15–17 × 1.5–1.7 mm, green with a narrow pale margin, oblong-lanceolate, reddish at the obtuse, corniculate apex. Ligules deep yellow, with a purple stripe on the outer face. Styles exserted, discoloured. Pollen absent. Achenes with body 2.8–3.0 mm, dark brown to purplish-brown, oblanceolate, with numerous, very narrow, sharp spiniform projections at the apex and sometimes to the middle; cone 0.7– 0.9 mm; beak 7.0–8.5 mm, pale brown; pappus 4–7 mm, whitish. Flowers 4–7. Agamosperm. 2n = 24. Native. Dry grassland, meadows, heaths and dunes. Local in England and Wales except for central and south-central England, rare and mainly coastal in Scotland. Throughout much of Europe except the south. 15. T. proximiforme Soest ex Lambinon & Soest Straw-fruited Dandelion Small to medium-sized, heterophyllous perennial herb with a stock-like rhizome. Leaves rather numerous, erect or spreading; lamina 5–25 × 1–4 cm, bright green on upper surface, green to faintly purplish on midrib, varying from entire, through dentate, to partly lobed, to divided almost to midrib; terminal lobe triangular-subsagittate or with the apex and basal corners elongate-linear and acute; lateral lobes 3–6 pairs, narrowly to broadly triangular, the apex sometimes elongate-linear and very acute, sometimes subacute to obtuse with a broad base, the upper sometimes dentate on the margin towards the proximal area of the leaf, spreading or slightly sloping down; interlobe area near the midrib usually with narrow teeth; glabrous or nearly so; petioles short to medium, dull purple, winged. Flowering stems 8–25 cm, erect, sometimes purplish, arachnoid-hairy. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 6–8 × 1.5–2.0 mm, green with a pale margin not obvious on either face, ovate or ovate-lanceolate, obtuse and scarcely corniculate at apex, spreading. Inner involucral bracts 15–17 × 1.5–1.7 mm, with a narrow, pale margin, oblong lanceolate, obtuse and slightly corniculate at apex. Ligules medium yellow, with a greyish-violet stripe on the outer face and purple apical lobes. Styles exserted, yellowish. Pollen present or absent. Achenes with body 2.8– 3.0 mm, straw brown, with spiniform projections at apex; cone 0.8–1.0 mm; beak 7–8 mm; pappus 4–7 mm, whitish. Flowers 4–5. Obligate agamosperm. 2n = 24.

34. Taraxacum Differs from T. proximum by its straw brown achenes, yellowish styles and purple ligule lobes. Native. Dry mesic grassland. A few scattered localities in southern and northern England, East Anglia and southern and eastern Scotland. Coasts of northern France, Belgium and Holland. It may not be conspecific with the Continental material. 16. T. pseudoproximum Soest Essex Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 15– 20 × 1.0–3.5 cm, bright green on upper surface, green to faintly purplish on midrib, elliptical or oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe triangular-subsagittate, acute at apex, usually entire; lateral lobes 3–5 pairs, triangular, more or less acute at apex, sometimes dentate on the margin towards the proximal area of the leaf, spreading or slightly sloping down; interlobe area near the midrib short and entire or with an occasional tooth; glabrous or nearly so; petioles short, green and unwinged. Flowering stems 15–25 cm, erect, arachnoid-hairy. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 6–8 × 3–4 mm, dark green suffused reddish-purple on both faces with a pale margin, ovate-lanceolate, obtuse and corniculate at apex, spreading. Inner involucral bracts 15–17 × 1.5–1.7 mm, green with a narrow, pale margin, oblong-lanceolate, reddish at the obtuse, corniculate apex. Ligules pale yellow, with a greyish-violet stripe on outer face. Styles exserted, yellow or discoloured. Pollen present. Achenes with body 1.8–3.0 mm, pale greyish-brown, with spiniform projections at apex; cone 0.7–0.9 mm; beak 7.0– 8.5 mm, pale; pappus 4–7 mm, whitish. Flowers 4–7. Native or possibly introduced. Known only from a few records from Essex. Originally described from Holland. 17. T. disseminatum G. E. Haglund Golden-fruited Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves rather few, erect or spreading; lamina 5–15 × 1.5–2.0 cm, pale green, green to faintly violet on midrib, divided to midrib; terminal lobe triangularsubsagittate, acute at apex, entire; lateral lobes 2–4 pairs, triangular, acute at apex, with several and sometimes large teeth on the margin towards the distal area of the leaf; interlobe area near the midrib narrow, sometimes with unequal teeth; glabrous or nearly so; petioles short to medium, violet, narrow and unwinged. Flowering stems 10–20 cm, green, rarely with violet tints, arachnoid-hairy below the capitulum. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 5–6 × 1.5–2.5 mm, green, sometimes suffused violet on both faces, with a pale margin, ovate-lanceolate, acuminate and somewhat corniculate at apex, recurved. Inner involucral bracts 11–12 × 1.5–2.0 mm, dark green with a pale margin, oblong, obtuse, sometimes violet and corniculate at apex. Ligules rather pale yellow, with a greyish-violet stripe on outer face. Styles blackish, exserted. Pollen present. Achenes with body 3.0– 3.2 mm, golden brown, subcylindrical with long, spiniform projections at apex; cone 0.7–0.8 mm; beak 7–8 mm;

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pappus 7–8 mm, whitish. Flowers 4–5. Obligate agamosperm. 2n = 24. Probably introduced. Short, dry grassland and lawns. A few scattered localities in the southern half of England and Wales. Western and central Europe and Scandinavia. 18. T. oxoniense Dahlst. Oxford Dandelion T. helvicarpum Dahlst. Small to medium-sized perennial herb with a stock-like rhizome. Leaves few to numerous, prostrate, spreading or erect; lamina 3–15(–25) × 1.0–3.0 cm, pure green on upper surface, green on midrib, oblong-oblanceolate in outline, varying from entire through dentate to shallowly lobed to deeply lobed, entire or with an occasional tooth; terminal lobe often long and subdivided, triangular, obtuse at apex; lateral lobes when leaf is deeply divided 3–7 pairs, triangular at base, with long-linear apex; interlobe area near the midrib narrow, with unequal, narrow teeth; glabrous or nearly so; petioles short to medium, bright purple, narrow and unwinged. Flowering stems 5–28 cm, purplish, decumbent to erect, slightly arachnoid-hairy under the capitulum. Capitulum 15–30 mm in diameter, opening. Outer involucral bracts 3–9 × 2–4 mm, dark green with a white margin on both faces, slightly pruinose on inner face, ovate, obtuse and scarcely corniculate at apex, ascending to erect. Inner involucral bracts 14–17 × 2.0–2.5 mm, oblong-lanceolate, dark green, with a narrow pale margin, obtuse and corniculate at apex. Ligules medium yellow, with a greyishviolet stripe on the outer face and yellow apical lobes. Styles exserted, slightly discoloured. Pollen present. Achenes with body 2.8–3.0 mm, cinnamon, with short, sharp spiniform projections at apex; cone 0.8–1.0 mm; beak 4.5– 5.0 mm, pale; pappus 5–7 mm, whitish. Flowers 3–6. Obligate agamosperm. 2n = 32. This species, the commonest in the section in England and Wales, despite its very variable dissected leaves is characterised by the contrasting bright green lamina and petiole and the erect, dark green, white-margined, ovate outer involucral bracts. Native. Dry, neutral or calcareous ground, especially downland, cliff-tops and sand-dunes and occasionally walls and waste places. Common throughout England and Wales, mainly coastal in Scotland and Ireland; Channel Islands. Western and central Europe. 19. T. fulviforme Dahlst. Green-bracted Dandelion T. simile auct. Small to medium-sized perennial herb with a stock-like rhizome. Leaves usually numerous, spreading; lamina 3– 10 × 1.0–3.0(–4.0) cm, dull green on upper surface, green to faintly tinted reddish on midrib, the interlobes near the midrib often blotched dark brownish-purple, oblongoblanceolate in outline, divided nearly to the midrib; terminal lobe well developed and larger than most in this section, triangular-subsagittate, with a more or less acuteapiculate apex, usually entire but sometimes with small teeth in larger plants; lateral lobes 3–6 pairs, often continuing as teeth on the petiole, spreading to somewhat recurved, narrowly triangular, long-acute at apex, with 1 or 2 spinulose teeth on either or both margins; interlobe area near the

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midrib narrow to rather wide, entire or with spinulose teeth; glabrous or almost so; petiole 10–30 mm, often reddish, without or with a narrow wing. Flowering stems usually few, 2–14 cm, erect or ascending, pale green, occasionally tinted brownish-purple, glabrous or slightly hairy at apex. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 5–7 × 2–3 mm, erect to spreading giving a starshaped involucre, green on both faces with a very narrow margin, scarcely pruinose, linear-lanceolate, more or less acute at apex, scarcely corniculate. Inner involucral bracts 12–16 × 1.5–2.0 mm, green with a narrow scarious border, sometimes tipped reddish, oblong-lanceolate, obtuse at apex, erect. Ligules yellow, with a sharply defined greyishviolet stripe on the outer face, the apical lobes tipped reddish. Styles discoloured. Pollen absent. Achenes with body 2.5–3.0 mm, cinnamon, oblong, with numerous spiniform projections at apex; cone 0.7–0.8 mm, cinnamon; beak 4– 5 mm, pale; pappus 6–7 mm, dirty white. Flowers 4–7. Obligate agamosperm. 2n = 32. The only cinnamon-fruited species in Great Britain and Ireland without pollen, which has green, erect to spreading outer involucral bracts. Native. Dry places, especially sand-dunes, cliff-tops and calcareous grassland, but also in paths and lawns. Scattered over Great Britain and locally frequent but absent from the Scottish Highlands; coastal in Ireland; in the Channel Islands. Also elsewhere in western Europe. 20. T. fulvum Raunk. Cinnamon-fruited Dandelion T. brachycranum (Dahlst.) A. W. Hill; T. fulvum subsp. brachycranum Dahlst. Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 3– 18 × 2.0–3.0 cm, dull green on upper surface, green to faintly purple on midrib, more or less oblong in outline, divided almost to midrib; terminal lobe triangularsubsagittate, acute at apex, entire; lateral lobes 4–5 pairs, triangular, acute at apex, toothed, spreading or slightly sloping down; interlobe area near the midrib narrow, unequally toothed; glabrous or nearly so; petioles usually short, purple, unwinged near the base. Flowering stem 6–15 cm, often suffused purplish, arachnoid-hairy above. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 6–7 × 1.5–2.0 mm, dark green on both faces with a pale margin, lanceolate, obtuse at apex. Inner involucral bracts 15–16 × 2.0–3.5 mm, dark green with a paler margin, oblong-lanceolate, abruptly narrowed to a long, narrow, obtuse, slightly corniculate apex. Ligules medium yellow, with a greyish-violet stripe on outer face and the apical lobes tipped reddish. Styles exserted, yellowish. Pollen absent. Achenes with body 2.8–3.0 mm, cinnamon, with sharp rather stout, spiniform projections at apex; cone 0.7–1.0 mm; beak 8–10 mm, pale; pappus 5–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 32. Native. On light, well-drained neutral to calcareous soils, often in species-rich grasslands, also on walls and cliffs. Scattered throughout England and Wales, mainly coastal in Scotland and in a few localities in Ireland. Western and central Europe and Scandinavia.

21. T. scoticum A. J. Richards Scottish Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 3– 18 × 2.0–3.0 cm, dull green on upper surface, green to faintly purple on midrib, more or less oblong in outline, divided almost to midrib; terminal lobe triangularsubsagittate, acute at apex, entire; lateral lobes 4–5 pairs, triangular, acute at apex, toothed, spreading or slightly sloping down; interlobe area near the midrib narrow, unequally toothed; glabrous or nearly so; petioles usually short, purple, unwinged near the base. Flowering stems 6–15 cm, often suffused purplish, arachnoid hairy above. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 5–6 × 2–3 mm, dark green on both faces with a pale margin, ovate, obtuse at apex. Inner involucral bracts 15– 16 × 2.0–3.5 mm, dark green with a paler margin, oblonglanceolate, abruptly narrowed to a long, narrow, obtuse, slightly corniculate apex. Ligules medium yellow, with a greyish-violet stripe on outer face. Styles exserted, yellowish. Pollen present. Achenes with body 3.2–3.5 mm, cinnamon, with sharp rather stout, spiniform projections at apex; cone 0.7–1.0 mm; beak 8–10 mm, pale; pappus 5–7 mm, whitish. Flowers 4–6. Native. Sandy places. Scattered records in the west and north of Great Britain from Somerset to Ross-shire. 22. T. falcatum Brenner Narrow-lobed Dandelion T. pectinosum G. E. Haglund; T. canulum G. E. Haglund Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 5– 15 × 5–7 cm, greyish-green on upper surface, the midrib green or faintly purple, oblong in outline, divided nearly to midrib; terminal lobe narrowly triangular with an elongated, linear, acute apex and basal corners; lateral lobes 6–8 pairs, linear, acute at apex, pointing forwards, spreading or sloping down; interlobe area near the midrib rather long with unequal teeth; glabrous or nearly so; petioles short or medium, purple, narrow and unwinged. Flowering stems 4–10 cm, green, ascending, arachnoid-hairy. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 5–7 × 1.8–2.0 mm, green on both faces with a slight, pale margin, broadly ovate-lanceolate, obtuse and somewhat corniculate at apex, spreading. Inner involucral bracts 12–15 × 1.5–2.0 mm, greyish-olive, oblong-lanceolate, narrowed but obtuse, dark purple and corniculate at apex. Ligules medium yellow, with a greyish-violet stripe on the outer face and yellow apical lobes. Styles discoloured to yellowish. Pollen present, if rudimentary. Achenes with body 2.7–3.5 mm, warm straw brown, with spiniform projections at apex; cone 1 mm; beak 9–10 mm, pale; pappus 9–10 mm, whitish. Flowers 4–5. 2n = 24. Probably introduced. An inconspicuous plant in dry grassland. A few, scattered records in England and Wales. Fennoscandia, Iceland, Switzerland and Belgium. 23. T. retzii Soest De Retz’s Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading; lamina 2–10(–25) × 1.0–4.0 cm, dull medium green on upper surface, green to purplish

34. Taraxacum on midrib, oblong in outline, divided nearly to the midrib; terminal lobe small, triangular-sagittate, subacute or with a narrow pointed apex, entire; lateral lobes 4–7 pairs, sloping down, narrowly linear, curved upwards, obtuse to acuteapiculate at apex, with long, spinulose teeth often on both margins, widened at base; interlobe area near the midrib narrow, with spinulose teeth; glabrous or with an occasional hair; petioles short, often dull reddish, unwinged or narrowly winged. Flowering stems 10–30 cm, greenish or suffused brownish-purple, usually ascending, often arachnoid-hairy above. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 6–8 × 1.5–2.0 mm, recurved, green on both faces with a narrow, pale margin, lanceolate, subobtuse, acute or acuminate at apex, not corniculate. Inner involucral bracts 14–16 × 1.5–2.0 mm, dull green with a narrow pale margin, oblong-lanceolate, obtuse at apex, erect. Ligules yellow, with a greyish-violet stripe on outer face, with pink apical lobes. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–4.0 mm, cinnamon, with numerous, short projections at apex; cone 1.0–1.2 mm; beak 6–7 mm, pale; pappus 8–11 mm, whitish. Flowers 4–6. Native. Dry sandy heaths and dunes. Very local in East Anglia and south-east England, north Wales and the Channel Islands. France. Named after Bernard Guy Gaston de Retz (b. 1910). 24. T. glauciniforme Dahlst. Many-toothed Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading; lamina 3–15(–20) × 1.0–3.0 cm, greyish-green on upper surface, green to suffused purplish on midrib, narrowly oblong in outline, divided nearly to midrib; terminal lobe triangular-sagittate, pointed at apex, entire; lateral lobes 6–12 pairs, spreading, linear to narrowly lanceolate, pointed at apex, slightly curved, entire or slightly toothed; interlobe area near the midrib fairly wide, with narrow teeth; glabrous or with a few hairs; petioles fairly short, usually green, sometimes tinted red, winged. Flowering stems 15–25 cm, green or suffused purplish, somewhat arachnoid-hairy at the apex. Capitulum 15–25 mm in diameter, opening. Outer involucral bracts 5.0–6.5 × 1.5 mm, spreading to recurved, greyish-green, sometimes suffused purple and somewhat pruinose on both faces, with a narrow pale margin, lanceolate or ovate, acute at apex, somewhat corniculate. Inner involucral bracts 8– 16 × 1.5–2.0 mm, dark green with a pale margin and slightly pruinose, lanceolate-oblong, purplish at the jagged, obtuse, corniculate apex, erect. Ligules rather deep dirty yellow, with a pale grey stripe on the outer face and purple apical lobes. Styles greenish to yellow. Pollen absent or rare. Achenes with body 2.5–2.8 mm, cinnamon to chestnut-puce, oblanceolate, with sharp spiniform projections at apex; cone 0.8–1.0 mm; beak 5.5–6.5 mm, white; pappus 4.0–4.5 mm, whitish. Flowers 4–6. Native. Grassland, waste places, walls, cliff-tops, heaths, gardens and lawns on light, sandy, well-drained, neutral to calcareous soils. Widespread and locally common in England and Wales, though mainly in the south; also in the Channel Islands, Isle of Man and Ayrshire. It also occurs in Belgium, France and Holland.

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25. T. wallonicum Soest Belgium Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves few to numerous, ascending or spreading; lamina 8–15 × 1.5–2.0 cm, medium green on upper surface, green or slightly tinted reddish on midrib, more or less oblong or oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe triangular-subsagittate, with an acute, somewhat elongated apex, entire; lateral lobes 4–8 pairs, triangular with a broad base, obtuse to acute at apex, dentate only on the margin towards the distal area of the leaf of the proximal lobes; interlobe area near the midrib short, often with 1–2 teeth; glabrous or nearly so; petioles green, short, narrow and unwinged. Flowering stem 8–20 cm, green or slightly tinted reddish, arachnoid-hairy under the capitulum. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 5–8 × 1.8–2.0 mm, rather pale green, flushed purple on both faces, with a narrow pale margin, lanceolate, obtuse and corniculate at apex, spreading or recurved. Inner involucral bracts 11–12 × 1.5–2.0 mm, dark green with a broad, pale margin, linear-lanceolate, broad below and narrowed to a purple, corniculate apex. Ligules rather pale yellow, with a greyish-purple stripe on the outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw-coloured, with minute spiniform projections at apex; cone 0.8–1.0 mm; beak 7–8 mm, pale; pappus 5–6 mm, whitish. Flowers 4–5. Possibly introduced. In a few places in London, Kent and Essex. Belgium. 26. T. degelii G. E. Hagland Degelius’s Dandelion Small to medium perennial herb with a stock-like rhizome. Leaves few to numerous, ascending or spreading; lamina 3–10(–15) × 2.0–3.5(–5.0) cm, medium green on upper surface, green to faintly tinted reddish on midrib, oblongoblanceolate in outline, divided nearly to base; terminal lobe triangular-subsagittate, obtuse, acute or acuminate at apex, entire; lateral lobes 5–8 pairs, spreading, sloping down or reflexed, triangular-based with a linear, acute apex, entire or with teeth at the base in larger plants; interlobe area near the midrib narrow to fairly wide, often with spinulose teeth; glabrous or nearly so; petiole medium, pinkish, often with spinulose teeth, narrowly winged. Flowering stem 5–15(–20) cm, green, erect, with arachnoid hairs above. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 7–9 × 1.8–2.0 mm, erect or spreading, dark green on both faces, with a conspicuous white or pinkish margin, ovate-lanceolate, subacute and somewhat corniculate at apex. Inner involucral bracts 15–17 × 1.5–2.0 mm, dark green, with a narrow, pale margin, narrowly oblonglanceolate, reddish at the obtuse to subacute, corniculate apex. Ligules rather pale yellow, striped grey on outer face. Styles exserted, discoloured. Pollen present or absent. Achenes with body 2–3 mm, greyish-brown, narrowly oblanceolate, with spiniform projections at apex; cone 1.3 mm; beak 7–8 mm, pale brown; pappus 6–7 mm, whitish. Flowers 5–7, 2n = 24. Native. Rocky places. Inland in mid-Wales, by the sea in north Wales, south-west England, south-west Scotland and western Ireland and one locality in east

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Ireland. Endemic. Named after Gunner Bror Fritiof Degelius (b. 1903). 27. T. acutum A. J. Richards Violet-striped Dandelion Small perennial herb with a stock-like rhizome. Leaves few to numerous, spreading; lamina 3–10 × 0.5–3.0 cm, dull dark green on upper surface, green to purplish on midrib, linear-oblanceolate to oblong-oblanceolate in outline, divided almost to the midrib; terminal lobe triangularsubsagittate or helmet-shaped, more or less acute at apex, entire or with a single tooth; lateral lobes 3–5 pairs, regular, spreading or sloping down, triangular, with an acute apex and a broad base, with one to several teeth mainly on the proximal lobes; interlobe area near the midrib short, narrow to fairly wide, sometimes with teeth; glabrous or with a few hairs; petioles short, green or purple, unwinged. Flowering stems 5–10 cm, green or tinted reddish, arachnoid-hairy above. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 6.0–7.5 × 1.5–2.0 mm, ascending-erect or spreading, dark green on both faces, sometimes with a narrow pale margin, lanceolate, acute at apex, not corniculate. Inner involucral bracts 15–18 × 1.5–1.7 mm, olive green with pale margins, oblong-lanceolate, reddish at the more or less acute apex. Ligules deep yellow, with a dark violet stripe on the outer face, the outer up to twice as long as the involucral bracts. Styles discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with numerous sharp spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, very pale brown; pappus 6–7 mm, dirty white. Flowers 4–6. 2n = 24. Native. Chalk and sandy grassland. In a few localities in south-east England and a solitary locality in Derbyshire. Endemic. 28. T. placidum A. J. Richards Alderney Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 6– 20 × 1–5 cm, pale green on upper surface, purplish on midrib, divided to midrib; terminal lobe triangular, acute at apex, entire; lateral lobes 5–8 pairs, deltoid, more or less acute at apex, entire or denticulate; interlobe area near the midrib usually short and narrow, entire; glabrous or nearly so; petioles long, bright purple, narrowly winged. Flowering stems 10–30 cm, purple, erect, arachnoid-hairy above. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 6–7 × 2.5–3.0 mm, green on both faces, with a white margin, ovate-lanceolate, more or less acute at the corniculate apex, spreading. Inner involucral bracts 15– 18 × 1.5–2.0 mm, dark green, oblong-lanceolate, obtuse at the corniculate apex. Ligules medium yellow, with a silver-grey stripe on outer face. Styles exserted, yellow. Pollen present. Achenes with body 2.8–3.0 mm, greyishbrown, with sharp spiniform projections at apex; cone 0.8 mm; beak 8–9 mm; pappus 8–9 mm, whitish. Flowers 4–5. 2n = 24. Native. Dry, grassy paths near St Annes, Alderney, Channel Islands. Central France and northern Spain.

29. T. tanylepis Dahlst. Orkney Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 7–15 × 2–3 cm, dull green on upper surface, green to somewhat purplish on midrib, oblanceolate to oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe triangular-sagittate, subacute at apex, entire; lateral lobes 4–6 pairs, triangular, sometimes with elongated apex, entire or filiform-dentate, spreading to slightly sloping down; interlobe area near the midrib short and narrow, toothed; glabrous or nearly so; petioles form one-quarter to half the length of the lamina, green or somewhat purplish, narrow, unwinged. Flowering stems 5–17 cm, green or somewhat coloured, ascendingerect. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 8–9 × 2.5–3.0 mm, dark green on both surfaces with a pale margin, linear-lanceolate, more or less acute at apex. Inner involucral bracts 15–18 × 1.5–2.0 mm, dark green, oblong, obtuse and corniculate at apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, yellow. Pollen absent. Achenes with body 3.0– 3.5 mm, straw brown (olive green when dried), with spiniform projections at apex; cone 0.5 mm, or more or less absent; beak 9–10 mm; pappus 9–10 mm, whitish. Flowers 5–6. Native. Stony pastures by the sea. Known only from Orkney. Endemic. 30. T. tortilobum Florstr¨om Twisted-lobed Dandelion Small perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 70–150 × 2–3 cm, medium green with the interlobe area near the midrib frequently blotched on the upper surface, green to faintly purplish on midrib, oblong or oblong-oblanceolate in outline, divided to midrib; terminal lobe flatly triangular, with an acute somewhat elongated apex, entire or slightly dentate; lateral lobes 3–5 pairs, unequally triangular, with a narrow, acute apex, forming a strongly sigmoid distal marginal hump, the proximal lobes dentate, spreading to sloping down and much twisted; interlobe area near the midrib short, frequently dentate; glabrous or nearly so; petioles rather short, purple, narrow and unwinged. Flowering stems 7–15 cm, pale green, often suffused purple, more or less glabrous. Capitulum 20–25 mm in diameter, opening. Outer involucral bracts 5–6 × 2.0–2.5 mm, greyish-green on both faces, often suffused with purple on inner face, pruinose and with a pale margin, lanceolate, more or less acute and strongly purple-corniculate at apex, erect. Inner involucral bracts 14–16 × 1.5–2.0 mm, dark green, oblong-lanceolate, acute and corniculate at apex. Ligules pale yellow, with a greyishpurple stripe on outer face and yellow apical lobes. Styles blackish. Pollen present. Achenes with body 2.5–3.4 mm, pale to straw brown, with short sharp spiniform projections at apex; cone 0.7–0.9 mm; beak 8–9 mm; pappus 8–9 mm, whitish. Flowers 4–5. 2n = 24. Strongly resembling T. lacistophyllum but with much twisted leaf-lobes and pale brown achenes. Native. Coastal and inland grasslands. Sussex, Surrey, Suffolk and Durham. North-western, western and southwestern Europe.

34. Taraxacum 31. T. pseudolacistophyllum Soest Recurved-bracted Dandelion T. affine G. E. Haglund, non Jord. Small perennial herb with a stock-like rhizome. Leaves 7– 20 × 2–4 cm, numerous, erect or spreading; lamina dark green with the interlobe area near the midrib sometimes blotched on the upper surface, green to faintly purplish on midrib, oblong or oblong-oblanceolate in outline, divided to midrib; terminal lobe with a linear, elongated apex and linear basal processes, obtuse to acute at apex, entire or slightly dentate; lateral lobes 3–5 pairs, unequally triangular, with a narrow, acute apex, sparsely subulate-dentate or entire; interlobe area near the midrib short to rather long, narrow and often dentate; glabrous or nearly so; petioles rather short, purple, narrow and unwinged. Flowering stems 7–20 cm, pale green, often suffused purple, slightly arachnoidhairy below the capitulum. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 6–7 × 2.0–2.5 mm, greyish-green on both faces, often suffused with purple on inner face, pruinose and with a conspicuous pale margin, lanceolate, more or less acute and not corniculate at apex, spreading to recurved. Inner involucral bracts 15– 18 × 1.5–2.0 mm, dark green, oblong-lanceolate, acute at apex. Ligules pale yellow, with a greyish-purple stripe on outer face and yellow apical lobes. Styles blackish. Pollen present. Achenes with body 2.5–3.0 mm, pale to straw brown, with spiniform projections at apex; cone 0.6–0.8 mm; beak 8–9 mm; pappus 8–9 mm, whitish. Flowers 4–5. Agamosperm. Probably introduced. A single plant was found in Surrey in 1995. Native of central Europe. Section 2. Obliqua (Dahlst.) Dahlst. Taraxacum taxon Obliqua Dahlst.; Taraxacum subsection Obliqua (Dahlst.) Schischk. Small perennial herbs. Leaves many, bright green, narrowly oblong, deeply divided, with many pairs of lateral lobes. Capitulum flat to almost closed. Outer involucral bracts 5–7 mm, appressed or erect-appressed, with a pale margin, more or less corniculate at apex. Ligules with a red stripe on outer face. Achenes with body greyish-brown; cone less than 0.5 mm. Obligate agamosperms. Open sandy turf by the sea, especially machair. Local and confined to the coasts of Great Britain, Ireland, Belgium, Holland and Scandinavia. Superficially similar to some species of the Section Erythrosperma from which they can only be distinguished with certainty by their achenes. They are perhaps more closely related to the high-alpine Section Dissecta Soest. 32. T. obliquum (Fr.) Dahlst. Many-lobed Dandelion Leontodon obliquum Fr. Small perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 3–8 × 0.4–2.5 cm, pale green on upper surface, with a green midrib, oblong in outline, divided to midrib; terminal lobe triangular or with the obtuse apex and basal corners elongate, entire; lateral lobes 7–10 pairs, triangular, obtuse at apex, entire or

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slightly toothed; interlobe area near the midrib narrow, often toothed; glabrous or nearly so; petioles short, green, narrow and sometimes slightly winged. Flowering stems 2–10 cm, green, slender, glabrous. Capitulum 10–15 mm in diameter, usually closed. Outer involucral bracts 5–6 × 1.8–2.5 mm, greenish-purple with a pale margin on both faces, ovate, obtuse and purplish-corniculate at apex, appressed. Inner involucral bracts 10–12 × 2.0–2.5 mm, olive green, with a narrow, pale margin, oblong-lanceolate, obtuse and corniculate at apex. Ligules orange-yellow, involute or flat, with a red stripe on outer face. Styles more or less exserted, yellowish. Pollen present. Achenes with body 2.8–3.0 mm, greyish-brown, with slender, sharp spiniform projections at apex; cone 0.3–0.4 mm; beak 4–5 mm, pale; pappus 5–6 mm, whitish. Flowers 5. Obligate agamosperm. 2n = 24. Native. Grey dunes and dune slacks. Coasts of northern England, Scotland and Ireland. Scandinavia. 33. T. platyglossum Raunk. Tongue-leaved Dandelion T. obliquum subsp. platyglossum (Raunk.) Nordh. Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 3–12 × 1.0–2.5 cm, medium green to dark green on upper surface, green to faintly purplish on midrib, oblong or elliptical-oblong in outline, divided almost to midrib; terminal lobe narrow with a subacute, elongate apex and the basal corners elongate and narrow, entire; lateral lobes 7–12 pairs, very narrowly triangular, with a long, narrow, acute apex, filiform-dentate on margin towards proximal area of leaf, sloping down; interlobe area near the midrib narrow and short to long, filiform-dentate; glabrous or nearly so; petioles short to medium, green or purplish, more or less winged proximally. Flowering stem 2–10 cm, slender, pale green, often suffused purplish, glabrous. Capitulum 25–30 mm in diameter, usually open, sometimes closed. Outer involucral bracts 6–7 × 1.5–2.0 mm, green on both faces, with a pale margin, lanceolate, with a green or purple corniculation at the acute apex, erect-appressed. Inner involucral bracts 10–12 × 2.0–2.5 mm, dark green, with a narrow, pale margin, oblong-lanceolate, obtuse at the reddish, corniculate apex. Ligules deep yellow, usually flat, but sometimes involute, with a red stripe on the outer face. Styles exserted, discoloured. Pollen present or absent. Achenes with body 3.0–3.5 mm, greyish-brown, with spiniform projections at apex; cone 0.3–0.4 mm; beak 5–7 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Native. Sand-dunes. On the coast of Scotland, Northern England, Isle of Man, Anglesey, Caernarvonshire, Somerset, Channel Islands and Co. Antrim. Scandinavia, north Germany and Baltic Russia. Section 3. Palustria (H. Lindb.) Dahlst. Taraxacum taxon Palustria H. Lindb. Medium-sized, slender perennial herbs. Leaves narrow, not or only shortly lobed. Outer involucral bracts 3–10 mm, with a broad, pale margin, ovate, appressed. Capitulum 20– 40 mm in diameter. Ligules striped greyish-violet to purple. Achenes with body 2.5–4.3 mm, with very few spiniform

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projections at apex; cone up to 1.0 mm. Probably all obligate agamosperm. Local in grassy fens and water meadows. Scattered throughout Europe except for high mountain regions, the Arctic and the Mediterranean islands. A natural section which possibly arose during an interglacial, by hybridisation between the paludal, but halophytic Section Leptocephala Soest and the Section Orientalia Hand.-Mazz.

low, with a reddish to blackish-red stripe on outer face, the apical lobes reddish. Styles exserted, more or less yellow to discoloured. Pollen absent. Achenes with body 4.5–5.0 mm, straw brown, smooth except near the apex which is tuberculate or with small projections; cone 0.8–1.0 mm; beak 6.5–7.5 mm, pale; pappus 5.5–6.5 mm, whitish. Flowers 5. Native. Rough grassland by turloughs subject to flooding. Co. Clare, western Ireland. Endemic. Named after David Allardice Webb (1912–95).

34. T. palustre (Lyons) Symons Fen Dandelion Leontodon palustre Lyons; T. limnanthes subsp. limnanthoides Soest; T. austrinum auct.; T. raii sensu Gray; ?T. pollichii auct.; T. lanceolatum Poiret; T. maritimum Hagendijk, Soest & Zevenbergen; T. westhoffii Hagendijk, Soest & Zevenbergen Small to medium-sized, slender perennial herb with a stocklike rhizome. Leaves few, usually erect, sometimes decumbent; lamina 2–15 × 0.5–2.0 cm, dull green on upper surface, green to faintly purplish on midrib, narrowly elliptical to narrowly oblanceolate, entire, denticulate or shallowly lobed, long attenuate at base; when lobed lateral lobes 2– 3 pairs, very short, triangular and entire; glabrous; petioles usually long, purplish, unwinged or narrowly winged. Flowering stem 4–20 cm, decumbent to erect, often purplish, sometimes sparsely arachnoid-hairy above. Capitulum 20– 40 mm in diameter, opening and flat. Outer involucral bracts imbricate, 6–8 × 3.0–3.5 mm, green or suffused with purple, with a broad, pale margin on outer face, ovate, acute at apex, appressed. Inner involucral bracts 15–22 × 2.5– 3.0 mm, dark green with a narrow, pale margin, oblonglanceolate, purplish at the obtuse, jagged apex. Ligules deep yellow, flat, with a purple stripe on the outer face. Styles exserted, greenish. Pollen absent or rarely present. Achenes with body 3.5–4.0 mm, straw-coloured, oblong, with few to numerous, very short spiniform projections at apex; cone 0.4–0.5 mm; beak 5–7 mm, pale; pappus 5.5–7.0 mm, whitish. Flowers 4–5. Obligate agamosperm. 2n = 32. Native. Hay meadows liable to seasonal flooding, less often in calcareous flushes, sometimes near the sea. Scattered through Great Britain and Ireland and in the Channel Islands. Also in Denmark, Belgium, Holland and France.

ˇ ep´anek 36. T. amarellum J. Kirschner & Stˇ Irish Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves more or less erect; lamina 4–9 × 0.4–1.0 cm, medium to dull green on upper surface, paler beneath, purplish on midrib, linear or narrowly elliptical, obtuse to acute at apex, remotely dentate, gradually narrowed at base, glabrous; petiole long, purple and narrow. Flowering stem 6–20 cm, green to brownish-red, erect, glabrous. Capitulum 20–25 mm in diameter, opening, flat. Outer involucral bracts more or less imbricate, 6.5–7.5 × (3.0–)3.5–4.5 mm, blackish-green, later sometimes suffused with purple near the apex, with a distinct whitish-green margin 0.5–0.8 mm wide, oblong-ovate to broadly lanceolate, with a narrow obtuse acumen at the sparsely ciliate apex, appressed. Inner involucral bracts 22–24 × 2–3 mm, dark green with a pale margin, with a narrow, but obtuse apex. Ligules medium yellow, with a dark greyish-red stripe on outer face, rarely greenish-red, lobes of inner ligules usually reddish. Styles exserted, stigmas greenish. Pollen absent. Achenes with body 4.5–5.5 mm, brown, oblong, with sparse, thin spiniform projections at apex; gradually narrowed to subconical cone 0.7–0.9(–1.3) mm; beak 7.0–8.5 mm, pale; pappus 7–8 mm, whitish. Flowers 4–5. Native. Flooded grassland near the shores of lakes, turloughs and rivers. Western Ireland. Endemic.

35. T. webbii A. J. Richards Webb’s Dandelion Small perennial herb with a stock-like rhizome. Leaves few, suberect or spreading; lamina 5–12 × 0.3–0.5 cm, medium green on upper surface, green to purplish on midrib, linear or narrowly oblanceolate, acute or subobtuse at apex, entire or with 2–3(–5) pairs of short teeth arising from a sinuate margin, glabrous or nearly so; petiole long, unwinged proximally, narrow and brownish-purple. Flowering stem 6–12 cm, pale green, glabrous or sparsely arachnoid-hairy at base. Capitulum 25–30 mm in diameter, opening, flat. Outer involucral bracts not distinctly imbricate, 5.5–6.5 × 3.0–4.5 mm, medium to blackish-green often suffused with purple near the apex and with a very distinct whitish margin about 0.9–1.2 mm wide on outer face, ovate or ovate-lanceolate, obtuse at apex, appressed. Inner involucral bracts 16–19 × 1.5–2.0 mm, olive green with narrow pale margin, oblong-lanceolate, obtuse at apex. Ligules bright yel-

37. T. ciliare Soest Channel Island Dandelion T. austrinum auct.; T. hoedicense Soest; T. sarniense A. J. Richards Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 8– 15 × 0.5–1.5 cm, medium green on upper surface, the midrib green or purplish proximally, narrowly elliptical or linear-oblanceolate, terminal lobe often elongated and acute at apex, entire or more usually with 3–5 pairs of distant, regularly spaced, short, triangular to deltoid teeth; glabrous; petiole long, proximally unwinged, narrow, purplish. Flowering stem 8–20 cm, usually suffused brownishpurple above, sparsely arachnoid-hairy at first beneath the capitulum. Capitulum 25–35 mm in diameter, opening flat. Outer involucral bracts imbricate, 3.0–6.5 × 1.5–5.5 mm, medium green on outer face with a darker midrib and a clearly demarcated white margin about one-quarter the area of the bract, the whole, especially in bud, sometimes suffused pink, ovate, acuminate with an obtuse point at apex, tightly appressed. Inner involucral bracts 12–15 × 1.5–2.5 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyish-violet stripe on outer face and purple apical lobes. Styles pale yellowish-green. Pollen

34. Taraxacum present. Achenes with body 2.5–4.2 mm, straw brown, more or less smooth with dense, short, spiniform projections at apex; cone 0.7–1.0 mm, cylindrical; beak 8–9 mm, pale; pappus 5–6 mm, whitish. Flowers 4–5. 2n = 32. Native. Grassy fens. Channel Islands. Common in western France and probably the only species of the section in Spain. Similar plants in Switzerland and Italy. Plants from the New Forest, Hampshire are intermediate between this species and T. palustre. 38. T. anglicum Dahlst. English Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, erect or spreading; lamina 6–20(–30) × 1–2 cm, medium, dull green, with small, scattered, punctate spots on upper surface, oblong or oblanceolate in outline, divided almost to midrib; terminal lobe triangular-subsagittate, large, acute at apex, entire; lateral lobes 2–3 pairs, triangular, acute at apex, margin towards the proximal area of the leaf filiform-dentate; interlobe area near the midrib short and narrow, sometimes filiformdentate; glabrous; petiole long, vivid purple, unwinged. Flowering stem 5–20 cm, erect, pale green, glabrous. Capitulum 25–40 mm in diameter, opening flat. Outer involucral bracts not imbricate, 7–10 × 3–5 mm, dark green with a very narrow pale margin on outer face, ovate, acuminate but with an obtuse point at apex, appressed. Inner involucral bracts 18–20 × 1.5–2.0 mm, olive green with a narrow, pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a leaden-grey, sometimes reddishbrown stripe on outer face, lobes of inner dirty reddish. Styles exserted, pale greenish. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with spinulose projections at apex; cone 0.8–1.0 mm; beak 6–8 mm, pale; pappus 8–9 mm, whitish. Flowers 4–5. 2n = 24. Native. Hay meadows liable to seasonal flooding. Southern and eastern England. Holland and France. Section 4. Spectabilia (Dahlst.) Dahlst. Taraxacum taxon Spectabilia Dahlst.; Taraxacum subsection Spectabilia (Dahlst.) Schischk.; Taraxacum section Euspectabilia M. P. Christ. nom. illegit. Small to medium-sized perennial herbs. Leaves spotted or blotched brownish-purple, unlobed or divided halfway to the midrib. Capitulum 20–50 mm in diameter. Outer involucral bracts 5–8 mm, without or with a narrow, pale margin, ovate or ovate-lanceolate, appressed or erect-appressed, not corniculate at apex. Ligules with a carmine or purplish-grey stripe on outer face. Achenes with body 3.5–5.0 mm, strawcoloured, with few to numerous spiniform projections at apex; cone 0.2–0.4 mm. Damp, usually acid places; montane or submontane. The section formally included the Sections Celtica and Naevosa, and many of the records for Taraxacum palustre in the Critical Supplement to the Atlas of the British Flora actually belonged to this former wider section. 39. T. faeroense (Dahlst.) Dahlst. Faeroes Dandelion T. spectabile var. faeroense Dahlst.; T. cimbricum Wiinst.; T. eximium auct.; T. spectabile auct.; T. reclinatum auct. Small to medium-sized perennial herb with a stock-like rhizome. Leaves rather few, erect or spreading; lamina

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5–25 × 1.0–2.5(–4.0) cm, dark green and often spotted and blotched brownish-purple on upper surface, brownish-purple on midrib, oblong-oblanceolate in outline, sometimes without lobes but dentate, sometimes divided one-quarter to halfway to the midrib; terminal lobe helmetshaped, more or less acute at apex and entire; lateral lobes 2–4 pairs, patent or slightly recurved, broad, acute at apex, entire; interlobe area near midrib broad and entire; glabrous or nearly so, but rough; petioles rather long, purplish, unwinged. Flowering stem 2–20 cm, pale green but often suffused purplish, more or less glabrous. Capitulum 20–40 mm in diameter, opening. Outer involucral bracts 6–8 × 2.5–3.5 mm, dark green, sometimes tinted purplish and without a pale margin on outer face, ovate-lanceolate, acute at apex, erect-appressed. Inner involucral bracts 15– 20 × 1.8–2.0 mm, dark green with a narrow, pale margin, oblong-lanceolate, obtuse at apex. Ligules bright yellow, with a carmine stripe on outer face. Styles exserted or more or less inserted, dirty yellow. Pollen absent. Achenes with body 3.5–4.0 mm, straw-coloured, oblong, almost lacking projections at apex; cone 0.2–0.3 mm; beak 5–8 mm, pale, stout; pappus 6–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 40. Easily recognised by its dark, often spotted leaves, which are often quite or nearly unlobed, purple petioles, appressed outer involucral bracts, carmine stripes on the ligules and large, pale achenes. Native. Wet places in hilly districts where it is very common, less frequently on lowland heaths and roadsides where it is perhaps introduced, ascending to 1,000 m in Scotland. Throughout Great Britain and Ireland and perhaps the most widespread of our dandelions, particularly in the north and west, scarce in fens in the south-east. Scandinavia, Faeroes and Iceland. 40. T. geirhildae (Beeby) R. Palmer & Walter Scott Shetland Dandelion T. spectabile subsp. geirhildae Beeby; T. ornatum auct. Medium-sized perennial herb with a stock-like rhizome. Leaves few to numerous, prostrate to widely spreading; lamina (6–)9–10(–14) × (2.2–)2.5–2.9(–3.0) cm, yellowishgreen to dark apple green and very sparingly and lightly spotted brownish-red on the upper surface, conspicuously reddish on midrib, broadly obovate or oblanceolate, obtuse at apex, unlobed, with up to 5 teeth or denticulations on either side; rough on upper surface with short, stout hairs; petioles typically very short and occasionally wanting. Flowering stems 5–18 cm, dull brownish-red, glabrous. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 7–8 × 2.8–3.0 mm, dark green with a narrow, pale margin on outer face, triangular-ovate, obtuse at apex, appressed. Inner involucral bracts 15–17 × 1.5–2.0 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow with a dark purplish-grey stripe on outer face. Styles discoloured. Pollen present. Achenes with body 4.3– 4.6 mm, straw brown, with short, spiniform projections at apex; cone about 0.4 mm; beak 5–6 mm, pale; pappus 6–7 mm, whitish. Flowers 5–6. Native. Known only from grassy ledges among rocky outcrops to the south-east of Lang Kl¨odie Loch and towards the

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end of Birka Water on Northmaven in the Shetland Islands. Endemic. 41. T. serpenticola A. J. Richards Serpentine Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves few, prostrate to spreading, thick and leathery; lamina (6–)9–11 × 2–3 cm, dark olive or brownish-green, with sparse irregular dark spots on the upper surface, the midrib and sometimes some of the lateral veins dark purplish-brown, obovate or oblanceolate, obtuse-mucronulate at apex, unlobed, with 1–3, large, irregular teeth on each side, or shallowly and irregularly lobed; glabrous to nearly so; petioles short, dark brownishpurple, unwinged. Flowering stem 6–11 cm, pale green suffused purplish, arachnoid-hairy just beneath the capitulum. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 5–7 × 2–4 mm, green suffused purple especially towards the apex on the outer face and with an obscure pale margin, ovate, acute at apex, appressed. Inner involucral bracts 15–17 × 1.5–2.0 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a deep carmine stripe on outer face and purple apical lobes. Styles more or less included, discoloured. Pollen absent. Achene with body 4.8–5.0 mm, straw brown, with short, spiniform projections at apex; cone about 0.4 mm; beak 5–6 mm, thick, pale; pappus 6–7 mm, white. Native. Grassy ledges among serpentine outcrops. Muckle Heog on Unst in the Shetland Islands. Endemic. Section 5. Taraxacum Medium-sized perennial herbs. Leaves smooth, bright green, unlobed or very narrowly lobed. Capitulum 30– 45 mm in diameter. Outer involucral bracts 7–14 mm, spreading to spreading-recurved or erecto-patent to erect, not corniculate at apex. Inner involucral bracts not corniculate at apex. Ligules deep yellow to orange-yellow. Achenes with body pale or straw brown or rust-coloured; cone nearly absent or up to 1.0 mm. Rare species of high mountain cliffs in Scotland and elsewhere in northern Europe. Taraxacum Wiggers (1780) is conserved against Taraxacum Zinn (1757). Its holotype species is Taraxacum officinale Wiggers, which is a new name for Leontodon taraxacum L. Linnaeus took his diagnosis unchanged from the earlier Flora Lapponica. Linnaeus’ specimen representing this species in his Flora Lapponica is now in the library of the Institut de France, Paris and has been selected as the lectotype of Leontodon taraxacum and therefore of Taraxacum officinale. It has been equated with Taraxacum campylodes G. E. Haglund which belongs to this section. 42. T. ceratolobum Dahlst. Bright Green Dandelion T. acidotum M. P. Christ.; T. croceum auct. Small to medium-sized perennial herb with a stock-like rhizome. Leaves rather few, erect or spreading; lamina 5–15 × 1.5–2.0 cm, pure green on upper surface, green on midrib, narrowly oblong in outline, divided nearly to midrib; terminal lobe very narrowly triangular with narrower spreading lobules at base, acute at apex, entire; lateral

lobes 5–8 pairs, regular, short and narrow, acute at apex, entire, spreading; interlobe area near the midrib narrow, entire or with an occasional small tooth; glabrous; petiole short, green, narrowly winged. Flowering stem 8–20 cm, pale green or slightly purplish, glabrous or slightly arachnoid-hairy. Capitula 40–50 mm in diameter, opening. Outer involucral bracts 6–7 × 2.0–2.0 mm, green on both faces, with a pale margin, linear-lanceolate, acute at apex, spreading. Inner involucral bracts 15–18 × 1.5– 2.0 mm, light green with a pale margin, oblong-lanceolate, at apex. Ligules deep orange-yellow, with a violet stripe on the outer face. Styles exserted, yellow. Pollen present or absent. Achene with body 3.8–4.0 mm, light brown, with dense, acute, often very sharp spiniform projections at apex; cone 0.4–0.7 mm; beak 7–9 mm, pale; pappus 8–10 mm, whitish. Flowers 5–7. 2n = 32, 33, 34, 35, 36, 37, 38, 39. Easily recognised by its bright green leaves with many, regular, narrow lateral lobes. Native. Basic rock-ledges on cliffs between 900 and 1,070 m. Mountains of central Scotland with an isolated locality in Ross-shire. Scandinavia, Iceland and northern Russia. 43. T. clovense A. J. Richards Clova Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, erect; lamina 10–28 × 2–3 cm, green or pale green on upper surface, green on midrib, oblanceolate or oblong-oblanceolate, unlobed and dentate to slightly lobed; terminal lobe large, triangular to helmetshaped, curved to a rather obtuse point, entire; lateral lobes 2–3(–4) pairs, shallow, acute at apex, entire and straightsided to somewhat concave on the margin towards the distal part of the leaf, slightly curved down; interlobe area near the midrib wide and entire; glabrous; petioles fairly long, green, narrowly winged. Flowering stem 10–20 cm, green, glabrous. Capitula 30–40 mm in diameter, opening. Outer involucral bracts 9–14 × 3–4 mm, green on outer face, with a pale green margin, pruinose on inner face, lanceolate, acute at apex, erecto-patent to suberect. Inner involucral bracts oblong-lanceolate, acute at apex. Ligules deep yellow. Styles exserted, discoloured. Pollen present. Achenes with body 4.8–5.3 mm, straw brown, almost smooth; almost imperceptibly narrowed into the short cone; beak 7–9 mm, pale; pappus 8–10 mm, whitish. Flowers 6–7. Native. Glen Dole in the Clova Mountains of Forfarshire. Endemic. 44. T. xiphoideum G. E. Haglund Parallel-leaved Dandelion T. hypochaeris auct. Small to medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, erect; lamina 5–20 × 1.5– 3.0 cm, medium green on upper surface, with a green midrib, oblong-oblanceolate, more or less acute or obtuse at apex, sparsely but regularly dentate, the teeth narrow and sharp; glabrous; petiole poorly differentiated, green, broadly winged. Flowering stem 6–25 cm, pale green or copper-coloured, nearly glabrous. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 7–9 × 2.0– 2.5 mm, green on both faces, with a pale margin, lanceolate,

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obtuse at apex, spreading. Inner involucral bracts 15– 17 × 2–3 mm, dark green with a pale margin, oblonglanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with spiniform projections in the upper part; cone 0.8–1.0 mm; beak 6–7 mm, pale; pappus 8–9 mm, whitish. Flowers 6–7. 2n = 24. Native. Known only from the Moy Corrie near Loch Laggan in Inverness-shire and by Loch Lyon in Argyllshire. Southern Iceland and south-west Norway.

Pollen present. Achenes with body 3.0–3.5 mm, rustcoloured with sharp spiniform projections at apex; cone 0.8–1.0 mm; beak 8–9 mm, pale; pappus 9–10 mm, whitish. Flowers 6–7. Obligate agamosperm. 2n = 32. The only species in our area with bright green leaves, rust-coloured fruits and deep yellow ligules. Native. Rare on calcareous schist at about 1,000 m. Known only in the south-west corrie of Ben Lawes in Perthshire. It is one of the notable relict species found on that mountain and is at least 10◦ latitude south of any other station in Europe. Faeroes, Iceland, Norway and Svalbard.

45. T. craspedotum Dahlst. Pale-leaved Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, erect; lamina 4–10 × 1–2 cm, rather pale bluish-green on upper surface, with a green midrib, oblong or oblong-oblanceolate in outline, divided three-quarters of the way to midrib; terminal lobe triangular, obtuse and often rounded at apex, entire; lateral lobes 3–4 pairs, deltoid, acute at apex, entire or toothed on the margin towards the proximal area of the leaf, short and spreading; interlobe area near the midrib thick, entire or with an occasional tooth; glabrous or nearly so; petiole short, green and winged. Flowering stem 5–10 cm, pale green, glabrous. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 6.5–7.5 × 1.8–2.2 mm, dark green with a conspicuous pale margin on outer face, lanceolate or oblong-lanceolate, abruptly narrowed to an obtuse apex, spreading. Inner involucral bracts 13–15 × 1.5–2.0 mm, dark green, with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a violet stripe on the outer face. Styles exserted, yellow. Pollen absent. Achenes with body 3.0–3.2 mm, pale brown, with spiniform projections at apex; cone 0.2–0.3 mm; beak 6–7 mm, pale; pappus 8–9 mm, whitish. Flowers 6–9. 2n = 32. Native. Acid but mineral-rich rock-ledges on cliffs from 900 to 1,200 m. Local in the mountains of central Scotland with a solitary disjunct locality in Ross-shire. Norway and Faeroes.

47. T. pycnostictum M. P. Christ. Small-spotted Dandelion T. stictophoreum M. P. Christ. Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 5– 15 × 0.7–2.5 cm, bluish-green with small spots on upper surface, with green midrib, oblong-oblanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe long-helmet-shaped, one-quarter to one-third the length of the lamina, subacute or mucronate at apex, entire or sometimes with a solitary tooth; lateral lobes 3–5 pairs, narrowly triangular, acute at apex, entire, margin towards the distal area of the leaf concave, spreading; interlobe area near the midrib fairly wide, with an occasional tooth; rough with short hairs; petiole fairly long, green, winged. Flowering stem 10–20 cm, pale green, erect, glabrous. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 8– 11 × 2–3 mm, glaucous and scarcely with a pale margin on outer face, stiff and thistle-like, oblong-lanceolate, obtuse at apex, erect to spreading. Inner involucral bracts 15– 17 × 1.5–2.5 mm, dark green with a pale margin, oblonglanceolate, obtuse at apex. Ligules deep yellow, with a greyish-purple stripe on the outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–3.8 mm, straw brown, with spiniform projections at apex; cone 0.4– 0.7 mm; beak 6–7 mm; pappus 7–8 mm, whitish. Flowers 6–7. Obligate agamosperm. 2n = 32. Easily recognised by its rough leaves with small spots, stiff, thistle-like outer involucral bracts and deep yellow ligules. Native. Very wet cliff ledges on calcareous schist from 600 to 900 m. Central Highlands of Scotland. Iceland and Faeroes.

46. T. cymbifolium H. Lindb. ex Dahlst. Ben Lawes Dandelion T. acromaurum Dahlst. Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 5–15 cm, pure bright green on upper surface, with green midrib, oblong or elliptical-oblong in outline, divided about three-quarters of way to the midrib; terminal lobe triangular to oblong-triangular, more or less acute at apex, entire; lateral lobes 3–4 pairs, narrowly triangular, obtuse or acute at apex, scarcely toothed, spreading; interlobe area near the midrib wide, with few teeth; glabrous; petiole short, green, winged. Flowering stem 5–10 cm, pale green, somewhat arachnoid-hairy near the capitulum. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 6–7 × 1.8–2.0 mm, green on both faces,without a pale margin, ovate-lanceolate, acute at apex, spreading-recurved. Inner involucral bracts 15–18 × 1.5–2.0 mm, dark green, linear-lanceolate, acute at apex. Ligules deep yellow, with a violet stripe on outer face. Styles exserted, discoloured.

Section 6. Naevosa M. P. Christ. Taraxacum subsection Naevosa (M. P. Christ.) A. J. Richards; Taraxacum taxon Eunaevosa M. P. Christ. nom. inval. Small to robust perennial herbs. Leaves typically rough, coarsely lobed and with large dark spots covering more than 10 per cent of the surface. Capitulum 20–60 mm in diameter. Outer involucral bracts 5–14 mm, mainly spreading to erect. Ligules with a grey, greyish-violet, greyish-purple or purple, rarely reddish stripe on outer face. Achenes usually straw-coloured, rarely brick red or orange. Mainly plants of western and northern regions. They may have arisen from hybridisation between species of the Sections Spectabilia and Ruderalia.

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48. T. naevosum Dahlst. Squat Dandelion T. subsimile Dahlst. Squat, very robust, medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 8–25 cm, dark greyish-green, usually with irregular spots on upper surface, purplish on midrib, sometimes also the other veins, narrowly oblong-elliptical or oblongoblanceolate in outline, divided three-quarters of the way to midrib; terminal lobe long helmet-shaped, obtuse at apex, entire or an occasional tooth; lateral lobes 5–7 pairs, rather crowded, triangular, acute at apex, often with a concave angle to the distal margin which is dentate, spreading; interlobe area near the midrib short, wide and dentate; rough with short, bristly hairs; petiole fairly long, dull purple, winged. Flowering stem 5–18 cm, thick, pale green or slightly pinkish, glabrous below, more or less arachnoidhairy above. Capitulum 50–60 mm in diameter, opening flat. Outer involucral bracts 10–12 × 3–5 mm, green with pink tips and scarcely pale margined on outer face, lanceolate, acute at apex, erect to spreading. Inner involucral bracts 15–22 × 1.5–2.0 mm, dark green,with pale margin, narrowly oblong-lanceolate, obtuse at the purplish apex. Ligules medium yellow, with a purple stripe on outer face. Styles exserted, discoloured. Pollen abundant. Achenes with body 3.7–4.0 mm, straw brown, with thick, spiniform projections above; cone 0.7–0.9 mm; beak 8–10 mm, pale; pappus 7–8 mm, whitish. Flowers 5–6. Obligate agamosperm. 2n = 32. Characterised by its large, rough leaves, broadly winged petioles, large capitula and abundant pollen. Native. Damp old hay meadows and sometimes on scars. Frequent in central and northern Scotland and northern England, with isolated records in southern Scotland, central England and north Wales. Fennoscandia, Faeroes and Iceland. 49. T. naevosiforme Dahlst. Wetland Dandelion T. hamatifrons Dahlst.; T. plicatum Dahlst.; T. acidodontum Dahlst. Robust, medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, erect or spreading; lamina 5–20 × 1.5–5.0 cm, yellowish-green or dark greyishgreen, spotted brownish-purple on upper surface, purplish on midrib, oblong-oblanceolate in outline, divided threequarters of the way to the midrib; terminal lobe triangularhelmet-shaped, acute at apex, entire; lateral lobes 4–7 pairs, triangular, the margin towards the distal area of the leaf convex, acute at apex, more or less dentate, recurved; interlobe area near the midrib short and thick, more or less dentate; rough with short, bristly hairs; petiole fairly long, purple, narrowly winged. Flowering stem 6–20 cm, erect, often purplish, somewhat arachnoid-hairy at apex. Capitulum 35–40 mm in diameter, opening. Outer involucral bracts 8–10 × 2.0–2.5 mm, dark green suffused with purple and with red tips on both faces and scarcely pale-margined, lanceolate, acute at apex, spreading. Inner involucral bracts 15–20 × 1.5–2.0 mm, dark green and pruinose with a pale margin, oblong-lanceolate, reddish and obtuse at apex. Ligules deep yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen abundant. Achenes

with body 3.0–3.5 mm, straw brown, with short spiniform projections at apex; cone 0.4–0.6 mm; beak 9–10 mm, pale; pappus 7–9 mm, whitish. Flowers 5–7. Obligate agamosperm. 2n = 32. Native. Wet grassland and cliffs. Fairly common in Scotland and northern England, scattered in Wales and central England; in a few localities of coastal areas of Ireland. Norway and Sweden. 50. T. rubellum M. P. Christ. Red-fruited Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves rather few, erect or spreading; lamina 15–20 × 1– 3 cm, dark green with brownish-purple spots on upper surface, purplish on midrib, narrowly oblong in outline, divided seven-eighths of the way to the midrib; terminal lobe triangular-hastate, acute and rather elongated at apex, entire; lateral lobes 6–9 pairs, triangular-deltoid with a linear, acute apex, but broader proximally to form a sigmoid distal margin when well developed, dentate only on proximal lobes, spreading; interlobe area near the midrib rather long and wide with unequal teeth; glabrous or nearly so; petioles short to fairly long, purple, narrow and unwinged. Flowering stem 15–20 cm, pale green, often purplish, somewhat arachnoid-hairy at apex. Capitulum 50–60 mm in diameter, opening. Outer involucral bracts 11–14 × 2.0– 3.5 mm, dark green on outer face, pruinose and with a pale margin, pale green on inner face, linear-lanceolate, acute at apex, loosely erect to spreading or recurved. Inner involucral bracts 15–17 × 2.0–2.5 mm, oblong-lanceolate, obtuse at apex. Ligules bright yellow, with a greyish-purple or green stripe on the outer face. Styles exserted, discoloured. Pollen abundant. Achenes with body 3.8–4.0 mm, brick red or orange, with acute spiniform projections at apex; cone 0.8–1.0 mm; beak 9–11 mm, pale; pappus 10–12 mm, whitish. Flowers 5–6. The only Scottish dandelion with brick red to orange achenes and spotted leaves. There is some question as to whether Scottish plants are identical with those from Iceland. Native. North coast of Scotland, Inner Hebrides, Orkneys and one locality inland in southern Scotland. Iceland. 51. T. euryphyllum (Dahlst.) Hjelt Wide-stalked Dandelion T. maculigerum subsp. euryphyllum Dahlst. Medium-sized to robust perennial herb with a stock-like rhizome. Leaves fairly numerous, erect or spreading; lamina 10–20 × 1–5 cm, pure green with irregularly shaped brownish-purple spots and often blotches on the interlobe area, purplish on midrib, oblong or oblong-oblanceolate in outline, divided up to two-thirds of the way to the midrib; terminal lobe triangular, obtuse at apex, entire or with an occasional tooth; lateral lobes 4–6 pairs, triangular, acuminate at apex, with a sigmoid distal margin when well developed, scarcely dentate or sometimes dentate on margin towards the distal area of leaf only, spreading; interlobe area near the midrib short and wide, sometimes toothed; rough with short, bristly hairs; petiole short, purplish or green at base, widely winged. Flowering stem 8–20 cm, erect, pale green, glabrous or nearly so. Capitulum 30–40 mm

34. Taraxacum in diameter, opening. Outer involucral bracts 9–12 × 2.8– 3.7 mm, bluish-green on outer face, pruinose and with a narrow, pale margin, pale green on inner face, lanceolate, acute at apex, spreading, recurved or spreading in all directions. Inner involucral bracts 16–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the reddish apex. Ligules yellow, with a reddish-purple stripe on outer face and purple apical lobes. Styles exserted, discoloured. Pollen very sparse in outer flowers only. Achenes with body 3.5–4.0 mm, straw-coloured, with acute spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 32. Native. Wet, rather sheltered, base-rich places and can be more weedy than other species in the section, especially on roadsides. Throughout Great Britain, but possibly more frequent in northern England with a solitary record in Co. Dublin in Ireland. Scandinavia, Holland and Germany. 52. T. hirsutissimum C. C. Haw. Hairy Dandelion Robust, medium-sized perennial herb with a stock-like rhizome. Leaves 5–20 × 2–5 cm, rather few, erect or spreading; lamina greyish-green and sometimes with a very few, faint spots on the upper surface, faintly tinted purplish on midrib, oblong in outline, divided nearly to midrib; terminal lobe triangular-sagittate, with an acute, elongated apex, entire; lateral lobes 4–7 pairs, narrowly triangular, drawn out to a linear, acute apex, spreading to somewhat recurved, the margin towards the distal area of the leaf curved or angled, not or scarcely dentate-spreading or slightly recurved; interlobe area near the midrib short and narrow, sometimes toothed; with numerous, short, bristly hairs; petioles short, purple, more or less winged. Flowering stem 10–20 cm, often purplish, arachnoid-hairy near the apex. Capitulum 20–30 mm in diameter, opening. Outer involucral bracts 10–12 × 2.0– 2.5 mm, bluish-green on outer face, pale green on inner face, linear-lanceolate, acute at apex, recurved. Inner involucral bracts 15–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules rather pale yellow, with a dark grey stripe on outer face and yellow lobes. Styles exserted, orange. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with spiniform projections at apex; cone 0.4–0.5 mm; beak 9–10 mm, pale; pappus 10– 12 mm, whitish. Flowers 5–6. Native. Roadsides, sand-dunes, coastal turf and sandy shore. South end of South Mainland, Shetland Islands. Endemic. 53. T. maculosum A. J. Richards Spotted Dandelion T. maculigerum auct. Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, erect or spreading; lamina 5– 20 × 1–6 cm, dull or dark green, with irregular brownishpurple spots on upper surface, purplish on midrib, oblongoblanceolate in outline, divided nearly to midrib; terminal lobe triangular-subsagittate, acute at apex, entire; lateral lobes 2–4 pairs but often more or less alternate, triangular, acute at apex, broader proximally to form a sigmoid margin when well developed, dentate on lobes towards the proximal area of the leaf; interlobe area near the midrib short and

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narrow, sometimes toothed; rough with short bristly hairs; petiole short, purple, more or less unwinged. Flowering stem 10–20 cm, decumbent to erect, pale green, glabrous. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 8–10 × 2.5–3.5 mm, dark green with hardly a pale margin and pruinose on outer face, linear-lanceolate, acute at apex, erect or spreading. Inner involucral bracts 12–25 × 1–2 mm, dark green with a pale margin, oblonglanceolate, obtuse to acute at apex. Ligules yellow, with a purple stripe on outer face. Styles exserted, discoloured. Pollen absent. Achenes with body 3.4–3.6 mm, straw brown, with rather few, acute, spiniform projections at apex; cone 0.6–0.7 mm; beak 7–9 mm, pale; pappus 7–9 mm, whitish. Flowers 5–6. Obligate agamosperm. 2n = 32. Native. Wet wood margins, streamsides, damp upland pasture, waysides and cliff-faces, usually on basic soils, perhaps introduced in the south-east. Fairly common in northern England and Scotland, scattered in the remainder of England and Wales; northern and western Ireland. Endemic. 54. T. pseudolarssonii A. J. Richards Spreading-bracted Dandelion T. edmondsonii A. J. Richards Medium-sized, very heterophyllous perennial herb with a stock-like rhizome. Leaves numerous, erect or spreading; lamina 10–20 × 2–3 mm, variable, dark green and very heavily spotted with purple or black spots on upper surface, often coalesced early on, the pigment water-soluble so the spots fade rapidly and are scarcely visible on lateseason plants and old leaves, purplish on midrib, oblong or oblong-oblanceolate in outline, divided almost to the midrib; terminal lobe often helmet-shaped, on first leaves subacute at apex, in later plants large and dentate; lateral lobes 3–5 pairs, narrowly triangular, acute at apex, neat and regular in young leaves, later becoming coarsely dentate on either margin, patent to somewhat recurved; interlobe area near the midrib short and fairly wide, often dentate; glabrous or nearly so; petiole rather short, purplish at first but fading, winged or unwinged. Flowering stem 10–20 cm, pale green, glabrous. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 9–12 × 2.0–2.5 mm, greyishgreen on both faces with a narrow pale margin, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 18–22 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with short, acute spiniform projections at apex; cone 0.4–0.5 mm; beak 4.5 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 32. Native. Meadows, lanesides, moorland tracks and sanddunes. Common and sometimes abundant in northern England with scattered records in Scotland and Wales and coastal England. Endemic. 55. T. subnaevosum A. J. Richards Pale-bracted Dandelion Medium-sized, rather delicate perennial herb with a stocklike rhizome. Leaves fairly numerous, spreading or erect; lamina 10–14 × 1–4 cm, dark green and sparsely but

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regularly spotted with small black spots on the upper surface, green on midrib, oblanceolate or oblong in outline, divided nearly to midrib; terminal lobe triangularsubsagittate, acute at apex, entire or denticulate; lateral lobes 3–4 pairs, narrowly triangular, acute at apex, entire or sometimes denticulate on the margin towards the distal area of the leaf, recurved to slightly spreading; interlobe area near the midrib short and narrow, denticulate; glabrous or nearly so; petioles short to fairly long, greenish or purplish, more or less dentate, unwinged. Flowering stem 10–16 cm, pale green, sparsely arachnoid-hairy. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 6– 9 × 1.5–2.2 mm, pale green with little sign of a pale margin, linear-lanceolate, acute at apex, recurved. Inner involucral bracts 17–19 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules pale yellow, with a greyish-purple stripe on outer face. Styles exserted, yellowish or occasionally darker. Pollen absent. Achenes with body 3.1–3.3 mm, straw brown, with short, spiniform projections at apex; cone 0.6–0.9 mm; beak 6–7 mm, pale; pappus 8–10 mm, whitish. Flowers 5–6. Native. Often in open and rather rocky areas. Widespread in some areas of northern England and Scotland and a few scattered records in the remainder of England and Wales. Endemic. 56. T. cornubiense A. J. Richards Cornish Dandelion Medium-sized, rather delicate perennial herb with a stocklike rhizome. Leaves few, spreading or suberect; lamina 10– 15 × 1–4 cm, dull green and sparsely covered with small, blackish spots on the upper surface, green to faintly purplish on midrib, oblong-elliptical in outline, divided nearly to midrib; terminal lobe triangular, often elongated at apex, often deeply divided on one side or entire; lateral lobes 4–5 pairs, triangular, acute and often elongate at apex, subconvex and denticulate on the margin towards the distal area of the leaf, subpatent or sloping down; interlobe area near the midrib short and narrow, denticulate; glabrous or nearly so; petioles rather long, dull purplish, narrow and unwinged, more or less dentate. Flowering stem 10–15 cm, pale green to purplish, arachnoid-hairy near the capitulum. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 7–9 × 1.7–2.6 mm, rather dark green on outer face with an indistinct pale margin, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 17–19 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.6–3.8 mm, straw-coloured, with tubercles or short spiniform projections at apex; cone 0.7–0.9 mm; beak 6–8 mm, pale; pappus 7–10 mm, discoloured. Flowers 3–4. 2n = 24. Native. Gardens, roadside verges and old railway lines. Widespread in Cornwall, especially on the Lizard. Endemic. 57. T. drucei Dahlst. Druce’s Dandelion T. lainzii auct.; T. maculigerum auct. Small to medium-sized, delicate perennial herb with a stock-like rhizome. Leaves few, erect or spreading; lamina 6–17 × 1.5–3.0 cm, medium green and usually with small, sparse, brownish-purple spots on upper surface, green to

fairly purplish on midrib, oblanceolate to obovate, little divided; terminal lobe large, broadly triangular or ovate, rounded at apex, entire; lateral lobes 2–5 pairs, shallow or sometimes only as teeth, shortly triangular, acute at apex and forming a sinuate edge on margin towards the distal area of the leaf; interlobe area near the midrib either not obvious or short, thick and entire; glabrous; petioles rather long, green to dull purple, unwinged or narrowly winged. Flowering stem 10–20 cm, erect, pale green, glabrous. Capitulum 35–40 mm in diameter, opening. Outer involucral bracts 6–8 × 1.5–2.0 mm, dark green on outer face with a narrow white margin, ovate, acute at apex, erect to spreading. Inner involucral bracts 16–25 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a grey stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, smooth, with short spiniform projections above; cone 0.4–0.5 mm; beak 6–10 mm, pale; pappus 8–10 mm, whitish. Flowers 3–5. Obligate agamosperm. 2n = 24. Native. Wet grassland, cliff ledges, rocks and in gorges especially on limestone. Western Great Britain and mainly coastal in Ireland. Spain and Portugal. Named after George Claridge Druce (1850–1932). 58. T. stictophyllum Dahlst. Stiff-leaved Dandelion T. calophyllum Dahlst.; T. laetifrons Dahlst. Small to medium-sized, squat perennial herb with a stocklike rhizome. Leaves few, stiff, prostrate to spreading, in a tight rosette; lamina 8–25 × 2–5 cm, greyish-green and usually with scattered, punctate spots on upper surface, pale to bright red on midrib, oblanceolate to oblong-oblanceolate in outline, much divided about three-quarters of way to midrib; terminal lobe large, triangular to helmet-shaped with the basal lobes pointing down, obtuse or rounded at apex, entire or dentate; lateral lobes 3–5 pairs, triangular, linear and acute at apex, with long, narrow teeth; interlobe area near the midrib short and thick, with narrow teeth; rough with short bristly hairs; petioles short to medium, pale green to purplish, broadly winged. Flowering stem 6–20 cm, pale green below, reddish above, somewhat arachnoid-hairy at apex. Capitulum 25–40 mm in diameter, opening. Outer involucral bracts many and crowded, 5–9 × 2.5–3.0 mm, dark green with a faint pale margin on outer face, ovate, obtuse at apex, curved upwards to give a rounded involucre. Inner involucral bracts 14–18 × 1.5–2.0 mm, medium green with a pale margin, oblong-lanceolate, obtuse at the reddish apex. Ligules yellow, striped reddish-purple on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.2 × 1.0–1.3 mm, straw brown, with short, sharp spiniform projections at apex; cone 0.8–1.0 mm, broad-based; beak 7–11 mm; pappus 6–7 mm, whitish. Flowers 4–6. Native. Wet shady rocks at low altitudes. Widespread in western and northern Great Britain. Faeroes, Iceland and Norway. 59. T. richardsianum C. C. Haw. Richards’s Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves spreading to suberect; lamina 15–20 cm, dark green with many, scattered, rather small, purplish-black spots on the upper surface, purplish on midrib, oblong in outline,

34. Taraxacum divided almost to midrib; terminal lobe large, triangularsubsagittate, subacute at apex, often subdivided with large teeth; lateral lobes 6–8 pairs, sloping down, crowded, regular, triangular, with rather long, narrow, pointed apices, margin towards the distal area of the leaf more or less straight, but sometimes sinuate and bearing a few subulate teeth; interlobe area near the midrib fairly wide, short and shortly dentate with filiform teeth; with fairly numerous simple eglandular hairs; petioles about one-fifth the length of the lamina, purplish, unwinged or scarcely winged. Flowering stems 15–20 cm, purplish, arachnoid-hairy. Capitulum 35–40 mm in diameter, opening. Outer involucral bracts 10–12 × 2–3 mm, pale green on inner face, medium green on the outer face with a distinct white margin, lanceolate, acute at apex, suberect to patent and somewhat arcuate. Inner involucral bracts 16–18 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a purple stripe on the outer face, with orange-red tips to those of the central flowers. Styles discoloured. Pollen absent or rarely with a few grains in the outer flowers. Achenes with body 3.2–3.7 mm, straw brown, with broad-based spiniform projections above; cone about 0.8 mm, conical; beak 6–7 mm; pappus 8–10 mm, white. Native. Moist grasslands and herb-rich meadows. Local in England and Wales, but frequent in some areas. Endemic. Named after Adrian John Richards (b. 1943). Section 7. Celtica A. J. Richards Medium-sized perennial herbs. Leaves medium green or bluish-green, not usually spotted, generally smooth and glabrous, midrib and petioles often bright purple or red. Capitulum (15–)20–50 mm in diameter. Outer involucral bracts 4–12 mm, spreading to erect. Ligules with greyishviolet, brownish-violet, greyish, brownish-purple or purple stripe on outer face. Achenes with body 2–4 mm, straw brown, greyish-brown or olive brown. Species of old hedgerows, woodland margins and riversides. Western, northern and central Europe. 60. T. gelertii Raunk. Gelert’s Dandelion T. adamii auct.; T. officinale subsp. gelertii (Raunk.) Dahlst. Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, flat, spreading; lamina 5–20 × 1–6 cm, dull green, often somewhat bluish on upper surface, with a pinkish midrib, smooth, oblong in outline, divided about three-quarters of way to the midrib; terminal lobe triangular, obtuse to subacute at apex, sometimes slightly mucronate, entire or with a single blunt tooth; lateral lobes 3–5 pairs, triangular-deltoid, convex to straightsided, the margin towards the distal area of the leaf not toothed or sometimes with a single tooth on the distal lobe, the proximal margin towards the proximal area of the leaf straight; interlobe area near the midrib short and fairly thick, sometimes with an occasional tooth, spreading to somewhat recurved; smooth and glabrous or nearly so; petioles fairly long, purple, narrowly winged. Flowering stem 5–20 cm, decumbent to erect, pale green to purplish, glabrous or nearly so. Capitulum 30–45 mm in diameter, opening. Outer involucral bracts 6–7 × 1.5–2 mm, dark green on outer face, with an indistinct whitish margin, ovate

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or ovate-lanceolate, obtuse at apex, erect. Inner involucral bracts 15–25 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, rather long, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen usually present, sometimes absent. Achenes with body 3.0–3.2 mm, brownish, rugose with numerous, large spiniform projections at apex; cone 0.4–0.5 mm; beak 6–8 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Variable in leaf-shape, T. gelertii is distinguished by its erect outer involucral bracts and rugose achenes with numerous, long, spiniform projections at their apex. Native. Semi-natural and natural grassy habitats on welldrained to wet, neutral to calcareous soils, often in speciesrich grasslands. Also, not infrequently, in man-made habitats such as walls, pavements, gardens and gravel pits. In the north and west it occurs on cliffs and in ravines. Throughout much of Great Britain especially south Wales and northern England and round the perimeter of Ireland. Western Europe, Czechoslovakia and Fennoscandia. Named after Otto Christian Leonor Gelert (1862–99). 61. T. bracteatum Dahlst. Dark-green Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 10–25 × 2–5 cm, dull dark green with the interlobe area sometimes with darker blotches on the upper surface, the midrib and veins bright red or purple to apex, oblong or oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe large, triangular to helmet-shaped, subacute or mucronate at apex, entire; lateral lobes 3–5 pairs, broadly triangular, acute at apex, the margin towards the distal area of the leaf somewhat convex or sinuate, sometimes filiform-dentate, slightly sloping down; interlobe area near the midrib short and fairly thick, sometimes filiform-dentate; glabrous or nearly so; petiole long, bright purple, unwinged. Flowering stem 5– 20 cm, often reddish, somewhat arachnoid-hairy just under the capitulum where there are sometimes distinct isolated bracts. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 8–10 × 2.5–3 mm, dark green on outer face without a pale margin, greyish-green often suffused with purple and pruinose on inner face, lanceolate, acute at apex, erect. Inner involucral bracts 14–18 × 1.5–2.0 mm, dark green with a pale margin, lanceolate, obtuse at apex. Ligules yellow, with a purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3– 4 mm, straw-coloured, with spiniform projections at apex; cone 0.7–0.9 mm; beak 6–8 mm, pale; pappus 6–8 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Easily recognised by its dark green leaf with contrasting purplish petiole, midrib and veining and its dark green, erect outer involucral bracts. Native. Damp grassy places and roadsides. Scattered through Great Britain with a few records round the perimeter of Ireland. Northern and central Europe. 62. T. orcadense Dahlst. Large-lobed Dandelion T. perlaciniatum Dahlst.; T. tanylepioides Dahlst. Medium-sized perennial herb with a stock-like rhizome. Leaves few, spreading to erect; lamina 5–20 × 2–5 cm, dark shining green on upper surface, deep pink to purple on

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midrib, oblanceolate, divided three-quarters of way to the midrib; terminal lobe large, helmet-shaped, rounded and usually apiculate at apex, often dentate; lateral lobes 2–4 pairs, triangular, acute at apex, margins straight and more or less dentate, spreading to more or less sloping down; interlobe area near the midrib short and fairly thick, often dentate; smooth and glabrous or nearly so; petiole rather long, deep purple, unwinged or winged. Flowering stem 4–20 cm, erect, often shorter than the leaves, pale green, glabrous. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 8–10 × 2.0–2.5 mm, blackish-green often suffused with purple on outer face, scarcely pale margined, pale green on inner face, lanceolate, acute at apex, erectspreading. Inner involucral bracts 15–18 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a brownish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with spiniform projections at apex; cone 0.5–0.6 mm; beak 6–8 mm, pale; pappus 6–8 mm, whitish. Flowers 5–6. Native. Grassy places. Known only from Mainland and Cava in the Orkney Islands where it is frequent. Endemic.

midrib short and rather thin, usually toothed; smooth and glabrous; petiole rather short, purple, narrow, unwinged proximally. Flowering stem 7–20 cm, often purplish, somewhat arachnoid-hairy near the capitulum. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 7–10 × 2–3 mm, blackish, pruinose and with distinct, but narrow whitish margin on outer face, ovate-lanceolate, acute at apex, erect to appressed. Inner involucral bracts 15– 18 × 2.0–2.5 mm, dark green with a pale margin, oblonglanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, blackish. Pollen present. Achene body 3.2–3.7 mm, brownish, with scarcely any projections at apex; cone 0.5–0.7 mm; beak 7–8 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Easily recognised by its blackish involucre and styles and achenes almost without projections. Native. On neutral to calcareous soils often in speciesrich grasslands. Western Great Britain and the perimeter of Ireland with a few scattered records elsewhere. Endemic.

63. T. nietoi A. J. Richards Flintshire Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves few, suberect to spreading; lamina 10–25 cm, dark green on upper surface, deep pink to purple interwoven with green strands on midrib, oblanceolate in outline, divided nearly to midrib; terminal lobe large, broadly hastate, subacute at apex, entire, with a solitary tooth or more deeply incised; lateral lobes 2–3 pairs, triangular, acute at apex, the margin towards the distal area of the leaf not or scarcely toothed, convex, sigmoid, straight-sided or concave-angled, the margin towards the proximal area of the leaf entire, spreading; interlobe area near the midrib fairly long and narrow, toothed; smooth and glabrous or nearly so; petiole rather long, purple, interwoven with green strands, unwinged. Flowering stem 10–25 cm, bronze-coloured, glabrous or nearly so. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 8–10 × 3–4 mm, dark green on outer face with a distinct white margin, ovate, acute at apex, erect. Inner involucral bracts 15–18 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyish-purple stripe on outer face and purple lobes at apex. Styles exserted, yellowish. Pollen present. Achenes with body 3.0–3.8 mm, greyish-brown, smooth, with short, spiniform projections at apex; cone 0.5–0.6 mm; beak 6–8 mm, pale; pappus 6–8 mm, whitish. Flowers 4. Native. Recorded only on a roadside in Flintshire. Spain.

65. T. subbracteatum A. J. Richards Dark-bracted Dandelion T. crispifolium auct. Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, prostrate to ascending; lamina 10– 25 cm, often crispate, dark green on the upper surface, the interlobe area near the midrib usually blotched on the lower side and occasionally on the upper side, deep pink to purple with interwoven green strands on midrib, often becoming green distally, oblong to oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe helmet-shaped or triangular, subacute at apex and mucronate, entire or with 1 tooth, the basal corners sometimes slightly elongated; lateral lobes 4–6 pairs, triangular, narrow and slightly elongated at apex, the margin towards the distal area of the leaf entire or with 1(−3) teeth and strongly sigmoid, sometimes more deeply incised; interlobe area near the midrib short and narrow, entire to slightly toothed; smooth and glabrous or nearly so; petiole short to medium, purple with interwoven green strands, narrowly winged. Flowering stems 8–20 cm, pale green and pinkish towards the apex or pinkish throughout, arachnoid-hairy in upper part. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 8–11 × 2.5– 3.0 mm, dark green and scarcely pruinose or pale-margined on outer face, pale green or tinged purple on inner face, lanceolate, acute at apex, spreading to suberect and arranged at the same level in a regular manner. Inner involucral bracts 15–18 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a dark greyish-violet to purple stripe on outer face, with purple apical lobes, flat. Styles dark. Pollen present. Achenes with body 3.0–3.3 mm, straw brown, smooth, with short spiniform projections at apex; cone 0.2–0.3 mm, conical; beak 6–7 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. T. subbracteatum can be distinguished from the closely allied T. britannicum by its different-coloured outer involucral bracts and presence of spiniform projections on the achenes.

64. T. britannicum Dahlst. British Dandelion T. hibernicum G. E. Haglund Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, flat, suberect; lamina 5–20 × 2–4 cm, bluish-green on upper surface, usually purple, sometimes green on midrib, oblong in outline, divided nearly to midrib; terminal lobe triangular-subsagittate, acute and slightly elongated at apex, entire; lateral lobes 4–6 pairs, triangular, acute at apex, filiform-dentate on the proximal ones, strongly recurved; interlobe area near the

34. Taraxacum Native. Throughout Great Britain and locally abundant in areas such as western Argyllshire, Kintyre, southern Wales and Devonshire. Endemic. 66. T. oellgaardii C. C. Haw. Øllgaard’s Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, ascending; lamina 10–20 × 2–5 cm, green or yellowish-green on upper surface, dull pinkish-purple interwoven with red and green strands on midrib, oblong or elliptical-oblong in outline, divided almost to the midrib; terminal lobe medium-sized, deltoid, subacute and sometimes apiculate at apex, entire; lateral lobes 4–6 pairs, broadly triangular, acute at apex, the lower lobes with many filiform teeth on the margin towards the distal area of the leaf which is frequently more or less sigmoid; interlobe area near the midrib short, narrow and more or less entire; smooth and glabrous or nearly so; petiole about one-third of the leaf length, dull pink to purple, usually winged, sometimes broadly so. Flowering stems 10–20 cm, ascending to erect, pale green flushed purplish, arachnoidhairy becoming glabrous. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 8–10 × 2–4 mm, pale green on inner face, the outer face dark green suffused purple and with a faint to conspicuous pale margin, ovatelanceolate, acute at apex, erect or suberect. Inner involucral bracts 15–18 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a purple stripe on the outer face. Styles discoloured. Pollen present. Achenes with body 2.6–3.0 mm, straw brown, with rather fine spiniform projections above and the rest smooth; cone 0.5–0.7 mm, conical; beak 6–8 mm; pappus 7–9 mm, white. Flowers 4–5. This species resembles T. bracteatum but its leaves are paler, less coriaceous and with more patent lobes. Native. A species of rich meadow communities. Recorded from Oxfordshire, Suffolk, Gloucestershire, Breconshire, Carmarthenshire, Wigtownshire and Stirlingshire. It is probably more common. Endemic. Named after H. Øllgaard (b. 1943). 67. T. duplidentifrons Dahlst. Double-toothed Dandelion T. raunkiaeri Wiinst.; T. duplidentiforme auct. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 5–20 × 1.5– 5.0 cm, often crispate, dull green on upper surface, dark, dull pinkish-red on midrib, oblong or elliptical-oblong in outline, divided four-fifths of the way to the midrib; terminal lobe triangular, acute at apex, usually entire; lateral lobes 3–5 pairs, broadly triangular, suddenly tapering to an acute or acuminate apex, the margin towards the distal area of the leaf with filiform and triangular teeth, spreading to slightly recurved; interlobe area near the midrib short and fairly wide, entire or toothed; with short, stiff, sparse hairs; petioles rather short, dull pink to purple, usually winged, sometimes broadly so. Flowering stem 5–20 cm, ascending to erect, pale green, arachnoid-hairy. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 8–9 × 2.0–2.5 mm, dark green with a faint pale margin on outer face, pale green on inner face, linear-lanceolate, acute at apex, suberect to spreading. Inner involucral bracts

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15–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyish-violet stripe on outer face, flat, the apical lobes of the inner blackish. Styles exserted, deeply discoloured or blackish. Pollen present. Achenes with body 2.8–3.0 mm, olive brown, with spiniform projections at apex; cone 0.4–0.7 mm; beak 9–10 mm, pale; pappus 10–11 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Native. A great variety of grassy and waste places, especially on well-drained base-rich soils and locally on sanddunes. Throughout Great Britain, especially in the north and west and scattered localities round the perimeter of Ireland. It is one of our commonest dandelions. Scandinavia, Holland, Belgium and Germany. 68. T. porteri C. C. Haw. Porter’s Dandelion Medium-sized to tall, fairly robust perennial herb with a stock-like rhizome. Leaves fairly numerous, ascending to erect; lamina 10–30 cm, dull dark green on upper surface, sometimes with a dark brownish-purple blotch on the interlobe area near the midrib, dull purple with interwoven green and coloured strands on the midrib, oblong to oblanceolate in outline, divided almost to the midrib; terminal lobe medium-sized, triangular or trilobate, acute and sometimes apiculate at apex, entire; lateral lobes 4–5 pairs, spreading, triangular or deltoid, with acute, filiform apex, often with a few filiform teeth on the margin towards the distal area of the leaf; interlobe area near the midrib narrow and fairly long, entire or with a few filiform teeth; smooth and glabrous or with a few hairs; petioles up to one-third the length of the leaf, purple, unwinged. Flowering stems 10–30 cm, dull green or purple, moderately arachnoid-hairy. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 8–12 × 2–4 mm, inner face green, outer face dark green suffused reddish towards the apex, somewhat pruinose and with a pale margin, ovate or lanceolate at apex, erect. Inner involucral bracts 15–22 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a purple stripe on the outer face. Styles discoloured. Pollen present. Achenes with body 3.0–3.2 mm, pale straw brown, more or less rugose throughout and with broad-based spiniform projections at the apex; cone 0.5–0.6 mm, cylindrical-conical; beak 9–11 mm; pappus 9–12 mm, white. Flowers 4–6. Distinguished from all other British and Irish dandelions by its combination of dark green, acute leaves with deltoid or triangular lateral lobes and dark, erect outer involucral bracts. Native. Grassy banks along hedgerows. Frequent in the southern half of Wales and recorded from northern Wales and the Welsh border counties. Endemic. Named after Michael Porter. 69. T. celticum A. J. Richards Celtic Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves few, spreading or ascending; lamina 12–24 cm, bright green on upper surface, the interlobe area more or less blotched on the under surface, deep pink to purple with interwoven green strands on the midrib, often becoming green distally, oblong to elliptic-oblong, divided nearly to

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the midrib; terminal lobe triangular, the apex elongated and obtuse-mucronate, the basal corners of the triangle elongated and subacute, all margins usually entire; lateral lobes 4–5 pairs, with a broad deltoid base with 1(–3) large, acute teeth on the margin towards the distal area of the leaf and a narrow, elongated apex, spreading or pointing forwards; interlobe area near the midrib short and narrow with long, narrow teeth; smooth and glabrous or nearly so; petiole short, purple with interwoven green strands, winged, sometimes broadly so. Flowering stem 10–24 cm, pale green or suffused pink, arachnoid-hairy just below the capitulum. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 9–12 × 3–4 mm, dark green, suffused purple, somewhat pruinose and scarcely pale-margined on outer face, suffused purple on inner face, lanceolate, obtuse at apex, spreading. Inner involucral bracts 17–20 × 2.0– 2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a dark greyish-violet stripe on outer face and purple apical lobes. Styles dark. Pollen sparse or absent. Achenes with body 3.5–3.8 mm, pale brown, smooth, with short spiniform projections at apex; cone 0.3–0.4 mm, conical; beak 9–10 mm, pale; pappus 9–11 mm, whitish. Flowers 4–6. Native. On light, well-drained neutral to calcareous soils, often in species-rich grassland, but also on walls and banks of lanes. Wales and Welsh border, and south-west England, two records in northern England and a single record from Co. Kerry in Ireland. Endemic. 70. T. hesperium C. C. Haw. Western Dandelion Usually robust perennial herb with a stock-like rhizome. Leaves usually numerous, crispate, erect; lamina 10– 30 × 2–6 cm, medium green on upper surface, very conspicuously purple-blotched, both above and below, on the interlobe area near the midrib, bright purple often interwoven with green strands on midrib, oblong or narrowly oblong-elliptical in outline, divided almost to the midrib; terminal lobe medium-sized, triangular, more or less obtuse at apex, irregularly dentate; lateral lobes 5–6 pairs but often irregularly arranged along the rhachis, narrowly triangularlanceolate, with rather obtuse, linear apices, with large subobtuse teeth which often divide the lobe, more or less patent; interlobe area near the midrib narrow and long, usually toothed; smooth and glabrous or nearly so; petiole up to one-quarter the length of the lamina, bright purple, unwinged or scarcely so. Flowering stems 10–30 cm, purple, sparsely arachnoid-hairy at first, becoming glabrous. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 10–12 × 2–3 mm, pale green on inner face, the outer face darker green and often with some purplish tinting, pruinose and without or with an inconspicuous pale margin, lanceolate, acute at apex, spreading and arcuate. Inner involucral bracts 16–20 × 2–3 mm, dark green, with a pale margin, oblong-lanceolate, obtuse at apex. Ligules bright yellow, striped purple on outer face, apical lobes bright purple. Styles exserted, discoloured. Pollen present but sometimes rather sparse. Achenes with body 2.8–3.0 mm, olive brown, with spiniform projections above; cone 0.3–0.4 mm, conical, but not clearly demarcated; beak 5.5–6.5 mm; pappus 7–8 mm, white. Flowers 4–7.

This species lies midway between the Sections Celtica and Hamata and is placed in the Section Celtica only on distribution. It has some resemblance to T. lamprophyllum in the Section Hamata, but differs in the irregular lobing of the leaves with fewer and larger teeth and narrower outer involucral bracts. Native. Grassy places. Western England, Wales and south-west Scotland with a few localities in south-east England. Endemic. [Plants with narrowly lanceolate leaves like T. excellens, but more dentate than T. hesperium and less strongly blotched than either, thus approaching T. lamprophyllum, have been recorded from Somerset, Staffordshire, Denbighshire, Flintshire, Cheshire and Lancashire. They probably need describing as a new species.] 71. T. excellens Dahlst. Purple-blotched Dandelion T. magnihamatum auct. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, prostrate to ascending, flat, in a dense rosette; lamina 5–20 × 2–4 mm, green on upper surface, the interlobe area near the midrib with conspicuous purple blotches or with dark margins, deep red to purple on midrib, oblong or elliptical-oblong in outline, divided half to two-thirds of the way to midrib; terminal lobe small and neat, triangular, obtuse but sometimes mucronate at apex, sometimes with a solitary tooth, sometimes incised; lateral lobes 4–6 pairs, regular, triangular, obtuse at apex, the margin towards the distal area of the leaf sometimes with large blunt teeth, convex, sigmoid, straight-sided or concave-angled, the margin towards the proximal area convex or concave; interlobe area near the midrib short and wide, toothed; smooth and glabrous or nearly so; petiole short, pink, winged, sometimes broadly so. Flowering stem 5–20 cm, pale green, glabrous. Capitulum 30–45 mm in diameter, opening. Outer involucral bracts 8–10 × 3.0–3.5 mm, blackish-green, pruinose and scarcely pale-margined on outer face, dark green and suffused with pink on inner face, lanceolate, acute at apex, spreading. Inner involucral bracts 16–20 × 2.0–2.5 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a dark, greyish-violet stripe on outer face, flat. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with short spiniform projections at apex; cone 0.4–0.5 mm; beak 12–13 mm, pale; pappus 12–14 mm, whitish. Flowers 4–6. 2n = 24. The multicoloured green, purple and red leaves are very distinctive. Native. Scattered throughout Great Britain, but commoner in the west; a solitary record in Co. Dublin, Ireland. Sweden. 72. T. inane A. J. Richards Pollenless Dandelion T. ardlense A. J. Richards Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 15–25 × 2–5 cm, dull dark green on upper surface, the interlobe area near the midrib often somewhat blotched, dull purple on midrib, oblong or long-oblanceolate in outline, divided seven-eighths of the way to the midrib; terminal lobe helmet-shaped or

34. Taraxacum triangular- subsagittate, obtuse to acute at apex, entire; lateral lobes 3–5 pairs, broadly triangular, tapering to a narrow, acute or acuminate apex with up to 7 long, filiform teeth on the margin towards the distal area of the leaf, patent to recurved; interlobe area near the midrib short and fairly wide, toothed; smooth and glabrous or nearly so; petiole fairly long, shining purple, winged. Flowering stem 15– 25 cm, erect, pale green. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts about 10 × 3 mm, dark green and scarcely pale-margined on outer face, pale green often slightly suffused purplish on inner face, lanceolate, acute at apex, spreading or recurved. Inner involucral bracts 8–10 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyish-purple stripe on outer face and apical lobes purple. Styles exserted or more or less inserted, discoloured. Pollen absent. Achenes with body 2.8–3.0 mm, straw brown, with spiniform projections at apex; cone 0.3–0.4 mm; beak 6–8 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Native. Locally frequent in northern England and Scotland with outlying records in south and north Wales and a solitary record in Co. Wexford in Ireland. Endemic. 73. T. fulgidum G. E. Haglund Bright Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, flat, spreading to erect; lamina 8– 20 × 2–5 cm, pure bright green on upper surface, sometimes with scattered, small, dark spots and always with marked blotches on the interlobe area near the midrib, shining purple on midrib, oblanceolate in outline, divided halfway to almost to the midrib; terminal lobe large, broadly ovate, rounded at apex, entire; lateral lobes 2–3 pairs, short and triangular, acute at apex, the margin towards the distal area of the leaf strongly toothed, spreading; interlobe area short and more or less wide, toothed; smooth and glabrous or nearly so; petiole long, shining bright purple, winged, the wings toothed. Flowering stem 8–25 cm, erect, purple at least at the base and below the capitulum. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 8–10 × 2–3 mm, dark green without a pale margin on outer face and often purple-tipped, pale green on inner face, lanceolate, acute at apex, spreading to erect. Inner involucral bracts 15–20 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a brownish-purple stripe on outer face, inner ligules with purple tips. Styles exserted, darkly discoloured. Pollen present. Achenes with body 0.3–3.5 mm, olive to straw-coloured, with spiniform projections at apex; cone 0.3–0.4 mm; beak 6–7 mm, pale; pappus 7–8 mm, whitish. Flowers 4–5. Native. Damp hay meadows. Scattered localities in southern and central England and recorded for the Channel Islands and Co. Antrim in Ireland. Fennoscandia, Holland and Belgium. 74. T. tamesense A. J. Richards Thames Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves rather few, erect; lamina 4–10 × 1–3 cm, dark green with irregular purple blotches especially in the interlobe area on the upper surface, deep pink to purple on midrib, oblong-oblanceolate in outline, divided nearly

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to midrib; terminal lobe about as long as broad, hastate or triangular, obtuse at apex, entire; lateral lobes 2–3 pairs, short and wide, triangular, acute to more or less obtuse at apex, the margin towards the distal area of the leaf on the upper lobes entire, convex, sigmoid and straight-sided or concave-angled, the margin towards the distal area of the leaf of the proximal lobes dentate or occasionally provided with a single large tooth, patent or recurved; interlobe area near the midrib short and narrow, entire or toothed; smooth and glabrous; petiole long, bright purple, unwinged. Flowering stem 4–20 cm, erect, slender, often purplish, glabrous. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 6–8 × 1.5–2.0 mm, dark green suffused with purple on outer face, lanceolate, acute at apex, spreading to erect. Inner involucral bracts 14–17 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a purple stripe on the outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw brown, with spiniform projections at apex; cone 0.5–0.6 mm; beak 6–7 mm, pale; pappus 7–8 mm, whitish. Flowers 4–5. Native. Water-meadows cut for hay. Scattered localities mainly in southern England, especially the Thames valley, with isolated records in Breconshire, Leicestershire and Cumberland. Fennoscandia. 75. T. texelense Hagend., Soest & Zevenb. Texel Dandelion Small perennial herb with a stock-like rhizome. Leaves rather few, ascending or erect; lamina 5–12 × 1–3 cm, dark green on upper surface, pink to purple on midrib, oblanceolate in outline, divided nearly to midrib; terminal lobe onequarter to one-third the length of the leaf, obtuse but with a slightly extended point, entire; lateral lobes 2–4 pairs, triangular-elongate, acute at apex, the margin towards the distal area of the leaf more or less dentate, convex, patent to recurved; interlobe area near the midrib short and narrow, dentate; smooth and glabrous or nearly so; petiole short, purple, unwinged. Flowering stem 6–15 cm, purplish, glabrous. Capitulum 20–30 cm in diameter, opening. Outer involucral bracts 8–10 × 3.0–3.5 mm, dark green, pruinose and with a paler margin on outer face, pale green on inner face, spreading. Inner involucral bracts 10–15 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a reddish-violet stripe on the outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw-coloured, with spinulose projections at apex; cone about 0.6 mm; beak 6–7 mm, pale; pappus 7–8 mm, whitish. Flowers 4–5. Perhaps native. Known only from an estuarine salt-marsh in Lancashire. Holland. 76. T. hygrophilum Soest Water-meadow Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves few, erect; lamina 5–10 × 1–3 cm, pure bright green on upper surface as is the midrib; narrowly oblanceolate in outline, divided nearly to midrib; terminal lobe about as long as broad, triangular, obtuse but apiculate or slightly drawn out at apex, entire; lateral lobes 3–4 pairs, crowded, narrowly triangular, more or less acute at

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apex, entire or sometimes dentate on the margin towards the distal area of the leaf, recurved; interlobe area near the midrib very short, sometimes dentate; glabrous and smooth; petiole whitish, winged, sometimes broadly so. Flowering stem 5–8 cm, erect, thin. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 4–5 × 2.0–2.5 mm, dark green with a clear white or rose margin on outer face, pale green on inner face, ovate-lanceolate, acute at apex, erect. Inner involucral bracts 10–12 × 1.5–2.0 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on the outer face. Pollen present. Achenes with body 2.8–3.0 mm, brown, with spiniform projections at apex; cone 0.2–0.3 mm; beak 0.3–0.5 mm, pale; pappus 4.0–4.5 mm, whitish. Flowers 4–5. Native. Water-meadows. Known only from near Stodmarsh in north-east Kent. Holland, where it grows to a larger size than in England. 77. T. haematicum G. E. Haglund ex H. Øllg. & Wiezell Blood-red Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 8–15 × 1–3 cm, dark pure green on upper surface, the interlobe area near the midrib usually with dark blotches, deep pink to bright purple on midrib, oblong to oblong-elliptical in outline, divided nearly to midrib; terminal lobe triangular, and often with a narrow, obtuse, elongated apex and elongated basal corners which slope down; lateral lobes 4–5 pairs, narrowly triangular with an elongated apex, the margin towards the distal area of the leaf concave or sigmoid, entire or filiform-dentate on the proximal lobes only, patent or sloping down; interlobe area near the midrib short and narrow, entire or filiform-dentate; glabrous or nearly so; petioles short to rather long, bright purple, unwinged. Flowering stem 8–17 cm, often purplish at base, arachnoid-hairy just below the capitulum. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 7–10 × 3–4 mm, dark, shiny green on outer face, pale green and often suffused purplish on inner face especially at the tips and margins, lanceolate, acute at apex, spreading to erect. Inner involucral bracts 15–18 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the reddish apex. Ligules deep yellow, with a dark brownish-purple stripe on the outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.2 mm, olive brown, with spiniform projections at apex; cones 0.3–0.4 mm, olive; beak 5– 6 mm, pale; pappus 8–9 mm, whitish. Flowers 4–5. Obligate agamosperm. 2n = 24. Native. Damp, rich, basic grassland, particularly those cut for hay. Central-southern England particularly in the river valleys, northern coastal England, north and south Wales, Isle of Man, Ayrshire and Aberdeenshire in Scotland and Co. Dublin in Ireland. It is the most widespread of the damp grassland species and should be looked for elsewhere. It is widespread in northern and central Europe. 78. T. akteum Hagend., Soest & Zevenb. Hampshire Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves rather few, often prostrate; lamina 8–15 × 1–3

cm, dark greyish-green on upper surface, brownish on midrib, oblong or oblanceolate in outline, divided nearly to midrib; terminal lobe in outer leaves shortly deltoid, obtuse at apex and sometimes dentate, in intermediate and inner leaves subhastate and obtuse at apex or sometimes triangular and dentate; lateral lobes 2–4 pairs, deltoid, subobtuse at apex, entire or shortly dentate; interlobe area near the midrib rather short and narrow, triangular-dentate; smooth and glabrous or nearly so; petiole narrow, pale green or tinted purplish. Flowering stem 7–12 cm, glabrous or arachnoidhairy just under the capitulum. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 7–10 × 2.5– 3.5 mm, dark green on outer face with a narrow, pale margin, ovate, acute at apex, more or less appressed. Inner involucral bracts 14–16 × 2.5–3.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a dark violet or purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with short, spiniform projections at apex; cones 0.2–0.3 mm; beak 7–8 mm, pale; pappus 4.0– 4.5 mm, white. Flowers 4–5. Native. Recorded only from a single locality in a wet meadow in Hampshire. Saline meadows in Holland. 79. T. beeftinkii Hagend., Soest & Zevenb. Beeftink’s Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves few to numerous, sometimes prostrate; lamina 8–25 × 1–3 cm, greyish-green on upper surface, purplish on midrib, oblong or oblanceolate in outline, divided nearly to midrib; terminal lobe helmet-shaped, obtuse at apex, entire; lateral lobes 2–3 pairs, claw-shaped to hamate, obtuse at apex, the distal entire or often with a single tooth, the proximal regularly denticulate on the margin towards the proximal area of the leaf; interlobe area near the midrib short and entire; smooth and glabrous or slightly arachnoidhairy; petiole narrow, wine-red. Flowering stem 8–25 cm, glabrous or slightly arachnoid-hairy. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 7–10 × 2.5– 3.5 mm, dark green with a pale purplish margin, more or less ovate, acute at apex, laxly appressed. Inner involucral bracts 10–12 × 2.5–3.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules bright yellow, with a dark violet stripe on the outer face. Styles discoloured. Pollen present. Achenes with body 3.5–4.0 mm, straw brown, with sharp, spiniform projections at apex; cone 0.3–0.4 mm, beak 6–7 mm, pale; pappus 4–5 mm, whitish. Flowers 4–5. Native. Known only from salt-marsh, grassland and roadsides subject to saline flushing. Recorded only from the Essex coast. Also in similar localities in south-west Holland. Named after W. G. Beeftink. 80. T. landmarkii Dahlst. Landmark Dandelion T. firmuliforme A. J. Richards nom. nud. Medium-sized, delicate perennial herb with a stock-like rhizome. Leaves numerous, the outer prostrate, the inner suberect; lamina 5–12 × 1.5–5.0 cm, dull green on upper surface, green on midrib, oblong or narrowly oblongelliptical in outline, divided almost to midrib; terminal

34. Taraxacum lobe small, triangular and with the apex and basal corners elongate, or medium-sized and hastate, entire; lateral lobes 4–5 pairs, narrowly triangular, narrowing sometimes abruptly to a narrow, acute, elongated apex, usually entire, the margin towards the distal area of the leaf concave, straight-sided or concave-angled, the margin towards the proximal area straight, patent; interlobe area near the midrib rather long and narrow, sometimes dentate; smooth and glabrous; petioles long, purple, unwinged. Flowering stem 5–15 cm, erect, green, smooth and glabrous or nearly so. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 8–10 × 2–3 mm, dark green suffused purple without a pale margin on outer face, lanceolate, acute at apex, erecto-patent. Inner involucral bracts 13–16 × 1.4– 2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a purple stripe on outer face. Styles exserted, discoloured. Pollen absent. Achenes with body 3.0–3.4 mm, warm straw brown, with sharp spiniform projections above; cone 0.5–0.6 mm, rather narrow; beak 6–7 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 32. Native. Streamsides, ravines, wet cliffs, shingle, wet meadows and pathsides. Western and northern Great Britain, where it is commonest in the Scottish Highlands, with occasional localities in eastern and central England; scattered round the perimeter of Ireland. Also found in Norway. 81. T. nordstedtii Dahlst. Nordstedt’s Dandelion T. cambriense A. J. Richards Small to robust perennial herb with a stock-like rhizome. Leaves numerous, very variable, spreading; lamina 4–16 × 1–3 cm, medium green on upper surface, sometimes suffused with purple, sometimes with spots but usually without, green, pink or purple on midrib, oblongoblanceolate in outline, divided two-thirds of the way to the midrib or deeper; terminal lobe medium-sized, usually slightly narrower than the adjacent lateral lobes, triangular, hastate or with the apex and basal corners elongate, obtuse to subacute at apex, entire; lateral lobes 4–5 pairs, triangular, obtuse at apex, the margin towards the distal area of the leaf not toothed, sigmoid and usually characteristically concave-angled, the margin towards the proximal area straight or convex, spreading to somewhat recurved; interlobe area near the midrib quite long and wide, sometimes toothed; usually glabrous and smooth; petiole rather long, purplish or green, usually winged. Flowering stem 4–20 cm, decumbent to erect, pale green to purplish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 6–10 × 1.5–3 mm, dark green, pruinose and hardly showing a pale margin on outer face, ovate-lanceolate, obtuse at apex, erect. Inner involucral bracts 14–20 × 1.5–2.0 mm, dark bluish-green with a pale margin, oblong-lanceolate, obtuse at purplish apex. Ligules short, deep yellow, with a brown or puce stripe on outer face. Styles exserted, discoloured. Pollen usually present. Achenes with body 3.2–3.5 mm, olive brown, with rather long spiniform projections above; cone 0.3–0.4 mm; beak 8–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 48.

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Very variable in amount of anthocyanin present, leaf shape and presence of pollen. Best recognised by its dark, pruinose outer involucral bracts with scarcely a pale margin and short ligules with a brownish stripe. This is the only known hexaploid dandelion in Great Britain and Ireland. Native. Wet places, in the south-east mostly confined to meadows, but in the west occurring on walls, banks, cliffs, roadsides and waste places. Common throughout Great Britain especially in the west and north and scattered over Ireland. Western Europe, Czechoslovakia and southern Sweden. Named after Carl Fredrik Otto Nordstedt (1838– 1924). 82. T. pseudonordstedtii A. J. Richards Teesdale Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, decumbent or prostrate; lamina 5– 13 × 1–3 cm, green or dark green on upper surface, often suffused purple, purple on midrib, oblong or oblongoblanceolate in outline, divided nearly to the midrib; terminal lobe medium-sized or small, usually slightly narrower than the adjacent lateral lobes, triangular, sagittate or rhomboidal, obtuse or subacute at apex, entire; lateral lobes 4–6 pairs, regular, triangular, acute at apex, usually entire, spreading to slightly sloping down; interlobe area near the midrib short and very narrow, often reduced to the midrib; glabrous and usually smooth; petiole short, purple, usually narrowly winged. Flowering stem 5–20 cm, erect, purplish. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 8–10 × 2–3 mm, dark green and hardly showing a pale margin on outer face, lanceolate, subacute at apex, erect. Inner involucral bracts 16–18 × 1.5– 2.0 mm, dark green, oblong-lanceolate, obtuse at the reddish apex. Ligules deep yellow, short, with a purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.5 mm, greyish-brown, shortly tuberculate above; cone 0.2–0.3 mm; beak 6–7 mm, pale; pappus 7–8 mm, whitish. Flowers 5–6. Obligate agamosperm. 2n = 32. Native. Calcareous flushes from 350 to 450 m. Centred on Upper Teesdale in Co. Durham with outlying records in western Cumberland and northern Lancashire. Endemic. 83. T. berthae C. C. Haw. Bertha’s Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, spreading; lamina 5–14 × 1– 3 cm, dull dark green with black spots on upper surface, purple on midrib, oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe usually large, up to one-third as long as lamina, subsagittate, subobtuse-mucronate at apex, entire or subdivided at base; lateral lobes 2–4 pairs, triangular, acute at apex, subconvex to straight or sigmoid and entire or with sparse, filiform teeth on the margin towards the distal area of the leaf, subpatent to slightly sloping down; interlobe area near the midrib short and narrow, entire to filiform-dentate or occasionally with a larger tooth; smooth and glabrous or nearly so; petiole rather short, purple, unwinged or narrowly winged. Flowering stem 5–15 cm, often purplish, with arachnoid hairs near the capitulum to

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nearly glabrous. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 7–9 × 1.5–4.0 mm, dark green often suffused purple at apex, but usually without a pale margin on outer face, pale green on inner face, ovate, subacute and somewhat corniculate at apex, erect to appressed. Inner involucral bracts 16–18 × 2.0–2.5 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules rather pale yellow, with a dark purple stripe on the outer face. Styles exserted, discoloured. Pollen absent. Achenes with body 3.6–3.9 mm, straw brown, with broad-based spiniform projections at apex, the rest smooth; cone 0.5–0.6 mm, conical; beak 6–7 mm, pale; pappus 7–8 mm, whitish. Flowers 5–6. Native. Grassy places and gardens. South-west Scotland, north-west England and Wales. Endemic. Named after the author’s wife, Bertha Haworth. 84. T. olgae A. J. Richards Olga’s Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves rather few, erect or spreading; lamina 3–12 × 1.2–4.0 cm, green to yellowish-green and rather sparsely spotted on the upper surface, the midrib dull pinkish-purple to green, oblong or lanceolate-oblong in outline, divided almost to the midrib; terminal lobe short, triangular-subsagittate, with a slightly elongated apex, entire; lateral lobes 3–4 pairs, recurved, lanceolate, obtuse to acute at apex, margin towards the distal area of the leaf subconvex, entire or somewhat denticulate; interlobe area near the midrib short to fairly long, narrow, dentate or with a single large tooth; glabrous or nearly so; petiole short to long, pink to purple, sometimes dentate distally, unwinged. Flowering stems 3–12 cm, purplish, ascending, arachnoid-hairy. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 6–9 × 2–4 mm, green, pruinose and with a conspicuous white margin on the outer face, erect. Inner involucral bracts 16–18 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a purple stripe on the outer face. Styles exserted, dark or even blackish. Pollen absent. Achenes with body 2.6–3.2 mm, straw grey, with broad-based spiniform projections above, the rest smooth; cone 0.5–0.8 mm; beak 6–7 mm; pappus 7–8 mm, whitish. Flowers 6–7. Native. Known only from a few localities in Kirkcudbrightshire and Cleartor in Cumberland. Endemic. Named after Olga Margaret Stewart (1920–98). 85. T. cambricum A. J. Richards Welsh Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, spreading; lamina 10–25 × 1–6 cm, bright green on upper surface, deep pink to purple interwoven with green strands on midrib, oblong or oblongoblanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe medium-sized, about as long as broad, helmet-shaped, obtuse but slightly mucronate at apex, entire, slightly narrower than the adjacent lateral lobes; lateral lobes 2–4 pairs, spreading to somewhat recurved, triangular-lanceolate, obtuse at apex, the margin towards the distal area of the leaf convex, not toothed, sigmoid and straight-sided or concave-angled, the margin towards the proximal area convex and occasionally provided with a single large tooth; interlobe area near the midrib

fairly long and wide, entire; glabrous or nearly so; petiole short to medium, purple with green strands, winged, sometimes broadly so. Flowering stems 10–14 cm, pale green, but pinkish at apex, arachnoid-hairy at first becoming glabrous. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 8–13 × 2.0–3.5 mm, pale green on inner face, blackish-green and heavily pruinose on outer face, and with an insignificant pale margin, mostly erect, the lowest mostly spreading. Inner involucral bracts 16–20 × 2.0– 2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a dark grey stripe on outer face, the apical lobes yellow. Styles exserted, discoloured. Pollen usually present, rarely absent. Achenes with body 3.8–4.0 mm, fulvous to straw brown, rugose at the apex, the rest smooth; cone 05.–0.6 mm, conical; beak 6–7 mm; pappus 6–8 mm, whitish. Flowers 4–5. Native. On light, well-drained neutral to calcareous soil, often in species-rich grasslands, but also on walls and waysides and in churchyards. South-west England, Wales, north-west England, south-west Scotland and solitary records from Caithness, and from Co. Dublin in Ireland. Endemic. 86. T. lancastriense A. J. Richards Lancashire Dandelion T. angliciforme Dahlst. nom. nud. Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, spreading; lamina 8–17 × 1–4 cm, pale green and sometimes more or less spotted on upper surface, green on midrib, oblong-oblanceolate in outline, divided almost to midrib; terminal lobe large, quarter to one-third the length of the leaf, hastate, subacute at apex, entire; lateral lobes 4–5 pairs, narrowly triangular, acute at apex, entire or slightly denticulate, the margin towards the distal area of the leaf sigmoid, straight-sided or concaveangled, the margin towards the proximal area straight, spreading; interlobe area near the midrib short and narrow to wide, often dentate; smooth and glabrous or nearly so; petiole short to medium, purple, unwinged. Flowering stem 8–20 cm, erect, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 7–9 × 3–4 mm, dark green or glaucous with a clear white margin on outer face, lanceolate, subacute at apex, appressed-erect. Inner involucral bracts 16–20 × 2.0–2.5 mm, dark green, oblonglanceolate, obtuse at apex. Ligules medium yellow, short, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen absent. Achenes with body 2.8–3.0 mm, pale brown, with short, spiniform projections at apex; cone 0.3–0.4 mm; beak 6–7 mm, pale; pappus 6–8 mm, whitish. Flowers 5–6. Native. Neutral to calcareous soils, often in species-rich grasslands, also on walls, lane banks and waste places. South Wales and northern England. Endemic. 87. T. palustrisquamum A. J. Richards False Fen Dandelion Dwarf to tall perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 10–25 × 1–4 cm, green with small, sparse, blackish spots on the upper surface, green on midrib, narrowly oblong to narrowly elliptical

34. Taraxacum in outline, nearly entire and denticulate to divided threequarters of the way to midrib; terminal lobe when present triangular, acute at apex, entire; lateral lobes when present 2–3 pairs, subdeltoid, acute at apex, entire, concave on the margin towards the distal area of the leaf, spreading; interlobe area near the midrib rather long and thick, entire; smooth and glabrous or nearly so; petiole up to half the length of the leaf, whitish, narrow, unwinged. Flowering stem 10–30 cm, green, glabrous. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 6–9 × 3–5 mm, rather dark green with a broad pale white or rose-coloured margin on outer face, ovate, subacute at apex, erect to subappressed. Inner involucral bracts 16–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a purple stripe on the outer face. Styles exserted, discoloured. Pollen absent. Achenes with body 3.1–3.5 mm, straw-coloured, with short, spiniform projections above, the rest smooth; cone 0.4–0.5 mm, conical; beak 6–8 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. This species simulates T. palustre, but its achenes suggest it is best placed in this section. Native. Grass fens. Norfolk, Lancashire, East Lothian and Perthshire. Endemic. 88. R. ostenfeldii Raunk. Ostenfeld’s Dandelion T. duplidens H. Lindb.; T. parvuliceps auct.; T. biforme auct.; T. officinale subsp. ostenfeldii (Raunk.) Dahlst. Small to medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, spreading; lamina 5–18 × 1– 4 cm; lamina rather dark green on upper surface, green or pinkish on midrib, oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe small, triangular, with a slightly elongated apex, entire; lateral lobes 3–5 pairs, triangular, acute at apex, the margin towards the distal area of the leaf convex and often with 1 tooth, spreading to slightly sloping down; interlobe area near the midrib short and narrow, sometimes toothed; smooth and glabrous or nearly so; petiole rather long, purple, unwinged. Flowering stem 5–20 cm, ascending to erect. Capitulum 25–30 mm in diameter, opening. Outer involucral bracts 7–9 × 1.8–2.0 mm, dark green on outer face, pale green and often pruinose on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–18 × 1.5–2.5 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, short, with a greyish-violet stripe on outer face. Styles exserted, yellow when both fresh and dried. Pollen absent. Achenes with body 2.0–2.8 mm, brown, with spiniform projections at apex; cone 0.5–0.6 mm; beak 6–7 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. 2n = 24. Probably native. Waste places, gardens, walls and paths. Scattered throughout Great Britain, but mainly coastal; not recorded for Ireland. Western and northern Europe. Named after Carl Emil Hansen Ostenfeld (1873–1931). 89. T. breconense C. C. Haw. Brecon Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 80–150 × 1–4 cm, medium green on upper surface, green on midrib, oblong in outline, divided nearly to midrib; terminal lobe triangular, acute at apex, much subdivided and more or less den-

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tate; lateral lobes 2–4 pairs, narrowly triangular, the apex linear-acute but broader proximally to form a convex or sigmoid edge of the margin towards the distal area of the leaf when well-developed, more or less dentate only on proximal lobes; interlobe area near the midrib short and often dentate; smooth and more or less glabrous; petiole rather long, green, unwinged. Flowering stem 8–10 cm, often purplish, somewhat arachnoid-hairy at apex. Capitulum 15–20 mm in diameter, opening. Outer involucral bracts 7–10 × 2.0–2.5 mm, dark green on outer face, sometimes with a white or pink margin, greyish-green often suffused with purple on inner face, lanceolate, subacute at apex, spreading to recurved. Inner involucral bracts 16– 20 × 2.0–2.5 mm, dark green with a pale margin, oblonglanceolate, obtuse at apex. Ligules rather pale yellow, involute. Styles exserted, yellow. Pollen absent. Achenes with body 3.0–3.5 mm, straw brown, with spiniform projections at apex; cone 0.6–0.7 mm; beak 7–8 mm, pale; pappus 7– 9 mm, whitish. Flowers 4–6. Native. Known only from limestone cliffs, walls and lane banks in Breconshire. Endemic. 90. T. caledonicum A. J. Richards Caledonian Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, prostrate; lamina 4–10 × 1–3 cm, dark green on upper surface, green to pink on midrib, oblong in outline, divided nearly to midrib; terminal lobe usually short, sometimes longer, triangular, subacutemucronate at apex, entire; lateral lobes 4–6 pairs, narrowly triangular, sometimes narrowed at apex, entire, the margin towards the distal area of the leaf convex, recurved; interlobal area near the midrib rather long and narrow, sometimes with teeth; nearly glabrous; petiole long, purple, unwinged. Flowering stem 2–15 cm, ascending or erect, pale green. Capitulum 15–20 mm in diameter, convex or closed. Outer involucral bracts 9–11 × 2–3 mm, very dark purplish-green, red-tipped and pruinose with a pale margin on outer face, erect. Inner involucral bracts 16–20 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a purple stripe on the outer face, flat or sometimes involute. Styles inserted, discoloured. Pollen absent. Achenes with body 3–4 mm, greyish-brown, with few, spiniform projections above; cone 0.1–0.2 mm; beak 7–9 mm, pale; pappus 8–9 mm, whitish. Flowers 6–7. Obligate agamosperm. 2n = 40. Native. Wet cliff-faces, usually in base-rich localities and over 650 m. The Scottish Highlands of Perthshire, Forfarshire, Aberdeenshire, Morayshire, Banffshire and Inverness-shire. Endemic. 91. T. fulvicarpum Dahlst. Brown-fruited Dandelion T. hamatifrons auct. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 10–20 × 1–5 cm, pale green sometimes lightly spotted near the green midrib on the upper surface, oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe sagittate or with the apex and basal corners elongate, divided and/or dentate; lateral lobes 3–5 pairs, narrowly triangular, acute at apex, coarsely and

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narrowly dentate, straight-sided or sigmoid, sloping down; interlobe area near the midrib short and sometimes quite thick, dentate; smooth and glabrous or nearly so; petiole short to medium, pink to purple, unwinged. Flowering stem 3–10 cm, decumbent to erect. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 7–9 × 2–3 mm, pale green suffused with purple, pruinose and with a more or less obvious pale margin on outer face, ovate, obtuse at apex, erect to spreading. Inner involucral bracts 15–20 × 1.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a violet stripe on the outer face. Styles exserted, discoloured. Pollen absent. Achenes with body 3.0–3.4 mm, cinnamon to brown, with spiniform projections at apex; cone 0.6–0.7 mm; beak 7–9 mm, pale; pappus 8–10 mm, whitish. Flowers 5–6. Obligate agamosperm. 2n = 32. Native. Wet grassland, wood borders, and dune slacks. In scattered localities in Wales, central and northern England and Scotland, and Co. Clare, Co. Galway and Co. Londonderry in Ireland. It is the commonest dandelion in the Outer Hebrides. Endemic. 92. T. unguilobum Dahlst. Claw-lobed Dandelion T. chloroleucophyllum Dahlst.; T. johnstonii Dahlst.; T. unguilobiforme Dahlst.; T. serratilobum Dahlst. Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, prostrate becoming erect late in the season; lamina 5–20 × 1–4 cm, medium to yellowish-green on upper surface, pink on midrib, oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe sagittate, acute at apex, entire; lateral lobes 4–6 pairs, narrowly triangular, acute at apex, the proximal lobes filiform-dentate on the margin towards the distal area of the leaf, recurved, sometimes strongly; interlobe area near the midrib short and fairly thick, filiform-dentate; smooth and glabrous or nearly so; petiole medium, pink, winged or unwinged. Flowering stem 5–25 cm, decumbent to erect, green or pink, glabrous or arachnoid hairy at apex. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 6–8 × 2–3 mm, dark green, pink-tipped and with a clear white margin on the outer face, pale glaucous green and pruinose on inner face, lanceolate, acute at apex, spreading to erect. Inner involucral bracts 14–20 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the pinkish apex. Ligules pale yellow, with a pink stripe on the outer face. Styles inserted, yellow. Pollen absent. Achenes with body 3.0– 3.2 mm, rust-coloured, with some acute spiniform projections at apex; cone 0.4–0.5 mm; beak 6–8 mm, pale; pappus 5–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 32. Native. Wet paths, rock-faces, heaths, walls, cliff-tops, meadows, streamsides, dunes, moorlands, waysides and flushes in hilly districts. Common in the west and north of Great Britain, rare and perhaps introduced in the south and east; scattered round the perimeter of Ireland. Also occurs in Norway. 93. T. luteum C. C. Haw. & A. J. Richards Pure Yellow Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, ascending to erect; lamina 5–20 × 1–3

cm, dull greyish-green on upper surface, purple on midrib, oblong or narrowly oblong-oblanceolate in outline, divided halfway to nearly to the midrib; terminal lobe medium-sized and triangular to large and subdivided, acute at apex, with a number of recurved teeth towards the base; lateral lobes 4–5 pairs, frequently alternate, narrowly triangular and tapering to a filiform apex, the margin towards the distal area of the leaf sigmoid or straight and entire or with a few filiform teeth, recurved; interlobe area near the midrib short and narrow, somewhat crisped, entire or with filiform teeth and occasionally with a single long, narrow tooth; arachnoidhairy on upper surface; petiole long, to one-third the length of the leaf, purple, unwinged. Flowering stem 10–20 cm, arachnoid-hairy below the capitulum, pale green and pink. Capitulum 35–40 mm in diameter, opening. Outer involucral bracts 8–10 × 2–4 mm, glossy green suffused reddish towards the tip and hardly with a pale margin on outer face, pale green on inner face, lanceolate or ovate, obtuse at apex, erect or appressed. Inner involucral bracts 16–20 × 1.5– 2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules luminous yellow without a stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–3.8 mm, straw brown, with a few, narrow, recurved spiniform projections at apex; cone 0.3– 0.4 mm, beak 10–11 mm, pale; pappus 10–12 mm, whitish. Flowers 4–6. Native. In damp grasslands and on walls and wayside banks. Apparently centred in Cumberland with scattered records elsewhere in northern England, south-west Scotland, south Wales and the perimeter of Ireland. Endemic. Section 8. Hamata H. Øllg. Mostly medium-sized perennial herbs. Leaves usually dark green with a greyish-blue tint, rarely yellowish-green, the midrib usually purplish interwoven with green strands, the lateral lobes of outer leaves more or less hamate, of inner leaves hamate or subhamate, more or less glabrous; petioles purplish interwoven with green strands, narrowly winged or unwinged. Capitulum 30–55 mm in diameter. Outer involucral bracts 8–14 mm, usually bluish-green and pruinose, erect to patent or slightly recurved. Achenes with body 2.5– 4.3 mm, straw-coloured. Great Britain, Ireland, Scandinavia, Finland, north Germany, Poland, Holland, Belgium, France and Estonia. 94. T. hamatum Raunk. Hook-lobed Dandelion T. pseudohamatum auct. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 8–30 × 2–6 cm, dark olive green, sometimes suffused purple on upper surface, the interlobe area near the midrib sometimes blotched, pink to purple in exposed localities with interwoven green strands on midrib, greenish in shadier locations, oblong in outline, divided nearly to midrib; terminal lobe rather small, triangular, more or less acute at apex, usually entire; lateral lobes 3–5 pairs, regular, narrowly triangular, acute or obtuse at apex, the margin towards the distal area of the leaf convex and entire or filiformdentate, the margin towards the proximal area of the leaf straight or concave, markedly recurved; interlobe area near the midrib narrow, sometimes toothed; glabrous or nearly

34. Taraxacum so; petiole medium to fairly long, purple interwoven with green strands, unwinged at least below. Flowering stem 8– 30 cm, green or pinkish. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 8–11 × 2.0–3.5 mm, forming a stellate involucre in bud, very dark green and pruinose without a pale border on outer face, pale green on inner face, lanceolate, acute at apex, mostly erect with the lower more or less spreading. Inner involucral bracts 17–20 × 1.5–2.0 mm, dark green with a pale margin, in bud inky-black at apex, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with short, sharp spiniform projections at apex; cone 0.5–0.6 mm; beak 6–7 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Native. Natural woodland, damp grassy places, waysides, scrub, gardens and rocky places. Widespread in Great Britain, locally common in the south, very common in lowland northern England and Scotland, scattered localities in Ireland. Northern and north-western Europe and Czechoslovakia. 95. T. hamatulum Hagend., Soest & Zevenb. Slender Hook-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves rather few, decumbent to erect; lamina 10–20 × 2–6 cm, dull dark green, on the upper surface, the interlobe area near the midrib sometimes blotched, deep pink to purple and interwoven with green strands on midrib, oblong or oblongelliptical in outline, divided nearly to midrib; terminal lobe medium-sized, triangular or helmet-shaped, obtuse at apex, entire or sometimes more or less subdivided; lateral lobes regular, on inner leaves 2–3 pairs, on outer up to 5 pairs, triangular, acute to obtuse at apex, the margin towards the distal area of the leaf convex and entire or filiform-dentate, the margin towards the proximal area straight or concave, recurved; interlobe area near the midrib short and narrow, entire or toothed; glabrous or nearly so; petiole medium, purple, interwoven with green strands, unwinged. Flowering stem 10–20 cm, erect, pinkish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 9–11 × 2.5– 3.5 mm, blackish-green and markedly pruinose with a very distinct, pale, ciliate margin on the outer face, pale green on inner face, lanceolate or ovate-lanceolate, acute at apex, spreading. Inner involucral bracts 14–17 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at purplish apex. Ligules medium yellow, with a dark violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.8–4.0 mm, straw brown, with short spiniform projections at apex; cone 0.2–0.3 mm; beak 7–8 mm, pale; pappus 8–9 mm, whitish. Flowers 4–6. Native. North and south Wales, scattered records in southern England, frequent in north-east England, with solitary records in Wigtownshire, Dumbartonshire and the Shetland Islands. Denmark, Belgium and France. 96. T. subhamatum M. P. Christ. Large Hook-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 10–30 ×

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2–8 cm, dull olive green on upper surface, the interlobe area near the midrib sometimes blotched, green to faintly pinkish with green strands on midrib, oblong or oblongelliptical in outline, divided two-thirds of the way to nearly all the way to the midrib; terminal lobe usually rather small, triangular or sagittate, usually with an elongated or apiculate apex, entire; lateral lobes 4–7 pairs, narrowly triangular, obtuse or narrowing to a long-acute apex, the margin towards the distal area of the leaf convex, straight, sinuose or concave and slightly dentate proximally, the proximal margin towards the proximal area of the leaf concave straight or convex; interlobe area near the midrib short and thick, dentate; glabrous or nearly so; petiole rather short, purple with interwoven green strands, narrowly winged. Flowering stem 10–30 cm, green or flushed pink, ascending. Capitulum 30–50 mm in diameter, opening. Outer involucral bracts 10–11 × 2.5–3.5 mm, dark green and densely pruinose on outer face with a slight pale margin, pale green lightly suffused purple on inner face, lanceolate or ovatelanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–18 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the purplish apex. Ligules deep yellow, with a greyish-violet stripe on outer face and orange lobes at apex. Styles exserted, discoloured. Pollen present. Achenes with body 3.1–3.7 mm, straw-coloured, with thick spiniform projections at apex; cone 0.4–0.5 mm; beak 6–7 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = ±24. Native. Grassy places, waysides, scrub, waste places, gardens and rocky places. Scattered over Great Britain and round the perimeter of Ireland. Denmark and Sweden. 97. T. marklundii Palmgr. Marklund’s Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, decumbent to erect; lamina 5– 20 × 1–5 cm, dull, often dark green on upper surface, pale brown, mauve or purple often interwoven with green strands on midrib, oblong in outline, divided nearly to midrib; terminal lobe usually rather small, sagittate, elongated at apex, often dentate; lateral lobes 3–5 pairs, regular, narrowly triangular, with a narrow, elongated apex, the margin towards the distal area of the leaf convex, usually filiform-dentate and often with 1 or 2 large teeth, the margin towards the proximal area of the leaf concave and usually recurved, sometimes acutely so, or patent at first and abruptly recurved at the apex; interlobe area near the midrib short and narrow, often toothed; glabrous or nearly so; petiole short to medium, purple with interwoven green strands, unwinged. Flowering stem 5–20 cm, often purplish, glabrous. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 8–10 × 2.0–2.5 mm, dark green suffused purple and without a pale margin on outer face, purplish on inner face, lanceolate, obtuse at apex, spreading to recurved. Inner involucral bracts 18–20 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face and orange lobes at apex. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw brown, with spiniform projections at apex; cone 0.4–0.5 mm; beak

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7–9 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Closely related to T. subhamatum and sometimes included in it. It is distinctive in its extreme form by its markedly recurved, regular, very dentate leaf-lobes but is connected to it by a range of intermediates. Native. Grassy places, water-meadows, tracks, roadsides, shingle and waste places. Scattered throughout Great Britain and in Co. Wexford in Ireland. Northern and western Europe. Named after Gunnar Georg Marklund (1892–1964). 98. T. hamiferum Dahlst. Dark Hook-lobed Dandelion T. atrovirens Dahlst. Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, spreading or erect; lamina 10– 20 × 1–5 cm, dark green on the upper surface, the interlobe area near the midrib sometimes blotched, deep pink to purple with interwoven green strands on midrib, oblong in outline, divided nearly to midrib; terminal lobe mediumsized, triangular or helmet-shaped, obtuse at apex, entire; lateral lobes of inner leaves 2–3 pairs, of outer leaves up to 5 pairs, regular, triangular-hamate, acute to somewhat obtuse at apex, the margin towards the distal area of the leaf convex and entire or filiform-dentate, the margin towards the proximal area of the leaf straight or concave, recurved; interlobe area near the midrib short and narrow, sometimes toothed; glabrous or nearly so; petiole short, purple, interwoven with green strands, unwinged proximally. Flowering stem 10–20 cm, tinted purplish, erect. Capitulum 30– 40 mm in diameter, opening. Outer involucral bracts 9– 10 × 2.0–2.5 mm, blackish-green, pruinose and with a slight pale margin on outer face, pale green on inner face, lanceolate, obtuse at apex, spreading. Inner involucral bracts not equally wide and usually irregularly connate, splitting as the capitulum opens, 18–20 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with spiniform projections at apex; cone 0.5–0.6 mm; beak 7–9 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. 2n = 24. Doubtfully native. Scattered records throughout Great Britain and mainly round the perimeter of Ireland. Denmark. 99. T. quadrans H. Øllg. Fleshy-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 5–25 × 1–4 cm, thick and fleshy, dull, dark bluish-green on upper surface, the interlobe area near the midrib sometimes blotched, deep pink to purple interwoven with green strands on midrib, oblongoblanceolate in outline, divided seven-eighths of the way to the midrib; terminal lobe medium-sized, triangular to helmet-shaped, obtuse and sometimes apiculate at apex, entire or with 1–2 teeth; lateral lobes 3–5 pairs, regular, triangular, acute to obtuse at apex, the margin towards the distal area of the leaf convex, entire or filiform-dentate and frequently and characteristically angled, the margin towards the proximal area of the leaf straight or concave, recurved;

interlobe area near the midrib short and rather thick, entire or filiform-dentate; glabrous or nearly so; petiole purple interwoven with green strands, unwinged. Flowering stem 5–25 cm, tinted pinkish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 9–11 × 3.0–3.5 mm, blackish-green and strongly pruinose with a distinct, narrow, pale margin on the outer face, pale green on inner face, spreading. Inner involucral bracts 13–16 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyish-violet stripe on the outer face. Styles exserted, faintly discoloured. Pollen present. Achenes with body 3.8–4.0 mm, straw brown, with spiniform projections at apex; cone 0.2–0.3 mm; beak 7–9 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Agamosperm. Probably not native. Scattered records throughout Great Britain with a few localities round the perimeter of Ireland. Denmark and Czechoslovakia. 100. T. pseudohamatum Dahlst. False Hook-lobed Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 15– 40 × 1–5 cm, dull olive green on upper surface, the interlobe area near the midrib sometimes blotched, purple interwoven with green strands on midrib, oblong in outline, divided nearly to midrib; terminal lobe triangular or sagittate, obtuse or subacute at apex, entire; lateral lobes 3–6 pairs, triangular, obtuse to acute at apex, the margin towards the distal area of the leaf more or less convex and sometimes filiform-dentate, the margin towards the proximal area of the leaf concave, straight or convex, spreading to recurved; interlobe area near the midrib short and fairly wide, more or less dentate; glabrous or nearly so; petiole medium, purple interwoven with green strands, more or less winged. Flowering stem 15–40 cm, purplish-tinted, glabrous. Capitulum 50–55 mm in diameter, opening. Outer involucral bracts 10–12 × 3.5–5.0 mm, dark green, pruinose and with a distinct pale margin on outer face, paler green on inner face, lanceolate or ovate-lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–20 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face and orange lobes at apex. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with spiniform projections at apex; cone 0.5–0.6 mm; beak 9–11 mm, pale; pappus 7–9 mm, whitish. Flowers 3–6. Agamosperm. Native. A weedy species on roadsides, in grassy places, scrub, waste land, gardens and rocky places. Throughout Great Britain and with scattered records in Ireland. Denmark, France and Czechoslovakia. 101. T. fusciflorum H. Øllg. Brown-striped Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 30–40 × 1–5 cm, dull greyish-green on upper surface, the interlobe area near the midrib sometimes blotched, purple interwoven with green strands on midrib, narrowly oblongelliptical in outline, divided nearly to midrib; terminal lobe

34. Taraxacum of outer leaves small, sagittate-cordate, obtuse at apex and often dentate, those of the middle and inner leaves larger and usually dentate; lateral lobes 5–6 pairs, triangular, obtuse to subacute at apex, the margin towards the distal area of the leaf convex, entire or on inner leaves dentate; interlobe area near the midrib short and fairly thick, more or less dentate; glabrous or nearly so; petiole medium, purple interwoven with green strands, not or narrowly winged. Flowering stem 30–40 cm, purplish, glabrous. Capitulum 40– 50 mm in diameter, opening. Outer involucral bracts 12– 14 × 4–5 mm, dark green and strongly pruinose with a pink tip and no pale margin on outer face, greyish-green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–20 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 4.0–4.3 mm, dark straw-coloured, with strong, recurved, spiniform projections; cone 0.6–0.7 mm; beak 7–10 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Scattered records in England, Wales and southern Scotland and two in the west of Ireland. Denmark, Finland, Germany and Czechoslovakia. 102. T. prionumHagend., Soest & Zevenb. Saw-leaved Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 15–25 × 1– 3 cm, dull dark green on upper surface, the interlobe area near the midrib sometimes blotched, purple with interwoven green strands on midrib, oblong in outline, divided nearly to midrib; terminal lobe small, triangular, with an elongated, subacute apex; entire; lateral lobes 4–5 pairs, broadly triangular or deltoid, subacute to acute at apex, the margin towards the distal area of the leaf straight or convex and sometimes filiform-dentate, the margin towards the proximal area of the leaf straight and sometimes dentate; interlobe area near the midrib fairly long and thick and more or less dentate; rather hairy; petiole medium, purple interwoven with green strands, winged at least below. Flowering stem 15–25 cm, erect, thickly arachnoid-hairy when young. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–13 × 3.5–4.5 mm, dark greyish-green with a distinct pale border on outer face, paler green on inner face, lanceolate, subacute at apex, spreading to recurved. Inner involucral bracts 15–17 × 1.5–2.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with spiniform projections at apex; cone 0.6–0.7 mm; beak 9–10 mm, pale; pappus 5– 6 mm, whitish. Flowers 4–6. Probably introduced. A few scattered records from England, Wales and southern Scotland. Holland and Belgium. 103. T. boekmanii Borgv. B¨okman’s Dandelion T. grandifolium G. E. Haglund, non H. Koidz. Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, decumbent to erect; lamina 15– 25 × 1–4 cm, dark green on upper surface, bright purple

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to the apex and usually with some red veining on midrib, oblong-oblanceolate, divided two-thirds of the way to the midrib; terminal lobe equalling or larger than the lateral lobes, helmet-like, obtuse to rounded at apex, entire; lateral lobes 3–5 pairs, regular, triangular, obtuse at apex, the margin towards the distal area of the leaf convex or straight and entire or filiform-dentate, the margin towards the proximal area of the leaf straight or concave, spreading or recurved; interlobe area near the midrib short and thick, entire or dentate; slightly hairy; petiole fairly long, brilliant purple, narrowly winged. Flowering stem 15–25 cm, tinted reddish. Capitulum 35–40 mm in diameter, opening. Outer involucral bracts 10–12 × 2.5–5.5 mm, dark blackish-green grading to pruinose without a pale border on outer face, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–4.0 mm, straw-coloured, with sparse spiniform projections at apex; cone 0.4–0.5 mm; beak 6– 8 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Agamosperm. 2n = ?24. Probably native. Grassy places, waysides, waste places, gardens and open places in natural woodlands. Scattered localities throughout Great Britain but rare in northern Scotland. Recorded for Co. Dublin and Co. Antrim in Ireland. Sweden and Czechoslovakia. Named after Krister B¨okman. 104. T. atactum Sahlin & Soest Narrow-bracted Dandelion Small to medium-sized perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 5– 20 × 1–4 cm, rather dark greyish-green on upper surface, the interlobe area sometimes blotched, greenish to faintly purple with interwoven green strands on midrib, oblongoblanceolate in outline, divided nearly to midrib; terminal lobe large, helmet-shaped, obtuse or acute at apex, with 1 or 2 distinct teeth; lateral lobes 2–3 pairs, regular, triangular, acute to subacute at apex, the margin towards the distal area of the leaf straight or convex and filiform-dentate proximally, the margin towards the distal area straight or concave, recurved; interlobe area near the midrib short and usually narrow, entire or dentate; glabrous or nearly so; petiole short to medium, purple with interwoven green strands, unwinged at least below. Flowering stem 5–25 cm, tinted reddish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 10–12 × 2–3 mm, very dark green grading to pruinose and without a pale margin on outer face, green to purplish on inner face, lanceolate, acute at apex, erect to spreading. Inner involucral bracts 16–20 × 2.0–2.5 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.2–3.5 mm, straw-coloured, with short, sparse spiniform projections at apex; cone 0.4–0.6 mm; beak 6– 8 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. 2n = 24. Possibly native in the south. Grassy places, waysides and waste places. Scattered localities throughout Great Britain

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and locally common; recorded from Co. Galway, Co. Dublin and Co. Sligo in Ireland. Holland, Denmark and Sweden. 105. T. sahlinianum Dudman & A. J. Richards Sahlin’s Dandelion Medium-sized perennial herbs with a stock-like rhizome. Leaves rather few, decumbent to erect; lamina 5–20 × 1–4 cm, dirty dark green on upper surface, deep pink to purple with interwoven green strands on midrib, oblongoblanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe medium-sized, about as long as broad to twice as long as broad, helmet-shaped, roundedobtuse-mucronate at apex, often with 1 or 2 large teeth; lateral lobes 2–4 pairs, triangular, subacute at apex, the margin towards the distal area of the leaf dentate, more or less convex or sometimes straight on the lower lobes, the lower lobes with filiform teeth, patent to somewhat recurved; interlobe area near the midrib short and fairly thick, dentate; glabrous or nearly so; petiole fairly long, purple with interwoven green strands, winged. Flowering stem 8–20 cm, ascending to erect. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 8–10 × 2–3 mm, dark green and scarcely with a pale margin on outer face, pale green on inner face. Inner involucral bracts 16–20 × 2.0–2.5 mm, dark green with pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a dark greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.8–4.0 mm, fulvous to straw brown, with spiniform projections at apex; cone 0.5–0.6 mm; beak 6–8 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Native. Scattered localities in western and northern Britain; recorded from Co. Wexford in Ireland. Endemic. Named after Carl Ingemar Sahlin (1912–90). 106. T. pruinatum M. P. Christ. Pruinose Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 15–30 × 1–4 cm, dull blue-grey-green on upper surface, green on midrib, oblong or narrowly oblong-elliptical in outline, divided nearly to midrib; terminal lobe sagittate, acute at apex, entire; lateral lobes 3–5 pairs, regular, narrowly triangular, acute at apex, the margin towards the distal area of the leaf straight or convex and entire or on proximal lobes filiform-dentate, the proximal margin straight or concave, patent or recurved; interlobe area near the midrib rather long and narrow, filiform-dentate; glabrous or nearly so; petiole long, very pale pink interwoven with green strands, unwinged at least below. Flowering stem 15–30 cm, sometimes pinkish. Capitulum 30–45 mm in diameter, opening. Outer involucral bracts 10–11 × 1.5–3.5 mm, pale green or whitish-green and with a strongly pruinose white margin, lanceolate, acute at apex, mostly erect but the lower more or less spreading. Inner involucral bracts 16–18 × 2.0– 2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with short spiniform projections at apex; cone 0.7–0.8 mm; beak 10–11 mm, pale; pappus 9–11 mm, whitish. Flowers 4–6.

Recorded only for Denbighshire, Flintshire and Cheshire. Denmark. 107. T. hamatiforme Dahlst. Asymmetrical Hook-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, spreading to erect; lamina 8– 20 × 1–4 cm, medium to dark bluish-green on upper surface, the interlobe area near the midrib often dark-blotched, purple with interwoven green strands on midrib, oblong or narrowly elliptic-oblong in outline, divided nearly to midrib; terminal lobe triangular-sagittate, with a more or less elongated apex, entire; lateral lobes 3–5 pairs, asymmetrical in arrangement, narrowly triangular-acute at apex, the margin towards the distal area of the leaf more or less straight or concave and dentate or filiform-dentate, the margin towards the proximal area straight or concave, patent or recurved; interlobe area near the midrib fairly long and narrow, often with teeth; glabrous or nearly so; petiole deep purple interwoven with green strands, unwinged at least below. Flowering stem 8–20 cm, sometimes tinted purplish. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 10–11 × 2–3 mm, very dark, shiny brownishgreen on outer face, with an indistinct pale margin, pale green to glaucous on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 15–20 × 2.0– 2.5 cm, dark green with a pale margin, oblong-lanceolate, obtuse at the dark purplish apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw-coloured, with spiniform projections at apex; cone 0.5–0.6 mm; beak 6–7 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 23, 24, 25, 26, 27. Native. Grassy places, waysides, waste places and hedgebanks. Throughout Great Britain and scattered records in Ireland. Northern and western Europe and Czechoslovakia. 108. T. spiculatum M. P. Christ. Sagittate-lobed Dandelion T. subditivum Hagend., Soest & Zevenb. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 8–20 × 1–4 cm, medium to dark bluish-green on upper surface, the interlobe area near the midrib often dark-blotched, purple to the apex with interwoven green strands on midrib, oblong in outline, divided nearly to the midrib; terminal lobe triangular-sagittate, with elongated apex, entire; lateral lobes 3–5 pairs, more or less regular, triangular, with a long, narrow, tapering acute apex, the margin towards the distal area of the leaf straight, sinuate to convex and entire to filiform-dentate, the margin towards the proximal area more or less straight, patent or recurved sometimes sharply so; interlobe area near the midrib long and narrow, sometimes toothed; glabrous or nearly so; petiole rather short, deep purple with interwoven green strands, unwinged at least below. Flowering stem 8–20 cm, sometimes pink tinted. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 10–11 × 2–3 mm, very dark, shiny brownish-green on outer face, with an indistinct pale margin, pale green to glaucous on inner face, lanceolate, obtuse at apex, spreading to

34. Taraxacum recurved, the lower often rudimentary or scaly. Involucral bracts 18–20 × 2.0–2.5 mm, dark green with a pale margin oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.2 mm, straw brown, with thick spiniform projections at apex; cone 0.5–0.6 mm; beak 5–6 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Native. Scattered records in western England and Dumfries-shire and Kincardineshire in Scotland. Denmark. 109. T. lancidens Hagend., Soest & Zevenb. Spear-toothed Dandelion Medium-sized, robust perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 8– 25 × 1–5 cm, medium green on upper surface, the interlobe area near the midrib often blotched, purple with interwoven green strands on the midrib, oblong in outline, divided nearly to midrib; terminal lobe sagittate, with an elongated apex, entire; lateral lobes 3–5 pairs, narrowly triangular, the apex long-acute or acuminate, the margin towards the distal area of the leaf straight or concave and frequently with 1 large tooth, the margin towards the proximal area straight or concave, sometimes dentate, patent or recurved; interlobe area near the midrib long and narrow, sometimes dentate; glabrous or nearly so; petiole deep purple interwoven with green strands, unwinged at least below. Flowering stem 6–25 cm, sometimes purplish. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 10–11 × 2–3 mm, dark olive green on outer face, without or with only a faint pale margin, pale greyish-green on inner face, lanceolate or ovate, acute at apex, spreading to recurved. Inner involucral bracts 15–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a dark brown stripe on the outer face and apices of inner tipped purple. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, pale straw brown, with dense, short spiniform projections at apex; cone 0.6–0.7 mm; beak 10–11 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Recorded for Dorset, Denbighshire, Flintshire and Cheshire and by the describing authors from Surrey and Kent. Holland, Germany and Czechoslovakia. 110. T. kernianumSoest, Hagend. & Zevenb. Kern’s Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 8–20 × 1–5 cm, medium green on upper surface, the interlobe area near the midrib blotched, purple with interwoven green strands on the midrib, oblong in outline, divided nearly to midrib; terminal lobe triangular with elongated apex and basal corners, frequently subdivided; lateral lobes 3–5 pairs, irregular, rapidly narrowing from a broad base, strongly toothed and often divided, sometimes deeply so, acute or acuminate at apex, the margin towards the distal area of the leaf straight or sinuate and entire or dentate, the margin towards the proximal area straight or concave, patent or recurved; interlobe area near the midrib long and fairly thick, usually toothed, the teeth sometimes large; glabrous or nearly so; petiole deep purple with interwoven green

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strands, unwinged. Flowering stem 8–20 mm, sometimes purplish. Capitulum 35–50 mm in diameter, opening. Outer involucral bracts 10–11 × 3–4 mm, dark green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 18–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a dark violet stripe on outer face. Styles exserted, discoloured. Pollen more or less present. Achenes with body 2.5–3.0 mm, pale straw-coloured, with spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. 2n = ±24. Introduced. Scattered records throughout Great Britain, but missing from the Scottish Highlands and Ireland. Holland and Belgium. Named after Johannes Hendrikus Kern (1903–68). 111. T. lamprophyllum M. P. Christ. Lustrous-leaved Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 18– 40 × 1–4 cm, dark green on upper surface, the interlobe area near the midrib usually heavily blotched, purplish with interwoven green strands on the midrib, oblong or narrowly oblong-elliptical in outline, divided almost to midrib; terminal lobe sagittate with an elongate apex or triangular and obtuse at apex, often dentate; lateral lobes 5–6 pairs, triangular-hamate, acute or acuminate at apex, obtuse when the lobe is entire, the margin towards the distal area of the leaf straight, concave or angled, usually very dentate and frequently with at least 1 large tooth, the margin towards the proximal area straight or concave and sometimes dentate; interlobe area near the midrib long and fairly thick, more or less dentate; glabrous or nearly so; petiole purple interwoven with green strands, unwinged at least below. Flowering stem 16–40 cm, sometimes purplish. Capitulum 40– 50 mm in diameter, opening. Outer involucral bracts 11– 13 × 3–5 mm, dark olive green on outer face with a pink tip and narrow, white margin, suffused purplish and pruinose on inner face, ovate-lanceolate at apex, spreading to recurved. Inner involucral bracts 17–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a dark brown stripe on the outer face. Styles exserted and discoloured. Pollen present. Achenes with body 3.0–3.6 mm, pale straw-coloured, with patent spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 8–11 mm, whitish. Flowers 4–9. 2n = ± 24. Agamosperm. Perhaps introduced. Scattered records throughout England and Wales, rather rare in Scotland, and the perimeter of north and east Ireland. Denmark and Czechoslovakia. ˇ ep´anek Section 9. Ruderalia Kirschner, H. Øllg. & Stˇ Taraxacum subsection Palustriformia Sahlin; Taraxacum Section Vulgaria (Dahlst.) Dahlst., non Series Vulgaria Hand.-Mazz. Usually robust perennial herbs. Leaves pale to dark green, sometimes blotched or spotted, deeply lobed, the lobes often complex and dentate, the midrib green or purplish; often hairy; petiole green or violet, often winged. Capitulum large. Outer involucral bracts recurved or reflexed. Achenes

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with body (2.5–)3.2–3.6(–4.3) mm, with short, broad-based cones, the beak usually 10–13 mm. Very common plants of grassland, waste places, paths, roadsides and walls. A few species are native and endemic, but most are weedy plants which are confined mainly to the northern part of the temperate region of Europe. Two sets of leaves are produced each year. Only the small overwintering ones which develop as the spring leaves at the time of flowering are reliable for classification. The second set which develop during the summer and are killed by autumn frost are unreliable for identification which should not be attempted after the end of May. 112. T. laeticolor Dahlst. Pale-stalked Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves rather few, flat, erect; lamina 8–20 × 1–4 cm, often pale to medium green on upper surface, green or brownish on midrib, oblong-oblanceolate in outline, divided threequarters of the way to the midrib; terminal lobe large, especially on inner leaves, helmet-shaped, obtuse to subacute at apex, entire or once or twice shallowly dissected, on outer leaves triangular and less obviously larger than lateral lobes; lateral lobes 4–6 pairs, subdeltoid to triangular, acute at apex, subentire or with a few large teeth, the margin towards the distal area of the leaf straight, slightly concave or sigmoid, the margin towards the proximal area more or less straight, patent or twisted in all directions; interlobe area near the midrib short and thick, entire or dentate; glabrous or nearly so; petiole short to medium, green, usually winged. Flowering stem 7–20 cm, pale green below, slightly purplish above, slightly arachnoid-hairy above. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 10–11 × 3–4 mm, dark green usually suffused purplish on outer face, pink-tipped, and hardly with a pale margin, ovate-lanceolate, acute at apex, erect to spreading. Inner involucral bracts 18–20 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the slightly purplish apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen sparse. Achenes with body 3.8–4.0 mm, straw-coloured, with short spiniform projections at apex; cone 0.6–0.8 mm; beak 8– 10 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Probably introduced. Grassy places and waysides. Scattered records in England and Wales, rare in Scotland. Belgium, Germany, and Fennoscandia. 113. T. macrolobum Dahlst. Incise-lobed Dandelion T. lacerabile Dahlst. Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, crispate, erect; lamina 10–30 × 1– 4 cm, medium to pale green on upper surface, green or brownish on midrib, the interlobe area near the midrib darkmargined, oblong-elliptical in outline, much divided nearly to midrib; terminal lobe narrowly triangular with elongated, obtuse to subacute, ligulate apex and basal corners larger and helmet-shaped, entire or subdivided; lateral lobes 4– 6 pairs, abruptly narrowed into linear, acute or swollen obtuse elongations, entire or subdivided, the proximal lobes dentate and sometimes with a few large teeth, the margin towards the distal area of the leaf sigmoid, the margin towards the proximal area more or less straight, spreading or

forward-pointing; interlobe area near the midrib short and narrow, entire or dentate especially at the base of the lateral lobes; glabrous or nearly so; petiole rather short, green, narrowly winged. Flowering stem 10–30 cm, green or slightly pink at base, slightly arachnoid-hairy above. Capitulum 50–55 mm in diameter, opening. Outer involucral bracts 10–11 × 3–4 mm, dark green usually suffused purplish on outer face, pink-tipped and hardly with a paler margin, paler green on inner face, lanceolate-ovate, acute at apex, erect to spreading. Inner involucral bracts 14–15 × 2.0–2.5 mm, dark green with pale margin, oblong-lanceolate, obtuse at the rose apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw brown, with spinulose projections at apex; cone 0.2–0.3 mm; beak 8–10 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Probably introduced. Grassy places and waysides. Scattered records in England, Wales and west Scotland and Co. Antrim in Ireland. Scandinavia and Belgium. 114. T. pannucium Dahlst. Green-stalked Dandelion T. protractifrons Dahlst. ex. M. P. Christ. & Wiinst. Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, flat, erect; lamina 15–30 × 1–4 cm, pale green on upper surface, green or faintly purplish on midrib, narrowly oblong-elliptical in outline, divided nearly to midrib; terminal lobe narrowly triangular, narrowly elongated at apex, sometimes subdivided; lateral lobes 4–6 pairs, abruptly narrowed from a broad base to a narrow, elongated, acute apex, the margin towards the distal area of the leaf concave or angled and denticulate, the margin towards the proximal area straight or convex, erect-spreading; interlobe area near the midrib short and narrow, usually dentate; glabrous or nearly so; petiole one-third to half the length of the lamina, green, narrowly winged to more or less unwinged. Flowering stem 15–30 cm, green. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 10–14 × 3–4 mm, dark green and sometimes violet-tipped on outer face and without a pale margin, paler green on inner face, lanceolate or ovate-lanceolate, acute at apex, erectspreading. Inner involucral bracts 18–22 × 1.5–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a brownish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–2.8 mm, straw brown, with slender, sharp spiniform projections at apex; cone 0.7–0.8 mm; beak 8.10 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. 2n = 24. Probably introduced. Grassy places, waysides, scrub and waste land. Scattered localities throughout Great Britain and locally common; perimeter of the northern half of Ireland. Fennoscandia, Belgium and Holland. 115. T. subexpallidum Dahlst. Tongue-lobed Dandelion T. linguatum Dahlst. ex M. P. Christ. & Wiinst.; T. sublaciniosum auct. Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, erect; lamina 10–30 × 2–7 cm, medium green on upper surface, green to slightly purplish on midrib, oblong or elliptic-oblong in

34. Taraxacum outline, divided nearly to midrib; terminal lobe of outer leaves triangular, somewhat waisted, with an elongated, lingulate, obtuse tip and sometimes subdivided, of inner leaves usually large, triangular or helmet-shaped, obtuse but frequently apiculate at apex, entire to dentate and sometimes subdivided and with a cordate base; lateral lobes 4– 7 pairs, deltoid or narrowly triangular, acute to obtuse at apex, both margins more or less straight or angled usually to give a hooked appearance but sometimes with a forward-pointing elongation, the margin towards the distal area of the leaf dentate to filiform-dentate or rarely entire, the margin towards the proximal area entire, patent to slightly recurved; interlobe area fairly long and narrow, often dentate; glabrous or nearly so; petiole short to medium, pale green or somewhat pink on inner leaves, unwinged at least below. Flowering stem 10–30 cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–14 × 2.5–4.5 mm, dark green on outer face without a pale margin, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 15–22 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, dark straw brown, with numerous, slender, sharp spiniform projections at apex; cone 0.8–1.0 mm; beak 8–10 mm, pale; pappus 6–10 mm, whitish. Flowers 4–6. Probably introduced. Grassy places, waysides, scrub and waste places. Scattered records throughout Great Britain and locally common; recorded from Co. Antrim in Ireland. Northern and western Europe and Switzerland. 116. T. corynodes G. E. Haglund Dense-lobed Dandelion Medium-sized to robust, squat perennial herb with a stocklike rhizome. Leaves numerous, highly crispate, erect; lamina 10–20 × 2–4 cm, medium to greyish-green on upper surface, green on midrib, oblong in outline, divided twothirds of the way to the midrib; terminal lobe usually small, triangular, obtuse to acute at apex, often subdivided; lateral lobes 4–8 pairs, crowded, triangular, narrowing from a broad base to a subacute to obtuse apex, sometimes subdivided, the margin towards the distal area of the leaf convex or sigmoid, often strongly humped and entire or dentate, the margin towards the proximal area concave or convex and often sinuate, patent and multidirectional; interlobe area near the midrib short and very thick giving a broad area along the midrib, often dentate; glabrous or nearly so; petiole rather long, green, broadly winged. Flowering stem 10–20 cm, pale green or pink-tinted. Capitulum 40–50 mm in diameter. Outer involucral bracts 11–15 × 4– 5 mm, dark green on outer face and more or less without a pale margin, greyish-green on inner face, lanceolate, acute at apex, spreading or recurved. Inner involucral bracts 18–20 × 3–5 mm, dark green with a pale margin, oblonglanceolate, obtuse at apex. Ligules medium yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with short spiniform projections at apex; cone 0.4–0.5 mm; beak 10–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm.

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Introduced. Grassy places, waysides and waste places. Northern and southern Wales and the Welsh border with outlying records in Devonshire, Derbyshire, Lancashire and Yorkshire. Native of Denmark, Sweden and Czechoslovakia. 117. T. undulatum H. Lindb. & Markl. Wavy-leaved Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, highly crispate, erect; lamina 10–30 × 4–10 cm, fleshy, waxy, dark green on upper surface, green on midrib, broadly oblong-oblanceolate in outline, divided one-third to two-thirds of the way to the base; terminal lobe small to medium, triangular, obtuse to acute at apex, often subdivided; lateral lobes 4–6 pairs, crowded, triangular, narrowed from a broad base to a more or less acute apex, frequently giving the effect of a high shoulder, the margin towards the distal area of the leaf convex and entire to dentate, the margin towards the proximal area concave, recurved; interlobe area near the midrib short and thick making a wide area of midrib, often dentate; glabrous or nearly so; petiole rather short, green, broadly winged. Flowering stem 10–40 cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–15 × 4– 5 mm, dark green on outer face, with a narrow, pale margin, greyish-green on inner face, lanceolate, more or less acute at apex, erect to spreading. Inner involucral bracts 15–20 × 2–3 mm, dark green with a pale margin, oblonglanceolate, obtuse at apex. Ligules medium yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with spiniform projections at apex; cone 0.4–0.5 mm; beak 7–10 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. 2n = 24. Probably native. Grassy places, waysides and waste land. Frequent in south Wales, scattered elsewhere in Wales and in east and north England; in Wigtownshire in Scotland and Co. Dublin in Ireland. Fennoscandia, Belgium, France and Czechoslovakia. 118. T. tenebricans (Dahlst.) Dahlst. Shiny-leaved Dandelion T. officinale subsp. tenebricans Dahlst. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, highly crispate, erect; lamina 10–20 × 1– 4 cm, medium shiny green on upper surface, and highly crisped, green on midrib, oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe small to mediumsized, more or less triangular, acute at apex, entire; lateral lobes 4–6 pairs, deltoid or triangular, narrowed from a broad base to a patent or forward-pointing, acute apex, often subdivided, the margin towards the distal area of the leaf convex or straight and entire or dentate, the margin towards the proximal area more or less straight; interlobe area near the midrib short and fairly thick, often dentate, glabrous or nearly so; petiole short, green, unwinged. Flowering stem 10–35 cm, green, erect. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–15 × 4–5 mm, dark green on outer face, without or with a narrow, pale margin, paler green on inner face, lanceolate, acute at

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apex, erect to appressed. Inner involucral bracts 15–20 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyishviolet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with short sharp spiniform projections at apex; cone 0.4–0.5 mm; beak 7–10 mm, pale; pappus 6–8 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Grassy places, waysides and waste places. A few scattered records in Great Britain from Somerset to the Shetland Islands. Fennoscandia, Belgium, Holland and Czechoslovakia. 119. T. dilaceratum M. P. Christ. Lacerate-leaved Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, crispate, erect; lamina 10– 25 × 2.5–6.0 cm, medium green on upper surface, green on midrib, narrowly elliptical-oblong in outline, divided seveneighths of the way to the midrib; terminal lobe mediumsized, triangular, the apex and basal corners sometimes elongated, acute at apex, entire or dentate and often subdivided; lateral lobes 4–6 pairs, narrowed from a broad base into an elongate, subacute to obtuse apex which is often forward pointing, sometimes subdivided, the margin towards the distal area of the leaf convex or sigmoid and entire to dentate, the margin towards the proximal area convex or often sinuate; interlobe area near the midrib short and thick, usually with long, narrow teeth or small lobes; glabrous or nearly so; petioles short to medium, green, narrowly to broadly winged. Flowering stem 10–25 cm, green. Capitulum 45– 50 mm in diameter, opening. Outer involucral bracts 11– 15 × 4–5 mm, dark green on outer face, with a narrow pale margin, greyish-green on inner face, lanceolate, acute at apex, more or less spreading. Inner involucral bracts 8–25 × 3–4 mm, dark green, with a pale margin, oblonglanceolate, obtuse at apex. Ligules medium yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with thick, short spiniform projections at apex; cone 0.4–0.5 mm; beak 10–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Probably introduced. Grassy and waste places and waysides. Scattered records throughout Great Britain. Denmark, Germany, Holland and Sweden. 120. T. alatum H. Lindb. Green Dandelion T. semiprivum Dahlst. Medium-sized perennial herb with a stock-like rhizome. Leaves rather numerous, erect; lamina 8–25 × 3–5 cm, shiny medium green on upper surface, green on midrib, narrowly oblong-oblanceolate, divided seven-eighths of the way to the midrib; terminal lobe triangular or helmetshaped, obtuse at apex, often subdivided; lateral lobes 3–5 pairs, regular, triangular, more or less acute at apex, the margin towards the distal area of the leaf more or less convex and entire or denticulate, the margin towards the proximal area straight or concave; interlobe area near the midrib short and thick, entire or denticulate; glabrous or nearly so; petiole up to half the length of the leaf, green, with characteristic, narrow, parallel, entire wings. Flowering stem 8–30 cm,

green, sometimes arachnoid-hairy above. Capitulum 45–60 mm in diameter, opening. Outer involucral bracts 10–12 × 3–4 mm, dark green on outer face, without a pale margin and pink-tipped, glaucous on inner face, lanceolate, obtuse-apiculate at apex, recurved but abruptly spreading at apex. Inner involucral bracts 18–20 × 2–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, olive brown, with spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Native. Roadsides, meadows and waste places. Throughout Great Britain and locally very common, Channel Islands and the coast of Co. Antrim and Co. Londonderry. North, west and central Europe. 121. T. horridifrons Rail. Prickly-leaved Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 8–25 × 4–6 cm, matt medium to dark greyish-green on upper surface, the interlobe area near the midrib sometimes blotched, green on midrib, oblong-oblanceolate in outline, divided seveneighths of the way to the midrib; terminal lobe on outer leaves small and triangular, on the inner leaves larger, helmet-shaped or lingulate, obtuse or subacute at apex and sometimes subdivided; lateral lobes 3–7 pairs, deltoid, acute and elongated at apex, sometimes subdivided, the margin towards the distal area of the leaf more or less convex and entire or dentate, the margin towards the proximal area straight or concave, patent or slightly recurved; interlobe area near the midrib short and fairly thick, sometimes dentate; evenly short hairy; petiole long, green, more or less broadly winged. Flowering stem 8–30 cm, green. Capitulum 45–50 mm in diameter, opening. Outer involucral bracts 12–15 × 3.5–5.0 mm, dark green on outer face, with or without a pale margin, green or flushed with purple on inner face, lanceolate, acute at apex, recurved or reflexed. Inner involucral bracts 18–20 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, blackish. Pollen present. Achenes with body 3.5– 4.0 mm, straw brown, with spiniform projections at apex; cone 0.6–0.8 mm; beak 9–10 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Very local, but sometimes abundant on roadsides. A few scattered records in England and Wales. Czechoslovakia. 122. T. densilobum Dahlst. Close-lobed Dandelion T. pericrispum M. P. Christ. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, flat, erect; lamina 8–20 × 3–5 cm, bright medium green on upper surface, green on midrib, narrowly elliptical-oblong in outline, divided nearly to midrib; terminal lobe triangular or hastate, obtuse at apex, sometimes subdivided; lateral lobes 3–6 pairs, regular, deltoid, elongated to a subacute or acute apex, frequently subdivided, the margin towards the distal area of the leaf more or less convex, entire or denticulate and frequently with

34. Taraxacum 1 larger tooth, the margin towards the proximal area straight or concave; interlobe area near the midrib short and narrow, more or less dentate; evenly short hairy; petiole short, green, narrowly to broadly winged. Flowering stem 8– 25 cm, green. Capitulum 45–50 mm in diameter, opening. Outer involucral bracts 11–13 × 3–4 mm, dark green on outer face, without a paler margin, green on inner face, oblong, acute at apex, recurved or reflexed. Inner involucral bracts 15–17 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, olive brown, with long, thick spiniform projections at apex; cone 0.6–0.8 mm; beak 7–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Introduced. Roadside verges. Scattered records in southern England, Breconshire in Wales, Cumberland in northern England and Co. Londonderry in Ireland. Fennoscandia and Belgium. 123. T. insigne Ekman ex M. P. Christ. & Wiinst. Remarkable Dandelion T. ordinatum Hagend., Soest & Zevenb. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, nearly flat, ascending; lamina 7–20 × 2–5 cm, bright medium green on upper surface, green to faintly purplish on midrib, oblong-oblanceolate in outline, divided seven-eighths of the way to the midrib; terminal lobe usually rather small, triangular or with apex and basal corners elongate, obtuse at apex, often subdivided; lateral lobes 4–7 pairs, regular, crowded, triangular, subacute to obtuse at apex, entire or with 1 or 2 large teeth, spreading or more or less pointing forwards; interlobe area near the midrib short and thick, dentate; glabrous or nearly so; petiole about one-third the length of the leaf, pale green or the inner ones faintly tinted pink, broadly winged. Flowering stem 7–25 mm, green. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 12–14 × 2.0–2.5 mm, dark green on outer face, without a pale margin, pale green, often suffused purple and violet-tipped on inner face, oblong, acute at apex, spreading and somewhat twisted in the horizontal plane. Inner involucral bracts 15–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on the outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with spinulose projections at apex; cone 0.8 mm; beak 8–10 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. 2n = 24. Native. Dry, open, grassy places, roadsides and waste land, locally common. Scattered throughout Great Britain and in Co. Laois and Co. Galway in Ireland. Scandinavia and north-west Europe south to Belgium and Germany. 124. T. laciniosum Dahlst. Divided-leaved Dandelion T. naeviferum Dahlst. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 15–30 × 4–6 cm, bright medium green on upper surface, the internode area near the midrib with dark blotches, green to faintly reddish on midrib, oblong in outline, divided to the midrib; terminal

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lobe rather small, narrowly to broadly sagittate, obtuse at apex, denticulate to dentate or shallowly lobulate; lateral lobes 4–6 pairs, sublinear, acute at apex and deltoid at base, spinulose-dentate, spreading or sloping down; interlobe area near the midrib quite long and narrow, with irregular acute teeth; glabrous or nearly so; petiole short to medium, pale green, more or less winged. Flowering stem 15–40 cm, pale green below, pinkish above, glabrous or nearly so. Capitulum 50–60 mm in diameter, opening. Outer involucral bracts 12–14 × 3.0–3.5 mm, dark green on outer face, without a pale margin, paler and sometimes tinted violet on inner face, lanceolate, acute at apex, spreading to reflexed. Inner involucral bracts 15–22 × 3.0– 3.5 mm, dark green with a pale margin, oblong-lanceolate, subacute at apex. Ligules yellow, with a greyish-violet stripe on the outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with sharp, short, spiniform projections at apex; cone 0.5– 0.8 mm, pale; beak 8–10 mm; pappus 7–9 mm, whitish. Flowers 4–6. Introduced. Known only from near Embleton and Lamplugh Green in Cumberland and Argyllshire. Native of Fennoscandia and northern Russia. 125. T. atonolobum Hagend., Soest & Zevenb. Recurved-lobed Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves few to numerous, nearly flat, ascending; lamina 7–25 × 2–7 cm, medium green on upper surface, green to faintly pinkish on midrib, oblong or oblong-oblanceolate in outline, divided nearly to midrib; terminal lobes hastate or sagittate, obtuse at apex, entire; lateral lobes 4–6 pairs, regular, deltoid, acute at apex, the distal margin more or less straight or sigmoid, the lowest sometimes denticulate or dentate, the proximal margin entire, spreading or often recurved; interlobe area near the midrib short, often with 1 tooth or denticulate; glabrous or nearly so; petiole about one-quarter to one-third the length of the leaf, green, winged. Flowering stem 10–30 cm, slightly arachnoid-hairy above, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 12–14 × 3.0–3.5 mm, dark green on outer face, without a pale margin, pale whitish-green, tinted purplish on inner face, linear-lanceolate or lanceolate, subacute at apex, spreading and recurved at apex. Inner involucral bracts 15–17 × 3.0–3.5 mm, dark green, oblonglanceolate, obtuse at apex. Ligules yellow, with greyishviolet striate on outer face or orange tips to the inner. Styles exserted, dirty yellow. Pollen present. Achenes with body 3.5–3.8 mm, straw-coloured with short spiniform projections at apex; cone about 0.3 mm, pale; beak 9.5–10.5 mm; pappus 5–6 mm, whitish. Flowers 4–5. Introduced. Known only from one record in Worcestershire and one in Denbighshire. Native of Holland and Belgium. 126. T. pannulatiforme Dahlst. Tar-blotched Dandelion Robust perennial herb with a stock-like rhizome. Leaves numerous, more or less crispate, decumbent to erect; lamina 15–30 × 3–7 cm, medium green on upper surface, the interlobe area near the midrib blotched, white or slightly reddish

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on midrib, narrowly elliptical-oblong in outline, divided nearly to midrib; terminal lobe triangular-sagittate, the apex and basal corners elongated, sometimes divided; lateral lobes 5–8 pairs, crowded, triangular to lingulate, sometimes narrowing abruptly from a broad base to give a convex angle to the margin towards the distal area of the leaf, acute or sometimes obtuse at apex, the margin towards the proximal area straight or concave, the margin towards the distal area usually dentate, sometimes with 1 or 2 large teeth or divided, patent, or recurved; interlobe area near the midrib short and fairly thick, dentate; glabrous or nearly so; petiole medium, white or pink on inner leaves, more or less broadly winged. Flowering stem 15–40 cm, green or slightly pinkish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–15 × 2–4 mm, green on inner face, with pink tips and without a pale margin, paler green on inner face, ovate-lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 18–22 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw brown, with short spiniform projections at apex; cone 0.3–0.4 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. 2n = 24. Native. Grassy and waste places and roadsides. Scattered localities in England and Wales, Dumfries-shire and Aberdeenshire in Scotland and Co. Dublin in Ireland. Scandinavia. 127. T. nigridentatum T. Edmondston Black-toothed Dandelion Medium-sized, robust perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 15– 25 × 2–6 cm, dark green on upper surface, the interlobe area near the midrib with dark blotches, pink on midrib, oblong-elliptical in outline, divided seven-eighths of the way to the midrib; terminal lobe slightly narrower than the adjacent lateral lobes, hastate or sagittate, with an elongated apex, more or less divided or with 1 or 2 teeth; lateral lobes 5–8 pairs, narrowly triangular, elongated to an acute apex, the margin towards the distal area of the leaf with more or less numerous dark or black teeth, occasionally with a single large tooth and convex or sigmoid, patent to somewhat recurved; interlobe area near the midrib short and fairly thick, dentate; glabrous or nearly so; petiole rather short, pink, winged, sometimes broadly so. Flowering stem 15–27 cm, green or pink. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–14 × 2.0–3.4 mm, dark green on outer face without a pale margin, paler green on inner face, lanceolate, acute at apex, mostly spreading, the lowest more or less recurved. Inner involucral bracts 16–18 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyish-brown stripe on the outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.0–2.2 mm, straw brown to greyish-brown, with large spiniform projections at the apex; cone 0.2–0.3 mm or almost absent; beak 5–8 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Native. Grassy places, walls and cliffs. Wales and the border counties, Somerset and Cumberland. Endemic.

128. T. laticordatum Markl. Decumbent Dandelion T. uncosum auct. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, nearly flat, usually decumbent, sometimes erect; lamina 10–20 × 3–5 cm, medium green on upper surface, green on midrib, oblong-oblanceolate in outline, divided three-quarters of the way to base; terminal lobe broad, triangular, subacute and often mucronate at apex, often divided; lateral lobes 4–7 pairs, regular, triangular, narrowed to a subacute apex, the margin towards the distal area of the leaf convex and entire or filiform-dentate, the margin towards the proximal area more or less straight, patent to recurved; interlobe area near the midrib short and thick, entire or dentate; glabrous or slightly hairy; petiole rather short, green, broadly winged. Flowering stem 10–25 cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–13 × 3.5–4.5 mm, green on outer face without a pale margin, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 15–27 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, narrowed to an obtuse apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–4.0 mm, straw-coloured, with spiniform projections at apex; cone 0.7–0.8 mm; beak 12–14 mm, pale; pappus 9–10 mm, whitish. Flowers 4–6. Agamosperm. Native. Grassy places, roadsides and waste places. Scattered localities throughout Great Britain and in Co. Dublin in Ireland. Baltic Russia and Czechoslovakia. 129. T. pallescens Dahlst. Pink-stalked Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, nearly flat, decumbent to erect; lamina 10–20 × 2–4 cm, pale yellowish- to medium green on upper surface, green on midrib, oblong-oblanceolate in outline, divided nearly to midrib; terminallobe triangular to hastate, acute at apex, sometimes divided; lateral lobes 4–7 pairs, regular, deltoid, drawn out to a narrow, acute apex, the margin towards the distal area of the leaf convex and entire or filiform-dentate, the margin towards the proximal area more or less straight and sometimes with 1 tooth, patent to recurved; interlobe area near the midrib short and narrow or absent, entire or dentate; glabrous or nearly so; petiole short, green, white or pink on inner leaves, broadly winged. Flowering stem 8–20 cm, green below, slightly pinkish above, sometimes slightly arachnoid-hairy above. Capitulum 45–50 mm in diameter, opening. Outer involucral bracts 11–13 × 3.5–4.5 mm, green or slightly suffused violet on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex, spreading. Inner involucral bracts 12–13 × 2–3 mm, dark green with a pale margin and sometimes pruinose, oblong-lanceolate, obtuse at the slightly violet apex. Ligules deep yellow, with a violetbrown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–2.7 mm, olive grey, with short spiniform projections at apex; cone 0.4–0.5 mm; beak 10–12 mm, pale; pappus 9–10 mm, whitish. Flowers 4–6. Agamosperm.

34. Taraxacum Probably introduced. Roadsides and occasionally old meadows. Scattered records throughout Great Britain. Northern and central Europe. 130. T. subhuelphersianum M. P. Christ. Light-green Dandelion Medium-sized to tall perennial herb with a stock-like rhizome. Leaves fairly numerous, more or less flat, erect; lamina 15–30 × 3–8 cm, pale green on upper surface, green to slightly pinkish on midrib, oblong-oblanceolate in outline, divided almost to midrib; terminal lobe much larger than lateral lobes, broadly triangular or hastate, acute at apex, entire or with a single tooth, rarely divided; lateral lobes 5–6 pairs, deltoid, acute at apex, the margin towards the distal area of the leaf straight to convex and dentate to filiform-dentate especially on the lower lobes, the margin towards the proximal area straight or concave and entire, patent to recurved; interlobe area near the midrib short and narrow, filiform-dentate; glabrous or nearly so; petiole medium, green to faintly pinkish, unwinged or narrowly winged distally. Flowering stem 15–35 cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 12–16 × 3.5–4.5 mm, dark green on outer face, with a pale margin, pale green suffused with violet on inner face, lanceolate, acute at apex, recurved to reflexed. Inner involucral bracts 18–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, yellow when fresh, discoloured when dry. Pollen absent. Achenes with body 3.0–3.5 mm, straw brown, with long spiniform projections at apex; cone 0.5–0.8 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5. Agamosperm. Introduced and casual. Recorded in Gloucestershire, Cheshire, Denbighshire and Northumberland, but not persisting. Denmark and Czechoslovakia. 131. T. necessarium H. Øllg. Dark-leaved Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 8–30 × 3–6 cm, bright, dark green, the interlobe area near the midrib usually blotched, green on midrib, narrowly oblong or oblong-oblanceolate, divided seven-eighths of the way to the midrib; terminal lobe sagittate, subacute to obtuse at apex, usually entire, sometimes divided, sometimes cordate at base; lateral lobes 5–7 pairs, regular, deltoid, narrowed to a more or less acute apex, the margin towards the distal area of the leaf straight to subconvex and entire to regularly dentate, the proximal margin straight or concave, sloping down or recurved; interlobe area short and thick, entire or dentate; petiole rather short, green, winged. Flowering stem 8–35 cm, green. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 12–14 × 3–4 mm, green without a pale margin on outer surface, paler green on inner surface, lanceolate, acute at apex, recurved to reflexed. Inner involucral bracts 14–16 × 3–4 mm, dark green, oblonglanceolate, obtuse at apex. Ligules yellow, with a greyishviolet stripe on outer face and the apex smudged blackish. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw brown, with dense, short spiniform

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projections at apex; cone 0.4–0.5 mm; beak 10–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Possibly introduced. Grassy places and roadsides. Scattered localities in Great Britain. Denmark, Finland, Germany and Sweden. 132. T. margettsii C. C. Haw. Margetts’s Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves fairly numerous, erect; lamina 10–20 × 4– 5 cm, medium fleshy shiny green, rather thick, green on midrib, oblong or spathulate in outline, divided seveneighths of the way to the midrib; terminal lobe of outer leaves rather large, helmet-shaped, rounded at apex and with up to 3 short, blunt teeth, those of the inner leaves very large and rounded at apex; lateral lobes of outer leaves 3–4 pairs, irregularly arranged, patent or sloping down, narrowly triangular, long-acute at apex and irregularly divided into broad-based, obtuse to subacute teeth, those of the inner leaves 1–3 pairs, very broad, triangular, acute at apex and patent to recurved; interlobe area of medium length and width with triangular teeth; glabrous or nearly so; petiole rather long, those of the outer leaves green at base, unwinged but becoming winged at the point it merges with the decurrent base of the lowest lobe, those of the inner leaves sometimes faintly pink. Flowering stems 15–20 cm, green or pink, glabrous. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 9–11 × 4–5 mm, dull green on outer face with a paler margin, greyish-pink and pruinose on inner face, spreading, lanceolate, subobtuse at apex. Inner involucral bracts 18–20 × 3–4 mm, dark green with a pale margin, lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on the outer face. Styles exserted, discoloured. Pollen present. Achene with body 3.0–3.5 mm, straw brown with short spiniform projections above but the rest smooth; cone 0.8–1.0 mm, conical; beak 10–11 mm, conical; pappus 7–8 mm, white. Flowers 4–6. Native. Known only from the Lizard peninsula, Cornwall. Endemic. Named after L. J. Margetts. 133. T. sublaeticolor Dahlst. Small-headed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, erect; lamina 5–20 × 1–5 cm, medium green on upper surface, the interlobe area near the midrib often blotched, green on midrib, oblongoblanceolate in outline, divided seven-eighths of the way to base; terminal lobe triangular, subacute at apex, often divided; lateral lobes 3–5 pairs, more or less regular, triangular or deltoid, more or less acute at apex, the margin towards the distal area of the leaf convex and entire or denticulate, the margin towards the proximal area more or less straight and entire, patent or sloping down; interlobe area near the midrib short and fairly thick, dentate; glabrous or nearly so; petiole rather short, green or rose at the base, narrowly winged below. Flowering stem 5–25 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 8–11 × 3.0–3.5 mm, dark, shiny green on outer face, without a pale margin, green suffused with pink on inner face, lanceolate, acute at apex, mostly erect-spreading to erect. Inner involucral bracts 12–16 × 2–3 mm, dark green with

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a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.2 mm, straw-coloured, with sharp spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Agamosperm. Native. Grassy places, water-meadows, roadsides and waste places. Widely scattered records throughout Great Britain. Fennoscandia and Czechoslovakia. 134. T. lepidum M. P. Christ. Pruinose-bracted Dandelion T. subpraticola auct. Medium–sized perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, erect; lamina 5–25 × 4– 6 cm, medium green on upper surface, the interlobe area near the midrib sometimes blotched, green on midrib, narrowly oblong-elliptical in outline, divided seven-eighths of the way to base; terminal lobe triangular to hastate, subacute at apex, often denticulate to dentate and sometimes divided; lateral lobes 3–5 pairs, more or less regular, triangular to deltoid, more or less acute at apex, the margin towards the distal area of the leaf convex and dentate, the margin towards the proximal area more or less straight and entire or denticulate, spreading or more or less recurved; interlobe area near the midrib short and fairly thick, dentate; glabrous or nearly so; petiole rather long, green or rose at base, unwinged at least below. Flowering stem 5–35 cm, pale green, slightly arachnoid-hairy. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 10–13 × 2.0– 3.0 mm, dark shiny green on outer face, without a pale margin, green suffused pink and pruinose on inner face, narrowly lanceolate, obtuse at apex, spreading to recurved. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, olive brown, with few, thick spiniform projections at apex; cone 0.6–0.7 mm; beak 6.0–8.5 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Possibly native. Roadsides and old meadows. Scattered records in Great Britain, mainly in the west, particularly Wales. Denmark. 135. T. expallidiforme Dahlst. Broad-stalked Dandelion T. oncolobum Dahlst. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, erect; lamina 8–25 × 2– 7 cm, rather pale green on upper surface, green on midrib, oblong or oblong-oblanceolate in outline, divided threequarters or more of the way to the midrib; terminal lobe triangular, shortly elongated at apex, sometimes divided; lateral lobes 3–5 pairs, rather crowded, deltoid, subacute to obtuse at apex, the margin towards the distal area of the leaf slightly convex and entire, denticulate or dentate, the margin towards the proximal area straight or concave, spreading to sloping down; interlobe area near the midrib short and wide, entire or dentate; glabrous or nearly so; petiole rather short, pale green or whitish, broadly winged. Flow-

ering stem 8–30 cm, green below, pinkish above, slightly arachnoid-hairy above. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 7–9 × 2.5–3.5 mm, dark green on outer face, without a pale margin, pale green often slightly suffused with violet on inner face, narrowly lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 12–20 × 2–3 mm, medium green, oblonglanceolate, obtuse at apex. Ligules deep yellow, with a purplish-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, olive brown, with acute spiniform projections at apex; cone 0.5–0.7 mm; beak 10–12 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Agamosperm. Native, or possibly introduced. Grassy places, roadsides, waste places and gardens. Widespread throughout Great Britain, Channel Islands and Co. Dublin and Co. Down in Ireland. Fennoscandia and Czechoslovakia. 136. T. subcyanolepis M. P. Christ. Reddish-bracted Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 8–23 × 3–6 cm, medium green on upper surface, green on midrib, oblong or oblongoblanceolate in outline, divided three-quarters or more of the way to the midrib; terminal lobe triangular-sagittate, more or less obtuse at apex, entire; lateral lobes 3–5 pairs, deltoid, subacute at apex, straight-sided or the margin towards the distal area of the leaf more or less convex, filiform-dentate, recurved; interlobe area near the midrib short and wide, entire or filiform-dentate; glabrous or nearly so; petiole rather short, green, narrowly winged. Flowering stem 5–25 cm, green or purplish, slightly arachnoid-hairy. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 10–12 × 2–3 mm, dark green on outer face, without a pale margin, suffused reddish-purple on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 12–16 × 2–3 mm, dark green with a pale margin, oblonglanceolate, obtuse at purplish apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.2 mm, straw brown, with thick spiniform projections at apex; cone 0.7–0.8 mm; beak 9–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Sexual or facultative agamosperm. 2n = 16, 17, 18, 20, 23, 24, 25, 26, 27. This species hybridises with pollen-bearing agamospermous species. Hybrids have been recorded with T. ancistrolobum, T. brachyglossum, T. oxoniense and T. hamatum and probably others. Diagnosing parentage depends on the knowledge of other species present. Its pollen grains are of uniform size. Probably native. Herb-rich grassland and sand-dunes. Scattered records throughout Great Britain and Ireland, but probably over-recorded. Belgium, Holland, France, Denmark and Sweden. 137. T. pallidipes Markl. Grey-bracted Dandelion T. pallidipes Markl. ex G. E. Haglund nom. nud. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, erect; lamina 8– 25 × 3–5 cm, rather pale green on upper surface, green on

34. Taraxacum midrib, oblong or oblong-oblanceolate, divided nearly to the midrib; terminal lobe triangular or hastate, obtuse at apex, rarely divided; lateral lobes 3–5 pairs, rather crowded, deltoid or triangular, subacute to obtuse at apex, the margin towards the distal area of the leaf slightly convex and entire, denticulate or dentate, the margin towards the proximal area straight or concave and entire or with 1 tooth, spreading to slightly recurved; interlobe area near the midrib short and narrow, often dentate; sometimes hairy; petiole rather short, pale green or whitish, narrowly winged. Flowering stem 8–30 cm, green. Capitulum 50–60 mm in diameter, opening. Outer involucral bracts 12–15 × 3.0–4.5 mm, dark green on outer face, with or without a pale margin, greyish-white pruinose on inner face, lanceolate, acute at apex, strongly recurved. Inner involucral bracts 18–20 mm, dark green with a pale margin, oblong-lanceolate, narrowed to the obtuse apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2–3 mm, straw-coloured, with acute spiniform projections at apex; cone 0.7–0.8 mm, pale; beak 10–12 mm; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Roadside verges and waste places often in rather shady sites. Scattered records in England and Wales and in Wigtownshire in Scotland. Denmark, Finland, Baltic Russia, Sweden and Czechoslovakia. 138. T. croceiflorum Dahlst. Orange-flowered Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, ascending; lamina 10–30 × 3–6 cm, medium green on upper surface, green on midrib, oblong in outline and divided threequarters of the way to the midrib; terminal lobe small, triangular, obtuse at apex, sometimes divided; lateral lobes 5–7 pairs, triangular to deltoid, abruptly elongated to a more or less acute apex, often divided into 1–2 lobules, the margin towards the distal area of the leaf more or less straight and markedly dentate, the margin towards the proximal area sometimes with a conspicuous tooth, pointing forward, spreading or recurved; interlobe area near the midrib short and wide, dentate; glabrous or nearly so; petiole short, green, broadly winged. Flowering stem 8– 18 cm, green below, sometimes purplish above, sometimes arachnoid-hairy below the capitulum. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 9–12 × 2.5– 3.5 mm, dark green on outer face, without a pale margin, paler green suffused with purple at least at tips on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 12–18 × 2.0–3.0 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the purplish apex. Ligules appearing orange-yellow due to the ligule lobes and inner flowers tipped orange-red, with a reddish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, brown, with few, narrow spiniform projections at apex; cone 0.5–0.7 mm; beak 10–11 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Native. Grassy and waste places and roadsides. Throughout Great Britain and common in places; a few records

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round the perimeter of Ireland with one locality in the centre. Northern and west-central Europe to Czechoslovakia. 139. T. lacerifolium G. E. Haglund Jagged-leaved Dandelion T. lacinulatum Markl. Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, ascending; lamina 10–30 × 4–6 cm, medium green on upper surface, green on midrib, oblong in outline and divided almost to midrib; terminal lobe small, triangular, obtuse at apex, often divided; lateral lobes 5–7 pairs, triangular to deltoid, more or less acute at apex, often divided into 1–2 lobules, the margin towards the distal area of the leaf more or less straight and more or less dentate, the margin towards the proximal area sometimes with conspicuous teeth, spreading or somewhat recurved; interlobe area near the midrib narrow, sometimes toothed; glabrous or nearly so; petiole medium, green to rather pink, more or less broadly winged. Flowering stem 8–35 cm, green. Capitulum 45–50 mm in diameter, opening. Outer involucral bracts 13–15 × 3.0– 4.5 mm, dark green on outer face, without a border, paler green more or less suffused with purple at least at the tips on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 14–16 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the reddish-violet apex. Ligules medium yellow, with a greyish-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, greyish-brown, with spiniform projections at apex; cone 0.2–0.3 mm; beak 8–10 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Probably introduced. Scattered records on roadsides and from other grassy places. Local in Wales and southern England north to Dumfries-shire. Belgium and Sweden. 140. T. stenacrum Dahlst. Linear-lobed Dandelion Robust perennial herb with a stock-like rhizome. Leaves numerous, crispate, erect; lamina 15–30 × 3–8 cm, pale to medium green on upper surface, green to slightly purplish on midrib, oblong in outline, crisped, divided to midrib; terminal lobe usually small and narrow, with a lingulate apex, more or less divided; lateral lobes 5–8 pairs, broadly linear, more or less clavate and more or less acute at apex, long-dentate at base, the margin towards the distal area of the leaf sometimes narrowing sharply from a broad base giving a conspicuous shoulder proximally, the margin towards the proximal area of the leaf straight, patent or forward pointing; interlobe area near the midrib very narrow, sometimes toothed; hairy; petiole medium, green, more or less unwinged and very narrow. Flowering stem 15–30 cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 12–21 × 2.5–3.5 mm, medium green on outer face, without a pale border, pale green somewhat suffused with pink on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 15–22 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with an olive brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw-coloured, with

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spiniform projections at apex; cone 0.5–0.6 mm; beak 9– 10 mm, pale; pappus 9–10 mm, whitish. Flowers 4–6. Native. Grassy places, roadsides and waste land. Scattered records through Great Britain and Co. Down and Co. Londonderry in Ireland. Endemic. 141. T. cherwellense A. J. Richards Cherwell Dandelion T. stenoglossum Dahlst., non Brenner Robust perennial herb with a stock-like rhizome. Leaves numerous, crispate, erect; lamina 15–20 × 3–5 cm, pale to medium green on upper surface, green to slightly purplish on midrib, oblong in outline, crisped, divided nearly to the midrib; terminal lobe very narrow, triangular or with narrow elongated apex and basal corners; lateral lobes 3–6 pairs sometimes 2 together on one side of pair, linear from a short, broad base, more or less acute at apex, the margin towards the proximal area of the leaf straight, spreading or forward pointing; interlobe area near the midrib fairly long and thin, sometimes dentate; glabrous or nearly so; petiole short, green, narrow and more or less unwinged. Flowering stem 4–15 cm, ascending-erect, green. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 8–10 × 1.5–2.0 mm, medium green on outer face, and without a pale margin, pale green on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 14–16 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen absent. Achenes with body 2.0–2.8 mm, brown, with short spiniform projections at apex; cone short; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5. Native. Gardens, paths, roadsides and grassland. Known only from a few localities in Wiltshire, Hampshire, Surrey, Oxfordshire andGloucestershire. Endemic. 142. T. porrigens Markl. ex Puol. Hump-lobed Dandelion Robust perennial herb with a stock-like rhizome. Leaves numerous, crispate, erect; lamina 15–30 × 3–5 cm, pale to medium green on upper surface, green to slightly purplish on midrib, broadly oblong in outline, divided nearly to the midrib; terminal lobe broadly triangular or shortly triangular with the apex and basal corners elongated into narrow projections; lateral lobes 5–8 pairs, broadly linear, more or less acute at apex, long-dentate at base, the margin towards the distal area of the leaf narrowing sharply from a broad base giving the lobe a conspicuous shoulder proximally, the proximal margin straight, usually patent, sometimes forward-pointing or occasionally slightly recurved; interlobe area near the midrib short and narrow, dentate; glabrous or nearly so; petioles short to medium, green or those of the inner leaves pinkish. Flowering stem 15–35 cm, green, arachnoid-hairy below the capitulum, otherwise glabrous. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 10–15 × 3.5–4.5 mm, medium green on outer face, without a pale margin, pale green somewhat suffused with pink on inner face, lanceolate, acute at apex, recurved or reflexed. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at

apex. Ligules yellow, with an olive brown stripe on outer face. Styles exserted, yellow to faintly discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw-coloured, with acute spiniform projections at apex; cone 0.5–0.6 mm; beak 8–10 mm, pale; pappus 6–7mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Only recently recorded from Somerset, Gloucestershire, Shropshire and Denbighshire. Probably under recorded being mistaken for T. stenacrum. Czechoslovakia. 143. T. broddesonii G. E. Haglund nom. nud. Broddeson’s Dandelion Robust perennial herb with a stock-like rhizome. Leaves numerous, more or less erect; lamina 12–40 × 2–8 cm, dark shining green on upper surface, green on midrib, fleshy, oblong in outline; terminal lobe triangular with rounded apex and the basal corners elongated; lateral lobes 2–5 pairs, triangular to deltoid, more or less acute at apex, with large teeth, spreading or backward pointing; interlobe area near the midrib fairly wide and dentate; glabrous or nearly so; petiole long, green, narrowly winged. Flowering stem 40–60 cm, ascending-erect, green. Capitulum 60–70 mm in diameter, opening. Outer involucral bracts 12–15 × 2– 3 mm, medium green on outer face, pale green on inner face, oblong-lanceolate, acute at apex, recurved. Inner involucral bracts 15–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate. Ligules yellow, with a greyish-purple stripe on outer face. Styles discoloured. Pollen absent. Flowers 4–5. Introduced. Waste places including a public house carpark. Flintshire, Derbyshire and Lancashire. 144. T. leucopodum G. E. Haglund White-stalked Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, decumbent to erect; lamina 8–20 × 3–4 cm, clear, rather dark green on upper surface, green on midrib, oblong-elliptical in outline, divided three-quarters of the way to the midrib; terminal lobe small and rounded with a markedly acute attenuate tip thus characteristically the shape of a German soldier’s helmet; lateral lobes 4–5 pairs, more or less hamate, more or less acute at apex, the margin towards the distal area of the leaf convex and entire to denticulate, the margin towards the proximal area straight or concave and often with a single large tooth towards the interlobe area near the midrib, recurved; interlobe area near the midrib short and wide, more or less dentate, often with 1 conspicuous tooth; petiole short to medium, whitish, winged. Flowering stem 8–23 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 15–16 × 3–4 mm, medium green on outer face, without a pale margin, pale green to whitish-pruinose on inner face, lanceolate, acute at apex, spreading to somewhat recurved, irregularly arranged to somewhat twisted. Inner involucral bracts 15–17 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with spiniform projections at

34. Taraxacum apex; cone 0.4–0.7 mm; beak 7–10 mm, pale; pappus 7– 8 mm, whitish. Flowers 4–5. Probably introduced. Grassy places, roadsides and waste land. Recorded only from Essex, Berkshire, Suffolk, Gloucestershire, Monmouthshire, Worcestershire, Breconshire, Radnorshire and Co. Dublin. Sweden. 145. T. obtusilobum Dahlst. ex G. E. Haglund Blunt-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, ascending to erect; lamina 5– 20 × 1–4 cm, fleshy greyish-green on upper surface, chalkwhite on midrib, oblanceolate or oblong-oblanceolate in outline, subentire, denticulate, dentate or shallowly lobed; terminal lobe broadly helmet-shaped, rounded-obtuse at apex, entire or denticulate; lateral lobes if present 3–8 pairs, very shallow, acute at apex, spreading; interlobe area near the midrib if present very short and very wide, dentate; glabrous or nearly so; petiole short, white to reddish, usually winged. Flowering stem 10–30 cm, pale green, slender. Capitulum 30–35 mm, opening. Outer involucral bracts 6– 8 × 2–3 mm, dark purple on both faces, without a pale margin, lanceolate, acute at apex, spreading-erect. Inner involucral bracts 16–18 × 2.0–2.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules pale yellow, with a red or grey stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes often poorly set, with body 2.5–3.0 mm, olive brown, with spiniform projections at apex; cone 0.3–0.4 mm; beak 9–10 mm, pale; pappus 7– 9 mm, whitish. Flowers 4–5. Sexual diploid. 2n = 16, 17. The only wholly sexual and self-incompatible species in northern Europe. It is usually accompanied with confusing hybrids with surrounding apomictic species, some of the hybrids being sexual and self-incompatible and some triploid and agamospermous. These sometimes persist after T. obtusilobum has disappeared. In Co. Durham the main species it hybridised with was T. polyodon. Introduced. Recorded from Kent, Gloucestershire, Co. Durham, Midlothian and Ross-shire, but not persisting and not seen since 1968. Native of south-east Sweden, and Denmark, Belgium and France. 146. T. undulatiflorum M. P. Christ. Dull-leaved Dandelion Medium-sized to tall perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, erect; lamina 15–30 × 4–6 cm, dull, rather dark green on upper surface, green to slightly purplish on midrib, oblong or oblongoblanceolate in outline, divided almost to midrib; terminal lobe triangular or hastate, acute at apex, with a more or less elongated tip, sometimes divided; lateral lobes 5–6 pairs, triangular at base, elongated into a narrow, linear, acute apex, the margin towards the distal area of the leaf straight to convex and dentate to filiform-dentate, the margin towards the proximal area straight or concave and entire or filiform-dentate, usually patent to recurved, sometimes forward-pointing; interlobe area near the midrib short and narrow, sometimes dentate; glabrous or nearly so; petiole short to medium, green to faintly purplish, unwinged or narrowly winged distally. Flowering stem 15–32 cm, green.

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Capitulum 45–55 mm in diameter, opening. Outer involucral bracts 12–14 × 2.5–4.0 mm, dark green without a pale margin on outer face, pale green suffused with violet on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, yellow when fresh, discoloured when dry. Pollen present. Achenes with body 2.5–3.5 mm, straw-coloured, with spiniform projections at apex; cone 0.4–0.5 mm; beak 9–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Roadsides and waste places. Scattered records throughout Great Britain and in Co. Dublin in Ireland. Native of Denmark and Czechoslovakia. 147. T. chloroticum Dahlst. Spiky-leaved Dandelion T. chloroticum Dahlst. ex Florstr¨om nom. nud. Medium-sized to tall perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, erect; lamina 15–30 × 3–6 cm, dull medium green to slightly tinged purplish, green to slightly purplish on midrib, oblong to oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe triangular or elongated at apex and basal corners, acute at apex; lateral lobes 5–6 pairs, triangular, narrowed to a very acute apex, the margin towards the distal area of the leaf straight to convex and dentate to filiformdentate, the margin towards the proximal area of the leaf straight or concave and entire or filiform-dentate, spreading to recurved; interlobe area near the midrib short and fairly wide, sometimes dentate; glabrous or nearly so; petiole short to medium, green to fairly purplish, unwinged or narrowly winged distally. Flowering stem 10–30 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 12–14 × 2.5–4.0 mm, dark green on outer face, without a pale green margin, pale green suffused with violet on inner face, lanceolate, obtuse at apex, recurved. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the purplishviolet apex. Ligules yellow, with a greyish-brown stripe on outer face. Styles exserted, yellow when flesh, discoloured when dry. Pollen present. Achenes with body 2.5–3.0 mm, with short, acute spiniform projections at apex; cone 0.7– 0.8 mm; beak 9–11 mm, pale; pappus 7–8 mm, whitish. Flowers 4–5. Introduced. On a few roadsides. Recorded from Surrey, Essex, Middlesex, Berkshire, Gloucestershire and Lancashire. Native of Finland, Sweden and Germany. 148. T. speciosum Raunk. Massive Dandelion T. officinale subsp. speciosum (Raunk.) Dahlst. Large perennial herb with a stock-like rhizome. Leaves numerous, erect, more or less flat; lamina 10–30 × 3–8 cm, dark green on upper surface, the interlobe area near the midrib blotched, green to slightly purplish on midrib, broadly oblanceolate in outline, divided nearly to midrib; terminal lobe broadly triangular or hastate, acute at apex with an elongated tip, entire or with 1 tooth, rarely divided; lateral lobes 5–6 pairs, deltoid or triangular, acute at apex, the margin towards the distal area of the leaf straight to convex and dentate to filiform-dentate (especially on the

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lower lobes), the margin towards the proximal area straight or concave and entire; interlobe area near the midrib short and rather narrow, dentate or filiform-dentate; glabrous or nearly so; petiole short or medium, green or faintly purplish, unwinged or narrowly winged distally. Flowering stem 20– 35 cm, green. Capitulum 50–70 mm in diameter, opening. Outer involucral bracts 12–16 × 3.5–4.5 mm, dark green on outer face, without a pale margin, pale green suffused with violet on inner face, lanceolate, acute at apex. Inner involucral bracts 18–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow to orange, with a greyish-violet stripe on outer face. Styles exserted, yellow when fresh, discoloured when dry. Pollen absent. Achenes with body 3.0–3.5 mm, straw brown, with few, acute spiniform projections at apex; cone 0.5–0.8 mm; beak 8–10 mm, pale; pappus 5–6 mm, whitish. Flowers 4–5. 2n = 24. Introduced. Three individual records from Surrey, Breconshire and Co. Durham. Fennoscandia and Belgium. 149. T. piceatum Dahlst. Leaden-bracted Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less crispate, decumbent to erect; lamina 8–20 × 2–6 cm, more or less crisped, dark green on upper surface, the interlobe area near the midrib with dark blotches, green near the base of the midrib but characteristically reddish towards the apex, oblong-oblanceolate, divided nearly to midrib; terminal lobe variable, usually large, triangular and subobtuse to acute at apex on inner leaves, small, subsagittate, with a short, elongated tip and sometimes divided or with a single tooth on outer leaves; lateral lobes 3–5 pairs, triangular, subacute to acute at apex, the margin towards the distal area of the leaf straight to convex and entire to dentate, the proximal margin more or less straight; interlobe area near the midrib short and narrow, sometimes dentate; glabrous or nearly so; petiole rather short, those of outer leaves green to whitish, those of inner pale but pinkish. Flowering stem 8–20 cm, green below, pinkish above, glabrous or slightly arachnoid-hairy above. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 11–12 × 3–4 mm, medium green on outer face, and without a pale margin, leaden-violet on inner face, oblong, acute at apex. Inner involucral bracts 14–15 × 3– 5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, olive, with acute, erecto-patent spiniform projections at apex; cone 0.5– 0.6 mm; beak 13–14 mm, pale; pappus 10–12 mm, whitish. Flowers 4–6. 2n = 24. Agamosperm. Probably introduced. Roadsides and waste places. Scattered records throughout Great Britain. North and west Europe and Czechoslovakia. 150. T. cyanolepis Dahlst. Bluish-bracted Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, decumbent to erect; lamina 8–20(–25) × 2–6 cm, pale green, the interlobe area by the midrib blotched, green to faintly pinkish near the apex of the midrib, oblong-oblanceolate in outline, more

or less crisped, divided nearly to midrib; terminal lobe triangular, usually larger on inner leaves with a subacute or obtuse apex, on the outer leaves subsagittate, smaller and sometimes divided or with 1 tooth; lateral lobes 3– 5 pairs, triangular, obtuse to subacute at apex, the margin towards the distal area straight to convex and entire or denticulate to dentate, the margin towards the proximal area of the leaf more or less straight and patent to recurved; interlobe area near the midrib short and rather narrow, often dentate; glabrous or nearly so; petiole rather short, the outer whitish to green, the inner pale but more or less reddish, narrowly winged. Flowering stem 8–20(–25) cm, green, glabrous or slightly arachnoid-hairy. Capitulum 50– 55 mm in diameter, opening. Outer involucral bracts 10– 12 × 2.8–3.5 mm, medium green on outer face, without a pale margin, bluish-purple on inner face, lanceolate, acute at apex. Inner involucral bracts 15–22 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the violet apex. Ligules deep yellow, with a reddish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, olive brown, with numerous, acute spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 7–12 mm, whitish. Flowers 4–6. Agamosperm. 2n = 24. Native at least in Scotland. Humid grasslands including those that are species-rich, walls, cliffs and roadsides. Scattered through Great Britain but absent from large areas; but abundant locally, especially in north-east Scotland; a few records round the perimeter of Ireland. Has apparently become less common since about 1970. Fennoscandia, Belgium, Holland and Czechoslovakia. 151. T. curtifrons H. Øllg. Variable-leaved Dandelion Medium-sized, heterophyllus perennial herb with a stocklike rhizome. Leaves numerous, erect; lamina 15–20 × 2–5 cm, bright green on upper surface, the midrib greenish or brownish, oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe rather large, sagittate to hastate, more or less acute at apex, entire; lateral lobes 3–5 pairs, those of the outer and intermediate leaves triangular, acute at apex, the margin towards the distal area of the leaf more or less straight and entire or filiform-dentate, the margin towards the proximal area straight, spreading, those of the inner leaves incompletely developed with wide, rounded interlobe spaces; interlobe area near the midrib short and rather narrow, and sometimes dentate; outer glabrous or nearly so, inner rather hairy; petiole rather short, green, winged. Flowering stem 15–25 cm, green. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 10–12 × 3–4 mm, dark green on outer face, without a pale margin, clearly violet-coloured on inner face, lanceolate, acute at apex, slightly recurved with erect tips. Inner involucral bracts 14–16 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with short, dense spiniform projections at apex; cone 0.3–0.4 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5.

34. Taraxacum Introduced. Roadsides and waste places. A few records from Berkshire, Oxfordshire, Gloucestershire, Leicestershire and Denbighshire. Denmark and Germany. 152. T. acutifrons Markl. Acute-leaved Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 8– 22 × 2–8 cm, very pale green on upper surface, the interlobe area near the midrib not obviously blotched, green to slightly pinkish on midrib, oblong in outline, divided nearly to midrib; terminal lobe more or less narrowly triangular or hastate, sometimes waisted, acute at apex, often with one characteristically rounded lobe; lateral lobes 4–6 pairs, regular, deltoid, abruptly narrowing to patent, linear elongations with an acute to very acute apex, the margin towards the distal area of the leaf angled and entire or filiform-dentate, the margin towards the proximal area more or less straight or convex and entire, more or less patent; interlobe area near the midrib short and rather narrow, often dentate; glabrous or nearly so; petiole rather short, green to faintly pink, unwinged. Flowering stem 8– 25 cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 10–15 × 3.5–5.0 mm, dark green on outer face, without a more or less pale margin, very deep violet and pruinose on inner face, lanceolate, acute at apex. Inner involucral bracts 15–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex, spreading. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with spiniform projections at apex; cone 0.5–0.6 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Introduced. Roadside verges. Recorded from Somerset, Gloucestershire, Herefordshire, Breconshire, Denbighshire, Cheshire and Lancashire. Finland. 153. T. chrysophaenum Rail. Orange-toothed Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, erect; lamina 10–25 × 2–5 cm, pale to medium green on upper surface, the interlobe area near the midrib not obviously blotched, green to slightly pinkish on midrib, oblong in outline, divided seven-eighths of the way to the midrib; terminal lobe triangular or helmet-shaped, acute or subacute and sometimes apiculate at apex, entire or sometimes divided, sometimes larger and helmet-shaped on inner leaves; lateral lobes 6–8 pairs, regular, crowded, deltoid or triangular, more or less straight, convex or angled to give a claw-shaped lobe, the upper entire and the lower dentate to filiform-dentate, the margin towards the proximal area of the leaf more or less straight, patent to slightly recurved; interlobe area near the midrib short and fairly wide, more or less dentate; glabrous or nearly so; petiole rather short, whitish, narrowly winged. Flowering stem 10–30 cm, green. Capitulum 35–40 mm in diameter, opening. Outer involucral bracts 14–17 × 2– 4 mm, medium green on outer surface, without a pale margin, pale green and pruinose on inner surface, lanceolate, acute at apex. Inner involucral bracts 18–20 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse

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at apex. Ligules deep yellow, with a greyish-violet stripe on outer face and reddish-orange lobes at apex of the inner ones. Styles exserted, discoloured. Pollen present. Achene body 3.0–3.5 mm, straw brown, with spiniform projections at apex; cone 0.8–1.0 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5. Introduced. Recorded from Devonshire, Oxfordshire, Gloucestershire, Monmouthshire and Carmarthenshire. Finland. 154. T. tumentilobum Markl. ex Puol. Swollen-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less crispate, decumbent to erect; lamina 10–25 × 4–6 cm, medium green on upper surface, more or less crisped, the interlobe area near the midrib with dark blotches, green to slightly pinkish on midrib, oblong or oblong-oblanceolate, divided almost to midrib; terminal lobe usually rather small and hastate on outer leaves, often larger on inner leaves and divided or with 1–2 teeth; lateral lobes 5–7 pairs, triangular, regular, subacute to subobtuse at apex, the margin towards the distal area of the leaf straight to convex and entire, dentate or filiform-dentate, the margin towards the proximal area more or less straight, spreading; interlobe area near the midrib fairly long and narrow, characteristically with a single large tooth or small lobe; glabrous or nearly so; petiole medium to long, pink, unwinged. Flowering stem 10–25 cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 10–15 × 3–4 mm, dark green without a pale margin on outer face, medium green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–20 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with numerous, acute, conical spiniform projections at apex; cone 0.5–0.6 mm; beak 10–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Probably introduced. Roadsides and grassy and waste places. A few scattered records in Great Britain. Fennoscandia and Baltic Russia. An unnamed allied species which has the outer leaves with an elongated tip to the terminal lobe which on the inner leaves is incise-dentate, the outer involucral bracts strictly erect but proximally recurved and the cone of the achene 0.8–1.0 has been recorded from Northumberland. 155. T. intumescens G. E. Haglund Red-tipped Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less crispate, decumbent to erect; lamina 8–20 × 4–6 cm, medium green on upper surface, the interlobe area near the midrib with dark blotches, green to slightly pink on midrib, oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe usually rather small, triangular to subsagittate, obtuse at apex, sometimes toothed or divided; lateral lobes 4–6 pairs, regular, triangular to deltoid, acute to subacute at apex, the margin towards the distal area of the leaf straight to convex, dentate on the upper lobes

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and filiform-dentate on the lower lobes, the margin towards the proximal area straight or convex, spreading; interlobe area near the midrib short and narrow to fairly wide, often with a single large distinct tooth; glabrous or nearly so; petiole short, green, unwinged to narrowly winged. Flowering stem 8–25 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 10–14 × 2.0–4.5 mm, dark green on outer face, without a pale margin, greyishgreen to lead-coloured on inner face, lanceolate, acute at apex. Inner involucral bracts 18–20 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the purplish apex. Ligules deep yellow with a greyish-violet stripe on outer face and red lobes at apex. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw-coloured, with small spiniform projections at apex; cone 0.5–0.6 mm; beak 9–11 mm, pale; pappus 6–8 mm, whitish. Flowers 4–6. Introduced. Roadsides and waste places. A few scattered localities in England and Wales north to Cumberland. Denmark. 156. T. angulare Hagend., Soest & Zevenb. Angular-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less crispate, erect; lamina 8– 20 × 4–6 cm, medium green on upper surface, the interlobe area near the midrib with dark blotches, green to slightly pink on midrib, oblong-oblanceolate, divided nearly to midrib; terminal lobe triangular to subsagittate, obtuse at apex, entire or with 1–2 teeth; lateral lobes 4–6 pairs, narrowly triangular, more or less acute at apex, the margin towards the distal area of the leaf straight to convex, dentate on the upper lobes and filiform-dentate on the lower lobes, the proximal margin straight or convex, spreading; interlobe area near the midrib short and fairly wide, often with a single large distinct tooth; glabrous or nearly so; petiole medium size, green, unwinged to narrowly winged. Flowering stem 8–25 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 10–16 × 3.0– 4.5 mm, dark green on outer face, without a pale margin, green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 18–20 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw brown, with spiniform projections at apex; cone 0.5–0.6 mm; beak 9–11 mm, pale; pappus 8–9 mm, whitish. Flowers 4–5. Introduced. Known only from records in Gloucestershire and Dumfries-shire. Holland. 157. T. ancistrolobum Dahlst. Few-lobed Dandelion Robust, squat perennial herb with a stock-like rhizome. Leaves numerous, usually decumbent; lamina 10–25 × 2– 5 cm, dull, dark green on upper surface, flat, the interlobe area near the midrib not obviously blotched, green on midrib but slightly pinkish distally, oblanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe large, subacute to obtuse at apex, entire, the base overlapping the first pair of lateral lobes; lateral lobes 2–3(–4) pairs,

regular, triangular, large and broad, subacute to obtuse at apex, the margin towards the distal area of the leaf convex and entire to filiform-dentate, the margin towards the proximal area straight or concave and entire; interlobe area near the midrib short and wide, entire; glabrous or nearly so; petiole short, green, winged. Flowering stem 10–30 cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 8–11 × 2.5–4.0 mm, green on outer face and without a pale margin, pale greyish-green more or less suffused with purple on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–18 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a reddishpurple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm and very wide (1.4 mm), straw brown, with spiniform projections at apex; cone 0.5–0.6 mm; beak 9–11 mm, pale; pappus 7–10 mm, whitish. Flowers 4–6. Agamosperm. Native. Grassy and waste places and roadsides. Widespread in England and Wales, locally common in the south and very common in lowland northern England, scarce in Scotland and only recorded from Co. Dublin in Ireland. Fennoscandia, Belgium, Holland, Czechoslovakia and Switzerland. 158. T. sellandii Dahlst. Selland’s Dandelion T. granvinense Dahlst. Robust perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, spreading to erect; lamina 10– 25 × 3–8 cm, dull, medium to dark green on upper surface, the interlobe area near the midrib not obviously blotched, discoloured on midrib and often bordered by black lines, oblong-oblanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe more or less triangular, obtuse to acute at apex, often divided; lateral lobes 3–6 pairs, triangular or deltoid, more or less acute at apex, the margin towards the distal area of the leaf convex or angled and those of the upper entire or with large teeth and the lower with large teeth, the margin towards the proximal area straight or concave, the angled margins frequently making the lobes characteristically arching, spreading to recurved; interlobe area near the midrib short and wide and dentate; glabrous or nearly so; petiole short, green, winged. Flowering stem 10–30 cm, green, slightly purplish, slightly arachnoid-hairy. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 10–14 × 3.5–6.0 mm, dark green on outer face, without a pale margin, dull green, pruinose, leaden or somewhat violet with pink tips on inner face, lanceolate, acute at apex, stiff and arching, spreading to recurved. Inner involucral bracts 18–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with spiniform projections at apex; cone 0.6–0.7 mm; beak 10–12 mm, pale; pappus 5–8 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24, 26, 28. Native. Grassy and waste places and roadsides. Widespread in Great Britain and one of the commonest species there; several localities in the northern part of

34. Taraxacum Ireland and two coastal localities further south. Norway, Sweden and Czechoslovakia. Named after Sjur Knutsen Selland (1867–1920). 159. T. altissimum H. Lindb. Tall Dandelion Tall perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, erect; lamina (8–)20–30 × 4–6 cm, dark to medium green on upper surface, the interlobe area near the midrib with dark blotches, green to slightly pinkish on midrib, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe usually long, helmetshaped, long acute at apex, dentate and divided; lateral lobes 3–5 pairs, regular, triangular to deltoid, acute at apex, the margin towards the distal area of the leaf convex and dentate, the margin towards the proximal area straight or concave, spreading to recurved; interlobe area near the midrib short and rather narrow, dentate; glabrous or nearly so; petiole long, green, unwinged below. Flowering stem 8–35 cm, green. Capitulum 50–60 mm in diameter, opening. Outer involucral bracts 11–15 × 4–5 mm, dark green on outer face with a pale margin, green on inner face, lanceolate, acute at apex. Inner involucral bracts 16–18 × 2–3 mm, oblonglanceolate, obtuse at apex. Ligules medium yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.0–2.5 mm, olive brown, with numerous, acute spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Grassy places, roadsides and wasteland. Scattered records throughout Great Britain, uncommon except perhaps in north-east England. Finland, Sweden and Czechoslovakia. 160. T. aequisectum M. P. Christ. Equal-lobed Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, usually decumbent to erect; lamina 8–22 × 3–5 cm, dull, dark green on upper surface, the interlobe area near the midrib with dark blotches, green to slightly pinkish on midrib, oblong in outline, divided nearly to the midrib; terminal lobe rather narrowly triangular, drawn out to the acute apex, often divided; lateral lobes 5–8 pairs, regular, deltoid, acute to long-acute at apex, the margin towards the distal area of the leaf straight to convex, dentate and sometimes with 1 or more large teeth or filiform-dentate, the margin towards the proximal area straight or concave and entire, spreading to recurved; interlobe area near the midrib short and narrow, dentate; glabrous or nearly so; petiole short or absent, green to faintly pink, winged. Flowering stem 8–25 cm, green. Capitulum 40–55 mm in diameter, opening. Outer involucral bracts 11–14 × 2.0–3.5 mm, dark green on outer face with a faint pale border, green with pink tips on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 18–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, yellowish. Pollen present. Achenes with body 2.8–3.5 mm, strawcoloured, with few, short spiniform projections at apex; cone

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0.5–0.6 mm; beak 10–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Probably introduced. Grassy places, roadsides and waste land. Scattered records throughout Great Britain, rather uncommon. Southern Sweden, Denmark, Belgium and Holland. 161. T. interveniens G. E. Haglund City Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, flat to slightly crispate, spreading to erect; lamina 8–20 × 3–5 cm, dull medium to dark green on upper surface, the interlobe area near the midrib sometimes with dark blotches, brown on midrib, flat to slightly crisped, narrowly oblong in outline, divided to the midrib; terminal lobe either long-helmet-shaped or rather small and sagittate, acute or very acute at apex, entire; lateral lobes 4–8 pairs, regular, deltoid or sigmoid, acute to acuminate at apex, the margin towards the distal area of the leaf sinuate and entire or filiform-dentate, the margin towards the proximal area, straight or concave recurved; interlobe area near the midrib short and very narrow, often dentate; slightly hairy or glabrous; petiole short or medium, pink, unwinged or narrowly winged. Flowering stem 8–20 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 10–15 × 4–5 mm, dark green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex. Inner involucral bracts 18–20 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with short spiniform projections at apex; cone 0.5–0.6 mm; beak 8–10 mm, pale; pappus 5–7 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Grassy and waste places and roadsides. Very scattered records in Great Britain. Locally abundant in the cities of London, Edinburgh and Glasgow, scarce elsewhere. Czechoslovakia and Sweden. 162. T. angustisquameum Dahlst. ex H. Lindb. Multilobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, decumbent to ascending; lamina 8–20 × 3–7 cm, dark to medium green on upper surface, the interlobe area near the midrib usually with dark blotches, green to slightly pinkish on midrib, oblong or narrowly elliptical-oblong in outline, divided nearly to the midrib; terminal lobe usually rather small and triangular or sagittate, acute at apex and sometimes mucronate, often divided; lateral lobes 4–8 pairs, deltoid, acute to apiculate at apex, the margin towards the distal area of the leaf straight to convex or somewhat sigmoid and entire to dentate, the margin towards the proximal area straight or concave and entire to dentate, recurved; interlobe area near the midrib short and narrow and sometimes with a distinct, acuminate tooth; glabrous or nearly so; petiole short, pink, unwinged below. Flowering stem 8–25 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 11–14 × 1.5–2.5 mm, medium green and without a pale margin on outer face, paler green and more or less pruinose on inner face, lanceolate, acute at apex,

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arcuate-spreading, becoming more recurved. Inner involucral bracts 17–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.0–2.5 mm, straw-coloured, with numerous, acute spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Probably introduced. Grassy and waste places and roadsides. Very scattered records through Great Britain. Northern Europe, Iceland, Belgium, France and Germany. 163. T. mimulum Dahlst. ex H. Lindb. Sharp-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, decumbent to ascending; lamina 8–20 × 3–7 cm, medium green on upper surface, the interlobe area near the midrib not obviously blotched, green to slightly pinkish on midrib, somewhat crisped, narrowly oblong in outline, divided nearly to the midrib; terminal lobe usually rather small and triangular or sagittate, acute and sometimes mucronate at apex, sometimes divided; lateral lobes 3–6 pairs, deltoid, acute to very acute at apex, the margin towards the distal area of the leaf straight to concave or somewhat sigmoid and entire to dentate, the margin towards the proximal area straight or concave and entire to dentate, spreading to recurved; interlobe area near the midrib often fairly long and narrow, sometimes with a distinct, acuminate tooth; glabrous or nearly so; petiole fairly long, pink, narrowly winged. Flowering stem 8– 25 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 11–14 × 3–4 mm, rather dark green on outer face, without a pale margin, paler green and more or less pruinose on inner face, lanceolate, acute at apex, recurved or reflexed. Inner involucral bracts 16–18 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.0–2.5 mm, straw brown, with spiniform projections at apex; cone 0.4–0.5 mm; beak 7–10 mm, pale; pappus 5–7 mm, whitish. Flowers 4–5. Introduced. Known only from a single record from improved chalk pasture at Chazey in Oxfordshire. Northern Europe, Belgium, France, Germany and Switzerland. 164. T. stereodes Ekman ex G. E. Haglund Hairy-stalked Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, decumbent to ascending; lamina 8–20 × 3–6 cm, medium green on upper surface, the interlobe area near the midrib not obviously blotched, green to slightly pinkish on midrib, oblong or narrowly elliptical-oblong in outline, divided nearly to the midrib; terminal lobe rather small and triangular or sagittate, acute and sometimes mucronate at apex, often divided; lateral lobes 4–8 pairs, deltoid, acute to very acute at apex, sometimes divided, the margin towards the distal area of the leaf straight to convex or somewhat sigmoid and entire to dentate, the margin towards the proximal area straight or concave and entire to dentate, recurved; interlobe area

near the midrib sometimes with a distinct, acuminate tooth; glabrous or nearly so; petiole fairly long, pink, unwinged below. Flowering stem 8–25 cm, arachnoid-hairy throughout especially when young, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 11–14 × 3– 4 mm, green on outer face and without a paler margin, paler green and more or less pruinose on inner face, lanceolate, acute at apex, spreading, becoming recurved. Inner involucral bracts 16–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.0–2.5 mm, straw brown, with spiniform projections at apex; cone 0.8– 1.0 mm; beak 8–10 mm, pale; pappus 5–7 mm, whitish. Flowers 4–6. Introduced. On the verges of main roads. Very scattered records in Great Britain. Fennoscandia and Germany. 165. T. adiantifrons Ekman ex Dahlst. Pretty-leaved Dandelion T. hemicyclum G. E. Haglund Medium-sized, heterophyllous perennial herb with a stocklike rhizome. Leaves fairly numerous, nearly flat, erect; lamina 8–30 × 2–5 cm, clear medium green on upper surface, green on midrib, nearly flat, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe variable, rather larger than adjacent lateral lobes, triangular with the area near the apex rhomboidal, or with a large tooth on either side and an elongated tip; lateral lobes 3–5 pairs, deltoid, acute at apex, the margin towards the distal area of the leaf straight or sinuate and entire or filiform-dentate, the margin towards the proximal area straight or more or less convex, spreading to recurved; interlobe area near the midrib short and narrow, often toothed; glabrous or nearly so; petiole short to medium, pale green or pink on outer leaves, clear pink on inner leaves. Flowering stem 8–35 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 12–14 × 3.0–4.5 mm, green on outer face, without a paler margin, paler green on inner face, lanceolate, acute at apex, recurved or reflexed. Inner involucral bracts 16–20 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Probably introduced. Grassy and waste places and roadsides. Scattered records throughout Great Britain and locally common. Norway. 166. T. retroflexum H. Lindb. Reflexed-lobed Dandelion Medium-sized heterophyllus perennial herb with a stocklike rhizome. Leaves numerous, nearly flat, erect; lamina 8– 30 × 3–7 cm, clear medium green on upper surface, green on midrib, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe variable, acute at apex; lateral lobes 3–5 pairs, deltoid, acute at apex, the margin towards the distal area of the leaf straight or sinuate and entire or filiform-dentate, the margin towards the proximal area

34. Taraxacum straight or more or less convex, spreading to recurved; interlobe area near the midrib short and rather wide, sometimes toothed; glabrous or nearly so; petiole short to medium, pale green on outer leaves, clear pink on inner leaves. Flowering stem 8–40 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 12–16 × 3.0– 4.5 mm, green on outer face, with a pale margin, paler green on inner face, lanceolate, acute at apex. Inner involucral bracts 18–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, yellowish when fresh, darker when dry. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with numerous, obtuse to acute spiniform projections at apex; cone 0.4–0.5 mm; beak 7–10 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. 2n = 24. Introduced. Road verges and wasteland. A few records in the London area, south Wales, northern England and Berwickshire. Northern Europe, Belgium, Germany and Switzerland. 167. T. semiglobosum H. Lindb. Hairy-ribbed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves fairly numerous, nearly flat, erect; lamina 8–30 × 3–7 cm, clear medium green on upper surface, green on midrib, oblong to oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe not obviously larger than lateral lobes, triangular, acute at apex; lateral lobes 3–5 pairs, deltoid or semilunate, very acute at apex, the margin towards the distal area of the leaf straight or sinuate and entire or filiform-dentate, the margin towardsthe proximal area straight or more or less convex, spreading to recurved; interlobe area near the midrib short and narrow, sometimes filiform-dentate; midrib with dense hairs; petiole short to medium, pale green on outer leaves, clear pink on inner leaves, narrowly winged. Flowering stem 8–35 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 12–14 × 3.0–4.5 mm, green on outer face, with a pale margin, paler green on inner face, lanceolate, obtuse at apex. Inner involucral bracts 16–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, acute at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with rather long, acute spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Road verges. Only known from Middlesex and Dumfries-shire. Fennoscandia, Germany and Czechoslovakia. 168. T. aequilobum Dahlst. Twisted-bracted Dandelion Robust, heterophyllus perennial herb with a stock-like rhizome. Leaves numerous, highly crispate, erect; lamina 15– 30 × 4–6 cm, dirty green on upper surface, the interlobe area near the midrib usually with dark blotches, slightly dirty pinkish on midrib, oblong or narrowly oblong-oblanceolate in outline, divided almost to the midrib; terminal lobe usually rather small and triangular, acute at apex or with an elongated tip; lateral lobes 5–9 pairs, rather crowded,

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regular, deltoid, with divergent, often forward-pointing, linear elongations at the acute or very acute apex, the margin towards the distal area of the leaf sigmoid, concave or convex and entire or filiform-dentate, the margin towards the proximal area more or less straight, spreading to recurved; interlobe area near the midrib short and narrow, usually dentate; glabrous or nearly so, or arachnoid-hairy on the midrib; petiole very variable in length, rose-purple, winged or more or less unwinged. Flowering stem 15–35 cm, erect, pale green above, purplish below. Capitulum 50–65 mm in diameter, opening. Outer involucral bracts 10–15 × 2– 4 mm, dark green on outer face, and without a pale margin, pale green, pruinose and often purple-tipped on inner face, lanceolate, acute at apex, twisted laterally, spreading and becoming recurved. Inner involucral bracts 18–20 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the reddish apex. Ligules pale ochre yellow, with a brownish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, olive brown, with few, rather obtuse spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Probably introduced. Grassy and waste places, roadsides and gardens. Scattered records throughout Great Britain and locally common; in Co. Antrim and Co. Londonderry in Ireland. Fennoscandia, Belgium, Holland, Switzerland and Czechoslovakia. 169. T. latissimum Palmgr. Broad-leaved Dandelion Robust, heterophyllous perennial herb with a stock-like rhizome. Leaves numerous, highly crispate, erect; lamina 15– 30 × 3–8 cm, medium green on upper surface, the interlobe area near the midrib usually with dark blotches, oblanceolate in outline, divided almost to the midrib; terminal lobe usually rather small and triangular, or with an elongate apex; lateral lobes 5–9 pairs, rather crowded, regular, deltoid or triangular, with divergent, often forward-pointing, linear, acute to very acute elongations at apex, the margin towards the distal area of the leaf sigmoid, concave or convex and entire or filiform-dentate, the margin towards the proximal area more or less straight, spreading to recurved; interlobe area near the midrib short and narrow, more or less dentate, sometimes with a large tooth; glabrous or nearly so; petiole very variable in length, rose-purple, unwinged. Flowering stem 15–35 cm, green. Capitulum 60–65 mm in diameter, opening. Outer involucral bracts 11–16 × 2–4 mm, dark green and often purple-tipped on outer face, without a pale margin, pale green and pruinose on inner face, lanceolate, acute at apex, twisted laterally, spreading. Inner involucral bracts 18–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules pale ochre yellow, with a brownish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, olive brown, with few, acute spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Introduced. Grassy and waste places and roadsides. In widely scattered localities throughout Great Britain but a scarce species. Fennoscandia.

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170. T. latens H. Øllg. Cut-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 10–20 × 4–6 cm, pale to medium green on upper surface, green to slightly pinkish on midrib, narrowly elliptical-oblong in outline, divided almost to base; terminal lobe of outer leaves small, triangular and with an elongated tip, of inner leaves larger, triangular or helmet-shaped, more or less acute at apex and sometimes divided; lateral lobes 5–6 pairs, regular, crowded, deltoid to triangular, more or less acute at apex, the margin towards the distal area of the leaf straight, more or less convex or sigmoid and entire or dentate, the margin towards the proximal area entire or sometimes with 1 tooth, spreading to recurved; interlobe area near the midrib short and narrow, sometimes dentate; glabrous or nearly so; petiole short or medium, pink, narrrowly winged or unwinged. Flowering stem 10– 25 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 10–15 × 3.0–3.5 mm, dark green on outer face, with a pale margin, paler green on inner face, lanceolate, acute at apex, recurved to reflexed. Inner involucral bracts 14–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw brown, with spinulose projections at apex; cone 0.5– 0.6 mm; beak 10–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Waste and grassy places. Five records from Essex, Shropshire, Denbighshire and Cheshire. Czechoslovakia, Germany, Holland and Denmark. 171. T. edmondsonianum H. Øllg. Edmondson’s Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, decumbent to erect; lamina 20–30 × 3–6 cm, yellowish-green on upper surface, the internode area near the midrib somewhat blotched, dark green to slightly pinkish on midrib, narrowly ellipticaloblong in outline, divided nearly to the midrib; terminal lobe usually triangular, more or less acute at apex, entire or with 1–2 teeth and sometimes divided; lateral lobes 4–6 pairs, regular, triangular or deltoid, acute at apex, the margin towards the distal area of the leaf more or less straight or sigmoid and entire or dentate, the margin towards the proximal area more or less straight and entire or with 1 tooth, spreading; interlobe area near the midrib short and fairly wide, moderately plicate and occasionally dentate; glabrous or slightly hairy, on inner leaves sometimes very hairy; petiole short or medium, pink to purple with a narrow green wing. Flowering stem 20–40 cm, green. Capitulum 45–55 mm in diameter, opening. Outer involucral bracts 14–15 × 4–5 mm, dark green on outer face, with a narrow pale border, pale bluish-green on inner face, lanceolate, acute at apex, strongly recurved. Inner involucral bracts 13–15 × 3–4 mm, dark green with a pale margin, oblonglanceolate, obtuse at apex. Ligules yellow, with a greyishviolet stripe on outer face. Styles exserted, yellow when fresh, discoloured when dry. Pollen present. Achenes with body 2.5–4.0 mm, straw brown, with spiniform projections

at apex; cone 0.4–0.7 mm; beak 10–12 mm, pale; pappus 7–9 mm, whitish. Flowers 4–6. Native. Recorded from Monmouthshire, Herefordshire, Cheshire, Lancashire, Cumberland and Midlothian; widespread around Chester. Denmark and Germany. Named after T. Edmondson. 172. T. acroglossum Dahlst. Broad-bracted Dandelion T. praeradians auct. Squat to robust perennial herb with a stock-like rhizome. Leaves numerous, flat, prostrate to ascending; lamina 12– 35 × 6–9 cm, dark greyish-green on upper surface, the interlobe area near the midrib often with dark blotches, green to slightly pinkish on midrib, narrowly oblong in outline, divided nearly to midrib; terminal lobe usually rather small, triangular or with elongated apex and basal corners; lateral lobes 4–8 pairs, regular, deltoid, sometimes narrowed to an elongated, linear, acute to very acute apex, the margin towards the distal area of the leaf straight, more or less convex or sinuate and entire to filiform-dentate, the margin towards the proximal area straight or concave, recurved to falcate; interlobe area often rounded, subentire near the midrib; shortly hairy on upper surface; petiole rather short, pink, with narrow, entire wings. Flowering stem 12–35 cm, green below, suffused violet above, arachnoid-hairy. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 12–14 × 4.0–5.5 mm, green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex. Inner involucral bracts 16–17 × 2–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achene with body 2.0–2.5 mm, straw-coloured, with short, acute, spiniform projections at apex; cone 0.3–0.4 mm; beak 13– 14 mm, pale; pappus 11–12 mm, whitish. Flowers 4–6. 2n = 24. Agamosperm. Probably introduced. Grassy and waste places and roadsides. Scattered records throughout Great Britain, but apparently uncommon. Native of Sweden and Czechoslovakia. 173. T. exsertum Hagend., Soest & Zevenb. Pale-green Dandelion Squat, sometimes robust perennial herb with a stock-like rhizome. Leaves numerous, prostrate to ascending; lamina 15–35 × 5–7 cm, pale green on upper surface and flaccid, the interlobe area near the midrib often with dark blotches, green to slightly pinkish on midrib, oblong to narrowly elliptic-oblong in outline, divided nearly to the midrib; terminal lobe usually rather small, triangular, with an elongated, acute apex and sometimes elongated basal corners; lateral lobes 4–8 pairs, regular, deltoid, sometimes elongated, linear, acute or very acute at apex, the margin towards the distal area of the leaf straight, more or less convex or sinuate and entire to filiform-dentate, the margin towards the proximal area straight or concave, recurved to falcate; interlobe area near the midrib more or less dentate, often rounded; hairy on midrib only; petiole short or medium, pink, with entire sometimes narrow wings. Flowering stem 15–35 cm, green, arachnoid-hairy. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts

34. Taraxacum 12–14 × 4.0–4.5 mm, green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex. Inner involucral bracts 18–22 × 2–4 mm, dark green, oblong-lanceolate and strongly callosed at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles sometimes yellow when fresh, discoloured when dry. Pollen present. Achene with body 2.0–2.5 mm, straw-coloured, with dense, short spiniform projections at apex; cone 0.3– 0.4 mm; beak 11–13 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Difficult to distinguish from T. acroglossum when dry and perhaps over-recorded. It differs in its pale green, more flaccid leaves and being hairy only on the midrib. Introduced. Grassy and waste places and roadsides. Scattered localities throughout much of Great Britain, the Channel Islands and Co. Roscommon and Co. Down in Ireland. Native of Holland. 174. T. exsertiforme Hagend., Soest & Zevenb. Erect-bracted Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves fairly numerous, prostrate to erect; lamina 10–25(–30) × 3–5 cm, pale to medium green on upper surface, the interlobe area by the midrib not obviously blotched, green to slightly pinkish on midrib, oblong in outline, divided nearly to the midrib; terminal lobe triangular to hastate, more or less acute at apex, entire; lateral lobes 5–7 pairs, deltoid, more or less acute at apex to occasionally obtuse, the apex often narrowed into a linear elongation which is sometimes forward-pointing, the margin towards the distal area of the leaf straight or sinuate, the margin towards the proximal area more or less straight, spreading; interlobe area near the midrib short and narrow, sometimes toothed; glabrous or nearly so; petiole short to medium, green on outer leaves, more or less reddish on inner leaves, narrowly winged. Flowering stem 10–30(–35) cm, green. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 10–15 × 3–4 mm, green on outer face, with a pale margin, paler green on inner face, lanceolate, acute at apex, erect to appressed or more or less spreading. Inner involucral bracts 18–20 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen absent. Achene with body 2.5– 3.0 mm, straw-coloured, with spiniform projections at apex; cone 0.4–0.5 mm; beak 11–13 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Agamosperm. This is the only species of the section with the combination of erect-spreading outer involucral bracts and no pollen, and is perhaps best placed in the Section Celtica. Introduced. Grassy and waste places. Recorded only from Somerset, Denbighshire, Flintshire and Cheshire. Holland and Czechoslovakia. 175. T. exacutumMarkl. Imbricate-bracted Dandelion T. spilophyllum auct. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, flat, erect; lamina 10–22 × 3–6 cm, clear dark green on upper surface, the interlobe area near the

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midrib sometimes with dark blotches, usually green on midrib, sometimes slightly pink, oblong in outline, divided nearly to midrib; terminal lobe triangular or with the apex and basal corners elongate, very narrowly and sharply acute or acuminate; lateral lobes 5–7 pairs, regular, deltoid, acute to acuminate at apex, often abruptly narrowed from a broad base, the margin towards the distal area of the leaf convex or sinuate and entire or filiform-dentate on the lower lobes, the margin towards the proximal area straight or concave, recurved; interlobe area near the midrib short and narrow, sometimes dentate; glabrous or nearly so; petiole short to medium, pink to dull purple, narrowly winged. Flowering stem 10–22 cm, green, very arachnoid-hairy when young. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 12–14 × 4.0–4.5 mm, green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex, imbricate, spreading or rarely recurved. Inner involucral bracts 18–22 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw-coloured, with spiniform projections at apex; cone 0.3–0.4 mm; beak 8–10 mm, pale; pappus 5–7 mm, whitish. Flowers 4–6. Probably introduced. Waste and grassy places and rubbish dumps. Scattered records, particularly in urban areas, throughout Great Britain and in Co. Tyrone in Ireland. Finland. 176. T. valens Markl. Reflexed-bracted Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 10–22 × 3–5 cm, very pale bright green on upper surface, usually green to slightly pinkish on midrib, flat, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe variable, often larger than lateral lobes, hastate, subacute to obtuse at apex, sometimes divided, sometimes long and waisted; lateral lobes 3–5 pairs, regular, deltoid, more or less acute at apex, the margin towards the distal area of the leaf convex or sinuate and entire or filiform-dentate on lower lobes, the margin towards the proximal area more or less straight, spreading to recurved; interlobe area near the midrib short and narrow, occasionally denticulate; glabrous or nearly so; petiole short to medium, pink to dull purple, narrowly winged or unwinged. Flowering stem 10–25 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 10–14 × 3–4 mm, green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex. Inner involucral bracts 14–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achene with body 2.5–3.0 mm, straw-coloured, with conical spiniform projections at apex; cone 0.3–0.4 mm; beak 10–12 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Waste and grassy places. Recorded from Somerset, Surrey, Herefordshire, Berkshire, Oxfordshire, Bedfordshire and Lancashire. Finland, Baltic Russia and Czechoslovakia.

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177. T. obtusifrons Markl. Obtuse-leaved Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, flat, erect; lamina 10–22 × 3–8 cm, medium green on upper surface, usually green to slightly pinkish on midrib, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe tending to be longer than the other lobes, triangular, subacute at apex, entire; lateral lobes 3–5 pairs, regular, deltoid, subacute to obtuse at apex, the margin towards the distal area of the leaf convex to sinuate and entire to filiform-dentate on lower lobes, the margin towards the proximal area straight or concave, spreading to recurved; interlobe area near the midrib short and narrow, often toothed; glabrous or nearly so; petiole short to medium, red to distinctly purple, unwinged or narrowly winged. Flowering stem 10–25 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 12–14 × 4.0–4.5 mm, green on outer face and sometimes with a distinct pale margin, paler green on inner face, lanceolate, acute at apex, erect to appressed or more or less spreading. Inner involucral bracts 18–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achene with body 2.5–3.0 mm, straw brown, with spinulose projections at apex; cone 0.7–0.8 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Waste and grassy places. A few scattered records from Devonshire to Northumberland, often in urban habitats. It has persisted at a single site in inner Newcastle for 14 years without spreading. Finland and Czechoslovakia. 178. T. leptodon Markl. Yellow-styled Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 15–30 × 5– 8 cm, medium to dark green on upper surface, the interlobe area near the midrib not obviously blotched, green to slightly pinkish on midrib, oblanceolate in outline, divided nearly to midrib; terminal lobe sagittate or helmet-shaped, on inner leaves the apex narrowly elongated, sometimes divided or with a single tooth, usually there are a few smaller leaves with characteristic larger ones; lateral lobes 3–5 pairs, regular, deltoid, more or less acute at apex, the margin towards the distal area of the leaf more or less straight and entire or filiform-dentate, the margin towards the proximal area straight to concave, spreading to somewhat recurved; interlobe area near the midrib short to fairly long and narrow, sometimes denticulate; hairy; petiole short to medium, red to distinctly purple, unwinged or narrowly winged. Flowering stem 15–30 cm, green. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 12–14 × 2.0–3.5 mm, green on outer face, sometimes with a slightly paler margin, paler green suffused with purple on inner face, lanceolate, acute at apex. Inner involucral bracts 13–18 × 2– 4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, yellowish. Pollen present. Achene with body 3.0–3.5 mm, straw brown, with thick spiniform projections at apex; cone 0.6–0.7 mm;

beak 12–13 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Introduced. Waste and grassy places. A few records from Surrey, Gloucestershire, Staffordshire and Cheshire. Fennoscandia and Germany. 179. T. pannulatum Dahlst. Brown-ribbed Dandelion T. tarachodum Hagend., Soest & Zevenb. Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, crispate, decumbent to erect; lamina 10–20 × 2–4 cm, greyish-green to medium green on upper surface, the interlobe area near the midrib somewhat dark blotched, brown on midrib, oblong-oblanceolate in outline, divided seven-eighths of the way to the midrib; terminal lobe sagittate, triangular-cordate or with falcate basal corners and lingulate apex; lateral lobes 5–8 pairs, variable, crowded, triangular or deltoid, acute to very acute at apex, the margin towards the distal area of the leaf straight, convex or sinuate and dentate at least on the lower lobes, the margin towards the proximal area straight or concave, recurved or more or less falcate; interlobe area near the midrib short and fairly wide, dentate; glabrous or nearly so; petiole short, pinkish, winged. Flowering stem 10–20 cm, green or pinkish, more or less arachnoid-hairy. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 15–17 × 3.5– 5.0 mm, green on outer face, without a pale margin, paler on inner face, linear-lanceolate, acute at apex. Inner involucral bracts 16–22 × 4–5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with short, broad, spiniform projections at apex; cone 0.3–0.4 mm; beak 12–14 mm, pale; pappus 11–12 mm, whitish. Flowers 4–6. 2n = 24. Probably introduced. Grassy and waste places and roadsides. Scattered records throughout Great Britain and in Co. Tyrone in Ireland. Northern Europe, Germany and France. 180. T. lingulatum Markl. Long-bracted Dandelion T. aequatum Dahlst.; T. subpallescens Dahlst. Medium-sized to large heterophyllous perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, spreading to erect, very variable; lamina 10–25 × 3–5 cm, dull green on upper surface, the interlobe area near the midrib sometimes with dark blotches, green to slightly pinkish on midrib, oblanceolate in outline, divided nearly to the midrib; terminal lobe usually rather small and triangular on outer leaves, larger and triangular on inner leaves, narrowed to an obtuse apex; lateral lobes 3–6 pairs, deltoid or triangular, more or abruptly narrowed from a broad base to an acute apex, the margin towards the distal area of the leaf convex to sinuate and entire to filiform-dentate on lower lobes, the margin towards the proximal area straight or concave, patent to recurved; interlobe area near the midrib rather long and narrow, sometimes dentate with a large tooth; glabrous or nearly so; petiole short to medium, that of the outer leaves pale green, that of the inner rose and winged, sometimes narrowly so. Flowering stem 10–25 cm, pale greenish. Capitulum 50–55 mm in diameter, opening. Outer involucral bracts 11–14 × 3.5–5.0 mm, dark green

34. Taraxacum on outer face, without a pale margin, paler green and pruinose on inner face, lanceolate, acute at apex. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the reddish apex. Ligules deep yellow, with a violet-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achene with body 2.7–3.4 mm, straw-coloured, with numerous, sharp spiniform projections at apex; cone 0.5–0.6 mm; beak 8–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. Native. Roadsides and waste and grassy places. Widespread in Great Britain where it is one of the commonest dandelions; in Co. Kildare, Co. Armagh and Co. Antrim in Ireland. Fennoscandia, Belgium, Holland, France, Czechoslovakia and Switzerland. 181. T. macranthoides G. E. Haglund Large-flowered Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, usually decumbent to erect; lamina 8–22 × 3–5 cm, medium to yellowish-green on upper surface, the interlobe area near the midrib not obviously blotched, the midrib green to slightly pinkish, oblong in outline, divided nearly to the midrib; terminal lobe lingulate to narrowly hastate, obtuse at apex, entire; lateral lobes 4–6 pairs, regular, deltoid, acute to very acute at apex, margin towards the distal area of the leaf straight to convex, and entire or rarely filiformdentate, the margin towards the proximal area straight or concave and entire, spreading to recurved; interlobe are near the midrib short, narrow and dentate; glabrous or nearly so; petiole short, green to faintly pink, narrowly or broadly winged. Flowering stem 8–28 cm, green. Capitulum 40–50 cm in diameter, opening. Outer involucral bracts 11–14 × 3.5–5.0 mm, dark green on outer face, with a slightly paler margin, paler geen on inner face, lanceolate, acute at apex. Inner involucral bracts 16–18 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligule yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with dense spiniform projections at apex; cone 0.5–0.6 mm; beak 9–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5. Agamosperm. Introduced. Recorded only for Chazey in Oxfordshire, Derbyshire and Lancashire. Sweden and Czechoslovakia. 182. T. rhamphodes G. E. Haglund Robust Dandelion Robust perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, erect; lamina 15–30 × 4–6 cm, medium green on upper surface, green to slightly pinkish on midrib, oblong or narrowly elliptical-oblong in outline, divided almost to midrib; terminal lobe variable, triangular, subacute at apex, sometimes divided, the lobules rounded; lateral lobes 5–6 pairs, regular, triangular or deltoid, more or less abruptly narrowed from a broad base to a more or less acute apex, the margin towards the distal area of the leaf more or less convex and entire or filiform-dentate on the lower lobes, the margin towards the proximal area straight or concave, spreading to recurved; interlobe area near midrib short and narrow, mostly entire,

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upturned at edges; glabrous or nearly so; petiole short or medium, pink, narrowly winged. Flowering stem 15–30 cm, greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 11–13 × 3.0–4.5 mm, dark green on outer face, without a pale margin, greyish-green and pruinose on inner face with pink tips, lanceolate, obtuse at apex, mostly recurved. Inner involucral bracts 13–16 × 3–4 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.0–2.5 mm, straw-coloured, with spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 7– 8 mm, whitish. Flowers 4–6. Introduced. Grassy and waste places and roadsides. Scattered records throughout England and Wales; Dumfries-shire, Lanarkshire, Berwickshire and Perthshire in Scotland. 183. T. procerisquameum H. Øllg. Large-headed Dandelion T. procerum auct. Large, robust perennial herb with a stock-like rhizome. Leaves numerous, nearly flat, decumbent to erect; lamina 15–30 × 4–8 cm, greyish-green on upper surface, often brownish on midrib, broadly oblanceolate, divided nearly to the midrib; terminal lobe of outer leaves rather large, sagittate to hastate, obtuse at apex and usually unilaterally 1-divided, on the inner leaves smaller with a longer more acute apex; lateral lobes 3–6 pairs, hamate, with a very long-acute apex, the inner leaves more dentate than outer; interlobe area near midrib short and narrow, entire or with a large tooth; shortly hairy; petiole fairly long, green and winged, sometimes widely. Flowering stem 15–35 cm, greenish. Capitulum 50–70 mm in diameter, opening. Outer involucral bracts 16–18 × 4.0–4.5 mm, green on outer face without a pale margin, paler green on inner face, lanceolateoblong, acute at apex, reflexed. Inner involucral bracts 18–22 × 2–3 mm, dark green, oblong-lanceolate obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, dark straw brown, with short, spiniform projections at apex; cone 0.6–0.8 mm; beak 14– 16 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Introduced. Waste places, mostly on road margins. Very scattered records in England, Wales and southern Scotland and Co. Meath in Ireland. Denmark, Finland, Sweden and Germany. 184. T. vastisectum Markl. ex Puol. Crowded-lobed Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, nearly flat, erect; lamina 15– 30 × 3–7 cm, medium green on upper surface, usually green to slightly pinkish on midrib, oblong-oblanceolate in outline, divided seven-eighths of the way to the midrib; terminal lobe triangular, obtuse to subacute at apex, sometimes divided; lateral lobes 5–6 pairs, regular, crowded in the centre of the leaf, deltoid to triangular, more or less acute at apex, the margin towards the distal area of the leaf straight, more or less convex or angled and entire

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or dentate, the margin towards the proximal area straight and entire or with 1 large tooth, spreading to recurved; interlobe area near midrib short and fairly wide, entire or dentate; glabrous or nearly so; petiole short to long, pink, with narrow, green wings or unwinged. Flowering stem 15– 30 cm, greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 11–13 × 3–5 mm, dark green, with a pale margin, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 15–18 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw-coloured, with numerous, narrow spiniform projections at apex; cone 0.5–0.6 mm; beak 9–10 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Grassy and waste places and roadsides. Scattered records through England and Wales and southern Scotland and in the Shetland Islands. Northern Europe, Belgium, France, Germany, Czechoslovakia, Holland and Switzerland. 185. T. remanentilobum Soest Falcate-lobed Dandelion Medium-sized to robust, heterophyllus perennial herb with a stock-like rhizome. Leaves numerous, nearly flat, erect; lamina 15–30 × 3–7 cm, medium green on upper surface, usually green to slightly pinkish on midrib, oblong or oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe sometimes larger than other lobes particularly on the middle leaves, triangular, acute and sometimes apiculate at apex, sometimes divided; lateral lobes 5–6 pairs, deltoid or more or less falcate, more or less acute at apex, the margin towards the distal area of the leaf convex and entire or rarely dentate, the margin towards the proximal area concave to straight and entire, recurved. Flowering stem 15–30 cm, greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 11–13 × 3.0–3.5 mm, dark green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex, mostly recurved. Inner involucral bracts 14–16 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw brown, with spiniform projections at apex; cone 0.6–1.0 mm; beak 8–10 mm, pale; pappus 5– 6 mm, whitish. Flowers 4–5. Introduced. Grassy and waste places. Recorded only from Somerset, Oxfordshire, Staffordshire and Breconshire. France. 186. T. cordatum Palmgr. Entire-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, spreading to erect; lamina 10–20 × 2–5 cm, rather pale, pure green on upper surface, green on midrib, elliptical-oblong in outline, divided nearly to midrib; terminal lobe triangular, obtuse at apex, entire or divided, cordate at base; lateral lobes 3–5 pairs, regular, deltoid, subacute to obtuse at apex, the margin towards the distal area of the leaf straight to somewhat concave and entire or occasionally dentate, or sometimes

filiform-dentate on the lower lobes, the margin towards the proximal area more or less straight or convex; interlobe area near the midrib short and narrow, entire or dentate; glabrous or nearly so; petiole medium, the outer green, the inner pink, with narrow green wings or unwinged. Flowering stem 10–20 cm, greenish, erect. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 10–12 × 3.0– 3.5 mm, dark green on outer face, without a pale margin, green suffused with violet on inner face, lanceolate, acute at apex, spreading. Inner involucral bracts 17–20 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the purplish apex. Ligules deep yellow, with a reddish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, olive brown, with numerous, unequal spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, pale; pappus 6–10 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 23, 24. Native. Cliffs, sand-dunes, waste and grassy places and roadsides. Common throughout much of Great Britain and with scattered localities in Ireland. Northern and western Europe and Czechoslovakia. 187. T. sagittipotens Dahlst. & R. Ohlsen ex G. E. Haglund Smooth Dandelion Medium-sized to large but rather slender perennial herb with a stock-like rhizome. Leaves rather few to numerous, erect; lamina 20–30 × 3–5 cm, clear medium green to greyish-green on upper surface, green to slightly pinkish on midrib, flat, oblong-oblanceolate in outline, divided threequarters of the way to the midrib; terminal lobe up to 50 mm, sagittate to hastate-sagittate, subacute to acute at apex and sometimes with the tip elongate, entire, often characteristically waisted; lateral lobes 3–4 pairs, regular, deltoid to triangular, acute at apex, the margin towards the distal area of the leaf more or less straight, or slightly convex, convex or sinuate and entire or occasionally filiform-dentate or with 1 tooth, the margin towards the proximal area straight or slightly convex; interlobe area near midrib short and wide, usually entire; glabrous or nearly so; petiole short or medium, those of outer leaves pale green, those of inner ones pink, the narrow wings green. Flowering stem 20–35 cm, greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 9–11 × 2.5–4.0 mm, dark green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 14–16 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules clear yellow, with a violet-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw-coloured, with spiniform projections at apex; cone 0.6–0.7 mm; beak 7–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. 2n = 24. Probably native. Grassy and waste places and roadsides. Scattered records through England and Wales particularly in the south, east and north-east, in Dumfries-shire, Lanarkshire and the Shetland Islands in Scotland and Co. Dublin in Ireland. Northern Europe and Germany. Smaller, neater plants with short, regular leaf-lobes recorded from Surrey, Middlesex, Bedfordshire, Co. Durham and Northumberland may be a distinct species.

34. Taraxacum 188. T. hexhamense A. J. Richards Hexham Dandelion Robust perennial herb with a stock-like rhizome. Leaves few, erect; lamina 20–30 × 3–5 cm, clear, often pale green on upper surface, clear reddish-purple on midrib, oblanceolate in outline, divided about half of the way to midrib; terminal lobe helmet-shaped, rounded-obtuse-apiculate at apex, often rather long, entire or divided; lateral lobes 3–4(–5) pairs, subdeltoid, acute or obtuse at apex, the margin towards the distal area of the leaf concave or sigmoid, the margin towards the proximal area convex, spreading; interlobe area, near midrib shallow, short and wide and entire or minutely and regularly denticulate; glabrous or nearly so; petiole a clear reddish-purple, the green wings up to one-third the length of the leaf. Flowering stem 20– 35 cm, greenish, flushed purplish at base, arachnoid-hairy above. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 8–11 × 2.4–3.5 mm, dark green on outer face scarcely with a pale margin, pale green sometimes suffused purplish on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 12–14 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a stripe on outer face. Styles exserted, yellowish or faintly discoloured. Pollen present. Achenes with body 2.7– 2.9 mm, reddish straw brown, with short, spiniform projections at apex; cone 0.3–0.4 mm, poorly demarcated; beak 8–10 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Native. Grows on banks, walls and shaded humid areas in a limited part of Hexham in Northumberland and has been recorded from Devonshire, Berkshire, Suffolk, Breconshire and Carmarthenshire. Endemic. 189. T. amplum Markl. Toothed Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, prostrate to ascending; lamina (15–)25–39 × 3–8 cm, very pale green on upper surface, green on midrib, narrowly elliptic-oblong in outline, divided nearly to the midrib; terminal lobe helmetshaped, hastate or triangular, often dentate and once divided; lateral lobes 4–6 pairs, regular, deltoid, acute at apex, the margin towards the distal area of the leaf straight, concave or convex and more or less dentate or denticulate, the margin towards the proximal area more or less straight; interlobe area near the midrib short and narrow, dentate; glabrous or nearly so; petiole short or medium, pink to purple, unwinged or with green wings. Flowering stem 15–30 cm, greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 11–14 × 3.5–4.5 mm, dark green on outer face, with or without a paler margin, paler green suffused with violet on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 18–20 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with greyish-violet stripe on outer face. Styles exserted, yellowish. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with sharp, irregular spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5. Agamosperm. Introduced. Grassy and waste places. Recorded only from Gloucestershire, Flintshire and Cheshire. Fennoscandia and Czechoslovakia.

189

190. T. ekmanii Dahlst. Ekman’s Dandelion T. connexum Dahlst. Medium-sized to robust, strongly heterophyllus perennial herb with a stock-like rhizome. Leaves numerous, crispate, spreading; lamina 15–30 × 3–8 cm, bright, pale, lettuce green on upper surface, green to more or less pinkish on midrib, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe shortly triangular, entire, in late leaves larger, dentate and irregularly divided; lateral lobes 3–6 pairs, on early leaves narrowly triangular with a lingulate apex, scarcely dentate, the margin towards the distal area of the leaf more or less sigmoid and the margin towards the proximal area more or less straight, on late leaves broad and highly dentate; interlobe area near the midrib short and narrow, entire or dentate; glabrous or nearly so; petiole short or medium, green on outer leaves, pale pink on inner leaves, with narrow to broad green wings. Flowering stem 15–35 cm, greenish. Capitulum 45–55 mm in diameter, opening. Outer involucral bracts 9–13 × 2–4 mm, dark green and suffused violet on outer face, with or without a pale margin, paler green suffused with violet on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 18–20 × 2–3 mm, medium green, oblonglanceolate, obtuse at the violet apex. Ligules deep yellow, with a violet-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw-coloured, with rather long, irregular spiniform projections at apex; cone 0.4–0.7 mm; beak 8–10 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. Probably native. Grassy and waste places and roadsides. Throughout Great Britain where it is common and locally abundant, and round the perimeter of Ireland. North and western Europe east to Austria, Czechoslovakia, Switzerland and northern Italy. Named after E. Ekman (1883– 1931). 191. T. ochrochlorum G. E. Haglund ex Rail. Winged Dandelion Medium-sized to robust, not strongly heterophyllus perennial herb with a stock-like rhizome. Leaves numerous, crispate, spreading to erect; lamina 15–30 × 7–10 cm, very pale, dull green on upper surface, green to somewhat pinkish or golden brown on midrib, oblong-elliptical-oblong, or lanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe in early leaves shortly triangular and entire, in late leaves much larger, dentate and irregularly divided; lateral lobes 3–6 pairs, on early leaves narrowly triangular, with a lingulate apex, the margin towards the distal area of the leaf more or less sigmoid and scarcely dentate, the margin towards the proximal area straight with a more or less conspicuous tooth, on late leaves with gross, few, very variable teeth; interlobe area near the midrib short and narrow or short and wide and dentate; glabrous or nearly so; petiole short, pale pink, with very broad, green wings. Flowering stem 15–35 cm, greenish, very arachnoid-hairy when young. Capitulum 45– 55 mm, opening. Outer involucral bracts 9–13 × 2–4 mm, green, suffused with violet on outer face, with or without a pale margin, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts

190

1 4 8 . A S T E R AC E A E

15–17 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a violet-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw brown, with spiniform projections at apex; cone 0.4–0.7 mm; beak 8–10 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Woodland banks, shaded road verges or grassy places. Very scattered records in Great Britain from Somerset to Aberdeenshire, but may be underrecorded because of its closeness to T. ekmanii. Fennoscandia, Baltic Russia and Czechoslovakia. 192. T. aurosulum H. Lindb. Tailed Dandelion Medium-sized to robust, strongly heterophyllus perennial herb with a stock-like rhizome. Leaves numerous, crispate, spreading to erect; lamina 15–30 × 3–6 cm, medium green, the interlobe area near the midrib sometimes with dark blotches, green to pinkish on midrib, oblong or ellipticaloblong in outline, divided nearly to the midrib; terminal lobe in early leaves small, triangular, often with an elongated apex and more or less entire or sometimes divided, in late leaves large, dentate and irregularly divided; lateral lobes 3–6 pairs, the early leaves narrowly triangular, lingulate at apex, and often narrowing into linear projections, often dentate, often divided, the margin towards the distal area of the leaf more or less sigmoid and the margin towards the proximal area more or less straight, the later leaves with gross, variable, few, large, broad teeth; glabrous or nearly so; petiole short, green on outer leaves, pale pink on inner leaves, with green, narrow to broad wings. Flowering stem 15– 35 cm, greenish, slightly arachnoid-hairy. Capitulum 45– 55 mm, opening. Outer involucral bracts 9–13 × 2–4 mm, green, suffused with violet on outer face, with or without a distinct pale margin, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 14–20 × 2–3 mm, dark green with a pale margin, oblonglanceolate, obtuse at the purplish apex. Ligules deep yellow, with a violet-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw brown, with short, acute spiniform projections at apex; cone 0.3–0.7 mm; beak 8–10 mm, pale; pappus 6–10 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Shady roadside verges and waste places. Widely scattered records in Great Britain, Channel Islands and Co. Galway and Co. Antrim in Ireland. Fennoscandia, Belgium, Holland, France, Switzerland and Czechoslovakia. 193. T. lunare M. P. Christ. Lunar-lobed Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, crispate, spreading to erect; lamina 15–30 × 3–8 cm, medium to dark green on upper surface, green to somewhat pinkish on the midrib, crisped, oblong or oblong-oblanceolate in outline, divided to the midrib; terminal lobe of early leaves shortly triangular and more or less entire, of later leaves large, dentate and irregularly divided; lateral lobes of early leaves 3–6 pairs, lunar-shaped with an elongated apex, the later leaves with very variable, few, large recurved teeth; interlobe area near the midrib short and narrow, dentate; glabrous or nearly so; petiole short, green, narrowly to broadly winged. Flowering stem 15–35

cm, greenish. Capitulum 45–55 mm in diameter, opening. Outer involucral bracts 12–16 × 3–5 mm, green, suffused with pale violet on outer face, with or without a pale margin, paler green on inner face, oblong-lanceolate, obtuse at apex, recurved. Inner involucral bracts 15–19 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a violet-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw brown, with spiniform projections at apex; cone 0.4–0.7 mm; beak 8–10 mm, pale; pappus 6– 7 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Grassy and waste places. Recorded only from Gloucestershire, Staffordshire, Breconshire, Carmarthenshire and Lancashire. Belgium, Denmark, Switzerland, Holland, Sweden and Czechoslovakia. 194. T. coartatum G. E. Haglund Irregular-bracted Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, spreading to erect; lamina 10–25 × 2–6 cm, medium green on upper surface, the internode area near the midrib not obviously blotched, green to slightly pinkish on the midrib, oblong or oblong-oblanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe not usually large, triangular, often equilaterally so, acute at apex, entire or sometimes divided; lateral lobes 4–6 pairs, regular, deltoid, acute to subacute at apex from a broad base, the margin towards the distal area of the leaf convex and often highshouldered, the margin towards the proximal area more or less straight; interlobe area near the midrib short and narrow, dentate, sometimes with a large tooth; glabrous or nearly so; petiole short to medium, pink, with green wings. Flowering stem 10–30 cm, distinctly arachnoidhairy throughout, greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 13–15(–18) × 3.5– 4.5 mm, green on outer face and not obviously pruinose, without a pale margin, paler green on inner face, lanceolate, acute at apex, irregularly arranged or twisted and spreading to recurved. Inner involucral bracts 18–20 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules pale yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw brown, with spiniform projections at apex; cone 0.4–0.5 mm; beak 8–10 mm, pale; pappus 6– 7 mm, whitish. Flowers 4–5. Agamosperm. Introduced. Grassy and waste places. Recorded from Devonshire, Gloucestershire, Herefordshire and Denbighshire. Sweden and Czechoslovakia. 195. T. aberrans Hagend., Soest & Zevenb. Apiculate-lobed Dandelion T. caudatulum auct. Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, erect; lamina 10–25 × 3–5 cm, dull medium green on upper surface, often drying yellowish-green, not obviously blotched and green to slightly pinkish on the midrib, oblong-oblanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe usually large, triangular or helmet-shaped, apiculate at apex, entire, dentate or sometimes divided,

34. Taraxacum sometimes cordate at base; lateral lobes 3–4 pairs, regular, deltoid, rather short with a broad base, more or less acute at apex, the margin towards the distal area of the leaf more or less sinuate and usually dentate or sometimes entire or filiform-dentate, the margin towards the proximal area more or less straight, spreading to slightly recurved; interlobe area near the midrib short and fairly wide, usually dentate; glabrous or nearly so; petiole fairly long, pink, unwinged at least below. Flowering stem 10–30 cm, greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–14 × 3.5–4.5 mm, green and pruinose on outer face, but without a paler margin, paler green on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 18–20 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present or absent. Achenes with body 2.8–3.0 mm, straw-coloured, with spiniform projections at apex; cone 0.4–0.5 mm; beak 12–14 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. 2n = 24. Probably introduced. Grassy and waste places and roadsides. Recorded from Devonshire, Gloucestershire, Carmarthenshire, Yorkshire, Northumberland, Workington in Cumberland and Kirkcudbrightshire but seems to be uncommon. Holland, Belgium and France. An unnamed, related species differs in having dark, dirtycoloured leaves bearing dark, acuminate teeth, rounded terminal leaf-lobes which are strongly recurved-dentate and divided and dark brown achenes with a body only about 2.5 mm. It is related to the Danish species T. severum M. P. Christ. It is recorded from grassy places in Devonshire, Gloucestershire, Monmouthshire, Breconshire and Lancashire and is possibly endemic. 196. T. pseudoretroflexum M. P. Christ. Spur-lobed Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, erect; lamina 10–30 × 3–8 cm, medium green on upper surface, the internode area near the midrib not obviously blotched, green to slightly pinkish on midrib, oblong or narrowly oblongelliptical in outline, divided nearly to midrib; terminal lobe triangular with a well-differentiated, elongated tip; lateral lobes 3–5 pairs, regular, deltoid, acute to very acute at apex, the margin towards the distal area of the leaf convex and entire to filiform-dentate, the margin towards the proximal area more or less straight, spreading to slightly recurved; interlobe area near the midrib short and narrow, more or less dentate; petiole short to medium, pink, unwinged at least towards the base. Flowering stem 10– 35 cm, greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–14 × 3.5–4.5 mm, green on outer face, with a pale margin, paler green on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 16–18 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with incurved spiniform projections at apex; cone 0.4–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm.

191

Introduced. Grassy and waste places. Recorded from Dorset, Breconshire, Denbighshire and Lancashire. Denmark and Czechoslovakia. 197. T. oblongatum Dahlst. Oblong-leaved Dandelion T. perhamatum Dahlst. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, erect; lamina 5–20 ×1– 6 cm, medium to olive green on upper surface, not obviously blotched, and green to slightly pinkish on midrib, oblongoblanceolate in outline, divided nearly to the midrib; terminal lobe usually rather small on outer leaves, larger on inner, rounded-ovate-cordate, sometimes minutely apiculate at apex, more or less entire; lateral lobes 3–5 pairs, regular, deltoid, subacute to acute at apex, the margin towards the distal area of the leaf more or less convex and entire or filiform-dentate, the margin towards the proximal area straight to sinuate; interlobe area near the midrib short and narrow to fairly wide, dentate; glabrous or nearly so; petiole medium to rather long, deep pink, unwinged or with narrow, green wings towards the base. Flowering stem 5–25 cm, erect, greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 9–12 × 2–3 mm, very dark green on outer face, without a paler margin, paler green and often suffused with purple on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 15–18 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured to blackish. Pollen present. Achenes with body 2.5–2.8 mm, straw-coloured, with spiniform projections at apex; cone 0.4–0.6 mm; beak 10–12 mm, pale; pappus 5–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Native. Grassy places, especially fertile, damp pastures. Widespread and locally common in England, Wales and southern Scotland, rather rare in northern Scotland; Channel Islands; in Co. Wexford and Co. Dublin in Ireland. Sweden, Finland and Holland, Czechoslovakia and possibly Norway. 198. T. cophocentrum Dahlst. Rounded-lobed Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves rather few, flat, erect; lamina 15–30 × 3–7 cm, dull dark green on upper surface, green to slightly pinkish on midrib, flat, oblanceolate, divided two-thirds of the way to the midrib; terminal lobe large, about one-quarter the length of the leaf, helmet-shaped, rounded, obtuse or subacute at apex, entire or dentate towards base; lateral lobes 1–3 pairs, regular, broadly deltoid or triangular, more or less acute at apex, the margin towards the distal area of the leaf more or less sinuate to convex and more or less dentate, the margin towards the proximal area more or less straight; interlobe area near the midrib short and wide, entire or dentate; glabrous or nearly so; petiole long, dull purple, with narrow, green wings or unwinged towards the base. Flowering stem 15–30 cm, erect, rather narrow, greenish. Capitulum 40–45 mm in diameter, opening. Outer involucral bracts 8–9 × 1.8–2.0 mm, very dark green on outer face, without a pale margin, paler and often becoming suffused with purple on inner face, lanceolate, acute at apex,

192

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spreading to recurved. Inner involucral bracts 12–15 × 2– 3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a brownish-purple stripe on outer face. Styles exserted, slightly discoloured. Pollen present. Achenes with body 3.5–3.7 mm, straw brown, with spiniform projections at apex; cone 0.4–0.6 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers; 4–6. Native. Grassy places, wood margins and scrub. Frequent in southern England, scattered records in Wales and further north; Channel Islands; Co. Galway and Co. West Meath in Ireland. Endemic. 199. T. pachymerum G. E. Haglund Dirty-leaved Dandelion Medium-sized, squat perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, prostrate to ascending; lamina 15–25 × 3–6 cm, dark to bluish-green on upper surface, centrally smudged with dirty markings, the interlobe area near the midrib with dark blotches, pinkish to dirty red on midrib, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe usually rather small, triangular, obtuse to subacute at apex, often dentate or divided, often cordate at base; lateral lobes 4–6 pairs, crowded, regular, deltoid, acute at apex, the margin towards the distal area of the leaf convex and more or less denticulate or dentate, the margin towards the proximal area straight, to convex, spreading to recurved; interlobe area near the midrib very short and sometimes just a point, the interlobe area V-shaped; glabrous or nearly so; petiole short to medium, pink to purple, with narrow to broad, green wings. Flowering stem 15–25 cm, greenish. Capitulum 30–40 mm in diameter, open. Outer involucral bracts 11–14 × 3.5– 4.5 mm, dark green on outer face, without a paler margin, paler green suffused violet on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 17–20 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.2–3.5 mm, straw-coloured, with short spiniform projections at apex; cone 0.5–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. 2n = 24. Probably introduced. Grassy and waste places and roadsides. Scattered records through England, Wales and southern Scotland; recorded for Co. Dublin in Ireland. Sweden. 200. T. dilatatum H. Lindb. Grassland Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, decumbent to erect; lamina 15–30 × 3–8 cm, dull medium green on upper surface, the margin, especially of the interlobe area, sometimes blackish, green to slightly pinkish on midrib, narrowly oblong-oblanceolate or oblong in outline, divided nearly to the midrib; terminal lobe on outer leaves rather small and triangular, on inner leaves large, helmet-shaped and often divided; lateral lobes 3–5 pairs, regular, deltoid or triangular and acute to very acute at apex, the margin towards the distal area of the leaf straight, convex or sinuate and often dentate, sometimes with large teeth, the margin towards the proximal area straight or concave, spreading to

recurved; interlobe area near the midrib short and narrow to fairly wide, often dentate with 1 long, broadbased, filiform tooth; glabrous or nearly so; petiole short to medium, clear red with narrow, parallel-sided green wings. Flowering stem 5–20 cm, greenish. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 11–13 × 3–5 mm, dark green on outer face, without a pale margin, paler and conspicuously brownish-purple on inner face, lanceolate, acute at apex, spreading. Inner involucral bracts 15–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse to subacute at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with short, sharp spiniform projections at apex; cone 0.4–0.5 mm; beak 6–10 mm, pale; pappus 6–10 mm, whitish. Flowers 4–6. 2n = 24. Related to some species in the Section Celtica and may be better placed in that section. Native. Grassy places, particularly rich grasslands, roadsides and waste places. Scattered records in western Great Britain from Devonshire to Lanarkshire. Northern Europe south to Germany and southern France.

201. T. diastematicum Markl. Bulbous-lobed Dandelion Medium-sized to large perennial herb with a stock-like rhizome. Leaves fairly numerous, slightly crispate, decumbent to erect; lamina 15–25 × 2–6 cm, dull medium green on upper surface, the margin, particularly of the interlobe area sometimes blackish, pale towards the base of the petiole and darkening towards the apex with striations, narrowly oblong in outline, divided nearly to the midrib; terminal lobe triangular or with the basal corners elongate, entire, obtuse at apex; lateral lobes 3–5 pairs, deltoid to triangular, acute to very acute at apex, the margin towards the distal area of the leaf straight, convex or sinuate, rapidly narrowing to give a characteristic high shoulder or hump and sometimes dentate or with large teeth, the margin towards the proximal area straight or concave, spreading or recurved; interlobe area near the midrib rather long and narrow, often dentate with 1 long, broad-based, filiform tooth; glabrous or nearly so; petiole rather long, pale green on the outer leaves, reddish on the inner leaves, with narrow, green wings. Flowering stem 5–20 cm, greenish. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 11–13 × 3–5 mm, dark green on outer face, without a pale margin, whitish or green or faintly pinkish, but not obviously pruinose on inner face, lanceolate, acute at apex, spreading. Inner involucral bracts 16–18 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.2–3.5 mm, straw brown, with spiniform projections at apex; cone 0.4–0.5 mm; beak 7–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5. Agamosperm. Introduced. Grassy and waste places. Recorded only from Northamptonshire, Breconshire, Flintshire and Cheshire. Sweden and Czechoslovakia.

34. Taraxacum 202. T. tanyphyllum Dahlst. Spreading-lobed Dandelion Medium-sized to tall perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, ascending to erect; lamina 8–30 × 2–5 cm, medium to rather dark green on upper surface, the interlobe area near the midrib sometimes with slight dark blotches, green or slightly pinkish and striate on midrib, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe more or less hastate, subacute at apex, more or less entire; lateral lobes 4–6 pairs, triangular or deltoid, drawn out to a parallel-sided, more or less acute apex, the margin towards the distal area of the leaf straight to convex and entire, dentate or filiformdentate, the margin towards the proximal area straight or concave, spreading or slightly recurved; interlobe area near the midrib rather long and narrow, often dentate; glabrous or nearly so; petiole rather long, of outer leaves sometimes green, of middle and inner leaves reddish, with narrow, green wings. Flowering stem 8–30 cm, pale greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 13–15 × 2–4 mm, green with a pale margin on outer face, paler green on inner face, lanceolate, acuminate at apex, recurved. Inner involucral bracts 18–20 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules pale yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–4.0 mm, straw brown, with short spiniform projections at apex; cone 0.6–0.7 mm; beak 7–10 mm, pale; pappus 6– 7 mm, whitish. Flowers 4–6. Introduced. A rare plant of roadsides and waste places. Widely scattered records in Great Britain from Cornwall to the Shetland Islands. Norway and Sweden. 203. T. sinuatum Dahlst. Sinuate-lobed Dandelion Medium-sized, squat perennial herb with a stock-like rhizome. Leaves numerous, crispate, spreading to erect; lamina 12–20 × 4–6 cm, shiny medium green on upper surface, pink on midrib, narrowly elliptical-oblong in outline, divided nearly to the midrib; terminal lobe usually rather small, obtuse to subacute at apex, divided; lateral lobes 3–5 pairs, irregular, oblong-lanceolate, subacute to obtuse at apex, more or less sinuate on both margins, frequently divided, frequently with large teeth and pointing in various directions; interlobe area near the midrib short and narrow to fairly wide and dentate; glabrous or nearly so; petiole short to medium, purple, unwinged. Flowering stem 5–25 cm, pale greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 9–11 × 2–4 mm, dark green on outer face, with a more or less pale margin on outer face, paler green and somewhat pruinose on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–18 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.0–2.5 mm, olive brown, with spiniform projections at apex; cone 0.3–0.4 mm; beak 7–10 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Agamosperm. 2n = 24. Native. Grassy places and roadsides. Scattered records through western England and Wales, the Middle Ebudes in

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Scotland and Co. Dublin in Ireland. Norway, Sweden and Czechoslovakia. 204. T. laciniosifrons Wiinst. Hairy-stemmed Dandelion Medium-sized to squat perennial herb with a stock-like rhizome. Leaves numerous, crispate, spreading to erect; lamina 12–20 × 2–5 cm, dull medium green on upper surface, near the interlobe area the midrib frequently with dark blotches, pink on midrib, broadly oblong to elliptical-oblong, divided almost to the midrib; terminal lobe usually rather small, obtuse to subacute at apex, usually divided; lateral lobes 3–5 pairs, irregular, broad at base and elongated at the more or less acute apex, sinuate on both margins, frequently divided, frequently long-dentate with an occasional large tooth on the proximal margin, pointing in all directions; interlobe area near the midrib short and thin to fairly wide, dentate; shortly hairy; petiole rather short, with purplishgreen wings. Flowering stem 12–25 cm, pale greenish, sometimes arachnoid-hairy throughout. Capitulum 40– 50 mm in diameter, opening. Outer involucral bracts 10– 17 × 3–5 mm, dark green on outer face, with a pale margin, paler green and somewhat pruinose on inner face, lanceolate, acute at apex, recurved or reflexed. Inner involucral bracts 18–22 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.2–2.5 mm, olive brown, with spiniform projections at apex; cone 0.3–0.4 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Probably native. Grassy places and roadsides. Scattered records through England and Wales, Dumfries-shire, Kirkcudbrightshire and Fifeshire in Scotland, and Co. Roscommon, Co. Down, Co. Antrim and Co. Londonderry in Ireland. Northern Europe, Belgium, Holland, Germany, Switzerland, and Czechoslovakia. 205. T. dahlstedtii H. Lindb. Dahlstedt’s Dandelion T. densiflorum M. P. Christ., non Brenner Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, ascending to erect; lamina 8–25 × 2–3 cm, medium to rather dark green on upper surface, the interlobe area near the midrib sometimes with dark blotches, green to slightly pink on midrib, slightly crisped, oblong-oblanceolate in outline, divided seven-eighths of the way to the midrib; terminal lobe large and helmet-shaped, more or less acute at apex, dentate or divided; lateral lobes 4–6 pairs, triangular or deltoid, more or less acute at apex, the margin towards the distal area of the leaf straight to convex and entire, dentate or filiform-dentate, the margin towards the proximal area straight or concave and sometimes with a single tooth, spreading to slightly recurved; interlobe area near the midrib short and rather wide, often dentate, somewhat hairy; petiole rather long and narrow, vivid crimson-purple, unwinged. Flowering stem 8–25 cm, pale greenish. Capitulum 30– 40 mm in diameter, opening. Outer involucral bracts 11– 15 × 1.5–3.0 mm, dark green on outer face, without a pale margin, paler green on inner surface, lanceolate, acute at apex, recurved. Inner involucral bracts 15–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse

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at apex. Ligules pale yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–4.0 mm, straw-coloured, with numerous, sharp spiniform projections at apex; cone 0.6–0.7 mm; beak 6–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24, 27. Native. Grassy and waste places and roadsides. Widespread through Great Britain and Ireland and locally common especially near the coast. Northern Europe south to France, Germany and Czechoslovakia. Named after Hugo Gustaf Adolf Dahlstedt (1856–1934). 206. T. obliquilobum Dahlst. Orange-tipped Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, decumbent to erect; lamina 8–25 × 4–6 cm, dull, olive green on upper surface, sometimes appearing to be blotched when dried, green to slightly pinkish on midrib, oblong-oblanceolate in outline, divided nearly to midrib or the inner leaves not lobed; terminal lobe triangular, obtuse to acute at apex, dentate or divided; lateral lobes 4–6 pairs, deltoid, more or less acute at apex, the margin towards the distal area of the leaf straight to convex and entire, dentate or filiform-dentate, the margin towards the proximal area straight or concave and sometimes with a single tooth, recurved; interlobe area short and narrow, often dentate; glabrous or nearly so or arachnoid-hairy on midrib; petiole fairly long and narrow, fairly red or pink, with narrow, green wings. Flowering stem 8–25 cm, pale greenish at base, often purplish below, more or less arachnoid-hairy. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 14–17 × 3.0– 5.0 mm, green on outer face, without a pale margin, paler green on inner face, lanceolate, obtuse at apex, the margins strongly recurved, spreading to recurved. Inner involucral bracts 14–15 × 2.0–3.5 mm, dark green with a pale margin, oblong-lanceolate, narrow and obtuse at the purplish apex. Ligules pale yellow, with a greyish-violet stripe on outer face, the inner with reddish-orange tips. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–4.0 mm, straw brown, with narrow, erect-patent spiniform projections at apex; cone 0.6–0.7 mm; beak 11–13 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Introduced. Waste places and roadsides. Scarce in scattered localities throughout Great Britain from Dorset to Morayshire. Northern Europe south to France and Switzerland. 207. T. pachylobum Dahlst. Thick-lobed Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, slightly crispate, ascending to erect; lamina 8–25 × 3–5 cm, medium to rather dark green on upper surface, the interlobe area near the midrib sometimes with slightly dark blotches, green to slightly pinkish on midrib, oblong to oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe triangular, more or less acute at apex, sometimes divided; lateral lobes 4–6 pairs, triangular or deltoid, more or less acute at apex, but some obviously rounded, the margin towards the distal area of the leaf straight to convex and entire, dentate or filiformdentate, the margin towards the proximal area straight or

concave and sometimes with a single tooth, spreading or slightly recurved; interlobe area near the midrib short and narrow or rather wide, often dentate; glabrous or nearly so, or sparsely arachnoid-hairy on midrib; petiole fairly short and narrow, vivid crimson-purple, with narrow, green wings. Flowering stem 8–25 cm, pale greenish above, purplish below, arachnoid-hairy above. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 11–15 × 1.5– 3.0 mm, green on outer face, without a pale margin, paler green on inner face, lanceolate, obtuse at apex, spreading. Inner involucral bracts 12–14 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, attenuate at the purplish apex. Ligules pale yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–4.0 mm, straw brown, with thick spiniform projections at apex; cone 0.6–0.7 mm; beak 12– 14 mm, pale; pappus 10–12 mm, whitish. Flowers 4–5. Introduced. Grassy places. Rare, recorded only from Breconshire and a sunken lane at Embleton in Cumberland. Northern Europe and Germany. 208. T. latisectum H. Lindb. Broad-lobed Dandelion T. subdilatatum Dahlst. nom. nud. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, more or less flat, erect; lamina 5–20 × 2– 8(–10) cm, bluish-green on upper surface, green or slightly pinkish on midrib, more or less flat, oblong-oblanceolate in outline, divided three-quarters of the way to the midrib; terminal lobe usually rather small on outer leaves, larger on inner, minutely apiculate at apex, more or less entire, cordate at base; lateral lobes 3–5 pairs, crowded, regular, deltoid, more or less acute, rarely obtuse at apex, the margin towards the distal area of the leaf more or less convex and dentate or filiform-dentate or rarely entire, the margin towards the proximal area straight or sinuate, recurved; interlobe area near the midrib short and fairly wide, usually entire; glabrous or nearly so; petiole short to medium rather dull pink, with narrow, green wings or unwinged towards the base. Flowering stem 5–25 cm, pale greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 9– 12 × 2.5–3.5 mm, very dark green on outer face, without a pale margin, paler and often becoming suffused with purple on inner face, lanceolate, more or less acute at apex, spreading to recurved. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face and suffused blackish at apex. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–2.8 mm, straw brown, with few, irregular, obtuse or acute spiniform projections at apex; cone 0.6–0.7 mm; beak 7–10 mm, pale; pappus 6–9 mm, whitish. Flowers 4–6. Native. Grassy places, especially fertile pastures. Scattered through England and Wales, in Dumfries-shire, Stirlingshire and the Shetland Islands in Scotland and Co. Antrim in Ireland. Fennoscandia and Baltic Russia. 209. T. huelphersianum Dahlst. ex G. E. Haglund H¨ulphers’s Dandelion Tall perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, decumbent to erect; lamina 8–30 × 3–8 cm, dull dark green on upper surface, the

34. Taraxacum

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Introduced. Grassy places and roadsides. A few scattered records in Great Britain from Devonshire to East Lothian. Norway. Named after Lars Fagerstr¨om (b. 1914).

interlobe area near the midrib with dark blotches, green to slightly pinkish on midrib, oblong or oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe of outer leaves small and triangular, that of the inner leaves larger, more or less acute at apex, entire to dentate and sometimes divided; lateral lobes 5–7 pairs, regular, deltoid, acute or tapering and very acute at apex, the margin towards the distal area of the leaf straight to convex, or angled with a heavy-shouldered effect and entire or dentate to filiform-dentate, the margin towards the proximal area more or less straight and entire or subulate-dentate, patent to recurved; interlobe area near the midrib short and narrow, sometimes dentate; somewhat hairy; petiole long, pale green on outer leaves, pink on inner leaves, with narrow, green wings. Flowering stem 8–30 cm, sometimes distinctly arachnoid-hairy. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 12–15 × 3–4 mm, dark green on outer face without a pale margin, pale green on inner face, lanceolate, acute at apex, spreading to form a star. Inner involucral bracts 18–20 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the violet apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw-coloured, with spiniform projections at apex; cone 0.5–0.7 mm; beak 11–13 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Grassy and waste places and roadsides. Scattered throughout England, Wales and southern Scotland. Fennoscandia, Baltic Russia and Czechoslovakia. Named after A. H¨ulphers.

211. T. hepaticum Rail. Regular-lobed Dandelion Tall perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, spreading to erect; lamina 8– 30 × 3–7 cm, bluish-green on upper surface, the interlobe area near the midrib unblotched, green or slightly tinted pinkish on midrib, oblong or narrowly oblong-lanceolate, divided much of the way to the midrib; terminal lobe often small, broadly triangular-hastate, obtuse at apex, entire or slightly dentate; lateral lobes 4–5 pairs, regular, acute or obtuse at apex, entire or slightly dentate, spreading or slightly recurved; interlobe area near the midrib short and narrow, often dentate; glabrous or nearly so; petiole rather short, rose-purple, with narrow to broad wings. Flowering stem 10–35 cm, pale green, densely arachnoid-hairy above. Capitulum 50–65 mm in diameter, opening. Outer involucral bracts 10–15 × 4–6 mm, dark green on outer face, without a paler margin, paler glaucous-green on inner face, lanceolate, acute at apex, reflexed. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblonglanceolate, obtuse at apex. Ligules pale ochre-yellow, with a brownish-purple stripe on outer face. Styles exserted, yellow. Pollen present. Achenes with body 3.5–4.0 mm, straw brown, with dense, slender spiniform projections at apex; cone 0.6–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Known only from Breconshire. Finland and Czechoslovakia.

210. T. fagerstroemii S˚altin Fagerstr¨om’s Dandelion T. sublaearifolium Hagend., Soest & Zevenb. Medium-sized to tall perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, decumbent to erect; lamina 8–30 × 3–7 cm, pale to medium green on upper surface, the interlobe area sometimes with a dark margin, green to slightly pinkish on midrib, oblong to narrowly elliptical-oblong in outline, divided nearly to midrib; terminal lobe of outer leaves small and triangular, that of the inner leaves larger, subacute to obtuse at apex, entire or dentate, divided, sometimes more than once; lateral lobes 5–7 pairs, regular, deltoid, more or less acute at the frequently elongated apex, the margin towards the distal area of the leaf straight to convex, or sinuate and denticulate to dentate, the margin towards the proximal area more or less straight and entire or subulate-dentate, spreading to recurved; interlobe area near the midrib short and narrow, sometimes dentate; glabrous or nearly so; petiole long, pink, with narrow green wings. Flowering stem 8–30 cm, glabrous, pale greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 12–15 × 3–4 mm, dark green on outer face, with a slightly paler margin, pale green on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 16–18 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with short spiniform projections at apex; cone 0.6–0.7 mm; beak 8– 10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6.

212. T. subundulatum Dahlst. Complex-leaved Dandelion T. amphiodon Dahlst. ex G. E. Haglund; T. hemipolyodon Dahlst.; ?T. similatum Dahlst.; T. bercheriense Dahlst. nom. nud. Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, crispate, decumbent to erect; lamina 10–25 × 3–6 cm, bright green on upper surface, the interlobe area near the midrib sometimes with darkish blotches, green to slightly pinkish on midrib, ellipticaloblong in outline, divided nearly to midrib; terminal lobe triangular, obtuse to subacute at apex, often dentate and divided; lateral lobes 4–7 pairs, irregular, triangular, acute at apex, often dentate on both margins, the margin towards the distal area of the leaf straight to concave, the margin towards the proximal area straight or convex, spreading or pointing forwards; interlobe area near the midrib short and narrow or fairly wide, more or less dentate; glabrous or nearly so; petiole short, bright rose pink, with narrow green wings towards the base. Flowering stem 10–30 cm, pale greenish. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 12–15 × 4–6 mm, dark green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex. Inner involucral bracts 18–20 × 2– 3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.2–3.5 mm, straw-coloured, with spiniform projections at apex; cone 0.6–0.8 mm;

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beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Native. Herb-rich waste meadows and lush roadsides. A characteristic component of the flora of the Thameside meads. Scattered throughout Great Britain from Hampshire to the Shetland Islands and in Co. Dublin in Ireland. Fennoscandia. 213. T. pulchrifolium Markl. Beautiful-leaved Dandelion Medium-sized to tall robust perennial herb with a stocklike rhizome. Leaves numerous, decumbent to erect; lamina 10–30 × 3–10 cm, very pale green on upper surface, green to slightly pinkish on midrib, narrowly ellipticaloblong in outline, divided nearly to midrib; terminal lobe hastate, from a broad base gradually elongate to a sometimes very thin, more or less acute tip, sometimes divided, rarely dentate; lateral lobes 4–6 pairs, regular, triangular, narrowing rapidly from a broad base to an elongated, acute, spreading or forward-pointing apex, the margin towards the distal area of the leaf straight to concave and dentate, the margin towards the proximal area straight or concave and sometimes dentate; interlobe area near the midrib shortish and narrow, entire or dentate; glabrous or nearly so; petiole short, bright rose pink at base, unwinged. Flowering stem 10–35 cm, pale greenish. Capitulum 35– 45 mm in diameter, opening. Outer involucral bracts 12– 15 × 4–6 mm, dark green on outer face, with a pale margin, paler green on inner face, lanceolate, acute at apex, spreading to erect. Inner involucral bracts 16–20 × 3– 4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes body 3.0–3.5 mm, straw brown, with few acute spiniform projections at apex; cone 0.7–0.8 mm; beak 10–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Scattered on waste ground in cities. It is abundant in the city of Dublin. Scattered records in England and Wales and Co. Dublin in Ireland. Finland, Baltic Russia and Czechoslovakia. 214. T. pectinatiforme H. Lindb. Pectinate-leaved Dandelion Medium-sized, rather delicate perennial herb with a stocklike rhizome. Leaves numerous, flat, erect; lamina 15– 30 × 3–7 cm, dark green on upper surface, pinkish-red on midrib throughout, elliptical-oblong in outline, divided nearly to the midrib; terminal lobe with elongated apex and basal corners or broader and helmet-shaped, divided and sometimes long-dentate; lateral lobes 6–12 pairs, rapidly narrowing from a broad base to a long, narrow, linear, forward-pointing acute apex, often deeply divided into parallel, linear lobules, the margin towards the distal area of the leaf of the basal being usually convex and narrowly dentate to filiform-dentate; interlobe area near the midrib short and narrow, dentate or filiform-dentate; somewhat hairy; petiole very short and poorly differentiated, reddish-purple. Flowering stem 15–30 cm, pale greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 10–12 × 2.5– 3.5 mm, green on outer face, without a pale margin, paler

green and sometimes suffused reddish on inner face, lanceolate, acute at apex. Inner involucral bracts 16–18 × 2–3mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules pale yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–4.0 mm, straw-coloured, with numerous, sharp, irregular spiniform projections at apex; cone 0.5–0.6 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. Probably introduced. Grassy and waste places and roadsides. Scattered throughout Great Britain from Somerset to the Shetland Islands and in Co. Antrim in Ireland. Northern Europe south to France, Germany, Switzerland and Czechoslovakia. 215. T. caloschistum Dahlst. Brilliant-stalked Dandelion Medium-sized, rather delicate perennial herb with a stocklike rhizome. Leaves numerous, flat, erect; lamina 15– 30 × 3–5 cm, medium green on upper surface, green to faintly pinkish on midrib, flat, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe with narrow, elongated apex and basal corners, or triangular, or helmet-shaped, obtuse at apex, divided or sometimes long-dentate; lateral lobes 6–10 pairs, triangular at base, rapidly narrowing to become a long, narrow, linear, forwardpointing, acute or obtuse apex, sometimes divided into parallel, linear lobules, the margin towards the distal area of the leaf of the basal part usually convex and narrowly dentate to filiform-dentate; interlobe area near the midrib short and fairly narrow, sometimes dentate; somewhat hairy; petiole very short and poorly differentiated, deep reddish-purple, unwinged. Flowering stem 15–30 cm, pale greenish, sparsely arachnoid-hairy. Capitulum 40– 50 mm in diameter, opening. Outer involucral bracts 10– 12 × 2.5–3.5 mm, green on outer face, without a pale margin, paler green sometimes suffused reddish on inner face, ovate-lanceolate or lanceolate, acute at apex, recurved or reflexed. Inner involucral bracts 13–15 × 2–3 mm, medium green, oblong-lanceolate, obtuse at the violet apex. Ligules pale yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–4.0 mm, straw-coloured, with rather few, short spiniform projections at apex; cone 0.5–0.6 mm; beak 10–12 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Grassy and waste places. Recorded from Flintshire, Cheshire, Stirlingshire and Perthshire. Northern Europe south to Germany, Switzerland and Czechoslovakia. 216. T. trilobatum Palmgr. Three-lobed Dandelion Robust, untidy perennial herb with a stock-like rhizome. Leaves numerous, decumbent to erect; lamina 15–30 × 3–8 cm, dull green on upper surface, the interlobe area near the midrib sometimes with dark blotches, more or less pinkish on midrib, strongly crisped, elliptical-oblong in outline, divided nearly to the midrib; terminal lobe, helmetshaped or sagittate, narrowed at apex into a lingulate, acute to very acute tip, entire or dentate; lateral lobes 5–7 pairs, rapidly narrowing from a broad base to form a long, narrow,

34. Taraxacum linear, often divided, acute or swollen-obtuse apex, the distal margin of the basal area concave and grossly dentate, the proximal margin straight or concave and entire or sometimes with a big tooth, spreading; interlobe area near the midrib short and narrow, sometimes denticulate; somewhat hairy; petiole short, shining violet-purple, with broad, green wings. Flowering stem 15–30 cm, pale greenish. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 12–15 × 2–3 mm, dark green on outer face, without a pale margin, paler green on inner face, linear-lanceolate, acute at apex, recurved. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, dark to blackish. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with large, recurved spiniform projections at apex; cone 0.8–1.0 mm, spinous at base; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Introduced. Grassy and waste places and roadsides. Scattered records through Great Britain, particularly near the coast from Devonshire to Aberdeenshire. Northern Europe south to France and Switzerland. 217. T. planum Raunk. Diverse-leaved Dandelion Squat, somewhat robust perennial herb with a stock-like rhizome. Leaves numerous, prostrate to ascending; lamina 12–35 × 3–9 cm, very pale green on upper surface, the interlobe area near the midrib often with dark blotches, green to slightly pinkish on midrib, oblong to narrowly ellipticoblong in outline, divided nearly to the midrib; terminal lobe of inner leaves sometimes larger than lateral lobes, helmet-shaped or triangular, long acute at apex or with an elongated tip and the basal corners elongated; lateral lobes 4–8 pairs, regular, crowded, deltoid, acute or very acute apex and sometimes elongated and linear, the margin towards the distal area of the leaf straight, more or less convex or sinuate and entire to filiform-dentate, the margin towards the proximal area straight, more or less concave, recurved to falcate; interlobe area near the midrib short and narrow and more or less dentate, upturned at margin; glabrous or nearly so; petiole rather short, pink or green on outer leaves, winged, sometimes narrowly so. Flowering stem 12–35 cm, green, glabrous. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 12–14 × 3.5–5.0 mm, green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex. Inner involucral bracts 18–20 × 2–3 mm, dark green with a pale margin, oblong-lanceolate at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achene with body 2.0–2.5 mm, straw-coloured, with spiniform projections at apex; cone 0.3–0.4 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Introduced. Verges of major roads. Very scattered records in Great Britain from Essex and Monmouthshire to Perthshire. Denmark. 218. T. polyodon Dahlst. Common Dandelion T. ardisodon Dahlst.; T. naeviferum auct. Medium-sized, heterophyllous perennial herb with a stocklike rhizome. Leaves numerous, more or less flat, spreading to erect; lamina 8–20(–30) × 4–6 cm, medium green on

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upper surface, the interlobe area near the midrib usually with dark blotches at first, green to slightly pinkish on midrib, more or less flat, oblong to elliptic-oblong in outline, divided nearly to the midrib; terminal lobe on outer leaves usually rather small and triangular, obtuse to subacute at apex, but sometimes with an elongated tip, usually entire, but sometimes divided, on inner leaves often much larger and dentate and divided; lateral lobes 5–6 pairs, regular, deltoid, more or less acute at apex, the margin towards the distal area of the leaf straight or convex and strongly dentate, the teeth often black-tipped, spreading to slightly recurved, on the later leaves often much larger and less regularly lobed often losing the black blotches and teeth; interlobe area near the midrib short and narrow to fairly wide, dentate; glabrous or nearly so or more or less arachnoid-hairy on the midrib; petiole rather short, pale to deep pink, unwinged or with narrow, green wings. Flowering stem 8–20(–30) cm, pale greenish and slightly purplish below, more or less arachnoid-hairy above. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 9–11 × 2–3 mm, dark green on outer face, without a pale margin, paler green suffused with purple, becoming purple on inner face, irregularly untidy and claw-like, spreading to recurved. Inner involucral bracts 14–22 × 2–5 mm, dark green with a pale margin, oblong-lanceolate, attenuate at the slightly purplish apex. Ligules yellow, with a purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.2 mm, straw-coloured, with irregular, short, broad spiniform projections at apex; cone 0.6– 0.7 mm; beak 10–12 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Agamosperm. T. polyodon is best distinguished from the other species with highly dentate leaves, blotched interlobe areas and pink petioles by its untidy, wispy, purple, claw-like outer involucral bracts. It is easily identified early in the year, but is problematical later. Native or possibly introduced. Grassy and waste places and roadsides. Throughout Great Britain and much of Ireland and one of our commonest dandelions. Northern Europe south to France, Germany and Czechoslovakia. 219. T. multicolorans Hagend., Soest & Zevenb. Many-coloured Dandelion Medium-sized to robust, heterophyllous perennial herb with a stock-like rhizome. Leaves numerous, crispate, spreading to erect; lamina 8–20(–30) × 3–6 cm, dark green on upper surface, rough, the interlobe area near the midrib with obvious dark blotches, pink on midrib, oblongoblanceolate in outline, divided nearly to midrib; terminal lobe on outer leaves usually small and long-triangular, obtuse to subacute at apex but sometimes with elongated tip, often denticulate to dentate and divided and cordate at base, on inner leaves often much larger, dentate and divided; lateral lobes 5–6 pairs, regular, deltoid, more or less acute at apex, the margin towards the distal area of the leaf straight or convex or concave and usually dentate, the teeth often black-tipped, the margin towards the proximal area more or less straight and usually entire, spreading or slightly recurved; interlobe area near the midrib short and narrow to fairly wide, dentate; glabrous or nearly so; petiole rather long, dark purple, unwinged or with narrow,

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green wings. Flowering stem 8–20(–30) cm, pale greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–14 × 3.0–3.5 mm, green on outer face, with a pale margin, paler green on inner face, lanceolate, acute at apex, spreading. Inner involucral bracts 16–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules medium to dark yellow, with a purple stripe on outer face. Styles exserted, yellowish. Pollen present. Achenes with body 3.0–3.2 mm, straw-coloured, with spiniform projections at apex; cone 0.6–0.7 mm; beak 7–10 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Introduced. Grassy and waste places. A few records for southern England and Monmouthshire and Lancashire. Holland. 220. T. nitidum Hagend., Soest & Zevenb. Shining Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 8–20 × 4–6 cm, medium to dark shiny green on upper surface, the interlobe area with a dark margin, pinkish on midrib, oblong or oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe of outer leaves usually rather small and triangular, obtuse to acute at apex but sometimes with an elongated tip and usually entire but sometimes divided, on inner leaves often much larger, dentate and divided; lateral lobes 5–6 pairs, regular, deltoid, more or less acute at apex, the margin towards the distal area of the leaf straight or convex and strongly dentate, the teeth often black-tipped, the margin towards the proximal area more or less straight or convex and usually entire, spreading to slightly recurved; interlobe area near the midrib short to rather long and narrow, dentate; glabrous or nearly so; petiole short to medium, dark purple, with narrow to broad green wings. Flowering stem 8–20 cm, pale greenish, arachnoid-hairy. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 9–11 × 3.0–3.5 mm, dark green on outer face, with a faint pale margin, paler green suffused with purple, becoming purple on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 17–19 × 3–4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.2 mm, straw brown, with spiniform projections at apex; cone 0.6–0.7 mm; beak 11–12 mm, pale; pappus 5–6 mm, whitish. Flowers 4–6. Introduced. A few scattered records in England and Wales from Somerset to Cumberland. Holland. 221. T. incisum H. Øllg. Incise-leaved Dandelion Medium-sized, rather delicate perennial herb with a stocklike rhizome. Leaves fairly numerous, crispate, spreading to erect; lamina 8–20(–30) × 4–6 cm, dark green on upper surface, the interlobe area near the midrib strongly dark blotched, green to slightly pinkish on midrib, ellipticoblong to oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe triangular, with elongated apex and basal corners, often divided and long-dentate; lateral lobes 5–8 pairs, distant, triangular to deltoid, rapidly narrowing from a broad base to a more or less acute apex,

strongly dentate on both margins, spreading to slightly recurved; interlobe area near the midrib fairly long and narrow, markedly dentate; glabrous or nearly so; petiole short to medium, pink, unwinged or rarely with narrow green wings. Flowering stem 8–20(–30) cm, pale greenish, tinted violet, more or less arachnoid-hairy. Capitulum 45–50 mm in diameter, opening. Outer involucral bracts 10–12 × 2.5– 4.5 mm, green on outer face, paler green becoming suffused with purple on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 15–16 × 3– 4 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–3.8 mm, straw-coloured, with spiniform projections at apex; cone 0.6–0.7 mm; beak 9–11 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Introduced. Grassy and waste places and roadsides. Scattered localities through Great Britain from the Thames basin to the Shetland Islands. Denmark. 222. T. xanthostigma H. Lindb. Ochre-styled Dandelion Medium-sized to robust, squat, more or less heterophyllous perennial herb with a stock-like rhizome. Leaves numerous, more or less crispate, erect; lamina 10–25 × 1–5 cm, medium to dark green on upper surface, the internode area near the midrib very obviously with dark blotches, green to slightly pinkish on midrib, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe on outer leaves small and triangular, sagittate or ovate-cordate, on inner leaves large, often hastate and entire or divided; lateral lobes 3–5 pairs, regular, deltoid or triangular, more or less acute at apex, the margin towards the distal area of the leaf straight or slightly angled, rarely convex or concave and entire or the lower lobes filiform-dentate, the margin towards the proximal area more or less straight, spreading to recurved; interlobe area near the midrib short and narrow, often triangular-dentate; glabrous or nearly so; petiole medium, purple, with narrow, green wings. Flowering stem 10–30 cm, erect, pale greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 10–13 × 3–4 mm, green on outer face, with a pale margin, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 12–18 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, yellowish-orange both when fresh and dried. Pollen present. Achenes with body 2.5–3.0 mm, straw-coloured, with few, broad, obtuse spiniform projections at apex; cone 0.4–0.5 mm; beak 6–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Agamosperm. 2n = 24. Possibly introduced. Grassy and waste places and roadsides. Widely scattered records over Great Britain and Ireland. Northern Europe south to Belgium, Germany, Switzerland and Czechoslovakia. An undescribed species similar to T. xanthostigma but a dark green, non-heterophyllous plant with whitish petioles and distinctive outer involucral bracts (the outer narrower than the inner) has been widely recorded in England and Wales from Devonshire to Cumberland.

34. Taraxacum 223. T. longisquameum H. Lindb. Elongate-bracted Dandelion T. adsimile Dahlst.; T. latispina Dahlst.; T. mucronatum auct. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 7–20(–25) × 3–9 cm, medium to dark green on upper surface, the interlobe area near the midrib usually with very obvious dark blotches, green to slightly pinkish on midrib, oblong or narrowly elliptical-oblong in outline, divided nearly to the midrib; terminal lobe narrowly triangular, more or less acute at apex and elongated to a lingulate tip, sometimes divided; lateral lobes 3–5 pairs, regular, deltoid, more or less acute at apex, the margin towards the distal area of the leaf straight to slightly angled, rarely convex or concave and entire or the lower lobes filiform-dentate, the margin towards the proximal area more or less straight, slightly recurved to nearly reflexed; interlobe area near the midrib short and narrow, sometimes dentate; sometimes hairy; petiole short to medium, purple, unwinged or almost so. Flowering stem 7–30 cm, pale greenish. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 8–10 × 3– 4 mm, dark green on outer face, with or without a pale margin, paler green and sometimes suffused purple on inner face, long-lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 18–22 × 3.0–3.5 mm, dark green with a pale margin, oblong-lanceolate, obtuse at the reddish apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, olive brown, with numerous, short, irregular spiniform projections at apex; cone 0.4–0.5 mm; beak 5–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Native. Grassy places and roadsides, often in rather natural habitats especially near the sea. Scattered localities throughout Great Britain and round the perimeter of Ireland. Northern Europe south to Belgium, Germany and Switzerland. 224. T. scotiniforme Dahlst. ex G. E. Haglund Deltoid-lobed Dandelion T. obscuratum G. E. Haglund, non Dahlst. Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 8–15 × 3–5 cm, dark or bluish-green, the interlobe area near the midrib heavily blotched black, green to slightly pinkish on midrib, oblongoblanceolate in outline, divided nearly to the midrib; terminal lobe triangular, obtuse to more or less shortly lingulate at apex, entire; lateral lobes 3–5 pairs, regular, deltoid, more or less acute at apex, the margin towards the distal area of the leaf convex and dentate to filiform-dentate, the margin towards the proximal area straight or concave and entire, spreading to slightly recurved; interlobe area near the midrib short and fairly wide, denticulate; glabrous or nearly so; petiole medium, pink, with narrow green wings. Flowering stem 8–15 cm, pale greenish, thickly arachnoid-hairy when young. Capitulum 30–35 mm in diameter, opening. Outer involucral bracts 10–15 × 4–5 mm, green without a pale margin on outer face, paler green on inner face, lanceolate,

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acute at apex, more or less recurved. Inner involucral bracts 14–18 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, yellow. Pollen present. Achenes with body 3.0–3.5 mm, straw brown, with spiniform projections at apex; cone 0.3–0.4 mm; beak 10–11 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Possibly introduced. Grassy and waste places. Scarce, in England from Kent to Cumberland, in Breconshire in Wales and in the Shetland Islands. Scandinavia and Switzerland. 225. T. maculatum Jord. Yellowish-green Dandelion Medium-sized to robust, squat perennial herb with a stocklike rhizome. Leaves numerous, erect; lamina 10–25(–30) × 3–6 cm, pale yellowish-green on upper surface, but darkening when dry, the interlobe area near the midrib with obvious dark blotches, green to slightly pinkish on midrib becoming darker-coloured towards the apex, oblong in outline, divided nearly to midrib; terminal lobe triangular or hastate, sometimes ligulate at the more or less acute apex, sometimes divided; lateral lobes 5–7 pairs, deltoid or triangular, acute at apex and often narrowing abuptly to form a linear elongation, entire or divided, the margin towards the distal area of the leaf of the basal part convex and entire or denticulate to dentate, the margin towards the proximal area more or less straight; interlobe area near the midrib short and narrow, often dentate; glabrous or nearly so; petiole medium, of outer leaves pale green, of inner leaves reddish, with narrow, green wings. Flowering stem 10–30(–35) cm, pale greenish. Capitulum 50–65 mm in diameter, opening. Outer involucral bracts 10–15 × 3–5 mm, dark green on outer face, without a pale border, paler green on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 17–19 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.8–4.0 mm, straw brown, with spiniform projections at apex; cone 0.4–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–5. Agamosperm. Introduced. Recorded only for Gloucestershire. Fennoscandia, western Europe and Czechoslovakia. 226. T. sublongisquameum M. P. Christ. Roadside Dandelion Medium-sized to robust, squat perennial herb with a stocklike rhizome. Leaves numerous, erect; lamina 10–15 × 3–6 cm, medium to dark green on upper surface, the interlobe area near the midrib sometimes with dark blotches, pink and becoming increasingly darker pink towards the apex of the midrib, oblong-oblanceolate in outline, divided nearly to the midrib; terminal lobe triangular to hastate, acute to apiculate at apex, entire; lateral lobes 3–5 pairs, regular, deltoid or triangular, very acute to apiculate at apex, the margin towards the distal area of the leaf straight or slightly angled, rarely convex or concave and entire or filiformdentate on the lower lobes, the margin towards the proximal area more or less straight, spreading to recurved; interlobe area short and narrow, often dentate; glabrous or nearly so; petiole one-quarter to one-third the length of the leaf, pale green or whitish, narrowly winged. Flowering stem

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10–30 cm, pale greenish. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 13–15 × 3.5–4.5 mm, dark green on outer face, without a pale margin, paler green on inner face, lanceolate, acute at apex. Inner involucral bracts 17–19 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.0 mm, straw brown or greyish-brown, with spiniform projections at apex; cone 0.4–0.7 mm; beak 8–10 mm, pale; pappus 6–7 mm, whitish. Flowers 4–6. Introduced. Roadside verges. Recorded from Surrey, Hertfordshire, Oxfordshire, Monmouthshire, Lancashire and Dunbartonshire. Denmark. 227. T. fasciatum Dahlst. Dense-bracted Dandelion T. sublatissimum Dahlst. Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 10– 25 × 3–5 cm, dark green on upper surface, the interlobe area near the midrib with obvious tar-coloured blotches, green on midrib, oblong in outline, divided almost to midrib; terminal lobe helmet-shaped to subsagittate, often slightly narrowed near the middle, obtuse at apex, but sometimes elongated; lateral lobes 3–5 pairs, regular, deltoid or triangular, acute at apex, the margin towards the distal area of the leaf straight or slightly angled, rarely convex or concave, or sigmoid and entire, or the lower lobes filiform-dentate, the margin towards the proximal area more or less straight, pointing forward, spreading or recurved; interlobe area near the midrib rather long and quite wide, often filiformdentate; glabrous or nearly so; petiole medium, pale rose to dull reddish-purple, with narrow, green wings. Flowering stem 10–30 cm, pale greenish or tinted pink. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 12–15 × 4–5 mm, crowded and imbricate, medium green on outer face, without a pale margin, paler green and somewhat suffused with purple on inner face, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–20 × 2.0–3.0 mm, dark green with a pale margin, oblonglanceolate, obtuse at the purplish apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, straw-coloured, with short, sharp spiniform projections at apex; cone 0.4–0.5 mm; beak 10–12 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Obligate agamosperm. 2n = 24. Possibly native. Grassy and waste places and particularly roadsides. Common in England and Wales, scattered records in Scotland, Co. Dublin and Co. Down in Ireland, and in the Channel Islands. Northern Europe south to Belgium, Germany and Czechoslovakia. 228. T. subxanthostigma M. P. Christ. ex H. Øllg. Purple-stalked Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 15–20 × 3–5 cm, dark green on upper surface, the interlobe area near the midrib tar-coloured, pink to purple on midrib, crisped, narrowly elliptical-oblong, divided three-quarters of the way

to the midrib; terminal lobe rather small, triangular to ovate-sagittate-cordate; lateral lobes 5–6 pairs, more or less triangular, acute at apex, the margin towards the distal area of the leaf more or less straight and entire, the margin towards the proximal area straight; interlobe area near the midrib short and wide, entire or rarely filiform-dentate; glabrous or nearly so; petiole short to medium, darkish pink to purple, with narrow, green wings. Flowering stem 15–25 cm, pale greenish. Capitulum 45–50 mm in diameter, opening. Outer involucral bracts 12–13 × 3.5–4.0 mm, dark green on outer face, without a pale margin, lanceolate, acute at apex, spreading to recurved. Inner involucral bracts 16–20 × 3–4 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.6 mm, straw brown, with spiniform projections at apex; cone 0.6–0.8 mm; beak 10– 12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Agamosperm. Difficult to distinguish from T. fasciatum except when well developed, when the more numerous leaf-lobes, dark purple petioles and long cone of the achene should be looked for. Introduced. Grassy and waste places. Recorded from Berkshire, Iffley in Oxfordshire, Breconshire and Lancashire. Czechoslovakia. 229. T. acutifidum M. P. Christ. Pointed-lobed Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, somewhat crispate, decumbent to erect; lamina 10–25 × 3–5 cm, dull, dark green on upper surface, the interlobe area near the midrib with dark blotches, green to slightly pinkish on midrib, oblongoblanceolate in outline, divided nearly to midrib; terminal lobe triangular or helmet-shaped, subacute to obtuse at apex, sometimes divided; lateral lobes 4–6 pairs, regular, deltoid, acute to very acute at apex, the margin towards the distal area of the leaf straight to convex and dentate with 1 or more large teeth or filiform-dentate, the margin towards the proximal area straight or concave and entire; interlobe area near the midrib short and narrow, often dentate, sometimes with 1 large tooth; glabrous or nearly so; petiole medium, green to faintly pink, with narrow, green wings or unwinged. Flowering stem 10–30 cm, clear green becoming darker below. Capitulum 40–50 mm in diameter, opening. Outer involucral bracts 11–14 × 3–5 mm, dark green on outer face, with a more or less slight pale margin, clear green on inner face, lanceolate, obtuse at apex, spreading to recurved. Inner involucral bracts 20– 22 × 3–4 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-purple stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.5–3.7 mm, straw brown, with short spiniform projections at apex; cone 0.4–0.6 mm; beak 10–12 mm, pale; pappus 7–8 mm, whitish. Flowers 4–6. Introduced. Grassy and waste places. Recorded from Staffordshire, Shropshire, Denbighshire, Flintshire, Cheshire and Lancashire. Denmark and Sweden.

35. Crepis 230. T. melanthoides Dahlst. ex M. P. Christ. & Wiinst. Bluish-leaved Dandelion Medium-sized perennial herb with a stock-like rhizome. Leaves numerous, spreading to erect; lamina 5–18(–25) × 3–5 cm, pale to medium bluish-green on upper surface, the interlobe area near the midrib with dark blotches, green to slightly pinkish on midrib, crisped, oblong to narrowly elliptical-oblong in outline, divided nearly to the midrib; terminal lobe more or less helmet-shaped or sagittate, sometimes elongate at the acute or very acute apex, sometimes divided, often cordate at base; lateral lobes 3–5 pairs, regular, deltoid, acute at apex, the margin towards the distal area of the leaf more or less straight or sigmoid and dentate, the margin towards the proximal area straight or concave, slightly recurved; interlobe area near the midrib fairly, long and narrow, dentate, crispate; hairy; petiole medium, pink to purple, with narrow, green wings. Flowering stem 15–23 (–30) cm, pale greenish. Capitulum 35–45 mm in diameter, opening. Outer involucral bracts 10–12 × 2–4 mm, dark green on outer face, with or without a pale margin, paler green on inner face, lanceolate, acute at apex, recurved. Inner involucral bracts 14–16 × 2–3 mm, dark green with a pale margin, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a dark purplish-brown stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.8–3.0 mm, brown, with spiniform projections at apex; cone 0.4–0.7 mm; beak 7–10 mm, pale; pappus 6– 7 mm, whitish. Flowers 4–6. Agamosperm. This species, with hairy, bluish, sparsely spotted leaves could be classified in the Section Naevaea or Borea Sahlin, but the outer involucral bracts and general facies are of the Section Ruderalia in which it is very distinct among the British and Irish species. Native. Water-meadows, especially those cut for hay, and old pastures. Scattered localities in south and north Wales and the Welsh border, Berkshire and Oxfordshire, northern England and Perthshire. Southern Scandinavia, Belgium, Holland and Czechoslovakia. 231. T. lucidum Dahlst. Large-bracted Dandelion Medium-sized to robust perennial herb with a stock-like rhizome. Leaves numerous, crispate, spreading to erect; lamina 8–22 × 3–5 cm, pale shining green on upper surface, the interlobe area near the midrib often with dark blotches, greenish-pink on midrib, oblong-oblanceolate, divided nearly to the midrib; terminal lobe long, more or less helmet-shaped, obtuse to subacute at apex, sometimes waisted, sometimes divided, sometimes more or less dentate; lateral lobes 2–3 pairs, regular, deltoid, acute at apex, the margin towards the distal area of the leaf more or less straight and dentate, the margin towards the proximal area straight to convex and entire, spreading to recurved; interlobe area near the midrib short and rather wide, denticulate to dentate; glabrous or nearly so; petiole short to medium, shining purple, unwinged, sometimes with filiform teeth. Flowering stem 8–30 cm, pale greenish or flushed purplish, often more or less arachnoid-hairy below the capitulum. Capitulum 30–40 mm in diameter, opening. Outer involucral bracts 10–15 × 3.5–4.0 mm, dark

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green on outer face, with a pale margin, faintly dark green on inner face, lanceolate, subacute at apex, spreading to erect. Inner involucral bracts 12–14 × 3–5 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 3.0–3.5 mm, straw-coloured, with few, very short spiniform projections at apex; cone 0.4–0.5 mm; beak 10–11 mm, pale; pappus 8–10 mm, whitish. Flowers 4–6. Agamosperm. 2n = 24. The involucral bracts and general coloration are highly suggestive of species of the Section Celtica, but when wellgrown it is much too gross, untidy and three-dimensional to be a member of that section. Probably introduced. Grassy and waste places and roadsides. Widely scattered localities in England from Essex to Cumberland. Northern Europe south to Belgium, Germany and Czechoslovakia. 232. T. sundbergii Dahlst. Sundberg’s Dandelion Robust perennial herb with a stock-like rhizome. Leaves numerous, erect; lamina 15–30 × 2–7 cm, medium green or slightly glaucous on upper surface, the interlobes near the midrib sometimes with dark blotches, green to slightly pinkish on midrib, oblong-oblanceolate in outline, divided nearly to midrib; terminal lobe hastate, subacute at apex, rarely divided; lateral lobes 3–5 pairs, regular, deltoid, abruptly more or less acute at apex from a broad base, the margin towards the distal area of the leaf more or less convex and entire or filiform-dentate on lower lobes, the margin towards the proximal area straight to concave, patent to recurved; interlobe area rather long and narrow, sometimes dentate; glabrous or nearly so; petiole rather long, pink, with narrow, green wings. Flowering stem 15–35 cm, pale greenish, sometimes suffused purplish, sparse to dense arachnoid-hairy above. Capitulum 45–45 mm in diameter, opening. Outer involucral bracts 14–16 × 4–6 mm, dark green on outer face, with a slightly pale margin, greyishgreen and pruinose with pink tips on inner face, lanceolate, acute at apex, mostly recurved. Inner involucral bracts 15–22 × 2–3 mm, dark green, oblong-lanceolate, obtuse at apex. Ligules deep yellow, with a greyish-violet stripe on outer face. Styles exserted, discoloured. Pollen present. Achenes with body 2.5–3.5 mm, straw brown, with few, short, acute spiniform projections at apex; cone 0.4– 0.5 mm; beak 9–10 mm, pale; pappus 8–9 mm, whitish. Flowers 4–6. Agamosperm. Introduced. Grassy and waste places. Recorded for Surrey, Bedfordshire, Breconshire and Northumberland. Northern Europe, Czechoslovakia and Portugal. Named after C. F. Sundberg. 35. Crepis L. Aracium Monnier; Soyeria Monnier; Hieraciodes Kuntze Annual to perennial herbs with latex and sometimes with short rhizomes, without stolons. Stems 1–many, usually branched, usually leafy, rarely all leaves basal. Leaves alternate, subentire to pinnately divided. Capitula solitary to numerous. Involucral bracts in 2 rows, the outer up to threequarters the length of the inner. Flowers bisexual. Corolla ligulate, yellow, sometimes with a red stripe on the outer

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face, with 5 lobes at apex. Anthers tailed, with elongate apical appendage; filaments smooth. Style with long, slender branches, with both shaft and branches hairy. Receptacle flat or convex, pitted, the pits often ciliate, rarely with scales. Achenes cylindrical, fusiform or oblong, usually narrowed above, sometimes beaked; pappus of 1–several rows of white, brownish or yellowish-white simple eglandular hairs. About 200 species chiefly in temperate and subtropical Europe and Asia, some in North and central Africa and North America and 1 each in South Africa and South America, but see under Hieracium. Crepis has been split into a number of genera, but we accept Babcock’s (1947) argument that it is not a practical classification. Crepis is best distinguished from Hieracium by its long and short rows of involucral bracts which form a ‘cup and saucer’. Once the species of Crepis are known in the field they should be easily recognisable at a glance by their different general appearance from all species of Hieracium. Main-flowering plants (early summer) of Crepis should not cause any problem in identification, but late summerand autumn-flowering plants can be very difficult to name as they often have smaller capitula and a different facies. To make things more difficult some late-flowering plants look like early-flowering ones, and in areas like roadsides, playing fields and amenity areas, where they are frequently cut back, secondary growth is an added problem. A large, mixed colony of C. biennis, C. capillaris, C. vesicaria subsp. taraxacifolia and possibly C. nicaeensis and C. tectorum flowering in September can produce a bewildering display of variation most of which is not typical. C. pulchra L., C. rubra L., C. sancta (L.) Babc. subsp. nemausensis (Gouan) Babc. and C. suberostris Coss. & Durieu ex Batt. have been recorded as rare casuals. Babcock, E. B. (1947). The Genus Crepis. Univ. Calif. Publ. Bot. 21 & 22. Grime, J. P. et al. (1988). Comparative plant ecology. London. [C. capillaris.] Halliday, G. (1990). Crepis praemorsa (L.) Tausch, new to western Europe. Watsonia 18: 85–87. Marshall, J. B. (1962). Notes on British Crepis. I. Introduction and key. B.S.B.I. Proc. 4: 398–403. Marshall, J. B. (1964). Notes on British Crepis. II. Variants of Crepis capillaris (L.) Wallr. B.S.B.I. Proc. 5: 325–333. Reveal, J. L. (1992). Re-establishment of a discarded name in Crepis L. (Asteraceae). Watsonia 19: 40–41. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [C. mollis.] Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [C. foetida, C. praemorsa.] 1. Stems leafless (but with tiny bracts) 4. praemorsa subsp. praemorsa 2. 1. Stems with obvious leaves 2. Receptacular scales present 6(c). foetida subsp. commutata 3. 2. Receptacular scales absent 3. Outer achenes with a short beak or beak absent, inner 4. achenes with a long slender beak 3. Achenes all the same, or inner and outer slightly 6. different, but grading into one another 9(b). vesicaria subsp. stellata 4. Flowering heads erect 5. 4. Flowering heads drooping

5. Outer involucral bracts up to 0.7 mm wide, linear-lanceolate, about half as long as inner; marginal 6(a). foetida subsp. foetida achenes 7–9 mm 5. Outer involucral bracts 1.0–1.5 mm wide, lanceolate, or oblong-lanceolate, about two-thirds as long as inner; marginal achenes 5–7 mm 6(b). foetida subsp. rhoeadifolia 7. 6. Achenes distinctly beaked 6. Achenes not beaked, but sometimes gradually narrowed 10. at apex 7. Upper part of plant with long, rigid, yellowish, 10. setosa bristle-like hairs 7. Upper part of plant without long, rigid, yellowish, 8. bristle-like hairs 5. tectorum 8. Achenes shortly beaked 9. 8. Achenes with a beak at least as long as body 9. Leaves sparingly hairy; capitula very shortly whitish hairy 9(a,i). vesicaria subsp. taraxacifolia var. taraxacifolia 9. Leaves densely hairy; capitula shortly whitish hairy and with short purplish or reddish glandular hairs 9(a,ii). vesicaria subsp. taraxacifolia var. rutilans 11. 10. Involucral bracts appressed-hairy on their inner faces 12. 10. Involucral bracts glabrous on their inner faces 11. Achenes 4.0–7.5 mm, equally attenuate at both ends, 3. biennis (10–)13–18(–20)-ribbed 11. Achenes (2.5–)3–4(–4.5) mm, strongly attenuate below, 5. tectorum scarcely beaked above, 10-ribbed 1. paludosa 12. Pappus hairs pale brownish-white, brittle 13. 12. Pappus hairs pure white, soft 13. Perennial arising from a short rhizome; achenes about 2. mollis 20-ribbed 14. 13. Annual or biennial with a tap-root; achenes 10-ribbed 14. Outer involucral bracts becoming lax; receptacle with fimbriate-ciliate achene pits; achenes 2.5–3.8 mm 7. nicaeensis 14. Outer involucral bracts appresssed to inner; receptacle with a few hairs around the achene pits; achenes 15. 1.5–2.5 mm 15. Stems few to numerous without an apparent main stem, ascending, diffuse or decumbent; involucral bracts 8(i). capillaris var. capillaris 4.5–7.0 mm 15. Stems solitary or with a main stem and subsidiary stems; involucral bracts 7–9 mm. 8(ii). capillaris var. agrestis

Section 1. Haplostephium (D. Don) Froel. Haplostephium D. Don; Crepis section Desiphylium Babc.; Geracium Rchb. Rhizomatous, usually hairy perennials. Stems stout, branched above. Lower leaves large, denticulate to shallowly lobed, petiolate, the upper becoming gradually smaller. Capitula 1–15, with many flowers. Ligules yellow. Receptacle glabrous or with a few hairs. 1. C. paludosa (L.) Moench Marsh Hawk’s-beard Hieracium paludosum L.; Hieracium paniculatum Gilib.; Aracium paludosum (L.) Monnier; Geracium paludosum (L.) Rchb.; Soyeria paludosa (L.) Godr.; C. caucasica C. A. Mey.; C. glabra Boiss.; C. ruprechtii Boiss.; C. rumicifolia Boiss.; Hieraciodes caucasicum (C. A. Mey.)

35. Crepis Kuntze; Hieraciodes paludosum (L.) Kuntze; Hieraciodes glabrum (Boiss.) Kuntze; Hieraciodes ruprechtii (Boiss.) Kuntze; Hieraciodes rumicifolium (Boiss.) Kuntze; Aracium attenuatum Opiz ex Domin Perennial herb with a short, stout, woody, densely fibrous rhizome. Stem solitary, 25–100 cm, pale green, often reddish near the base, striate, glabrous, erect, branched above. Leaves dark green on upper surface, paler beneath, thin; basal few, soon disappearing, the lamina 8–28 × 3–5 cm, oblanceolate, acute at apex, denticulate to sinuate-dentate or shallowly lobed, attenuate at base to a winged petiole; lower cauline similar but sometimes sessile; middle cauline with lamina lanceolate to ovate, acute to acuminate at apex, entire to dentate, rounded-auriculate, sessile and amplexicaul at base; uppermost with lamina linear or bract-like; all glabrous. Capitula 1–15, 12–25 mm in diameter, in a corymbose cyme, campanulate; peduncles 5–100 mm, slender to rather stout, arcuate, glabrous to slightly tomentose, sometimes with a few glandular hairs. Involucral bracts in 2 rows, dark green, usually with more or less numerous, long and short, black or greenish glandular hairs, rarely glabrous; inner 8–12 × 1.5–2.0 mm, linear-lanceolate, narrowed to an acute apex, ciliate at apex, glabrous on inner face; outer up to one-third as long as the inner, lanceolate, acuminate at apex. Flowers all ligulate, 13–17 mm, the ligules yellow, with 5 lobes at apex. Styles greenish-black. Receptacle flat, pitted, glabrous, without scales. Achenes 4.0–4.5 mm, pale brownish-yellow, cylindrical and only slightly narrowed upwards, 10-ribbed, smooth; pappus 6–7 mm, of 1 row of stiff, brittle, pale brownish hairs. Flowers 7–9. Visited by bees and flies. 2n = 12. Very uniform in most of Europe, but becoming very variable in the Caucasus region. Often mistaken for a Hieracium species from which it can be distinguished by the arrangement of the involucral bracts and the cylindrical achenes. Native. Streamsides and wet woodland, meadows and fens up to 915 m in Scotland. Frequent in northern Great Britain, extending southwards to Glamorganshire and Worcestershire; throughout Ireland. It has declined at the southern edge of its range. Europe from Spain, central France, northern Italy and the north Balkan peninsula northwards to 70◦ 30 N in Norway; western Russia. A member of the European Boreo-temperate element. Section 2. Hieracioides Froel. Crepis section Mesomeris Babc. Rhizomatous, usually hairy perennials. Stems slender, branched from or above the middle. Leaves entire or denticulate, the lower petiolate. Capitula 1–6, with many flowers. Ligules yellow. Receptacle glabrous. 2. C. mollis (Jacq.) Asch. Northern Hawk’s-beard Hieracium molle Jacq.; Hieracium succisifolium All.; C. hieracioides Waldst. & Kit.; Hieracium croaticum Waldst. & Kit.; Hieracium sternbergii Hoppe ex Hornem.; C. succisifolia (All.) Tausch; C. croatica (Waldst. & Kit.) Froel.; Hieraciodes molle (Jacq.) Kuntze; C. velenovskyi

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Domin; C. mollis subsp. hieracioides (Waldst. & Kit.) Domin; C. mollis subsp. velenovskyi (Domin) Domin Perennial wintergreen herb with a premorse rhizome bearing fleshy fibres. Stem solitary, 30–75(–90) cm, pale green, erect, slender, terete, fistulose, glabrous or more or less hairy below, branched from or above the middle. Leaves yellowish-green on upper surface, paler beneath; basal few, the lamina 4–27 × 1.5–5.0 cm, elliptical to oblanceolate, obtuse or acute at apex, entire or denticulate, gradually or abruptly attenuate into a long or short, winged petiole; lower cauline similar or sessile; middle cauline few, remote, mostly with lamina lanceolate, acute at apex, sessile, amplexicaul; uppermost cauline reduced and bractlike; glabrous or with fine, yellow simple eglandular hairs. Capitula 1–6, 20–30 mm in diameter, in a terminal corymb, campanulate; peduncles 10–90 mm, slender, with few to numerous, fine, short or unequal glandular hairs and finely tomentose, thickened below the capitula. Involucral bracts in 2 rows, dark green, with more or less numerous, dark or yellow glandular hairs; inner 8–13 × 1.0–1.5 mm, linearlanceolate, acute or acuminate at apex, white-ciliate at apex, becoming carinate and spongy-thickened dorsally in fruit, glabrous on inner face; outer unequal, up to half as long as inner, linear-lanceolate, acuminate and white-ciliate at apex. Flowers all ligulate, up to 14 mm, the ligules yellow, with 5 short lobes at apex. Styles dark green. Receptacle flat, pitted, obscurely fimbriate, glabrous. Achenes 3.0–4.5 × 0.5–0.7 mm, reddish-brown; inner straight, gradually attenuate at both base and apex, about 20-ribbed, the ribs smooth and nearly equal but 2–4 wider than rest; outer curved; pappus 5–6 mm, of 2 rows of soft, white hairs, the outer shorter and finer and united at base. Flowers 6–8. Visited by bees. 2n = 12. Native. Herb-rich grassland and wood pasture on shallow, slightly flushed, base-rich soils, often on north-facing slopes, and by upland burns, which are typically associated with intrusive rocks. It usually occurs between 150 and 400 m, but descends to 90 m in Northumberland and reaches 670 m in Forfarshire. Very local, but plentiful in some localities in eastern Great Britain. It has disappeared from some places, but has been recently found in many new ones in Northumberland, Co. Durham and the Scottish borders. Central Europe from the Pyrenees, northern Italy and southern Russia northwards to Germany, Poland and central Russia. A member of the European Temperate element. Section 3. Crepis Berinia Brign.; Crepis section Berinia (Brign.) Babc.; Brachyderea Cass. Hairy biennials with a woody tap-root. Stems branched above. Leaves denticulate to pinnatifid, the lower petiolate. Capitula few to numerous, with many flowers. Ligules yellow. Receptacle with shortly ciliate pits. 3. C. biennis L. Rough Hawk’s-beard Hedynois biennis (L.) Huds.; C. muricata Gilib.; C. lodomeriensis Bess.; C. gmelinii Schult.; Barkhousia pinguis Rchb.; C. maritima Boucher; C. glandulosa

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Bastard ex M¨ossler, non Guss.; C. sabauda Balb. ex DC.; Hieracium bienne (L.) Karsch Biennial herb with a woody, elongated root, forming a rosette of leaves in the first year. Stem 20–120 cm, pale green, sometimes reddish below, erect, slender to very robust, striate to sulcate, more or less stiffly hairy, branched above. Leaves yellowish-green on upper surface, paler beneath, basal with lamina 5–25 × 1.5–7.5 cm, oblanceolate, acute at apex, denticulate, dentate, runcinate-pinnatifid or pinnatisect, the terminal lobe triangular-ovate, acute at apex and entire or denticulate, the lateral lobes lanceolate, acute at apex, with acute or mucronate and spreading or curved-down teeth, attenuate at base into a narrowly winged petiole; lower cauline similar to basal; middle and upper cauline with lamina lanceolate to linear, acute, acuminate or caudate-acuminate at apex, pinnatifid to entire; uppermost bract-like; all with numerous fine, yellow, stiff simple eglandular hairs on both surfaces. Capitula 30–45 mm in diameter, cylindrical becoming campanulate in fruit, arranged in compound corymbs or panicles with 1–6 per branch; peduncles 5–160 mm, strict, erect, becoming arcuate, striate or sulcate, more or less thickened near the capitula, canescenttomentulose or with yellow or black, stiff simple eglandular hairs. Involucral bracts in 2 rows; inner 10–13 × 1–2 mm, green with scarious margins, linear-lanceolate, acute and white-ciliate at apex, appressed-hairy on inner face; outer nearly equal, up to two-thirds as long as the inner, linearlanceolate, acute at apex, glabrous or tomentulose, becoming lax and scarious. Flowers all ligulate, 12–18 mm, the ligules slightly ragged, yellow, with 5 short lobes at apex. Styles yellow or rarely green. Receptacle flat, pitted, with a ring of short hairs round each pit, without scales. Achenes 4.0–7.5 × 0.6–1.0 mm, yellowish-brown, equally attenuate to both ends or more gradually attenuate upwards but not beaked, (10–)13–18(–20)-ribbed; pappus 5–7 mm, of 2–3 rows, of soft, white hairs. Flowers 6–9. Visited freely by insects, especially bees and hoverflies, but probably often apomictic. 2n = 40. This species is easily distinguished from C. vesicaria in its summer flowering period by its large, erect flower heads, but autumnal-flowering forms have much smaller heads and do not look greatly different from autumnal-flowering C. vesicaria, but can be distinguished by their achenes. Some of these small autumnal forms are also very similar to C. tectorum, when the number of ribs on the achenes has to be considered. Native. Rough grassy places, waysides and neglected arable land. Rather local in Great Britain and Ireland north to central Scotland; frequent in south-east and south-central England. Central Europe from central Spain, north Italy, central Balkan peninsula and central Russia northwards to Denmark, southern Scandinavia and Estonia. A member of the European Temperate element. Section 4. Intybellioides Froel. Intybellia Monnier, non Cass.; Crepis section Intybellia Babc.; Intybus Fr., non Zinn More or less hairy perennials with rhizomes. Leaves entire to repand-dentate, the margin narrowly revolute, petiolate.

Capitula few to many, with few to many flowers in a narrow, elongated cyme. Ligules yellow. Receptacle with glabrous pits. 4. C. praemorsa (L.) Walther Leafless Hawk’s-beard Hieracium praemorsum L.; Hieracium spicatum Gilib.; Geracium praemorsum (L.) Rchb. ex M¨ossler; Intybus praemorsus (L.) Fr.; Intybellia praemorsa (L.) Monnier; Hieracium lactucaceum Schrank ex DC.; C. fistula Fisch. ex Besser; C. racemosa Cariot & St-Lag.; C. racemiformis Cariot & St-Lag.; Hieraciodes praemorsum (L.) Kuntze Perennial herb with a slender, premorse, fibrous rhizome up to 3 cm. Stem 15–75 cm, erect, stout or slender, striate, glabrous or with few to dense, short, curly hairs. Leaves yellowish-green on upper surface, paler beneath, all basal except for small bracts at the branching angles of the inflorescence, or rarely 1 cauline leaf near the middle; lamina 5–20 × 1.5–5.5 cm, mostly oblanceolate, sometimes oblong, elliptical or lanceolate, obtuse to acute and sometimes apiculate at apex, entire or obscurely denticulate to repand-dentate and with a narrowly revolute margin, abruptly or gradually contracted into a short, winged petiole; all with few to dense, very short, curly, pale simple eglandular hairs or becoming glabrous. Capitula few to many, 15–20 mm in diameter, forming a raceme-like, simple or compound cyme; peduncles 10–25 mm, slender, whitish tomentose and/or with short, curly simple eglandular hairs. Involucral bracts in 2 rows; 4–12 × 0.7–1.0 mm, dark green, the inner paler, with scattered, pale, short simple eglandular hairs particularly on the margin and apex; inner oblong or oblong-lanceolate, acute at apex; outer up to half as long as inner, lanceolate or ovate, obtuse to acute at apex, glabrous on inner face. Flowers all ligulate, 11– 12 mm, the ligules yellow, with 5 short lobes at apex. Styles yellow. Receptacle flat, pitted, glabrous. Achenes 3–4 mm, pale brown, fusiform, about equally attenuate at both ends, about 20-ribbed, the ribs narrow and smooth; pappus 4.5– 5.0 mm, of soft, white hairs in 2 rows. Flowers 6. 2n = 8. Native. Discovered in 1988 in limestone grassland, only slightly grazed, in northern Westmorland. Most of Europe eastwards from south-east Norway and south-east France to central Siberia, but absent from the Mediterranean region. A member of the Eurosiberian Temperate element. Our plant is subsp. praemorsa, which occurs through much of the range of the species. Section 5. Mesophylion Babc. Hairy annuals with slender, tapering roots. Leaves denticulate to 2-pinnatisect, the lower petiolate. Capitula many, with few to many flowers. Ligules yellow. Receptacle with ciliate pits. 5. C. tectorum L. Narrow-leaved Hawk’s-beard C. murorum S. G. Gmel.; C. dioscoridis Pollich; C. varia Moench; C. lachenalii Gochnat; C. integrifolia Vest; C. angustifolia d’Urv.; Hieracium tectorum (L.) Hornem.; Hieraciodes tectorum (L.) Kuntze; C. tectorum subsp. barckhausioides Rouy Annual herb sometimes overwintering, with a slender, tapering root. Stem 30–100 cm, pale green, erect, striate

35. Crepis or sulcate, more or less tomentulose, sometimes with short simple eglandular hairs. Leaves medium yellowish-green on upper surface, paler beneath, glabrous; basal in a rosette, few to numerous, the lamina 10–15 × 1.0–4.0 cm, lanceolate or oblanceolate, acute at apex, denticulate, dentate or runcinate-pinnatifid or lyrate-pinnatifid, or bipinnately parted with remote, unequal, lanceolate or linear, acute lobes, tapered into a winged petiole; lower cauline similar; middle cauline regularly decreasing in size, the lamina lanceolate or linear, acute at apex, sessile, usually acutely auriculate; uppermost with lamina linear with revolute margins or bract-like; all more or less tomentulose or with few to numerous, short, fine glandular or simple eglandular hairs on both sides. Capitula 15–20 mm in diameter, in corymbs or panicles, sometimes with branches from the base, but usually from above the middle; peduncles 3–70 mm, slender or very slender, slightly thickened beneath the capitula, more or less tomentulose and sometimes with rigid simple eglandular hairs. Involucral bracts in 2 rows, 7–10 × 0.8–1.2 mm, yellowish-green; inner with broad membranous margins, lanceolate, acuminate and white-ciliate at apex, appressedhairy on inner face, becoming dorsally keeled, more or less shortly hairy; outer unequal, up to one-third as long as inner, linear-lanceolate, acute at apex, tomentulous and with simple eglandular hairs. Flowers all ligulate, 12–15 mm, the ligules yellow, with 5 short lobes at apex. Styles dark green with yellow on inner face. Receptacle flat, pitted, ciliate round the pits. Achenes (2.5–)3–4(4.5) mm, dark purplishbrown, fusiform, strongly attenuate below, scarcely beaked, 10-ribbed, finely spiculate especially near the apex; pappus 4–5 mm, of 1 row of soft, white hairs. Flowers 5–8. Visited by flies and bees. 2n = 8. Introduced. Casual grain and grass-seed alien on roadsides, disturbed soil and reseeded verges and amenity areas. Scattered records in England, Wales and Scotland. Native in much of Europe except the Mediterranean region; Siberia and Caucasus and eastwards to Manchuria and Kamchatka; introduced in North America and Australia. Our plant is subsp. tectorum, which occurs throughout the range of the species. Section 6. Barkhousia (Moench) Gaudin Barkhousia Moench; Wibelia P. Gaertn., B. Mey. & Scherb.; Hostia Moench nom. illegit.; Crepis section Hostia Babc.; Anisoderis Cass. nom. illegit.; Rodigia Spreng.; Phalacroderis DC. Mainly annual to perennial herbs with slender roots. Leaves denticulate to 2-pinnatifid, lower usually petiolate. Capitula solitary or in a corymb, with few to many flowers. Ligules yellow. Receptacle with ciliate pits and sometimes with linear scales. 6. C. foetida L. Stinking Hawk’s-beard Annual, rarely biennial or short-lived perennial herb, foetid when fresh, with slender roots. Stems 10–50 cm, pale green, erect, more or less stiffly hairy, branched above or from near the base. Leaves 2–16 × 0.6–4.0 cm, yellowishgreen; basal with lamina oblanceolate, obtuse at apex, denticulate to bipinnate, petiolate; cauline with lamina

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elliptical, ovate, lanceolate or linear, obtuse at apex, runcinate to deeply pinnatifid with linear lobes, more or less laciniate near the sessile, auriculate base; all more or less stiffly hairy. Capitula few to many, 15–20 mm in diameter, drooping or erect in bud, terminal on long peduncles; peduncles somewhat thickened or inflated towards the apex, with glandular and simple eglandular hairs. Involucral bracts in 2 rows, greyish-green, hairy on inner face, with grey, curled, short hairs, usually some glandular hairs and some simple eglandular hairs; inner 10–13 × 0.7–1.5 mm, linearlanceolate, more or less acute at apex, later hardening and enclosing the outer achenes, finally spreading starwise, outer up to two-thirds as long as inner, linear to lanceolate, becoming lax. Flowers all ligulate, 9–19 mm, the ligules yellow and reddish-purple on outer face of marginal flowers, with 5 lobes at apex. Styles yellow. Receptacle flat, with densely ciliate pits or with linear scales. Achenes of 2 kinds, the outer stout, shortly and coarsely beaked or beakless, the inner longer, slender and finely beaked; pappus 3–7 mm, of 2 rows of dirty white hairs. Flowers 6–8. In their native habitats the three subspecies hybridise and intergrade and many local variants occur. The species includes plants with and without receptacular scales, a character which usually separates genera in the Asteraceae. For a discussion of their evolution see Babcock (1947). (a) Subsp. foetida C. barbata Mill.; Picris foetida (L.) Lam.; Barkhousia foetida (L.) F. W. Schmidt; Wibelia graveolens P. Gaertn. B. Mey. & Scherb. nom. illegit.; Hostia foetida (L.) Moench; C. interrupta Sibth.; Barkhousia graveolens (P. Gaertn. B. Mey. & Scherb.) Link; C. foetens Link ex Buchanon; C. glandulosa Guss.; Barkhousia glandulosa (Guss.) C. Presl; Barkhousia candollei Spreng.; Barkhousia prostrata Dumort.; Barkhousia zacynthia Margot & Reut. ex DC.; C. insularis Moris & De Not.; C. graveolens Schrad. ex Steud.; Anisoderis foetida (L.) Fisch. & C. A. Mey.; Wibelia foetida (L.) Sch. Bip.; Barkhousia triangularis K. Koch; C. fallax Boiss.; Hieraciodes foetidum (L.) Kuntze; Barkhousia supina Rouy; Barkhousia gracilis Lej. ex Rouy; C. foetida subsp. glandulosa (Guss.) Hayek; C. foetida subsp. maritima (Boiss.) Hayek; C. foetida subsp. zacintha (Margot & Reut. ex DC.) Hayek Outer involucral bracts up to 0.7 mm wide, linearlanceolate, about half as long as the inner, sometimes with predominantly glandular hairs. Receptacle without scales. Marginal achenes 7–9 mm, the inner 12–17 mm. 2n = 10. ˇ (b) Subsp. rhoeadifolia (M. Bieb.) Celak. C. rhoeadifolia M. Bieb.; Barkhousia rhoeadifolia (M. Bieb.) Fisch. & C. A. Mey.; Barkhousia byzantina DC.; C. echioides Ledeb.; Barkhousia rodigioides Sch. Bip. ex W. D. J. Koch; C. foetida var. hispida Bisch.; C. strybrnyi Velen.; Barkhousia foetida var. subdivisa Schur; C. nestmeiri F. Herm. & Degen Outer involucral bracts 1.0–1.5 mm wide, lanceolate, about two-thirds as long as inner with mostly or entirely simple eglandular hairs. Receptacle without scales. Marginal achenes 5–7 mm, the inner 12–16 mm. 2n = 10.

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(c) Subsp. commutata (Spreng.) Babc. Rodigia commutata Spreng.; Seriola commutata (Spreng.) Less.; Phalacroderis coa DC.; Barkhousia hirta W. D. J. Koch; Rodigia bulgarica Velen.; Barkhousia gracilis Freyn & Sint.; C. brachypappa Bornm. Outer involucral bracts 0.5–0.7 mm wide, linearlanceolate, about half as long as inner, with glandular and simple eglandular hairs. Receptacle with 2 scales subtending each achene. Marginal achenes 5.0–9.5 mm, the inner 10–14 mm. 2n = 10. The species is a member of the Eurosiberian Southerntemperate element. Subsp. foetida is native and formerly occurred on waysides and rough ground in scattered localities in south-east England and was introduced north-west to Worcestershire, but is now only on shingle at Dungeness in Kent. It also occurs in west and southern Europe from Spain and Portugal to Belgium, Germany and Yugoslavia. Subsp. rhoeadifolia has been recorded as a casual at Bristol and is native of south-east Europe and south-west Asia. Subsp. commutata has been recorded as a casual previous to 1930 and is native of south-eastern Europe. The total variation of this species has been included to facilitate better understanding of our plants. Section 7. Alethocrepis Bisch. Crepis section Phytodesia Babc. Hairy annual or biennial herbs with slender roots. Leaves denticulate to pinnate, the lower petiolate. Capitula many, with few to many flowers. Ligules yellow, usually reddish on the outer face of the outer flowers. Receptacle without scales, but with ciliate pits. 7. C. nicaeensis Balb. French Hawk’s-beard C. scabra DC., non Willd.; Barkhausia nicaeensis (Balb.) Link; C. adenantha Vis.; C. agrestis Fr. ex Bisch.; Brachyderea nicaeensis (Balb.) Sch. Bip.; Berinia nicaeensis (Balb.) Sch. Bip. Annual or biennial herb. Stem 25–110 cm, pale green and often reddish below, erect, terete, sulcate, with dense, rigid simple eglandular hairs below, branched above. Leaves medium yellowish-green on upper surface, paler beneath; basal few to many, the lamina up to 19 × 4 cm, oblanceolate, obtuse at apex, runcinate-pinnatifid, dentate or finely and remotely denticulate, attenuate below into a short, winged petiole with an amplexicaul base; lowest cauline similar, but with longer petioles; other cauline mostly with lamina lanceolate, sessile, broad or narrow at base and usually with short or long, acuminate, pendulous auricles; all with more or less numerous, short, yellow simple eglandular hairs. Capitula 10–20 mm in diameter, campanulate, arranged in a cymose corymb; peduncles rather slender, slightly thickened just below the capitulum, more or less white-tomentose and with long and short glandular or simple eglandular hairs. Involucral bracts in 2 rows, 8–10 × 1.0–1.5 mm, green; inner linear-lanceolate, attenuate to the obtuse, ciliate apex, with short arachnoid-hair and sometimes also with long, rather stiff simple eglandular hairs and shorter glandular hairs, glabrous on inner face, dorsally strongly carinateindurate in fruit, partly enclosing marginal achenes; outer

up to half (to two-thirds) as long as inner, more or less linear, acute at apex, becoming lax. Flowers about 11 mm, all ligulate, the ligules yellow, often red-tipped, with 5 short lobes at apex. Styles dark green. Receptacle convex, with ciliate achene pits. Achenes 2.5–3.8 × about 0.6 mm, golden brown, fusiform, shortly attenuate to the narrow apex, the marginal curved, narrowed at base, 10-ribbed, the ribs broad and rounded, faintly rugulose below and finely spiculate near the apex; pappus 4–5 mm, of 3 rows of soft white hairs united at the base. Flowers 5–8. 2n = 8. Introduced. Occasional casual of grain and grass-seed on roadsides, disturbed soil and reseeded verges, thought to be becoming rare but may be missed as C. capillaris var. agrestis. Scattered records in England and Scotland. Native of south Europe from southern France to the north Balkan peninsula; Caucasus. Introduced in central Europe and North America. 8. C. capillaris (L.) Wallr. Smooth Hawk’s-beard Lapsana capillaris L.; Crepis virens L.; C. umbellata auct.; C. diffusa DC.; C. polymorpha Willd.; C. humilis Gaudin; C. gaditana Boiss.; Malacothrix crepoides A. Gray ex J. G. Cooper; C. cooperi A. Gray Annual or biennial herb with a tap-root. Stems 1–several, 20–90 cm, pale green, sometimes tinted reddish-purple, solitary, erect and branched above, or profusely branched from the base upwards, or central axis short with numerous, suberect, nearly equal stems, or stems diffuse, ascending or decumbent, striate, shortly and stiffly hairy throughout or at the base only. Leaves yellowish-green on upper surface, paler beneath, occasionally tinted reddish-purple; basal in a rosette, numerous, with lamina 5–30 × up to 4.5 cm, lanceolate or oblanceolate, obtuse to acute and sometimes mucronate at apex, denticulate, dentate, runcinate-pinnatifid or lyrately, pinnately or bipinnately parted, with remote, unequal, oblanceolate or linear lobes, attenuate into a long or short, winged petiole with a clasping base; lower cauline similar to basal; middle and upper cauline with lamina lanceolate, acute at apex, sessile, amplexicaul, acutely auriculate; uppermost with lamina linear or bract-like; all glabrous or with short, stiff, yellow simple eglandular hairs on both sides. Capitula 10–13(–25) mm in diameter, erect in bud, in lax terminal corymbs; peduncles 5–65 mm, slender or very slender, not much enlarged below the capitulum, tomentose, or glabrous or sometimes with a few black glandular hairs. Involucral bracts in 2 rows, green or pale and scarious, tomentose, glabrous or sparsely glandular-hairy, glabrous on inner face; inner 4.5– 9.0 × 0.5–1.0 mm, linear-lanceolate, more or less acute at apex, becoming dorsally thickened in fruit; outer up to half as long as inner, linear, acute at apex, appressed to inner. Flowers 7.5–12.0 mm all ligulate, the ligules deep yellow, reddish on the outer face of the outer, shortly 5lobed at apex. Styles yellow with greenish barbs or greenish. Receptacle flat, pitted, without scales. Achenes 1.5–2.5 mm, brownish-yellow, pale brown or dark brown, fusiform or oblong, curved or straight, abruptly narrowed at both ends, 10-ribbed, the ribs narrow, rounded, smooth or finely spiculate at apex; pappus 3–4 mm, of soft, white hairs in 2 rows. Flowers 5–9. Freely visited by flies and bees. 2n = 6.

35. Crepis A very variable species which can be divided into two main varieties as below. Some plants are difficult to place in either variety. (i) Var. capillaris C. uniflora Thuill.; C. polymorpha var. diffusa Wallr. Stems few to numerous, ascending, diffuse or decumbent, but without an apparent main stem. Involucral bracts 4.5–7.0 mm. (ii) Var. agrestis (Waldst. & Kit.) Dalla Torre & Sarnth. C. agrestis Waldst. & Kit.; C. virens var. agrestis (Waldst. & Kit.) Bisch.; C. capillaris var. glandulosa Druce; C. virens var. elatior Cariot & St Lag.; C. capillaris var. anglica Druce & Thell.; C. druceana Murr Stems solitary or with a main stem and some subsidiary stems, erect, more or less branched. Involucral bracts 7–9 mm. Native. Grassland, heaths, walls, waste and rough places. Common throughout the British Isles. Most of Europe northwards to Denmark and southern Sweden; Canary Islands; introduced in North America. A member of the European Temperate element. Var. agrestis is the commonest plant in the north, but both varieties are general in distribution. Section 8. Lepidoseris (Rchb.) Benth. Lepidoseris Rchb. Hairy annual to perennial herbs, usually with a long, woody root. Basal leaves denticulate to pinnatifid, petiolate. Capitula numerous, with numerous flowers. Ligules yellow, usually reddish or purplish on the outer face of the outer flowers. Receptacle with ciliate pits. 9. C. vesicaria L. Beaked Hawk’s-beard Hieraciodes vesicarium (L.) Kuntze Annual or biennial herb with a fairly stout tap-root. Stems 1–several, 30–80 cm, pale yellowish-green, often deeply reddish-purple towards the base, fairly robust, erect, markedly striate, with arachnoid and short, rigid simple eglandular hairs, branched in upper half and sometimes from below. Leaves 4–30 × 1–6 cm, medium yellowishgreen, often reddish-purple tinted; basal in a neat rosette, basal and lowest cauline with lamina oblanceolate to spathulate, obtuse at apex, sinuately or retrorsely denticulate, dentate or runcinate-pinnatifid, or pinnately or bipinnately cleft or parted, the terminal segment often large, ovate and hastate or truncate, the lateral segments few or many, remote or close, the interlobe area winged or not, with a winged usually reddish-purple petiole; upper cauline similar, but sometimes with lamina oblong, sessile and auriculateamplexicaul; uppermost bract-like; all glabrous or with few to numerous short, pale simple eglandular hairs on both sides, but sometimes restricted to the veins. Capitula numerous, 15–40 mm in diameter, erect in bud, cylindrical to narrowly campanulate, in corymbs terminating the main stem and branches; peduncles with some arachnoid hairs and few to numerous short simple eglandular and sometimes glandular hairs. Involucral bracts in 2 rows, 8–12 × 1.5–2.0 mm, dark green with paler margins, with short and sometimes

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some longer, stiff simple eglandular hairs and sometimes numerous short to long glandular hairs, appressed-hairy on inner face; inner linear-lanceolate, obtuse and whiteciliate at apex, thickened in fruit; outer up to half as long as inner, lanceolate or ovate-lanceolate, subacute at apex, not imbricate, spreading. Flowers (9–)11–12 mm, all ligulate, the ligules yellow, the outer with reddish-purple on the outer face, shallowly 5-lobed at apex. Styles greenish or rarely yellow. Receptacle flat, the pits densely white-ciliate. Achenes (5–)6–8(–9) mm, pale brown, subterete, constricted at the narrow base, all gradually attenuate into a fine beak equal to or slightly longer than the body, 10-ribbed, the ribs narrowly, spiculate; pappus 4–6 mm, of 1–2 rows of soft white hairs, or the outer achenes narrowed at apex but not beaked, or only very shortly beaked, and half enclosed in the inner involucral bracts. Flowers 5–9. 2n = 8, 16. (a) Subsp. taraxacifolia (Thuill.) Thell. C. taraxacifolia Thuill.; C. polymorpha Pourr.; C. umbellata Thuill.; C. ruderalis Bouch.; C. praecox Balb.; C. taurinensis Willd.; C. scabra Willd.; C. cinerea Desf.; Barkhousia taraxacifolia (Thuill.) DC.; C. rubicaulis Pers.; C. intybacea Brot.; C. recognita Hall. fil.; Barkhousia laciniata Lowe; Barkhousia haenseleri Boiss.; Barkhousia heterocarpa Boiss.; C. haenseleri (Boiss.) F. W. Schultz; C. laciniata (Lowe) F. W. Schultz; C. numidica Pomel; C. hackelii Lange; Hieraciodes taraxacifolium (Thuill.) Kuntze; C. vesicaria subsp. haenseleri (Boiss.) P. D. Sell Achenes more or less uniform, all with long beaks up to at least half as long as body. (i) Var. taraxacifolia (Thuill.) B. Boivin Leaves sparingly short hairy. Involucral bracts with dense, very short, whitish hairs and some very short, dark glandular hairs. (ii) Var. rutilans (Lacaita) P. D. Sell C. rutilans Lacaita Leaves densely hairy on both surfaces. Involucral bracts with dense, very short, whitish hairs and short, purple, red and dark glandular hairs. (b) Subsp. stellata (Ball) Babc. C. taraxacifolia subsp. stellata Ball Involucral bracts with dense, long and short, greenish or dark glandular hairs. Achenes biform, the outer attenuate at apex and obscurely ribbed, the inner with a long beak. Introduced. Grassy places, waysides, walls and rough ground. Common, particularly on disturbed roadsides, in south and central-east England, with scattered records north to northern England and in Ireland and the Channel Islands. The species, which is part of a large complex of infraspecific taxa, occurs throughout Europe and North Africa and is introduced in North America, New Zealand and Australia. Our common plant is subsp. taraxacifolia var. taraxacifolia which also occurs in western Europe. Var. rutilans seems to occur in a number of coastal localities. It was originally described from Corfu in Greece, and no original material has been seen. Our plants, however, seem to exactly fit the original description. We have used the name subsp. stellata

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for plants found in Cambridgeshire, again without confirming its identification. It is native of Morocco and will run down in most floras to C. foetida. Section 9. Nemauchenes (Cass.) Benth. Nemauchenes Cass. Hairy annual herbs with slender roots. Basal leaves denticulate to pinnatifid, petiolate. Capitula with many flowers. Ligules yellow, usually reddish or purplish on the outer face of the outer flowers. Receptacle with ciliate pits. 10. C. setosa Haller fil. Bristly Hawk’s-beard C. hispida Waldst. & Kit.; Barkhausia setosa (Haller fil.) DC.; C. banatica Willd.; Wibelia setosa (Haller fil.) R¨ohl.; Hieraciodes setosum (Haller fil.) Kuntze; C. glabrata Porcius; Barkhousia hispida (Waldst. & Kit.) Link Annual herb with a slender, tapering root. Stem 8–80 cm, pale green, erect, terete, striate or sulcate, with more or less numerous, stiff simple eglandular hairs, with long, spreading or strict branches. Leaves yellowish-green on upper surface, paler beneath; basal few to many, the lamina 6–30 × 2–8 cm, oblanceolate, obtuse to acute at apex, denticulate, dentate, runcinate-pinnatifid or pinnately parted, the terminal lobe usually large, the lateral lobes triangular to lanceolate, acute and more or less dentate, attenuate at base into a narrowly or broadly winged petiole with a broader base; cauline with lamina mostly lanceolate, acuminate at apex, dentate to deeply laciniate near the base, amplexicaul-auriculate, the auricles acuminate; all with pale simple eglandular hairs. Capitula 10–20 mm in diameter, in cymose panicles; peduncles slender, erect, with rigid, pale simple eglandular hairs. Involucral bracts in 2 rows, 8–10 × 1.0–1.5 mm, pale green, with yellow, rigid simple eglandular hairs, appressed-hairy on inner face; inner linear-lanceolate, acuminate at apex, becoming strongly carinate, membranous at margin; outer up to half as long as inner, linear, acuminate at apex, carinate, lax. Flowers 10–13 mm, all ligulate, the ligules yellow, with or without red on the outer face, shortly 5-lobed at apex. Styles dark green. Receptacle flat, the pits shortly ciliate. Achenes 3.2–5.0 × 0.3–0.6 mm, pale brown, fusiform, abruptly or gradually attenuate to a fine beak 1.0–2.5 mm, 10-ribbed, the ribs more or less prominent, strongly spiculate near the base of the beak; pappus 2.5–5.0 mm, of 1 row of soft white hairs. Flowers 7–9. 2n = 8. Introduced. Occasional casual with crops, grass seed or wild flower seed or on waste ground. Scattered records in Great Britain and northern Ireland. Native of south and south-east Europe from southern France to the Crimea; west Asia; introduced further north in Europe, North and South America and New Zealand. Our plant is subsp. setosa. 36. Tolpis Adans. Annual herbs with latex, without rhizomes or stolons. Stems usually solitary, simple or branched, leafy. Leaves alternate, dentate to slightly pinnately divided. Capitula 1– several. Involucral bracts in 2–3 rows, the outer curved and not appressed, the inner erect. Flowers all bisexual. Corolla ligulate, lemon yellow, or the inner purplish, turning green when dry, with 5 lobes at apex. Anthers tailed,

with elongate apical appendage; filaments smooth. Style with short branches, with both shaft and branches with short hairs. Receptacle flat, without scales. Achenes neither narrowed or beaked above, of 2 kinds, the outer oblong with a pappus about 0.3 mm, of rigid hairs much shorter than the achene, the inner narrowly oblong, with a pappus of short bristles mixed with several longer bristles, much longer than the achene which are expanded at base. About 20 species in the Mediterranean region and Atlantic islands extending south to tropical Africa. 1. T. barbata (L.) Gaertn. Tolpis Crepis barbata L.; T. umbellata Bertol. Annual herb. Stem 6–90 cm, erect, slender, sulcate, slightly arachnoid-hairy, usually branched. Leaves medium yellowish-green on upper surface, paler beneath; basal with lamina 2–12 × 1–4 cm, obovate, oblong-obovate or lanceolate, acute or obtuse at apex, entire to coarsely dentate or more or less pinnatifid, tapering at base to an ill-defined petiole; cauline leaves with lamina 2–4 × 0.2–0.6 cm, linear to lanceolate, entire to denticulate, sessile; slightly arachnoid-hairy. Capitula 15–35 mm in diameter, campanulate, in a lax, dichasial cyme; peduncles slender, slightly swollen near the apex and bearing numerous, filiform, subapical bracts 5–8 mm, equalling or exceeding the involucral bracts. Involucral bracts in 2–3 rows, 6–15 × 1.0– 1.5 mm, outer as long as inner, dark green, linear-lanceolate at apex, slightly arachnoid-hairy, the outer not appressed, the inner erect. Flowers 8–9 mm, all ligulate, the ligule lemon yellow, the central ones often purplish, usually turning greenish when dry, with 5 lobes at apex. Receptacle flat, pitted, without scales or hairs. Achenes of 2 kinds; outer about 1.6 × 0.6 mm, dark brown, oblong, truncate at apex, sometimes curved and shortly hairy, the pappus about 0.3 mm, of numerous, short, whitish bristles; inner about 1.5 × 0.4 mm, dark brown, narrowly oblong, longitudinally ribbed, minutely papillose-scabridulous, the pappus about 4 mm, of (0–)3–4(–5) scabridulous bristles expanded at base, with a row of very short bristles at their base. Flowers 4–7. 2n = 18. Introduced. A wool, bird-seed and grain casual. In a few scattered localities in Britain. Native of southern Europe and the Mediterranean region. 37. Andryala L. Annual to perennial herbs with latex, without stolons. Stems usually solitary, branched, leafy. Leaves alternate, entire to deeply pinnatisect. Capitula few to numerous. Involucral bracts in several rows. Flowers bisexual. Corolla ligulate, yellow, the outer sometimes with a reddish stripe on the outer face, 5-lobed at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long branches, with both shaft and branches with short hairs. Receptacle flat, pitted, the margin of the pits laciniatedentate with long hairs. Achenes oblong or obconical, truncate at apex, not beaked; pappus of greyish-white simple hairs, falling entire. About 20 species in the Mediterranean region, southern Europe, North Africa, Canary Islands and Middle East.

38. Pilosella 1. A. integrifolia L. Common Andryala Annual to perennial herb. Stems 12–80 cm, dull green, striate, with few to dense stellate and simple eglandular hairs and sometimes with glandular hairs above, with numerous leaves, branched from halfway up. Leaves yellowish-green on upper surface, scarcely paler beneath; lamina 2–8 × 0.3– 3.0 cm, linear, oblanceolate, lanceolate or ovate, obtuse to acute at apex, entire to deeply pinnatisect, the lower usually narrowed at base, the upper broader and sometimes amplexicaul, the basal usually few, the cauline numerous and sometimes aggregated towards the base; with dense stellate and short simple eglandular hairs. Capitula 14– 15 mm in diameter, in cymes with long lower branches; peduncles pale green, with dense stellate and short simple eglandular hairs. Involucral bracts in several rows, 7– 11 × 1.0–1.2 mm, linear-lanceolate, obtuse to acute at apex, with dense stellate, short simple eglandular and longer glandular hairs. Flowers 9–12 mm, all ligulate, the ligules lemon yellow, deeply 5-lobed at apex. Receptacle flat, pitted, the pits laciniate-dentate with long minutely denticulate hairs on the margin. Achenes 1.2–1.5 mm, pale brown, oblong or obconical, strongly ribbed; pappus 6–8 mm, greyish-white, of minutely denticulate hairs, falling entire. Flowers 8–10. 2n = 18. Very variable in all its parts and almost certainly an aggregate species. Introduced. A casual introduced with shoddy. Native of the Mediterranean region and south-west Europe. 38. Pilosella Hill Hieracium subgen. Pilosella (Hill) Gray Perennial herbs with latex, rhizomes and stolons. Stems 1–numerous, without or with a few, alternate leaves, sometimes branched. Leaves entire or slightly denticulate, gradually attenuate at base but never distinctly petiolate. Capitula 1–numerous. Involucral bracts in numerous rows. Flowers bisexual. Corolla ligulate, yellow, often with a red stripe on the outer face, or reddish, glabrous, with 5 lobes at apex. Anthers tailed, with an elongate apical appendage; filaments smooth. Style with long branches, with both shaft and branches with short hairs. Receptacle flat, pitted, without scales. Achenes up to 2.5 mm, each rib shortly projecting above to form a crenulate apex; pappus hairs in 1 row with a few shorter than the rest. This genus is easily distinguished from Hieracium by its stoloniferous habit, leaves without a distinct petiole, corolla often with a reddish stripe or sometimes reddish and the small achenes with projecting ribs at the apex and pappus hairs mainly in one row. This must be amongst the most difficult of all genera to classify, but in Great Britain and Ireland we only have three native species and the rest are introduced. At least 18 easily recognised species are known in the genus and a great many infraspecific taxa have been described. Hybrids are frequent, are often fertile and can spread vegetatively. Pilosella is found throughout Europe and in western Asia. It has been introduced in North America, Australia and New Zealand where it has sometimes become a troublesome weed. For South American plants that have

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been put in this genus see discussion under Hieracium (p. 219). In Great Britain and Ireland the Pilosella taxa have never been treated taxonomically as they have been in Continental Europe by H. Dahlstedt (1890), C. N¨ageli and A. Peter (1885), S. O. F. Omang (1936) and K. H. Zahn (1923) who have called all taxa species or subspecies as in Hieracium. They do not fall clearly into apomictic taxa as they do in Hieracium and this is because of their variable breeding mechanisms. Diploids (2n = 18) and tetraploids (2n = 36) are sexual and self-incompatible and the chromosomes pair regularly at meiosis. Hybrids between them are triploid (2n = 27) and sterile, but can spread by stolons. The higher polyploids are usually apomictic, but their pollen is fertile, and as they are usually sympatric over wide areas with the diploids and tetraploids, can hybridise with them. The offspring are apomictic and can spread by stolons, but are often closer morphologically to one or other of the parents. Skalinska found that in some capitula of P. aurantiaca both sexual and apomictic flowers occurred. It is not known how often this phenomenon occurs. When preparing the account of Pilosella (as Hieracium subgenus Pilosella) for Flora Europaea, Sell and West (1976) had available the 500 voucher specimens of G. & B. Turesson (1972), but could not identify taxa which tied up with chromosome number, nor could they define taxa as they could in Hieracium subgenus Hieracium. They came to the conclusion that they could recognise species in the same way as they did normal sexual species in other genera, and that these agreed with the main species of K. H. Zahn (1923). Hybrids seemed to exist between all these species in the way one normally recognised hybrids. As regards the recognition of infraspecific taxa, we felt that the broad groups recognised by K. H. Zahn (1923) had some useful meaning, both geographically and ecologically, and that they were worth calling subspecies. We felt that in Hieracium apomixis had reached a state of virtual completeness, while in Pilosella it was still in a state of flux. Bate-Smith, E. C., Sell, P. D. & West, C. (1968). Chemistry and taxonomy of Hieracium L. and Pilosella Hill. Phytochemistry 7: 1165–1169. Bishop, G. F. & Davy, A. J. (1994). Hieracium pilosella L. (Pilosella officinarum F. Schultz & Schultz-Bip.) in Biological flora of the British Isles. Jour. Ecol. 82: 195–210. Bowman, R. P. & Sell, P. D. (1993). Pilosella × floribunda (Wimmer & Grab.) Arvet-Touvet (Asteraceae) in the British Isles. Watsonia 19: 187–188. Br¨autigam, S. & E. (1996). Determination of the ploidy level in the Genus Hieracium subgenus Pilosella (Hill) S. F. Gray by flow cytometric DNA analysis. Folia Geobot. Phytotax. 31: 315–321. Chater, A. O. (1998). A hybrid Pilosella. B.S.B.I. News 78: 88. Dahlstedt, H. (1890). Bidrag till syd¨ostra Sveriges Hieracium-flora. I. Piloselloidea. Kungl. Svenska Vet.-Akad. Handl. 23(15): 1– 134. Gadella, T. W. J. (1972). Biosystematic studies in Hieracium pilosella L. and some related species of the Subgenus Pilosella. Bot. Not. 125: 361–369. Gadella, T. W. J. (1981). Het Schellings havikskruid (Hieracium peleterianum M´erat). Gorteria 10: 120–129.

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Gadella, T. W. J. (1984). Cytology and the mode of reproduction of some taxa of Hieracium subgenus Pilosella. Proc. Kon. Ned. Akad. Wetensch. Ser. C, 87: 387–399. Gadella, T. W. J. (1984). Voortplantingswijze, hybridisatie en cytologie van het weidehavikskruid Hieracium caespitosum Dumort. Gorteria 12: 108–109. Gadella, T. W. J. (1987). Sexual tetraploid and apomictic pentaploid populations of Hieracium pilosella (Compositae). Pl. Syst. Evol. 157: 219–245. Gadella, T. W. J. (1988). Some notes on the origin of polyploidy in Hieracium pilosella aggr. Acta Bot. Neerl. 37: 515–522. Gadella, T. W. J. (1991). Variation, hybridization and reproductive biology of Hieracium pilosella L. Proc. Kon. Ned. Akad. Wetensch. 94: 455–488. Gadella, T. W. J. (1991). Reproduction, variation and interspecific hybridisation in three species of Hieracium section Pilosellina (Compositae). Polish Bot. Studies 2: 85–103. Gadella, T. W. J. (1992). Notes on some triple and inter-sectional hybrids in Hieracium L. and Pilosella (Hill) S. F. Gray. Proc. Kon. Ned. Akad. Wetensch. Ser. C, 95: 51–63. Gadella, T. W. J. & Kleiphuis, E. (1968). Enige opmerkingen over Hieracium pilosella L. in Nederland. Gorteria 4(2): 17– 26. Gadella, T. W. J. & Kleiphuis, E. (1968). Een merkwaardige Verm van Hieracium pilosella L. gevonden bij Winterswijk. Gorteria 4(5): 57–60. N¨ageli, C. von & Peter, A. (1885). Die Hieracien Mittel-Europas. Vol. 1. Piloselloiden. M¨unchen. Omang, S. O. F. (1936). Die norwegisch Unterarten des Hieracium pilosella L. Nytt. Mag. Naturvidensk. 75: 33–127. Ostenfeld, C. H. (1906). Castration and hybridisation experiments with some species of Hieracia. Bot. Tidssk. 27: 225–248. Pugsley, H. W. (1921). On Hieracium aurantiacum L. Jour. Bot. (London) 59: 60–69. Pugsley, H. W. (1948). A prodromus of the British Hieracia. Jour. Linn. Soc. London (Bot.) 54: 312–319. Rotreklova, O., Krahulcov´a, A., Vankova, D., Peckert, T. & Mraz, P. (2002). Chromosome numbers and breeding systems in some species of Hieracium subgen. Pilosella in Central Europe. Preslia 74: 27–44. Scott, W. (1968). Pilosella flagellaris (Willd.) Sell & C. West subsp. bicapitata Sell & C. West. in Zetland. Proc. B.S.B.I. 7: 192–193. Sell, P. D. (1988). Pilosella Hill in Rich, T. C. G. & M. D. B. Plant Crib. London. Sell, P. D. & West, C. (1976). Hieracium subgenus Pilosella in Tutin, T. G. et al. (Edits.) Flora Europaea, Vol. 4, pp. 366–376. Cambridge. Skalinska, M. (1967). Cytological analysis of some Hieracium species, subgen. Pilosella from mountains of southern Poland. Acta Biol. Cracov. Ser. Bot. 10: 127–141. Skalinska, M. (1969). Apomixis in Hieracium aurantiacum L. Genet. Polon. 10: 91–93. Skalinska, M. (1970). Further cytological studies in natural populations of Hieracium aurantiacum L. Acta Biol. Cracov. Ser. Bot. 13: 111–117. Skalinska, M. (1971). Further studies in apomixis in Hieracium aurantiacum L. Genet. Polon. 12: 209–212. Skalinska, M. (1971). Experimental and embryological studies in Hieracium aurantiacum L. Acta Biol. Cracov. Ser. Bot. 14: 139– 152. Turesson, G. & B. (1960). Experimental studies in Hieracium pilosella L. I. Reproduction, chromosome number and distribution. Hereditas 46: 717–736. Turesson, G. & B. (1972). Experimental studies in Hieracium pilosella L. II. Taxonomy and differentiation. Bot. Not. 125: 223–240.

Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [P. flagellaris subsp. bicapitata, peleterana] Zahn, K. H. (1923). Hieracium L. subgenus Pilosella in A. Engler, Das Pflanzenreich 82(IV. 280): 1147–1705. 1. Ligules orange-brown or orange-red, often turning 2. purplish when dry 1. Ligules yellow, sometimes with a red stripe on the outer 5. face 2. Capitula 20–30 mm in diameter, often solitary on long 3. peduncles; involucral bracts 10–13 mm 2. Capitula 10–20 mm in diameter, peduncles usually short but sometimes with long lower branches; involucral 4. bracts 5–11 mm 3. Involucral bracts with simple eglandular as well as glandular hairs 2(a). × stoloniflora nothosubsp. stoloniflora 3. Involucral bracts with numerous unequal glandular hairs, without simple eglandular hairs 2(b). × stoloniflora nothosubsp. schuriana 4. Stolons short, plant not spreading easily; involucral 1(a). aurantiaca subsp. aurantiaca bracts 8–11 mm 4. Stolons long, plant spreading easily; involucral bracts 1(b). aurantiaca subsp. carpathicola 5–8 mm 5. At least some flowering stems with more than 6. 1 capitulum 12. 5. All flowering stems with only 1 capitulum 7. 6. Involucral bracts (8–)9–12 mm 8. 6. Involucral bracts 5–9 mm 7. Peduncles with simple eglandular hairs 2–3 mm; involucral bracts with simple eglandular hairs up to 7(a). flagellaris subsp. flagellaris 1.5 mm 7. Peduncles with simple eglandular hairs up to 7.5 mm; involucral bracts with simple eglandular hairs up to 7(b). flagellis subsp. bicapitata 2.5 mm 8. Flowering stems usually not more than 35 cm; capitula 9. 1–7 10. 8. Flowering stems up to 70 cm; capitula numerous 9. Flowering stems 10–35(–45) cm; basal leaves 5–18 × 0.6–2.0 cm; involucral bracts 6–9 × 1.0–1.5 mm 4. × floribunda 9. Flowering stems (6−)10–20(−45) mm; basal leaves 2–7(−9) × 0.3–1.5 cm; involucral bracts 6. lactucella subsp. lactucella 5–7 × 0.5–1.0 mm 10. Stem and leaves with numerous simple eglandular hairs; capitula 10–20 mm in diameter; involucral bracts 3. caespitosa subsp. colliniformis 1.0–2.0 mm wide 10. Stem and leaves glabrous or with few simple eglandular hairs, rarely with numerous hairs; capitula 7–10 mm in 11. diameter; involucral bracts 0.7–1.0 mm wide 11. Stolons very short or absent; involucral bracts and peduncles with more or less numerous glandular hairs, without or with few simple eglandular hairs 5(a). praealta subsp. praealta 11. Stolons long and slender; involucral bracts with numerous glandular hairs, without or with few to numerous simple eglandular hairs 5(b). praealta subsp. thaumasia 12. Involucral bracts with numerous glandular hairs, without or with very occasional simple eglandular hairs 13.

38. Pilosella 12. Involucral bracts with numerous to dense simple eglandular hairs, without or with few to numerous 14. glandular hairs 13. Glandular hairs on involucral bracts more or less equal in length, mostly not more than 0.5 mm 8(a). officinarum subsp. micradenia 13. Glandular hairs on involucral bracts markedly unequal, 8(b). officinarum subsp. euronota up to 1 mm 14. Involucral bracts with more or less numerous simple eglandular hairs and more or less numerous glandular 15. hairs 14. Involucral bracts with numerous to dense simple eglandular hairs, glandular hairs absent or few and not 16. obvious 15. Involucral bracts with pale, sometimes dark-based simple eglandular hairs 8(c). officinarum subsp. officinarum 15. Involucral bracts with dark, sometimes darker-based simple eglandular hairs 8(d). officinarum subsp. trichosoma 16. Involucral bracts with moderately dense, dark simple 17. eglandular hairs 16. Involucral bracts with dense, pale simple eglandular 18. hairs (sometimes with black bases) 17. Simple hairs of flowering stems up to 2 mm 8(f). officinarum subsp. melanops 17. Simple hairs of flowering stems up to 5 mm 8(g). officinarum subsp. trichoscapa 18. Stolons long and slender; involucral bracts 7–11 × 0.5–1.5 mm, with dense stellate hairs 8(e). officinarum subsp. tricholepia (P. officinarum × peleterana may key out here, but will tend to have thicker stolons and broader bracts, and to be found also where both parents occur) 18. Stolons short and thick; involucral bracts 1.3–3.0 mm 19. wide, usually with few stellate hairs 19. Rosette leaves 2.1–8.7 × 0.9–2.0 cm, mostly oblanceolate or elliptical, cuneate or shortly attenuate at base; flowering stems up to 12(–18) cm; involucre 11–15 × 12–30 mm, the bracts 1.5–3.0 mm wide 9(a). peleterana subsp. peleterana 19. Rosette leaves (2–)4–11(–15) × 0.4–1.2(–1.6) cm, lanceolate or oblanceolate, attenuate at base; flowering stems (6–)10–30(–35) cm; involucre 10–15 × 20. (9–)10–17 mm, the bracts 1.5–2.0 mm wide 20. Involucre 10–15 × 12–17 mm, the bracts 1.5–2.0 mm 9(b). peleterana subsp. subpeleterana wide, lanceolate 20. Involucre 10–12(–13) × (9–)10–12(–14) mm, the bracts about 1.5 mm wide, linear-lanceolate 9(c). peleterana subsp. tenuiscapa (P. officinarum × peleterana may key out here but it is a generally more slender plant)

1. P. aurantiaca (L.) F. W. Schultz & Sch. Bip. Fox-and-cubs Hieracium aurantiacum L. Perennial herb with fibrous roots and stolons above or below ground, when above ground with leaves 1–5 × 0.3–2.0 cm, elliptical or obovate, rounded at apex, entire or denticulate, narrowed at base to a short, winged petiole or sessile, with numerous, medium to long, pale, more or less stiff simple eglandular hairs. Leaves yellowish-green or

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slightly glaucous on upper surface, paler beneath; basal few to numerous, in a rosette, the lamina 6–20(–30) × 1.2– 6.0(–7.0) cm, usually lanceolate or oblanceolate, sometimes oblong or narrowly elliptical, usually rounded, sometimes subacute at apex, entire or remotely denticulate, long attenuate at base; cauline 1–4, like those of the rosette or linear, acute at apex and bract-like; all with numerous medium to long, pale, stiff simple eglandular hairs on both surfaces and the margin. Flowering stems 20–40(–65) cm, pale yellowish-green, erect or ascending, slender to robust, striate, with sparse stellate hairs, numerous, dark simple eglandular hairs 1–6 mm and few, shorter, dark glandular hairs. Inflorescence with 2–12(–25) capitula, cymose-corymbose, often with long lower branches; peduncles usually short, with dense stellate hairs, numerous, medium and long, dark simple eglandular hairs and numerous, short, dark glandular hairs. Capitula 10–20 mm in diameter, rounded at base. Involucral bracts in numerous rows, 5–11 × 1.5–3.0 mm, olive or blackish-green, with paler margins, often flushed reddish at apex, lanceolate or linear-lanceolate, obtuse to acute at apex, with more or less numerous stellate hairs, numerous, medium to long, dark simple eglandular hairs and fewer, short, dark glandular hairs. Ligules orangebrown or orange-red, turning purplish when dry, glabroustipped, with 5 lobes at apex. Styles discoloured. Receptacle pits with margins more or less dentate. Achenes 2.0–2.5 mm, purplish-black. Flowers 5–7(–9). Sexual and apomictic. 2n = 27, 36, 45, 54, 63. (a) Subsp. aurantiaca Stolons rather short and mostly more or less underground, the plant not spreading very easily. Rosette leaves 10–20(– 30) × 2.2–6.0(–7.0) cm, oblanceolate or oblong. Involucral bracts 8–11 mm. 2n = 36. (b) Subsp. carpathicola (N¨ageli & Peter) Soj´ak Hieracium aurantiacum subsp. carpathicola N¨ageli & Peter; Hieracium brunneocroceum Pugsley; P. aurantiaca subsp. bunneocrocea (Pugsley) P. D. Sell & C. West Stolons often long and leafy, usually above ground, the plant spreading in large mats. Rosette leaves 6–10(–16) × 1.2– 2.0(–3.0) cm, lanceolate or narrowly elliptical. Involucral bracts 5–8 mm. Introduced. Naturalised garden escape in rough ground, on walls, waysides and railway banks, in grassy places and cemetaries. Frequent throughout Great Britain and Ireland. Native of northern and central Europe, mainly in the mountains, extending locally southwards to south-central France and Bulgaria; widely cultivated and naturalised elsewhere in Europe. Subsp. aurantiaca is mainly in the Alps and west Carpathians. Subsp. carpathicola occurs throughout the range of the species. Subsp. aurantiaca seems to have been the first subspecies introduced into British gardens, is the most handsome of the two subspecies and does not spread rapidly. Subsp. carpathicola seems to be a later introduction which spreads rapidly over open ground. 2. P. × stoloniflora (Waldst. & Kit.) F. W. Schultz & Sch. Bip. Hybrid Fox-and-cubs P. aurantiaca × officinarum Hieracium stoloniflorum Waldst. & Kit.

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Perennial herb with fibrous roots and long stolons which bear leaves 2–7 × 0.7–1.5 cm, oblanceolate or narrowly elliptical, rounded-obtuse at apex, entire and long attenuate at base, with long, pale stiff simple eglandular hairs. Leaves yellowish-green on upper surface, paler beneath; basal in a rosette, the lamina 5–14 × 1–3 cm, oblong, elliptical, oblanceolate-oblong or oblanceolate, obtuse to subacute at apex, entire, long-attenuate at base; cauline 0–3, lower sometimes similar to basal, but usually linear or bractlike; with medium to very long, pale, stiff simple eglandular hairs on both surfaces and the margins. Flowering stems 13–30 cm, pale yellowish-green, striate, ascending or erect, slender, with few stellate hairs below but more or less dense in the upper part, few to numerous, dark or darkbased, medium to long simple eglandular hairs particularly above and few to numerous, short, dark glandular hairs, or glandular hairs absent. Inflorescence very variable, sometimes a solitary capitulum at the end of a stem or stolon, sometimes with a few capitula grouped like P. aurantiaca but the capitula larger, but most often a number of large heads on very long peduncles; peduncles with dense stellate hairs, few to numerous, medium to very long, dark or dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs, or glandular hairs absent, or eglandular hairs absent. Capitula 20–30 mm in diameter, broadly rounded at base. Involucral bracts in numerous rows, 10–13 × 1–2 mm, blackish-green with pale margins, often reddish at the apex, linear-lanceolate, obtuse to acute at apex, with more or less numerous stellate hairs, numerous to dense, medium to long, dark or dark-based simple eglandular hairs and few to numerous, short to medium, dark glandular hairs. Ligules brownish-orange, orange-red, yellowish-orange or yellow with various amounts of red, glabrous, with 5 lobes at apex. Styles discoloured. Receptacle pits with the margins more or less dentate. Achenes 2.0–2.5 mm, purplish-black, apparently a high percentage fertile. Flowers 5–7. (a) Nothosubsp. stoloniflora Involucral bracts with simple eglandular as well as glandular hairs. (b) Nothosubsp. schuriana (N¨ageli & Peter) P. D. Sell Hieracium × stoloniflorum subsp. schuriana N¨ageli & Peter Involucral bracts with numerous, unequal glandular hairs, without simple eglandular hairs. Introduced. Roadsides, railway banks, walls and waste places. Nothosubsp. stoloniflora occurs in scattered localities from Guernsey in the Channel Islands to northern Scotland. It is sometimes grown in gardens and may escape, or can form hybrids where the two parents grow together. At some sites such as a railway bank at Cromdale Station in Morayshire and at Northwich in Cheshire, almost every possible intermediate in morphology has occurred. It is widespread in central Europe and north-west Russia where it shows the same sort of variation as at Cromdale and Northwich. Nothosubsp. stoloniflora is most likely to be P. aurantiaca subsp. carpathicola × one of the subspecies of P. officinarum with simple eglandular hairs on its involucral bracts. Nothosubsp. schuriana is known only in the chapel

graveyard at Tan-y-groes, Cardiganshire, where it has probably arisen from its supposed parents, P. aurantiaca subsp. carpathicola and P. officinarum subsp. euronota which grow with it. It was originally described from Transsilvania. 3. P. caespitosa (Dumort.) P. D. Sell & C. West Yellow Fox-and-cubs Hieracium caespitosum Dumort.; Hieracium pratense Tausch Perennial herb with an oblique rootstock, fibrous roots and long stolons which bear leaves 2–6 × 0.5–2.0 cm, narrowly elliptical to oblanceolate, rounded-obtuse at apex and gradually narrowed at base. Leaves yellowish-green on upper surface, paler beneath; basal (3.5–)5–20 × 0.7– 2.5 cm, oblong, linear-oblong, oblanceolate or oblongspathulate, rounded-obtuse to acute at apex, entire or rarely minutely denticulate, long-attenuate at base; cauline 1–3, like those of the basal, linear-oblanceolate or bract-like; with few to numerous, medium to long, pale, rather stiff simple eglandular hairs on the surfaces and the margins and especially the midrib beneath. Flowering stems (20–)30– 50(–80) cm, erect, robust, striate, with sparse stellate hairs, numerous to dense, short to long, pale, spreading simple eglandular hairs and a few, short, dark glandular hairs in the upper part. Inflorescence with 3–50 capitula, cymosecorymbose, often compact and crowded; peduncles with more or less dense stellate hairs, numerous, short, dark glandular hairs and a few, medium to long, dark-based simple eglandular hairs. Capitula 10–20 mm in diameter, rounded at base. Involucral bracts in numerous rows, (5–)7–9 × 1.0– 1.2 mm, olive or blackish-green, the inner with paler margins, linear-lanceolate, usually obtuse at apex, with stellate hairs mainly only at the base, numerous medium to long, pale or dusky, dark-based simple eglandular hairs and numerous, short, dark glandular hairs. Ligules pale yellow, glabrous, with 5 lobes at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 1.7–1.8 mm, purplish-black. Flowers 5–7. Sexual or apomictic. 2n = 18, 27, 36, 45. Introduced. Naturalised garden escape on rough ground, walls, railway banks, river banks, fixed dunes and waysides. In scattered localities in Great Britain and southern Ireland. Native of north, central and eastern Europe and south-west Asia. Our plant is subsp. colliniformis (Peter) P. D. Sell & C. West (Hieracium collinum subsp. colliniforme Peter; Hieracium caespitosum subsp. colliniforme (Peter) P. D. Sell; Hieracium colliniforme (Peter) Roffey). It is native mainly in northern and eastern Europe. 4. P. × floribunda (Wimm. & Grab.) Arv.-Touv. Dense-flowered Fox-and-cubs P. caespitosum × lactucella Hieracium floribundum Wimm. & Grab.; Hieracium floribundum subsp. helveolum Dahlst.; Hieracium helveolum (Dahlst.) Pugsley; Pilosella lactucella subsp. helveola (Dahlst.) P. D. Sell & C. West Perennial herb with a rather thick rootstock, sometimes producing stolons which have leaves 2–8 × 0.5–1.4 cm, linear-oblanceolate, obtuse to acute at apex, entire and gradually attenuate at base, with long, pale simple eglandular hairs on the midrib beneath and sometimes on the

38. Pilosella margin. Leaves glaucous-green with a pale midrib on the upper surface, paler beneath, basal numerous, in a rosette with lamina 5–18 × 0.6–2.0 cm, narrowly elliptical or narrowly oblanceolate, the outermost rounded at apex, the inner obtuse or subacute and mucronulate at apex, long attenuate at base; cauline 0–3, similar to basal but smaller; all with few to numerous, long (–4 mm), pale simple eglandular hairs on or near the margin, and similar ones with reddish bases along the midrib, sometimes occasional hairs are found on the surfaces and sometimes small stellate hairs are present on the lower surface. Flowering stems 10–35(–45) mm, erect, rather slender, green below, darker above, with numerous, long (–7 mm), pale, dark-based simple eglandular hairs throughout, numerous to dense, small stellate hairs and numerous, short, dark glandular hairs in the upper part. Inflorescence with 3–7 capitula, compactly cymose-corymbose; peduncles 3–25 mm, with dense, white stellate hairs, numerous, short, dark glandular hairs and few to numerous, long simple eglandular hairs. Capitula 15– 20 mm in diameter, rounded at base. Involucral bracts in numerous rows, 6–9 × 1.0–1.5 mm, dark green with whitish margins, linear-lanceolate, rounded-obtuse at apex, with numerous very small stellate hairs particularly on the margin, numerous, short, dark glandular hairs down the centre, and few to numerous, long, whitish, dark-based simple eglandular hairs. Ligules bright yellow, paler beneath, glabrous, with 5 lobes at apex. Styles yellow or slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 2–3 mm, purplish-black. Flowers 5–7. 2n = 27, 36, 45. Introduced. Naturalised in grass heath with Calluna vulgaris–Erica tetralix tussocks between Stephill Bottom and Pig Bush in the New Forest, Hampshire. Formerly naturalised in an old limestone quarry at Cave Hill, Belfast between 1897 and 1910. In its many forms native in north and central Europe, southwards to northern Switzerland and the eastern Carpathians. 5. P. praealta (Vill. ex Gochnat) F. W. Schultz & Sch. Bip. Small-flowered Fox-and-Cubs Hieracium praealtum Vill. ex Gochnat Perennial herb with an oblique rootstock, without or with long, slender stolons which have leaves 1–3.5 × 0.2– 0.5 mm, linear-oblanceolate, obtuse at apex, entire and narrowed at base, usually with long, pale simple eglandular hairs on the margin and midrib beneath. Leaves pale green or glaucous-green on upper surface, paler beneath, basal in a rosette, the lamina 3–8 × 0.5–1.2(–2.0) cm, oblanceolate, spathulate or narrowly elliptical, obtuse to acute at apex, entire, narrowed at base; cauline 1–3(–9), like the basal but mostly smaller; all glabrous or with few to numerous, long, subrigid, pale simple eglandular hairs on the margin and sometimes on the surfaces and sometimes with stellate hairs beneath. Flowering stems 30–70 cm, pale yellowishgreen, often purplish below, erect, rigid, striate, glabrous or with few to numerous, medium to long, pale, dark-based, subrigid simple eglandular hairs and a few, short, dark glandular hairs. Inflorescence with few to many capitula, usually in lax to compact cymose-corymbose clusters, but sometimes with long branches; peduncles with more or less dense stellate hairs, and few to numerous simple eglandular and

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dark glandular hairs. Capitula 7–10 mm in diameter, narrowed or rounded at base. Involucral bracts in numerous rows, 5.0–8.5 × 0.7–1.0 mm, olive or blackish-green with paler margins, linear-lanceolate, more or less acute at apex, with more or less numerous stellate hairs, and few to numerous, dark or dark-based, medium to long simple eglandular hairs and short, dark glandular hairs in various proportions. Ligules yellow, the outer sometimes with a red stripe on the outer face, glabrous, with 5 lobes at apex. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 1.5–2.0 mm, purplish-black. Flowers 5–7(–9). (a) Subsp. praealta Hieracium florentinum subsp. praealtum (Vill. ex Gochnat) N¨ageli & Peter Stolons very short or absent. Involucral bracts and peduncles with more or less numerous glandular hairs, without or with few simple eglandular hairs. 2n = 36, 45. (b) Subsp. thaumasia (Peter) P. D. Sell Hieracium magyaricum subsp. thaumasium Peter; Hieracium praealtum subsp. thaumasium (Peter) P. D. Sell; P. thaumasia (Peter) Dost´al; Hieracium magyaricum subsp. arvorum N¨ageli & Peter; Hieracium bauhinii subsp. arvorum (N¨ageli & Peter) Zahn; P. praealta subsp. arvorum (N¨ageli & Peter) P. D. Sell & C. West; Hieracium spraguei Pugsley; P. praealta subsp. spraguei (Pugsley) P. D. Sell & C. West Stolons long and slender. Involucral bracts with numerous glandular hairs, without or with few to numerous simple eglandular hairs. Introduced. An established garden escape on roadsides, walls and railway banks. Scattered localities in England and Wales north to Cheshire and in West Lothian. Subsp. praealta, first recorded in 1899, occurs in a number of localities in Devonshire, in Middlesex near Elstree, Hanslope in Buckinghamshire and Plumley in Cheshire. It is widespread in Continental Europe from Belgium and France to Russia. Subsp. thaumasia occurs at Great Bedwyn in Wiltshire, Hungerford in Berkshire, Carriden, Bo’ness in West Lothian and on the border of Hertfordshire and Buckinghamshire. The plant on the border of Hertfordshire and Buckinghamshire was named Hieracium spraguei on the slight difference in the indumentum of the involucral bracts. Subsp. thaumasia also occurs in central and eastern Europe. The species extends eastwards into western Asia. 6. P. lactucella (Wallr.) P. D. Sell & C. West Glaucous Fox-and-cubs Hieracium lactucella Wallr.; Hieracium auricula auct. Perennial herb with a slender rootstock and numerous, long, prostrate stolons which sometimes root at the nodes and bear leaves 0.5–2.0 × 0.3–0.7 cm, spathulate, rounded at apex, entire, narrowed at base. Leaves glaucous on upper surface, paler beneath; basal with lamina 2–7(−9) × 0.3–1.5 cm, spathulate or narrowly elliptical, usually rounded-obtuse at apex, entire, gradually narrowed below; cauline 0–2(−3), like the basal but smaller and sessile and semiamplexicaul; all glabrous or with pale, stiff simple eglandular hairs up to 4 mm on the margin or midrib beneath.

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Flowering stems (6–)10–20(−45) cm, pale green, with minute stellate hairs, numerous minute glandular hairs and sometimes occasional simple eglandular hairs. Inflorescence with 1–5 capitula, irregularly cymose; peduncles with dense stellate hairs, numerous, dark glandular hairs and usually some simple eglandular hairs. Capitula 12–15 mm in diameter, rounded at base. Involucral bracts in numerous rows, 5–7 × 0.5–1.0 mm, blackishgreen with pale green margins, linear-lanceolate, obtuse at apex, with scattered stellate hairs, numerous, long and short, dark glandular hairs and sometimes with numerous simple eglandular hairs. Ligules yellow, sometimes tipped red on outer surface, glabrous, with 5 lobes at apex. Styles yellow. Receptacle pits with margins irregularly dentate. Achenes 1.5–1.7 mm, purplish-black. Flowers 6–7. 2n = 18, 27, 36. Introduced. Found in a pasture at Keevil in Wiltshire in 1904 in a small patch which did not flower well, and was almost certainly not native. A persistent weed in a garden at Harrow in Middlesex. There is a specimen in the Liverpool herbarium labelled at the foot of Ingleborough Hill, Yorkshire, 2 June 1818. Native of most of Europe and extending into western Asia. Our plant is subsp. lactucella which occurs throughout most of the range of the species. 7. P. flagellaris (Willd.) P. D. Sell & C. West Few-headed Mouse-ear Hawkweed Hieracium flagellare Willd. Perennial herb with an oblique rootstock and long, thick hairy stolons which bear leaves 1–5(–7) × 0.3–2.0 cm, narrowly elliptical, oblanceolate or spathulate, obtuse to acute at apex, entire, gradually narrowed at base, hairy. Leaves pale green on upper surface, paler beneath; basal with lamina 3–13 × 0.5–2.5 cm, oblanceolate, spathulate or narrowly elliptical, obtuse to more or less acute at apex, entire or very obscurely denticulate, gradually narrowed below; cauline 0–3, small, narrowly oblanceolate or linear, acute at apex; all with long, pale, stiff, often bulbousbased simple eglandular hairs on both surfaces and the margins and few to numerous, minute stellate hairs beneath. Flowering stems 12–40 cm, often furcate, pale yellowishgreen, striate, with numerous stellate and short, dark glandular hairs and few to numerous, pale simple eglandular hairs. Inflorescence lax, with 1–6 capitula; peduncles often very long, with dense stellate hairs, numerous short, dark glandular hairs and few to numerous, medium to long, pale simple eglandular hairs. Capitula 15–25 mm in diameter, rounded at base. Involucral bracts in numerous rows, (8–)9–12 × 1.0–1.2 mm, dark olive green, the inner with paler margins, linear-lanceolate, obtuse to acute at apex, with few to numerous stellate hairs, numerous, short, dark glandular hairs and few to dense, medium to long pale simple eglandular hairs. Ligules yellow, glabrous, with 5 lobes at apex. Styles yellow or discoloured. Receptacle pits with margins incise-dentate. Achenes 1.7–2.0 mm, purplishblack. Flowers 5–9. (a) Subsp. flagellaris Hieracium pilosella var. viride F. Hanb.; Hieracium stoloniflorum auct.; H. pilosella var. stoloniflorum auct.

Flowering stems up to 40 cm. Capitula 2–4(–7); peduncles with more or less numerous simple eglandular hairs 2–3 mm. Involucral bracts with few to numerous simple eglandular hairs up to 1.5 mm. 2n = 36, 45. (b) Subsp. bicapitata P. D. Sell & C. West Hieracium flagellare subsp. bicapitatum (P. D. Sell & C. West) P. D. Sell Flowering stems up to 18 cm. Capitula (1–)2(–4); peduncles with numerous simple eglandular hairs up to 7.5 mm. Involucral bracts with dense simple eglandular hairs up to 2.5 mm. 2n = 54. Native and introduced. The species has a European Boreo-temperate range. Subsp. bicapitata is native and endemic to limestone rocks at White Ness from north of Pund to south of Brugarth in Central Mainland; on granulitic gneiss at West Burrafirth in West Mainland; and crumbling granite between The Kirk and Slocka on the north side of Ronas Voe in North Mainland; all in the Shetland Islands. Subsp. flagellaris is naturalised on grassy roadsides and railway banks and was first recorded in 1869. It occurs in scattered localities in south and central England and is abundant on railway banks in the Edinburgh area. It is native of north, west and central Europe and just into western Asia. 8. P. officinarum F. W. Schultz & Sch. Bip. Mouse-ear Hawkweed Hieracium pilosella L.; Hieracium sarmentosum Salisb. nom. illegit.; Hieracium repens Gray, non Willd. Perennial herb with a generally more or less slender rootstock and long, prostrate branched stolons which bear leaves 1–6 × 0.3–1.5 cm, spathulate to narrowly elliptical, obtuse to acute at apex, entire and narrowed at base with medium to long, stiff, pale simple eglandular hairs on both surfaces and the margins and dense stellate hairs beneath, the stolons terminating in rooting rosettes of overwintering leaves. Leaves dull or dark green on upper surface, grey or whitish beneath; basal in a rosette, the lamina 1–12 × 0.5–2.0 cm, oblanceolate, spathulate or narrowly elliptical, obtuse to acute at apex, entire or rarely obscurely denticulate, long-attenuate at base; cauline absent except sometimes for 1 minute bract; with numerous, medium to long, stiff, pale simple eglandular hairs on both surfaces and the margin and with dense stellate hairs beneath. Flowering stems 5–30(–50) cm, pale green, with dense stellate hairs and simple eglandular and glandular hairs in various proportions. Inflorescence a solitary capitulum at the end of the stem, or rarely at the end of stolons. Capitula 15–25 mm in diameter, rounded at base. Involucral bracts in numerous rows (6–)8–12(–15) × 0.5–1.5(–2.0) mm, greyish to blackish-green, the inner with broad, pale, margins and occasionally reddish-tipped, linear-lanceolate, acute at apex, with dense stellate hairs, with few to numerous or without dark glandular hairs, without or with few to dense simple eglandular hairs. Ligules pale lemon yellow, the outer usually marked with red on the outer face, glabrous with 5 lobes at apex. Styles yellow. Receptacle pits with margins incise-dentate. Achenes about 2 mm, purplish-black. Flowers 5–8. Sexual or apomictic. In natural habitats usually only one of the following subspecies is to be found, but in disturbed areas up to three

38. Pilosella have been found together. Intermediates also occur. The frequent records from roadsides and by railways suggest it can be distributed by traffic. (a) Subsp. micradenia (N¨ageli & Peter) P. D. Sell & C. West Hieracium pilosella subsp. micradenium N¨ageli & Peter; Hieracium pilosella var. concinnatum F. Hanb.; P. officinarum subsp. concinnata (F. Hanb.) P. D. Sell & C. West; Hieracium pilosella var. concolor Tausch; Hieracium concinnatum (F. Hanb.) Roffey Flowering stems usually 8–20 cm, with numerous stellate and glandular hairs and sometimes a few pale simple eglandular hairs. Involucral bracts 8–11 mm, with numerous, short, pale or dark, more or less equal glandular hairs mostly not more than 0.5 mm and dense stellate hairs, without simple eglandular hairs. 2n = 18, 36, 45. (b) Subsp. euronota (N¨ageli & Peter) P. D. Sell & C. West Hieracium pilosella subsp. euronotum N¨ageli & Peter; P. euronota (N¨ageli & Peter) Dost´al Flowering stems up to 30 cm, with numerous stellate and glandular hairs and sometimes a few, pale simple eglandular hairs. Involucral bracts (8–)10–12(–15) mm, with numerous dark, markedly unequal glandular hairs up to 1.0 mm and dense stellate hairs, without simple eglandular hairs. 2n = 45. (c) Subsp. officinarum Hieracium pilosella L.; Hieracium pilosella subsp. latiusculum N¨ageli & Peter; Hieracium latiusculum (N¨ageli & Peter) Roffey; Hieracium pilosella subsp. subvirescens N¨ageli & Peter; Hieracium subvirescens (N¨ageli & Peter) Roffey Flowering stems (5–)10–30(–40) cm, with stellate, pale simple eglandular and glandular hairs in various proportions in upper part. Involucral bracts 7–10(–12) mm, with numerous stellate and pale, sometimes dark-based simple eglandular hairs and with more or less numerous dark glandular hairs. 2n = 36. (d) Subsp. trichosoma (Peter) P. D. Sell & C. West Hieracium pilosella subsp. trichosomum Peter; Pilosella trichosoma (Peter) Dost´al; Hieracium pilosella var. nigrescens Fr.; Hieracium pilosella subsp. nigrescens (Fr.) P. D. Sell & C. West; Hieracium pernigrescens (Zahn) Roffey; Hieracium pilosella subsp. pernigrescens Zahn; Hieracium pilosella var. angustifolium Tausch; Hieracium pilosella subsp. angustius N¨ageli & Peter Flowering stems (5–)10–25 cm, with dark, sometimes darker based simple eglandular, dark glandular and stellate hairs in upper part. Involucral bracts 7–10(–12) mm, with numerous stellate and dark or dark-based simple eglandular hairs and with more or less numerous dark glandular hairs. 2n = 36, 45, 54, 63. (e) Subsp. tricholepia (N¨ageli & Peter) P. D. Sell & C. West Hieracium pilosella subsp. tricholepium N¨ageli & Peter; Hieracium pilosella var. tricholepium (N¨ageli & Peter)

215

Pugsley; Hieracium tricholepium (N¨ageli & Peter) Roffey; P. tricholepia (N¨ageli & Peter) Dost´al Flowering stems 8–25 cm, with stellate, pale simple eglandular and glandular hairs in upper part. Involucral bracts 7–11 mm, with dense stellate and dense, pale simple eglandular hairs, without or with inconspicuous glandular hairs. 2n = 36. (f) Subsp. melanops (Peter) P. D. Sell & C. West Hieracium pilosella subsp. melanops Peter; Pilosella melanops (Peter) Dost´al Flowering stems 6–30 cm, with stellate hairs and dark simple eglandular and glandular hairs in the upper part. Involucral bracts 10–12(–15) mm, with numerous stellate and moderately dense, dark simple eglandular hairs up to 2 mm, without or with inconspicuous glandular hairs. 2n = 36, 45. (g) Subsp. trichoscapa (N¨ageli & Peter) P. D. Sell & C. West Hieracium pilosella subsp. trichoscapum N¨ageli & Peter; P. trichoscapa (N¨ageli & Peter) Dost´al; Hieracium pseudopilosella auct.; Hieracium pilosella var. pseudopilosella auct. Flowering stems 6–30 cm, with stellate hairs, dense, dark simple eglandular hairs up to 5 mm and glandular hairs at the apex of the stem. Involucral bracts 10–12(–15) mm, with numerous stellate and dense, dark simple eglandular hairs, without or with inconspicuous glandular hairs. 2n = 45, 54. Native. Short grassland on well-drained soils, heaths, banks, cliff ledges, dunes and shingle. Locally common throughout Great Britain and Ireland, but absent from the Shetland Islands. Throughout Europe to western Siberia and Caucasus. A member of the European Temperate element. Subsp. micradenia is probably our commonest subspecies and is found throughout Great Britain and Ireland especially in lowland areas. It is often abundant on sandy and chalk heaths and on shingle, dunes and cliff-tops by the sea. It occurs throughout most of Europe and in Turkey and the Caucasus. Subsp. euronota is recorded from scattered localities in western and northern Britain from Cornwall to Sutherland. It occurs in native habitats on dunes, rocks, river banks and grassland in upland areas such as Teesdale and by the sea in Kintyre. It also occurs on railway banks as far apart as Cornwall, Derbyshire, Westmorland and Perthshire. In Europe it occurs mainly in the north and on mountains further south. Subsp. officinarum probably occurs throughout Great Britain and Ireland but is mainly a plant of the lowlands. It is particularly common in chalk and limestone grassland and also occurs near the sea. It may be introduced in some places where it occurs on roadsides and railways. It is found throughout most of Europe. Subsp. trichosoma is the common plant of the mountain areas, but occurs sparingly in other places. In Europe it is mainly in the north and east. Subsp. tricholepia occurs on heaths, grassland, quarries, golf courses, roadsides, sea cliffs and railway banks in scattered localities throughout Great Britain and seems to be quite common in Ireland. It is found throughout most of Europe. Subsp. melanops occurs in scattered localities in Great Britain particularly on dunes and cliffs by the coast and is recorded for Galway in Ireland.

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39. Hieracium

217

subulate-dentate

dentate

fimbriate-dentate Receptacle pits

entire

denticulate

dentate

serrate

mammiform dentate

Leaf teeth

glandular plumed

simple eglandular Hairs

stellate

aquiline mammiform

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It is widespread in Europe, particularly in upland regions. Subsp. trichoscapa is a scarce plant of northern Britain. It is widespread in Europe, but mainly in upland areas. Much more recording needs to be made to get the true distribution of the subspecies, bearing in mind that they may be introduced by roads and railways and such places as golf courses and quarries. 9. P. peleterana (M´erat) F. W. Schultz & Sch. Bip. Shaggy Mouse-ear Hawkweed Hieracium peleteranum M´erat Perennial herb with a thick rootstock and short, curved, stolons with very long hairy (−15 mm) and bearing leaves 1–5 × 0.4–1.5 cm, narrowly elliptical or oblanceolate, obtuse or acute at apex, entire, attenuate at base and with very long, pale simple eglandular hairs. Leaves in a rosette, the lamina 2.1–8.7(–15) × 0.4–2.0 cm, deep green above, greyish-white beneath, oblanceolate or elliptical, obtuse to acute at apex, entire, more or less attenuate at base; with numerous to dense, long(–12 mm), pale, rigid, bulbousbased simple eglandular hairs on both surfaces and the margin and dense stellate hairs on the lower surface. Flowering stems 1–several, (6–)10–30(–35) cm, lateral from the axils of the rosette leaves, with 0–2 linear bracts, with dense stellate hairs, more or less numerous, short, dark glandular hairs and more or less numerous, long (–5 mm), pale, spreading simple eglandular hairs. Capitulum solitary, 25–45 mm in diameter. Involucral bracts in numerous rows, 8–15 × 1.5–3.0 mm, greyish-green, sometimes reddish-tipped, lanceolate, acute at apex, with dense, long (– 6 mm), pale, shining, whitish, black-based simple eglandular hairs, usually few, to rarely more numerous stellate hairs and few glandular hairs completely obscured by the simple eglandular ones. Ligules pale yellow, at least the outer strongly marked with red on the outer face, glabrous, with 5 lobes at apex. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 2.0–2.5 mm, purplish-black. Flowers 5–7. Sexual, but apomictic plants occur in mainland Europe. (a) Subsp. peleterana Rosette leaves 2.1–8.7 × 0.9–2.0 cm, mostly oblanceolate or elliptical, cuneate or shortly attenuate at base. Flowering stems up to 12(–18) cm. Involucre 11–15 × 12–20 mm; bracts 1.5–3.0 mm wide at base, lanceolate, with dense, long simple eglandular hairs, rarely with a few, minute glandular hairs, usually with few stellate hairs. 2n = 18. (b) Subsp. subpeleterana (N¨ageli & Peter) P. D. Sell Hieracium peleteranum subsp. subpeleteranum N¨ageli & Peter Rosette leaves (2–)4–11(–15) × 0.6–1.2 cm, mostly narrowly elliptical, attenuate at base. Flowering stems (6–)10–20(–30) cm. Involucre 10–15 × 12–17 mm; bracts 1.5–2.0 mm wide at base, lanceolate, usually with dense, long simple eglandular hairs, with few glandular hairs and usually few stellate hairs. 2n = 18. (c) Subsp. tenuiscapa (Pugsley) P. D. Sell & C. West Hieracium peleteranum var. tenuiscapum Pugsley; Hieracium peleteranum subsp. tenuiscapum (Pugsley) P. D. Sell

Rosette leaves (3–)4–7(–13) × 0.4–1.2(–1.6) cm, mostly oblanceolate, long-attenuate at base. Flowering stems (6–)12–30(–35) cm. Involucre 10–12(–13) × (9–)10– 12(–14) mm; bracts about 1.5 mm wide at base, linearlanceolate, with more or less dense, long simple eglandular hairs, usually with few glandular hairs and scattered stellate hairs. Native. Short open grassland on well-drained soils, dunes and rock ledges. In a few scattered localities in the southern half of Britain. It occurs in north, west and west-central Europe and is a member of the Suboceanic Boreo-temperate element. Subsp. peleterana occurs in the Channel Islands and at Freshwater in the Isle of Wight. It also occurs in western Europe, mainly in coastal areas. Subsp. subpeleterana is known in Britain from Craig Breidden in Montgomeryshire, Portland in Dorset and Jersey in the Channel Islands. In Europe it is mainly the plant of the mountains. Subsp. tenuiscapa occurs in Jersey, Wetton Mill in Staffordshire, Dovedale in Derbyshire and Linton Falls in Yorkshire. In Europe it probably occurs throughout the range of the species. × officinarum = P. × longisquama (Peter) Holub Hieracium × longisquamum Peter; Hieracium pachylodes N¨ageli & Peter nom. illegit.; P. pachylodes Soj´ak nom. illegit. This is a sexual hybrid which is difficult to distinguish, especially from P. officinarum subsp. tricholepia, except in the field when growing with its parents. It is most like P. peleterana with dense, pale simple eglandular hairs on the rather broad involucral bracts, but differs in its longer stolons and smaller capitula. From P. officinarum it differs in its thicker stolons and dense simple eglandular hairs of the involucral bracts. P. officinarum subsp. tricholepia is a tall more slender plant with long, slender stolons. 2n = 27. Native. At Wetton Mill in Staffordshire the parents are P. peleterana subsp. tenuiscapa and P. officinarum subsp. officinarum. This combination of parents is also found in Jersey. A similar plant was found near Folkestone in 1890, but P. peleterana is not known there. A plant found in Jersey in 1933 may have P. officinarum subsp. micradenia as one of its parents. Hybrids from Guernsey are P. peleterana subsp. peleterana × P. officinarum subsp. officinarum. In all cases they appear to be individual plants scattered among their parents. P. × longisquama is widespread in north and west-central Europe. 39. Hieracium L. Perennial herbs with an erect or oblique, branched stock, never stoloniferous. Stems usually solitary, usually branched, leaves absent to numerous. Leaves alternate, entire to laciniate-dentate, at least the basal usually distinctly petiolate. Capitula 1–numerous. Involucral bracts in numerous rows. Flowers bisexual. Corolla ligulate, usually yellow, rarely green, sometimes hairy, with 5 lobes at apex. Anthers tailed, apical appendage elongate; filaments smooth. Style slender, with long, slender branches, the branches with short hairs. Receptacle flat, pitted, without scales. Achenes 2.5–5.0 mm, narrowly obconical or cylindrical with a narrowed base, truncate at apex, the 10 ribs

39. Hieracium apically confluent into an obscure ring; pappus hairs in 2 rows, simple, both long and short intermixed. Throughout Europe, northern Asia, north-west Africa and North America, but most diverse in Europe, especially in the mountains. The South African species should be placed in Tolpis or in a distinct genus. The Central and South American species do not belong to Hieracium. Their involucral bracts are in 2 rows, some species have ligules other than yellow, and some species have their achenes narrowed at the apex. All these characters fit Crepis better than Hieracium. By these same characters they are different from Pilosella, though some of them have stolons as in that genus. It is possible they should be put in a distinct genus, but it would be difficult to define it from Crepis, though easy to define it from Hieracium. It is difficult to tell how many species there are in Hieracium as a species is defined here. Possibly 10,000, but much depends on how narrow the species limit, as the wider the area which is covered, the more difficult it becomes to define the species. Even in Great Britain and Ireland the botanist working only on his own county will find things much easier than when dealing with the whole area. History P. D. S. has dealt with the history of the study of hawkweeds in Great Britain and Ireland at some length (Sell, 1987). James Backhouse jun. (1825–90) was to start this study with his monograph in 1856. He only dealt critically with the Section Alpina however. He was followed by F. J. Hanbury (1851–1938), A. Ley (1842–1911), E. F. Linton (1848–1928), W. R. Linton (1850–1908) and E. S. Marshall (1858–1919). At the same time S. O. I. Almquist (1844– 1923), M. M. W. Brenner (1843–1930), G. A. H. Dahlstedt (1856–1934), C. J. Lindeberg (1815–1909), M. Elfstrand (1859–1927) and J. P. Norrlin (1842–1917) were studying them in Fennoscandia and C. J. M. Arvet-Touvet (1841– 1913) and H. Sudre (1862–1918) in France and the Alps. In 1921–2, a German schoolmaster, K. H. Zahn (1865– 1940), published a world monograph of the genus in three volumes in the Pflanzenreich series. He used major aggregate species with large numbers of subspecies, which are the equivalent of the species in this flora. Unfortunately his large herbarium at Berlin (B), on which he mainly based his account, was destroyed in the Second World War. Fortunately most of the specimens he used as regards British and Irish species are duplicated in our herbarium (CGE). J. Roffey (1860–1927) in 1925 attempted to bring all the British and Irish taxa in line with Zahn’s monograph, but he treated Zahn’s subspecies as species, creating numerous new combinations. In the 1920s and early 1930s G. C. Druce (1850–1932) and H. H. Johnston (1856–1930) sent specimens to the ageing Dahlstedt, who added many new names to our flora without really understanding the genus in Great Britain. It took the critical and logical mind of H. W. Pugsley (1868–1947) to put chaos into order in his Prodromus (1948). This monograph, much of which was written during the war when books and specimens were not easily obtained, cannot be praised too highly. Between 1950 and 1966 Cyril West and P. D. S. carried out a major revision of the genus resulting in the account

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in the Critical Supplement to the Atlas of the British Flora in 1968. They looked at some 39,000 specimens, used the Cambridge herbarium (CGE) as their main source, and borrowed material from many public herbaria and private collections. The great majority of type specimens were examined and both saw a large number of the species in the field. Quite a lot of the species defined were aggregates in which not enough fresh material was available to do the necessary splitting. Much new collecting, particularly by Archie Kenneth in Kintyre, Argyllshire and especially Ross-shire, made some of these species very unsatisfactory, but West and Sell were not prepared to alter their earlier decisions without a major review of at least all the material in the Cambridge herbarium. A start was made on the Section Alpina, but the death of C. W. and P. D. S.’s long illness meant that this was not finally finished, with the major help of David Tennant, until 1995. The account of Hieracium in this flora is based on a major revision of the 20,000 specimens in the Cambridge herbarium, all of which have been examined by P. D. S. under the microscope. He has seen 320 of the species described here in the field and grown over 300 clones in the Botanic Garden at Cambridge, 33 of which are additional to those seen in the field. Morphology All the species of Hieracium are perennials, though in cultivation at least they will flower in their first year, the rhizomatous stock then branching freely and the plant living for many years (at least 30 years in the Cambridge Botanic Garden). How long-lived they are in nature is not known, but they probably live for some time if there is no competition. The stems are very short in some species and very tall in others; branching is usually above the middle. They are slender to robust. Indumentum varies from almost glabrous to very hairy and consists of stellate, glandular and simple eglandular and rarely plumose hairs. Simple eglandular hairs are sometimes minutely denticulate. All kinds of hair may be present on a plant, or only one. They are very helpful in diagnosing species, but are mostly not characteristic of any one section. Leaves may form a basal rosette (phyllopodous), or be entirely cauline (aphyllopodous), or have the basal dying before flowering (hypophyllopodous), or have the cauline congested near the base (pseudophyllopodous). This habit is one of the features on which the major grouping of the genus is based. The basal leaves are few to numerous. The first formed, the primordial, are small and often subrotund. They were described for every species by H. W. Pugsley in his monographic treatment but we think this is unnecessary. In some species the basal leaves are all very similar, in others they are very variable and in some the outer are distinctly different from the inner. Shape, apex, toothing and base are very important in the definition of species. The petiole is only rarely an important character. The number of cauline leaves is a sectional character, as is whether they are petiolate, sessile or amplexicaul. In a few species the angle in which they come from the stem is diagnostic. Hair clothing is very important and may be different on upper and lower surfaces and the margin. The rigidity of the simple

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eglandular hairs is diagnostic of two sections, but is variable in others. Glandular hairs on the leaves occur only in three sections. Plumose hairs are found only in two introduced sections.

Achenes Only occasionally is achene size useful for species definition, and colour seems to be of no use at all except for one small section.

Inflorescence The arrangement of the capitula, together with the number of cauline leaves, are the basis for the main classification. Most of the species of the Section Alpina usually only have one capitulum, rarely more, except in cultivation. The remaining sections either have a furcate-corymbose inflorescence in which the peduncles are more or less erect and straight or a paniculate-corymbose inflorescence in which the peduncles are more or less curved. Their indumentum, its density, amount of glandular, simple eglandular and stellate hairs is important.

Colour The subtle tones of ligule and leaf colour in the field are an important way of recognising the species, but it is difficult to pass this information from one botanist to another, especially as each of us probably sees colour in a different way and certainly each uses different terminology.

Capitulum There are two things which will give you the size of the capitulum, the length of the longest involucral bracts and the diameter of the open head. The base of the capitulum is usually either rounded or subtruncate and is rarely cuneate. The involucre in the Section Alpina is distinctive in shape and colour, but these characters have not been dealt with in other sections. Involucral bracts The colour, shape, apex laxity, dimensions and indumentum of the involucral bracts are possibly the most important set of characters when defining a species. They seem to remain constant under all conditions except very rarely in autumn-flowering forms in the Section Hieracium. Whether these bracts are curved in at the apex (incumbent) or erect (porrect) in bud is a helpful character to define some sections. The length of the bract is measured from the base of the capitulum to the tip of the bract. Measurements have mostly been made from pressed specimens. Ligules The colour of the ligules and whether they are hairy or not is a useful character for defining species. In the Section Alpina, where David Tennant had all the species in cultivation, use has been made of their density, width, angle and whether they overlap and length by which the hairs project beyond the apex. This information has not been recorded for most of the species in the other groups. Styles The colour of the style is sometimes helpful in the definition of a species but really needs to be decided while the plant is fresh and in good condition. Pure yellow styles are a very good character, but there are all grades of discoloured ones. Often the stigma is a slightly different colour from the style. David Tennant has been more precise about this character in Section Alpina. Receptacle pits Whether the receptacle pits are shortly dentate, incisedentate, subulate-dentate or fimbriate-dentate is useful both for species and section definition.

Hair clothing Because of the importance of hair clothing in this genus, the details given in the glossary are strictly adhered to. It should be noted that in some herbaria the application of corrosive sublimate as an insect repellant has destroyed the hairs, particularly the stellate ones. Cultivation P. D. S. and Cyril West grew over 300 clones of Hieracium species in the Cambridge Botanic Garden. Their most constant characters in cultivation were the shape of the involucral bracts and the total indumentum of the plant. Leaf shape, apex and dentation were constant at first flowering and some species retained these characters in laterflowering plants. However, other plants, particularly in the Section Hieracium, looked like a completely different species when flowering in the autumn, but they never looked like any other known species. Their colour tended to become glaucous rather than green and they lost their characteristic leaf toothing. Achenes taken from these plants and planted immediately flowered with similar characters the following autumn, but the year after that they flowered in their characteristic spring form. All descriptions in this account refer to plants in nature and to the early (spring) form and not the later (autumnal) one. The least constant of all characters in cultivation was colour. Height and size of plants varied according to the amount of rain or watering they received and possibly the amount of light, but in any one year all plants of a clone looked exactly alike. No species in this account are accepted purely from cultivated plants. David Tennant grew all the species of the Section Alpina from collected achenes, and by growing them under controlled conditions in soils from their natural habitat was able to produce plants almost indistinguishable from those in nature in most cases. Collecting Haphazard collecting of hawkweeds, especially in all the well-known localities, is not to be encouraged as many species are rare, several being among the rarest of the British and Irish plants and herbaria are already full of specimens from those areas. Examination of the Critical Supplement to the Atlas of the British Flora will show large areas where there are no records at all and these are the areas which need attention. Inadequate specimens should not be collected and if possible a typical plant in a colony should be selected. Notes of leaf, ligules and style colour should be made and the

39. Hieracium direction of the involucral bracts when in bud. Fruiting specimens alone are usually not adequate. Photographs are also usually not adequate for identification. The higher mountain species can be collected at any time within the flowering period but lowland plants need to be collected at first flowering. In practice this means the species with one stem leaf in May and June, sometimes July, those with a few stem leaves in July and August and the very leafy ones at any time. Care should be taken with the very leafy ones that they are not secondary growth, having been cut off, or grazed, and then grown up again. Poor specimens gathered at the wrong time of the year do not help anyone, and may be destroying a rare plant. The species The species of Hieracium are regarded as apomictic because they have poor or practically no pollen, have a very high percentage of good seed and offspring are exactly alike. R. A. Finch has found that triploids and tetraploids form only univalents in the mother cell at meiosis. Many botanists believe that you can group apomicts to form the equivalent of sexual species. Experience suggests that such a group species is very difficult to define and does not show the interesting ecological and geographical distribution shown by the microspecies. Moreover the group species is likely to include plants of entirely different origins. The larger the microspecies group the greater the number of plants that will fall between groups. In actual fact the International Code of Botanical Nomenclature does not recognise either group species or microspecies and anything which has been given a binomial must be regarded as a species. Thus, Hieracium species as defined in this account are as distinct as any other species. What bothers botanists is the vast number of them. Some species of Hieracium present more problems than others. A large proportion of the species show almost no variation. They are not all of local distribution. H. hypochaeroides for example, occurring in widely different areas, seems to show almost no variation at all. H. duriceps, although showing great variation in size, is usually recognisable at a glance throughout its range. Species of the Sections Alpina and Alpestria are very local and there is usually little trouble in knowing when to split or when to lump. Other species such as H. vulgatum, H. schmidtii and H. sabaudum, which occur over a large area, have within their compass variants which, unless you have a wide range of material available, would appear to be distinct species. It is thought better to have a broader concept for these species than to split them up and ignore intermediates as is sometimes done in apomictic genera. Sometimes complexes are only difficult in certain areas. For example the H. argenteum, caledonicum, nitidum, scoticum complex in the north of Sutherland does not present the same problems elsewhere. Origin of the species When the Hieracia moved south in front of the last glaciation and then north again after the glaciers receded, they were probably still sexual and would have come in contact with species they were previously isolated from and hybridised with them. Selection could then have taken place for particular habitats or distribution and at some period,

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probably after they moved north again, they all became apogamous. As far as is known the only sexual diploid Hieracium in Great Britain and Ireland is H. umbellatum. All the remainder which have had their chromosomes counted, with one exception, are either tetraploid or triploid and apogamous. The one exception, in the Section Alpina, is pentaploid. New species may arise as sports. In nature when the species are on cliffs or streamsides this probably does not happen very often as there are usually not many niches in which seedlings may grow. Where plants of the Section Hieracium arrive in recently disturbed open ground however, in a few years they can multiply into large numbers, as capitula produce almost 100 per cent fertile achenes. In such situations, as Nils Hylander found in Swedish grasslands (1943), a large number of closely allied species can be found. Hylander pointed out to P. D. S. several closely similar plants growing intermixed on a roadside bank at Royston (actually in Cambridgeshire), which he grew in the Cambridge Botanic Garden where their differences remained constant over many years. C. H. Ostenfeld (1921) grew all the seeds from one plant and then all the seeds from successive generations until he finally got a sport, which being apomictic reproduced itself exactly. Some of the most wide-ranging variable species may be those that had not sorted themselves into geographical or ecological populations when they became apomictic. Many of the local species may well have originated as sports of an allied more widespread species. Suggestions about this possibility in specific cases are mentioned under the accounts of sections or the species themselves. Intermediates sometimes occur between H. vagum and H. sabaudum. In Great Britain R. A. Finch found that both species and the intermediates were triploid apomicts. On the other hand, V. Ferakova in Czechoslovakia found that H. vagum was diploid yet was an exact match morphologically for our triploid apomictic H. vagum. Where species have poor or no pollen, but produce a high percentage of good seed they have been labelled apomictic. If looked at in detail by R. A. Finch, that has been indicated. Almquist, S. O. I. (Nov.–Dec. 1881). Studier o¨ fver sl¨agtet Hieracium. Stockholm. Arvet-Touvet, C. J. M. (1873). Monographe des Pilosella et des Hieracium du Dauphin´e suivie de l’analyse de quelques autres plantes. Grenoble. Supplement (1877) in Bull. Soc. Bot. France 1–39. Additions (1879) in Bull. Soc. Bot. France 1–20. Arvet-Touvet, C. J. M. (Oct. 1880). Essai de classification sur les genres Pilosella & Hieracium principalement pour les esp`eces et les formes de la r´egion Sud-Ouest de l’Europe. Bull. Soc. Dauphin 1880: 278–292. Arvet-Touvet, C. J. M. (Oct. 1881). Spicilegium rariorum vel novum Hieraciorum praecipue americanorum et europaeorum. Grenoble. Arvet-Touvet, C. J. M. (1888). Les Hieracium des Alpes franc¸aises ou occidentales de l’Europe. Lyon, Gen`eve, Bˆale. Arvet-Touvet, C. J. M. (1913). Hieraciorum praesertim Galliae et Hispanicae catalogus systematicus. Paris. Babington, C. C. (June 1856). Manual of British botany. Ed. 4. London. Backhouse, J. jun. (1853). Alpine British plants, particularly Hieracium. Phytologist (Newman) 4: 804–808. Backhouse, J. jun. (1855). Hieracium L. in Hooker, W. J. & Arnott, G. A. W. The British flora, ed. 7, pp. 216–227. London.

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Backhouse, J. jun. (Apr.–May 1856). Monograph of the British Hieracia. York. Baker, J. G. (1879). On a variety of Hieracium caesium from the Great Ormes Head. Jour. Bot. (London) 17: 360–362. Beeby, W. H. (1908). On the flora of Shetland. Ann. Scot. Nat. Hist. 17: 110–117. Bevan, J. (1982). Section Amplexicaulia Zahn. Hier. Group Notes 2: 4–8. Bevan, J. (1983). Section Amplexicaulia Zahn. Hier. Group Notes 4: 3–4. Bevan, J. (1990). Hieracium britannicum F. J. Hanb. in Wales. Watsonia 18: 199–213. Bladon, J. (1847). Description of Hieracium heterophyllum, a species new to science. Phytologist (Newman) 2: 961–963. Boreau, A. (1857). Flore du centre de la France. Ed. 3. Paris. Chater, A. O. (1984). Recent collections of Hieracium in v.c. 46. Hier. Group Notes 6: 1–3. Dahlstedt, H. (1893). Bidrag syd¨ostra Sveriges Hieracium-flora. II. Kungl. Svenska Vet.-Akad. Handl. 25(3): 1–201. Dahlstedt, H. (1893). Bidrag syd¨ostra Sveriges Hieracium-flora. III. Kungl. Svenska Vet.-Akad. Handl. 26(3): 1–266. Dahlstedt, H. (1894). Scandinaviens Hieracium-flora. Acta Hort. Berg. 2(4): 1–146. Dahlstedt, H. (1903). The Hieracia from the Faeroes in Warming, E. (Edit.) Botany of the Faer¨oes, vol. 2, pp. 625–659. Dahlstedt, H. (1927). Hieracium L. Rep. Bot. Soc. Exch. Cl. Brit. Isles 8: 28–29. Dahlstedt, H. (1930). Hieracium L. Rep. Bot. Soc. Exch. Cl. Brit. Isles 9: 24–26. Dahlstedt, H. (1933). Hieracium L. Rep. Bot. Soc. Exch. Cl. Brit. Isles 10: 23–25. Druce, G. C. (1928). British plant list. Ed. 2. Arbroath. Elfstrand, M. (1890). Botaniska utflygter i sydvestra Jemtland och angr˚ansande del af S¨odra Trondhjems amt sommeren 1889. Bihang till Kungl. Svenska Vet.-Akad. Handl. 16, Afd. III, no. 7. Elfstrand, M. (1893). Hieracia alpina aus den Hochgebirgsgegenden des mittleren Scandinaviens. Upsala. Fries, E. M. (1847–1848). Symbolae ad historium Hieraciorum. Nova Acta Reg. Soc. Sci. Uppsala 13: 383–416 (1847); 14: 1– 220 (1848). Fries, E. M. (1862). Epicrisis generis Hieraciorum. Upsala. Gibson, S. (Mar. 1844). Note on Hieracium hypochoeroides, in reference to Mr Watson’s notes. Phytologist (Newman) 1: 907– 908. Gremli, A. (1896). Exkursionsflora f¨ur die Schweiz. Ed. 8. Aarau. Hanbury, F. J. (1888). Notes on some Hieracia new to Britain. Jour. Bot. (London) 26: 204–206. Hanbury, F. J. (1889). Further notes on Hieracia new to Britain. Jour. Bot. (London) 27: 73–76. Hanbury, F. J. (1889–1898). An illustrated monograph of the British Hieracia. London. Hanbury, F. J. (1892). Further notes on Hieracia new to Britain. Jour. Bot. (London) 30: 129–134; 165–170; 206–209; 255–261; 366–370. Hanbury, F. J. (1893). Further notes on Hieracia new to Britain. Jour. Bot. (London) 31: 16–19. Hanbury, F. J. (1894). Notes on British Hieracia. Jour. Bot. (London) 32: 225–233. Hanbury, F. J. (1894). A tentative list of British Hieracia. Jour. Bot. (London) 32: post 224. Hanbury, F. J. & Thompson, R. F. (1904). Hieracia L. in Babington, C. C. Manual of British botany. Ed. 9. London. Hieracia Group Notes (1982–1989). 1–15. Hylander, N. (1943). Die Grassameneink¨ommlinge schwedischer Parke mit besonderer ber¨ucksichtingung der Hieracia silvaticiformia. Sym. Bot. Upsal. 7(1).

Johansson, K. & Samuelsson, G. (1923). Dalarnes Hieracia Silvaticiformia. Leipzig. Johansson, K. & Samuelsson, G. (1923). Dalarnes Hieracia Vulgatiformia. Leipzig. Johnston, H. H. (1929). Additions to the flora of Orkney. Edinburgh. Johnston, H. H. (1932). Additions to the flora of Orkney. Edinburgh. Johnston, H. H. (1933). Additions to the flora of Orkney. Edinburgh. Kenneth, A. G. & Stirling, A. McG. (1970). Notes on the hawkweeds (Hieracium sensu lato) of western Scotland. Watsonia 8: 97–120. Ley, A. (1895). Hieracium diaphanum Fr. var. cacuminum, n. var. Jour. Bot. (London) 33: 86–87. Ley, A. (1898). Two new forms of Hieracium. Jour. Bot. (London) 36: 6–7. Ley, A. (1899). Two new Hieracium forms. Jour. Bot. (London) 37: 35–36. Ley, A. (1900). Some Welsh hawkweeds. Jour. Bot. (London) 38: 3–7. Ley, A. (1901). Notes on Welsh hawkweeds. Jour. Bot. (London) 39: 167–168. Ley, A. (1907). Hieracium notes. Jour. Bot. (London) 45: 108–112. Ley, A. (1909). Brecon and West Yorkshire hawkweeds. Jour. Bot. (London) 47: 8–16; 47–55. Ley, A. (1910). Three South Wales hawkweeds. Jour. Bot. (London) 48: 326–327. Lindeberg, C. J. (1868–1878). Hieracia Scandinavica exsiccata. 1: nos. 1–50 (May 1868); 2: nos. 51–100 (1 June 1872); 3: 101–150 (July 1878) (with Latin descriptions of new species). Linton, E. F. (1891). Some British hawkweeds. Jour. Bot. (London) 29: 271–273. Linton, E. F. (1897). Hieracium ogweni, sp. nov. Jour. Bot. (London) 35: 407–408. Linton, E. F. (1911). Some new forms of Hieracia. Jour. Bot. (London) 49: 353–356. Linton, E. F. & W. R. (1893). British hawkweeds. Jour. Bot. (London) 31: 145–149; 177–182; 195–202. Linton, E. F. & W. R. (1896–1906). Set of British Hieracia. Fascicle 1 (May 1896); 2 (1896); 3 (Oct. 1897); 4 (Oct. 1898); 5 (Oct. 1899); 6 (Jan. 1901); Supplement (1906). Linton, W. R. (1890). Hieracium holophyllum sp. nov. Jour. Bot. (London) 28: 376. Linton, W. R. (1905). An account of the British Hieracia. London. Marshall, E. S. (1892). A new British Hieracium. Jour. Bot. (London) 30: 18–19. Marshall, E. S. (1892). ‘Hieracium anfractiforme’. Jour. Bot. (London) 30: 183. Marshall, E. S. (1894). New variety of Hieracium dovrense Fries. Jour. Bot. (London) 32: 215–216. Marshall, E. S. (1913). Two new Scottish hawkweeds. Jour. Bot. (London) 31: 119–122. Merx¨uller, H. (1975). Diploide Hieracien. Anal. Inst. Bot. Cavanilles 2: 189–196. Mills, J. N. (1968). A new species of Hieracium in Derbyshire. Watsonia 7: 40–42. Mills, J. N. & Stace, C. A. (1974). Chromosome numbers of British plants. Watsonia 10: 167–168. Morton, J. K. (1974). Chromosome numbers of British plants. Watsonia 10: 169. Norrlin, J. P. (1906). Suomen Keltanot. Appendix to Melan, A. J. Suomen Kasvio, pp. 613–746. Omang, S. O. F. (1935). Die Hieracien Norwegens. Oslo. Omang, S. O. F. (1938). Monographische Bearbeitung der Hieracien Islands. Skr. Vid.-Akad. Oslo 1938(3): 1–67. Omang, S. O. F. (1940). Hieracium L. in Nordhagen, R. Norsk Flora, pp. 712–744. Oslo. Omang, S. O. F. (1949). Descriptiones specierum novarum e stirpe Hieracii alpini. I. Nytt. Mag. Naturvidensk. 87: 95–195.

39. Hieracium Omang, S. O. F. (1951). Descriptiones specierum novarum e stirpe Hieracii alpini. II. Nytt. Mag. Naturvidensk. 88: 87–152. Omang, S. O. F. (1954). Descriptiones specierum novarum e stirpe Hieracii alpini. III. Nytt. Mag. Bot. 2: 61–100. Oskarsson, I (1966). Synopsis and revision of Icelandic Hieracia. Visind. Islend. 37: 1–142. Ostenfeld, C. H. (1921). Some experiments on the origin of new forms in Genus Hieracium subgenus Archieracium. Jour. Genet. 11: 117–122. Pugsley, H. W. (1920). Notes on British hawkweeds. Jour. Bot. (London) 58: 281–285. Pugsley, H. W. (1925). Review of Hieracium list in the London Catalogue of British Plants. Jour. Bot. (London) 63: 302–304. Pugsley, H. W. (1941). New species of Hieracium in Britain. Jour. Bot. (London) 79:177–183; 193–197. Pugsley, H. W. (1946). List of British species of Hieracium L. Jour. Ecol. 33: 345–347. Pugsley, H. W. (1948). A prodromus of the British Hieracia. Jour. Linn. Soc. London (Bot.) 54: 1–356. Purchas, W. H. (1895). Hieracium murorum var. pachyphyllum, n. var. Jour. Bot. (London) 33: 114–115. Purchas, W. H. (1899). Hieracium cymbifolium, sp. n. Jour. Bot. (London) 37: 421–422. Rich, T. C. G. (2000). Distribution and conservation of Hieracium snowdoniense, Snowdonia Hawkweed. Unpublished report. National Museums and Galleries of Wales. Rich, T. C. G. (2000). Distribution and conservation of Hieracium holosericeum, Beautiful Hawkweed, in Wales. Unpublished report. National Museums and Galleries of Wales. Rich, T. C. G. (2001). Distribution and conservation of Hieracium cyathis, Chalice Hawkweed. Unpublished report. National Museums and Galleries of Wales. Rich, T. C. G. (2001). Distribution and conservation of Hieracium asteridiophyllum, Llangatt´ock Hawkweed. Unpublished report. National Museums and Galleries of Wales. Rich, T. C. G. (2001). Distribution and conservation of Hieracium cillense, Craig y Cilau Hawkweed. Unpublished report. National Museums and Galleries of Wales. Rich, T. C. G. (2002). Conservation of Britain’s biodiversity: Hieracium asteridiophyllum and H. cillense (Asteraceae). Watsonia 24: 101–106. Roffey, J. (1925). Hieracium L. in The London catalogue of British plants. Ed. 11. London. Roffey, J. (1925). The Hieracia of the London Catalogue. Jour. Bot. (London) 63: 315–322. Scott, W. & Palmer, R. C. (1987). The flowering plants and ferns of the Shetland Islands. Lerwick. Scott, W. & Palmer, R. C. (1995). A new Shetland Hieracium of the section Alpestria [Fries] F. N. Williams. Watsonia 20: 282–284. Sell, P. D. (1987). Hieracium westii sp. nov. Watsonia 16: 283– 286. Sell, P. D. (1987). Hieracium pellucidum Laest., H. lucidulum (A. Ley) Roffey and H. asteridiophyllum P. D. Sell & C. West. Watsonia 16: 287–289. Sell, P. D. (1987). Typification of Hieracium amplexicaule L. Watsonia 16: 336–337. Sell, P. D. (1987). Obituary of Cyril West (1887–1986). Watsonia 16: 358–360. Sell, P. D. (1987). An introduction to the study of the British Hieracia. I. History and classification. Watsonia 16: 365–371. Sell, P. D. (2000). Hieracium kentii sp. nov. (Asteraceae). Watsonia 23: 216–217. Sell, P. D. & West, C. (1955). Notes on British Hieracia. I. Watsonia 3: 233–236. Sell, P. D. & West, C. (1958). List of Hieracium L. in Dandy, J. E. List of British vascular plants, pp. 120–130. London.

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Sell, P. D. & West, C. (1962). Notes on British Hieracia. II. The species of the Orkney Islands. Watsonia 5: 215–223. Sell, P. D. & West, C. (1965). A revision of the British species of Hieracium section Alpestria [Fries] F. N. Williams. Watsonia 6: 85–105. (Illustrations of all the species.) Sell, P. D. & West, C. (1967). Hieracium L. in Sell, P. D., Notes on the British flora. Watsonia 6: 303–313. Sell, P. D. & West, C. (1968). Hieracium L. in Perring, F. H. & Sell, P. D. (Edits.) Critical supplement to the atlas of the British flora, pp. 75–131. London. Sell, P. D. & West, C. (1975). in Heywood, V. H. (Edit.) Flora Europaea. Notulae systematicae ad floram Europaeam spectantes. No. 19. Bot. Jour. Linn. Soc. 71: 256–267. Sell, P. D. & West, C. (1976). Hieracium L. in Tutin, T. G. et al. (Edits.) Flora Europaea, vol. 4, pp. 358–410. Cambridge. Sell, P. D. & West, C. (1980). Hieracium zygophorum Hyl., new to the British Isles. Watsonia 3: 27–29. Sell, P. D., West, C. & Tennant, D. J. (1995). Eleven new British species of Hieracium L. section Alpina (Fries) F. N. Williams. Watsonia 20: 351–365. Shi, Y., Gornall, R. J., Draper, J. & Stace, C. A. (1996). Intraspecific molecular variation in Hieracium sect. Alpina (Asteraceae), an apomictic group. Folia Geobot. Phytotax. 31: 305–313. Stace, C. A., Gornall, R. J., Squirrel, J. & Shi, Y. (1995). Chromosome numbers in Hieracium L. section Alpina (Fries) F. N. Williams. Watsonia 20: 367–377. Stenstr¨om, K. O. E. (1889). V¨arml˚andska Archieracier. Upsala. (All the taxa with an asterisk in this account are subspecies.) Sudre, H. (1902). Hieracium du centre de la France. Albi. Syme, J. T. B. (1879). Description of Hieracium dewari, a new species. Trans. Bot. Soc. Edinb. 13: 211. Tennant, D. J. (1986). Further dicoveries of Hieracium backhousei F. J. Hanb. Watsonia 16: 86–87. Webster, M. McC. (1978). Flora of Moray, Nairn and East Inverness. Aberdeen. (A key to Hieracium species by P. D. Sell and C. West.) Williams, F. N. (1902, 1903). Hieracium L. in Prodromus Florae Britannicae. Brentford. Zahn, K. H. (1921–2). Compositae-Hieracium in Engler, H. G. A. (Edit.) Das Pflanzenreich IV. 280. Leipzig. Heften 75, 76 and 77 (1921); 79 (1922).

Key To make a key to 412 species of Hieracium is very difficult. It is not helpful to divide it into sections as two of the sections particularly contain species of hybrid origin between other sections and the characters that define the sections are the most difficult to understand. The leafy species from break 1 to 61 vary considerably in the number of their cauline leaves and many have also had to be keyed out again with the Section Vulgata at breaks 253 to 356. It is felt the appearance of species of the Section Alpina will be quickly understood and they have been keyed out together from breaks 64 to 108. One or two species have been keyed out again elsewhere. Section Cerinthoidea forms a natural group and has been keyed out between breaks 111 and 147. Included with species of that group are some of the Section Subalpina whose origin is probably a cross between a species of the Section Cerinthoidea and a species of the Section Alpina. The Section Oreadea and the Section Stelligera have been keyed out together between breaks 148 and 252. Their species form a natural group, but are hard to define in words though no other way has been found of dealing with them than the description at break 148. The

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Section Vulgata is keyed out between breaks 253 and 356. It is a well-defined group but unfortunately the key has to include many smaller plants of the leafy groups. It includes all species with between 2 and 12 cauline leaves, except those species in the Section Oreadea. The last huge group from break 357 onwards deals with the very large Section Hieracium plus the Section Subalpina. It contains all those species with 0–1(–2) cauline leaves, except for those in the Section Stelligera. Specimens which will key out best are usually the largest in a colony. It should be remembered that the length of an involucral bract is from the base of the capitulum to the tip of the longest bract. Sections with cauline leaves 12 or more 1. Cauline leaves more than 12; basal leaves usually absent, rarely forming a rosette, sometimes forming a 2. false rosette 1. Cauline leaves less than 12; basal leaves present or not 61. 3. 2. Medium and upper leaves more or less amplexicaul 25. 2. Leaves not amplexicaul 3. Involucral bracts without or with very few glandular 4. hairs or microglands 3. Involucral bracts with numerous glandular hairs and or 7. microglands 4. Leaves mostly less than 4 times as long as broad, 56. subtruncatum broadly elliptical, lanceolate or oblong 4. Many of leaves more than 4 times as long as broad, 5. oblong, narrowly elliptical, lanceolate or linear 12. maritimum 5. Stems robust; leaves 12–40 6. 5. Stems slender; leaves (7–)9–16(–20) 6. Leaves abruptly narrowed to a pointed apex; styles 54. northroense discoloured 6. Leaves gradually narrowed to a pointed apex; styles 57. dilectum yellow when fresh 7. Involucral bracts with numerous simple eglandular hairs 8. as well as numerous glandular hairs 7. Involucral bracts with numerous glandular hairs, 12. without or with few simple eglandular hairs 19. melanoglochin 8. Styles yellow 9. 8. Styles discoloured 10. 9. Leaves 3–4 cm wide, ovate or broadly elliptical 9. Leaves up to 2.5(–3.0) cm wide, oblong, elliptical, 11. lanceolate or oblong-ovate 10. Involucral bracts 10–13 × 1.0–1.3 mm, with numerous 50. carpathicum stellate hairs, densely so on the margins 10. Involucral bracts 9–10 × about 1.0 mm, with scattered stellate hairs towards the base and on the margins 51. dewari 11. Leaves with stellate hairs on lower surface only; involucral bracts with numerous very short and short dark glandular hairs, intermixed with few to numerous, 16. strictiforme short to medium simple eglandular hairs 11. Leaves with stellate hairs on both surfaces; involucral bracts with very short to medium glandular hairs and dense microglands intermixed with a few medium 20. drummondii simple eglandular hairs 13. 12. Peduncles with numerous glandular hairs 12. Peduncles without or with only occasional glandular 16. hairs

14. 13. Median and upper leaves less than 2 cm wide 13. At least some of the median and upper leaves more than 15. 2 cm wide 14. Stems with scattered simple eglandular hairs 53. vinicaule throughout; styles discoloured 14. Stems glabrous or nearly so; styles yellow 55. klingrahoolense 15. Peduncles without or with occasional simple eglandular hairs; involucral bracts 0.8–1.3 mm wide, dark olive 47. prenanthoides green 15. Peduncles with numerous simple eglandular hairs; involucral bracts 1.0–1.5 mm wide, blackish 48. lanceolatum 17. 16. Styles discoloured 18. 16. Styles yellow 17. Leaves less than 2 cm wide; involucral bracts 1.0–1.3 13. pycnotrichum mm wide 17. At least some leaves more than 2 cm wide; involucral 21. subcrocatum bracts 1.2–1.5 mm wide 18. At least some of the teeth on the leaves more than 3 mm 19. melanoglochin 19. 18. Teeth on leaves all less than 3 mm 19. Leaves mostly ovate, tending to be rounded towards the base or at least less drawn out below than at the apex 22. latobrigorum 19. Most of the leaves narrow in outline and drawn out at 20. both base and apex 21. 20. Inflorescence narrow in outline 23. 20. Inflorescence widely spreading 21. Leaves 1.5–2.5 cm wide; involucral bracts 1.2–1.5 mm 23. tavense wide 21. Leaves 1–2 cm wide; involucral bracts 1.0–1.2 mm wide 22. 22. Leaves 15–30(–40), gradually decreasing in size 14. bakeranum upwards, mostly obtuse at apex 22. Leaves 10–19, rapidly decreasing in size in upper half of 55. klingrahoolense stem, mostly acute at apex 23. Leaves mostly only denticulate; involucral bracts with glandular hairs mostly down the median line 15. subumbellatiforme 23. Leaves often more sharply toothed; involucral bracts 24. with glandular hairs spread over the surface 24. Leaves 8–14 cm, with teeth few and fairly widely spread 17. reticulatiforme 24. Leaves 5–10 cm, with teeth numerous and close together 18. reayense 25. Involucral bracts more or less glabrous or with an occasional glandular or simple eglandular hairs or both 26. 25. Involucral bracts with more or less numerous microglandular and glandular hairs, or simple 41. eglandular hairs, or both 27. 26. At least some leaf-teeth more than 5 mm 28. 26. Leaf-teeth not exceeding 5 mm 27. Leaves lanceolate or ovate to oblong-lanceolate, deeply 4. prominentidens laciniate 27. Leaves linear to oblong, with distant, long curved teeth 11(a,iii). umbellatum subsp. umbellatum var. coronopifolium 29. 28. Many leaves more or less ovate 28. Leaves longer, being oblong, linear or elliptical-oblong 33.

39. Hieracium 29. Inflorescence umbellate 11(c,viii). umbellatum subsp. bichlorophyllum var. latifolium 30. 29. Inflorescence paniculate 30. Involucral bracts not more than 10 mm 38. cambricogothicum 31. 30. Longest involucral bracts more than 10 mm 31. aphyllopodioides 31. Involucral bracts 1.5–2.0 mm wide 32. 31. Involucral bracts not more than 1.5 mm wide 1(1). vagum forma vagum 32. Styles discoloured 1(2). vagum forma croceostylum 32. Styles yellow 33. Leaves with margins not recurved and ciliate; 34. inflorescence paniculate, not umbellate 33. Leaves with margins recurved and scaberulous; 35. inflorescence more or less umbellate 3. salticola 34. Leaves bluish-green; styles discoloured 40. lissolepium 34. Leaves yellowish-green; styles yellow 36. 35. Dwarf coastal plants up to 30 cm 38. 35. Taller inland plants up to 100 cm 36. Leaves rigid, acute at apex, usually less than 10 mm wide 11(a,i). umbellatum subsp. umbellatum var. dunense 36. Leaves flaccid or fleshy, more or less obtuse at apex, 37. usually some more than 10 mm wide 37. Stem slender; leaves up to 1.5 cm wide, narrowly elliptical 11(c,v). umbellatum subsp. bichlorophyllum var. curtum 37. Stem robust and fleshy; leaves up to 5 cm wide, thick and fleshy, narrowly to broadly elliptical 11(c,vi). umbellatum subsp. bichlorophyllum var. sarniense 38. Leaves mostly not more than 10 mm wide, narrowly linear and sharply pointed 11(a,ii). umbellatum subsp. umbellatum var. umbellatum 38. Many leaves more than 10 mm wide, oblong and often 39. more or less obtuse 39. Leaves pale green and flaccid, teeth often blunt 11(c,vii). umbellatum subsp. bichlorophyllum var. taylori 40. 39. Leaves darker green and stiff, teeth often sharper 40. Leaves numerous and close together; outer involucral bracts reflexed 11(a,iv). umbellatum subsp. umbellatum var. commune 40. Leaves more widely spaced; outer involucral bracts not 11(b). umbellatum subsp. ogwenii reflexed 41. Involucral bracts with more or less numerous glandular hairs or microglands, without or with very occasional 42. simple eglandular hairs 41. Involucral bracts with more or less numerous simple 46. eglandular hairs, with or without glandular hairs 32. scullyi 42. Leaves not more than twice as long as broad 43. 42. Many leaves more than twice as long as broad 5. virgultorum 43. Leaves long attenuate at base 44. 43. Leaves rounded or shortly narrowed at base 44. Involucral bracts with long, spidery glandular hairs; margins of receptacle pits sharply dentate 33. scabrisetum 44. Involucral bracts with numerous, short glandular hairs or microglands; margins of receptacle pits 45. fimbriate-dentate 45. Leaves dark green, sharply serrate or serrulate; inflorescence often in a narrow panicle; involucral bracts with numerous very short to short, unequal glandular 2. rigens hairs

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45. Leaves medium to dark green, less sharply toothed; inflorescence a spreading corymbose panicle; involucral bracts with numerous to dense glandular hairs and 8(2). sabaudum forma bladonii microglands 46. Involucral bracts with numerous simple eglandular hairs 47. and few or no glandular hairs 46. Involucral bracts with numerous simple eglandular hairs 49. and numerous glandular hairs 47. Leaves laciniate with slender teeth; styles discoloured; 36. charitodon receptacle pits fimbriate-dentate 47. Leaves toothed but not laciniate; styles yellow; 48. receptacle pits subulate-dentate 48. Stem throughout and both surfaces of the leaves with 35. substrigosum numerous simple eglandular hairs 48. Upper part of stem and upper surface of leaves with few 37. gothicoides or no simple eglandular hairs 50. 49. Leaf-teeth blunt and more or less mammiform 51. 49. Leaf-teeth pointed and not mammiform 26. cantianum 50. Involucral bracts without stellate hairs 50. Involucral bracts with numerous stellate hairs towards 27. acamptum the base 51. Leaves long lanceolate, oblong-lanceolate or narrowly 6. salicetorum elliptic-oblong, attenuate at both ends 51. Leaves broader and not so attenuate, particularly at the 52. base 53. 52. Margins of receptacle pits subulate-dentate 56. 52. Margins of receptacle pits fimbriate-dentate 53. Capitula 25–35 mm in diameter; involucre up to 12 mm 54. wide; involucral bracts up to 10 mm 53. Capitula 35–45 mm in diameter; involucre up to 15 mm 55. wide; involucral bracts up to 12 mm 54. Leaves up to 3 cm wide, elliptical, ovate, elliptical28. trichocaulon oblong or lanceolate 54. Leaves up to 2 cm wide, narrowly elliptical, lanceolate 29. calcaricola or linear-lanceolate 55. Leaves ovate or lanceolate, the lower at least narrowed at both ends, dentate, the teeth long and narrow up to 6 mm; involucral bracts 1.0–1.3 mm wide, linearlanceolate, gradually narrowed to an obtuse apex 24. ornatilorum 55. Leaves mainly ovate, often more narrowed at the apex than base, often entire or minutely toothed, but can develop larger, sharper teeth; involucral bracts 1.0– 1.6 mm wide, broadly linear-lanceolate, obtuse at apex 30. stewartii 39. uiginskyense 56. Styles yellow 57. 56. Styles discoloured 25. eboracense 57. Involucral bracts up to 10 mm 58. 57. Longest involucral bracts exceeding 10 mm 58. Leaves elliptical, equally tapered at both ends, broadest in the middle and with fine, sharp or spinulose teeth up 7. argutifolium to 6 mm 58. Leaves mostly broadest below the middle and teeth 59. although sometimes sharp are not spinulose 10. dumosum 59. Involucral bracts with broad pale margins 59. Involucral bracts blackish-green and nearly concolorous 60. 60. Leaves lanceolate, ovate-lanceolate, oblong-lanceolate or ovate; capitula 25–35 mm in diameter 8(1). sabaudum forma sabaudum

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60. Leaves larger and more frequently ovate with larger capitula up to 40 mm in diameter on a widely spreading 9. eminentiforme inflorescence 61. Leaves with numerous obvious medium, stout glandular 62. hairs 61. Leaves without glandular hairs or with few to fairly 64. numerous very short or minute glandular hairs

72. Leaves dark green, strongly suffused with purple, 91. einichense elliptic-oblong, cuneate at base 73. Involucral bracts with numerous simple eglandular and numerous glandular hairs 75(2). hanburyi forma hanburyi

Section Amplexicaulia

74. At least some of the basal leaves more than 20 mm 75. wide, often broadly elliptical or ovate 74. Basal leaves not more than 20 mm wide, usually narrow 89. and often drawn out at base 75. Styles pure yellow when fresh, sometimes becoming 76. slightly discoloured when dry 75. Styles not pure yellow but discoloured when fresh often 81. turning even darker when dry 76. Leaves subentire or with an occasional small tooth; 100. calvum ligules with few hairs 76. Leaves obviously toothed; ligules with numerous hairs 77. 78. 77. Leaves short, rarely more than 50 mm 79. 77. Leaves long and narrow, up to 110 mm

72. amplexicaule 62. Leaves with glandular hairs only 62. Leaves with simple eglandular hairs as well as glandular 63. hairs 63. Cauline leaves gradually decreasing in size upwards but staying broad, with additional ones on the branches, the 71. speluncarum upper broadly ovate 63. Cauline leaves gradually decreasing in size upwards and becoming narrower, oblong-ovate, oblong-obovate or 73. pulmonarioides oblong-oblanceolate

Section Alpina 64. Dwarf mountain plants up to 35(–75) cm, with a rosette of few to numerous leaves and small cauline ones which are often bract-like, both often bearing minute glandular hairs; capitula relatively large, often solitary, sometimes 2–4, rarely with stellate hairs on the involucre; ligules often with hairs on the backs as well as the tips (once one of this group of species has been seen the group 65. usually becomes easily recognisable) 64. Often taller plants with larger basal leaves and larger cauline leaves when they are present; minute glandular hairs sometimes present on leaves; capitula often more numerous with an open panicle, stellate hairs often present on involucre; ligules glabrous or if hairy usually 109. only on tips 65. Basal leaves with stellate hairs on both surfaces, usually 66. densely so beneath 65. Basal leaves without stellate hairs or with a few on the 67. under surface 66. Leaves entire, denticulate or with a few shallow teeth; 83. optimum styles dark olive-grey 66. Leaves usually irregularly dentate or serrate-dentate with more or less mammiform teeth; styles pure yellow 84. grovesii 68. 67. Glandular hairs on involucral bracts very obvious 67. Glandular hairs on involucral bracts few or more or less 74. obscured by simple eglandular hairs 74. perscitum 68. Involucral bracts not exceeding 13 mm 69. 68. Involucral bracts usually much exceeding 13 mm 70. 69. Leaves mainly long attenuate at base 71. 69. Leaves mainly shortly contracted or cuneate at base 70. Involucral bracts 1.1–1.8 mm wide, broadly linear-lanceolate, rather broadly acute; ligules palish yellow; styles pure yellow to dirty greenish-yellow with 94. milesii yellow stigmas 70. Involucral bracts 0.9–1.5 mm wide, linear-lanceolate to linear, narrowly acute at apex; ligules deepish yellow; 95. memorabile styles olive-grey, rarely yellow 72. 71. Leaves mostly subentire or with a few small teeth 73. 71. At least some of the leaves with large teeth 72. Leaves usually pale green, ovate or subrotund, rounded, broadly cuneate or truncate at base 75(1). hanburyi forma pusillum

73. Involucral bracts with numerous unequal glandular hairs, without or with occasional simple eglandular hairs 75(3). hanburyi forma atraticeps

86. deargicola 78. Leaves ovate, lanceolate or subrotund 78. Leaves narrowly elliptical or oblong-elliptical 89. graniticola 79. Leaves with sharp almost spinulose teeth, upper surface 97. backhousei glabrous or nearly so 79. Leaves with blunt or rather pointed mammiform teeth, 80. more or less hairy on upper surface 80. Capitulum solitary, 45–65 mm in diameter; involucral bracts 12–19 × 1.0–1.2 mm, lax, very narrow at apex 96(1). insigne forma insigne 80. Capitula 1–3, 25–50(–55) mm in diameter; involucral bracts 10–16 × 1.0–1.6 mm, appressed but narrow at 96(2). insigne forma celsum apex 82. 81. Ligules with few hairs in upper half 85. 81. Ligules with numerous hairs in upper half 83. 82. Leaves mostly less than 50 mm long 84. 82. Many leaves over 50 mm long 83. Capitula 30–48 mm in diameter; inner involucral bracts 76. kennethii 11–15 mm, subacute at apex 83. Capitula 35–58 mm in diameter; inner involucral bracts 79. pensum 12–20 mm, sharply acute at apex 84. Leaves often blotched, subrotund, ovate, elliptical or lanceolate, often pointed at apex, cuneate or sometimes 77. completum truncate at base 84. Leaves not blotched, obovate, oblanceolate, oblong-elliptical or elliptical, often rounded at apex, 78. probum attenuate at base 86. 85. Whole plant densely long-hairy throughout 87. 85. Plant much less densely hairy and hairs shorter 80. calenduliflorum 86. Achenes 3.7–5.2 mm, purplish-black 86. Achenes (2.5–)3.0–4.0 mm, reddish-brown, never 81. notabile purplish-black 87. Leaves mostly more or less acute at apex, with sparse simple eglandular hairs on upper surface 82. subgracilentipes 87. Leaves mostly more or less obtuse at apex, usually with numerous simple eglandular hairs at least towards the 88. margins on upper surface

39. Hieracium 88. Leaves usually without reddish-violet tinge, the cauline 1–3; capitula 30–45 mm in diameter; ligules medium yellow, very discrete, usually curving upwards; styles 92. tenuifrons light olive-grey 88. Leaves usually with some reddish-violet tinge, the cauline 3–5(–9); capitula up to 55 mm in diameter; ligules deeper yellow, hardly discrete, usually straight; 93. subtenuifrons styles deep olive-grey 89. Styles pure yellow when fresh, sometimes slightly 90. discoloured when dry 99. 89. Styles discoloured, often becoming darker when dry 91. 90. Leaves not usually more than 50 mm long 95. 90. Some leaves more than 50 mm long, up to 120 mm 91. Leaves often ovate, obovate, subrotund or broadly elliptical with hairs scattered over the upper surface 86. deargicola 91. Leaves often narrow and more elongated without hairs 92. in the central area of the upper surface of the leaf 92. Teeth of leaves mammiform or cusped-mammiform 89. graniticola 92. Leaves frequently more or less entire or with small 93. sharp teeth 93. Capitulum characteristically nearly spherical in bud; ligules densely arranged and barely discrete 88. globosiflorum 93. Capitulum cylindrical or narrowly campanulate in bud; 94. ligules few and usually very discrete 94. Involucral bracts with dense simple eglandular hairs up 99(1). marginatum forma marginatum to 2.5 mm 94. Involucral bracts with dense simple eglandular hairs up 99(2). marginatum forma chaetocephalum to 4 mm 95. Most leaves long and narrow, spathulate, narrowly obovate or oblong, more or less obtuse at apex, teeth if 96. present usually small 95. Most leaves usually elliptical, ovate or lanceolate, some 97. usually acute at apex, teeth when present often large 96. Leaves deep green; inner involucral bracts lanceolate-subulate, hairs of involucre shaggy but not hanging at the base like a beard; outer ligules straight, 106. alpinum often recurving 96. Leaves pale green; inner involucral bracts usually linear or linear-lanceolate, hairs of involucre hanging at base like a beard; outer ligules curving upwards 107. holosericeum 97. Leaves glabrous on upper surface; ligules with a few to 100. calvum several hairs on the backs and tips 97. Leaves with very numerous hairs uniformly distributed on upper surface; ligules with more or less numerous 98. hairs on the backs and tips 98. Capitula 25–45 mm in diameter; inner involucral bracts (8–)10–15(–17) mm; achenes (3.0–)3.7–5.3 mm 102. macrocarpum 98. Capitula 30–50 mm in diameter; inner involucral bracts 9–20 mm; achenes 3.0–4.6 mm 103(2). eximium forma tenellum 100. 99. Ligules with few hairs in upper half 103. 99. Ligules with numerous hairs in upper half 100. Inner leaves spathulate, narrowly elliptical or oblanceolate; capitulum narrowed at base 105. pseudopetiolatum

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100. Inner leaves elliptical or narrowly obovate; capitulum 101. not narrowed at base 101. Leaves with numerous hairs on upper surface 90. pseudocurvatum 101. Leaves without hairs on upper surface except near the 102. margin 102. Involucre greenish-grey, its hairs with short black bases 85. larigense 102. Involucre blackish, its hairs with long black bases 87. subglobosum 103. Leaves with hairs on upper surface mostly only towards the margin; capitulum narrowed at base 105. pseudopetiolatum 103. Leaves with hairs all over upper surface; capitulum not 104. narrowed at base 104. Leaves drawn out at base to long petioles 103(1). eximium forma eximium 104. Leaves cuneate to shortly attenuate to a normally short 105. petiole 101. leptodon 105. Some leaves with long spinulose teeth 106. 105. Leaves subentire or with short teeth 98. mundum 106. Hairs of involucre up to 2.5 mm 107. 106. Hairs of involucre up to 6.0 mm 107. Leaves mostly elliptical and acute at apex 104. pentaploideum 108. 107. Leaves frequently elongated and obtuse at apex 108. Leaves usually without reddish-violet tinge, the cauline 1–3; capitula 30–45 mm in diameter; ligules medium yellow, very discrete, usually curving upwards; styles 92. tenuifrons light olive-grey 108. Leaves usually with some reddish-violet tinge, the cauline 3–5(–9); capitula up to 55 mm in diameter; ligules deeper yellow, hardly discrete; usually straight; 93. subtenuifrons styles deep olive-grey

Sections Lanatella and Andryaloidea 110. 109. Some of the hairs on the plant plumose 109. None of the hairs on the plant plumose, but they may 111. be minutely denticulate 110. Leaves spotted, blotched and marbled with brownish154. rionii purplish, with large mammiform teeth 110. Leaves not spotted, blotched or marbled, entire or 155. lanatum denticulate

Sections Cerinthoidea and Villosa 111. Stems usually more or less robust, with numerous long simple eglandular hairs especially below and sometimes with glandular hairs above; leaves often more or less glaucous, the basal forming a rosette, the cauline up to 6, usually more or less amplexicaul, with long simple eglandular hairs; capitula fairly large, often with long peduncles, usually with long simple eglandular hairs on the involucre, sometimes also with glandular hairs; involucral bracts incumbent in bud; ligules hairy at tips; styles discoloured; main flowering June and July. (The Section Cerinthoidea and some species in the Section Subalpina whose origin is 112. probably a hybrid with a species of Section Alpina.) 148. 111. Not with the above combination of characters 112. Leaves with numerous stellate hairs on the lower 113. surface and sometimes also on the upper surface

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112. Leaves without stellate hairs or with a few on the lower 117. surface 114. 113. Some involucral bracts over 13 mm 115. 113. Involucral bracts not more than 13 mm 114. Stem 30–60 cm; leaves erect, with stellate hairs on both surfaces; peduncles and involucral bracts with few 171. flocculosum to fairly numerous glandular hairs 114. Stem 15–50 cm; leaves spreading, with stellate hairs only beneath; peduncles and involucral bracts with 173. shoolbredii numerous glandular hairs 115. Leaves not more than 2.5 cm wide; involucral bracts with few to numerous glandular hairs 153. pseudanglicoides 115. Some leaves more than 2.5 cm wide; involucral bracts 116. without or with few glandular hairs 116. Leaves elliptical, broadly elliptical or subrotund; styles 170. flocculosiforme yellow 116. Leaves elliptical, ovate-lanceolate or lanceolate; styles 172. flocculipubens discoloured 118. 117. Leaves entire or with an occasional tooth 133. 117. Leaves with numerous teeth, sometimes small 118. Glandular hairs on involucral bracts more or less 119. obvious 118. Glandular hairs absent or not obvious on involucral 126. bracts 120. 119. Involucral bracts up to 16 mm 122. 119. Involucral bracts not more than 13 mm 120. Whole plant with long dense simple eglandular hairs, 156. pilosum without glandular hairs 120. Plant with less dense and shorter simple eglandular 121. hairs and at least some glandular hairs 121. Leaves 2–8(–11) × 1.5–3.0(–4.0) cm 141. pseudanglicum 161. magniceps 121. Leaves 3–15 × 2–7 cm 123. 122. Involucre broad 124. 122. Involucre narrow 123. Leaves predominantly ovate; peduncles often long; 164. anglicum ligules pale yellow 123. Leaves predominantly elliptical; peduncles shorter; 169. hebridense ligules medium yellow 148. eustales 124. Involucral bracts with dense stellate hairs 124. Involucral bracts with only some stellate hairs along 125. the margin or at the base 147. fissuricola 125. Leaves elliptical or oblong-elliptical 144. glandulidens 125. Leaves mainly lanceolate or ovate 126. Involucral bracts with numerous to dense stellate hairs, 127. particularly on the margin 126. Involucral bracts with few stellate hairs mostly on the 129. margin or towards the base 140. petrocharis 127. Leaves mostly obtuse at apex 128. 127. Leaves mostly acute at apex 150. arranense 128. Leaves 1.5–5.5 × 1–3 cm, often spotted 128. Leaves 3–12 × 2.0–4.5 cm, not spotted 162. langwellense 130. 129. Cauline leaves absent or bract-like 131. 129. Cauline leaves 1–4 and amplexicaul 130. Leaves 2.5–7.0 × 1.5–3.0 cm, ovate to ovate-oblong, often with small teeth near the base, often cuneate at 145. sannoxense base 130. Leaves 2–5 × 1.5–2.5 cm, elliptical or ovate, almost 149. ovaliforme entire, rounded at base

131. Leaves frequently obovate; involucral bracts 160. hartii 1.5–2.0 mm wide, green 131. Leaves not obovate; involucral bracts 1.0–1.5 mm 132. wide, blackish-green 151. thalassinum 132. Involucral bracts 4–10 × 1.0–1.4 mm 132. Involucral bracts 4–13 × 1.2–1.5 mm 159. cerinthiforme 134. 133. Leaves more or less attenuate at base 133. Leaves abruptly contracted or shortly cuneate at base 140. 138. aristidens 134. Leaves with fine spinulose teeth 135. 134. Leaves with broader teeth 135. Involucral bracts not more than 12 mm 151. thalassinum 136. 135. Involucral bracts more than 12 mm 136. Cauline leaves with a characteristic tuft of hairs at their 137. base 138. 136. Cauline leaves without a tuft of hairs at their base 137. Stem (10–)30–60 cm, robust; leaves 3–17 × 1.5–5.0 157. iricum cm, hairy on upper surface; capitula 1–20 137. Stem 10–35 cm, slender; leaves 2–9 × 2–4 cm, more or less glabrous on upper surface; capitula 2–6 158. scarpicum 138. Involucre very broad; bracts 1.5–2.0 mm wide 161. magniceps 139. 138. Involucre narrow; bracts 1.0–1.5 mm wide 139. Leaves usually green, with few teeth; ligules full yellow 162. langwellense 139. Leaves usually bluish-green, with numerous teeth; 163. oxybeles ligules pale yellow 141. 140. Ligules poorly developed (stylose) 142. 140. Ligules fully developed 141. Leaves 3.5–12 × 2–6 cm; involucral bracts 6–12 mm 165. jaculifolium 141. Leaves 4–10 × 2.5–4.0 cm; involucral bracts 6–15 mm 168. ampliatiforme 143. 142. Involucral bracts up to 15 mm 144. 142. Involucral bracts less than 13 mm 143. Peduncles with few glandular hairs; inner involucral 166. ampliatum bracts similar to outer 143. Peduncles with more numerous glandular hairs; inner involucral bracts gradually narrowed and more or less 167. cyclicum acute 145. 144. Leaves mostly rounded at apex 146. 144. Leaves mostly pointed at apex 145. Involucral bracts with numerous stellate hairs 142. oenophyllum 145. sannoxense 145. Involucral bracts with few stellate hairs 146. Involucral bracts with stellate hairs all over 150. arranense 146. Involucral bracts with stellate hairs only on margin and 147. at apex 147. Leaves mostly elliptical; involucral bracts porrect in 143. cumbriense bud 147. Leaves mostly ovate; involucral bracts incumbent in 144. glandulidens bud

Sections Oreadea and Stelligera 148. Leaves with rigid hairs especially on the margin, but sometimes also on the upper surface; inflorescence usually with few capitula on long peduncles; involucral bracts often porrect; styles usually yellow; ligules 149. usually glabrous (Sections Oreadea and Stelligera)

39. Hieracium 251. 148. Not with the above combination of characters 149. Cauline leaves 2 or more (Section Oreadea and some 150. species of Section Stelligera which overlap) 176. 149. Cauline leaves 1, rarely 2 193. chloranthum 150. Ligules green 151. 150. Ligules yellow 152. 151. Leaves hairy on upper surface 151. Leaves glabrous on upper surface or the hairs of the margin sometimes encroaching a short way on to the 165. surface 153. 152. Involucral bracts acute at apex 159. 152. Involucral bracts obtuse at apex 154. 153. Involucral bracts with numerous stellate hairs 155. 153. Involucral bracts with few or no stellate hairs 154. Involucral bracts with numerous glandular hairs 188. ascendentidens 154. Involucral bracts with few or no glandular hairs 185. proximum 156. 155. Involucral bracts with numerous glandular hairs 157. 155. Involucral bracts with few glandular hairs 156. Involucral bracts up to 14 mm, blackish; ligules with 114. senescens short hairs at apex. 156. Involucral bracts up to 12 mm, dark green, the inner with paler margins; ligules glabrous at apex 186. subrude 157. Leaves 2–5 cm, ovate, lanceolate or elliptical, more or 190. griffithii less spotted or blotched 157. Leaves 1.5–12.0 cm, elliptical, lanceolate or 158. oblong-lanceolate, not spotted or blotched 158. Involucral bracts drawn out to a filamentous apex 192. filisquamum 158. Involucral bracts not drawn out to a filamentous apex 191. orimeles 159. Involucral bracts with more or less numerous to dense 160. stellate hairs 162. 159. Involucral bracts with few or no stellate hairs 182. scottii 160. Involucral bracts up to 14 × 2 mm 161. 160. Involucral bracts not more than 10 × 1.3 mm 161. Basal leaves often dentate, abruptly contracted or 178. pusillifolium cuneate at base 161. Basal leaves usually subentire or denticulate, attenuate 183. farrense at base 162. Leaves often with big teeth; peduncles with numerous 187. deganwyense dark glandular hairs 162. Leaves rarely with big teeth; peduncles with few dark glandular hairs and numerous simple eglandular hairs 163. 163. Most leaves less than 6 cm, abruptly contracted or 189. beebyanum cuneate at base 163. Many leaves more than 6 cm, often attenuate at base 164. 180. breconicola 164. Cauline leaves mostly ovate 191. orimeles 164. Cauline leaves narrower and more oblong 165. Involucral bracts with numerous stellate hairs 166. especially along the margins 169. 165. Involucral bracts with few stellate hairs 176. apheles 166. Leaves with fairly remote, but sharp teeth 166. Leaves subentire, denticulate or if a few teeth, the teeth 167. blunt 167. Involucral bracts 1.0–1.2 mm wide, narrowly 184. angustisquamum linear-lanceolate

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167. Involucral bracts 1.2–1.7 mm wide, broadly 168. linear-lanceolate 168. Leaves broadly ovate, broadly elliptical or 195. subrubicundum oblong-elliptical, up to 13 × 6 cm 168. Leaves mostly ovate, up to 11 × 5 cm 201. rubicundiforme 170. 169. Basal leaves present at anthesis 173. 169. Basal leaves usually dead at anthesis 170. Leaves with sharp teeth; peduncles with occasional 171. glandular hairs 170. Leaves with small or blunt teeth; peduncles with 172. numerous glandular hairs 171. Basal leaves and leaves just above the base numerous, ovate, narrowly elliptical, lanceolate or oblong179. aggregatifolium lanceolate 171. Basal leaves few, broadly elliptical or ovate 181. pseudozetlandicum 202. boswellii 172. Leaves dentate 196. hesperium 172. Leaves subentire or denticulate 173. Leaves narrowly elliptical or oblong and not more than 194. cacuminum 2.5 cm wide 173. At least some leaves broadly elliptical or ovate and 174. some more than 2.5 cm wide 174. scoticum 174. Some leaves usually dentate 175. 174. Leaves subentire or denticulate 177. sowadeense 175. Leaves elliptical, lanceolate or ovate 175. Leaves oblong-elliptical or oblong-lanceolate 175. subscoticum 177. 176. Leaves with dense stellate hairs beneath 176. Leaves without or with very few stellate hairs beneath 183. 177. None of the leaves oblong or oblong-lanceolate, mostly 178. ovate, subrotund or elliptical 177. At least some of the leaves oblong, oblong-lanceolate 179. or oblong-elliptical 178. Leaves 2–11 × 1.2–6.5 cm; ligules hairy at tips 247. basalticola 252. fratrum 178. Leaves 3–6 × 2–3 cm; ligules glabrous 180. 179. At least some of the leaf-teeth long and sharp 179. Leaves mostly entire or denticulate, if teeth longer 181. rounded at apex 180. Involucral bracts with numerous short glandular hairs 260. stenopholidium 180. Involucral bracts without or with few small glandular 230. grampianum hairs 181. Leaves with stellate hairs only beneath; involucral bracts with stellate hairs mostly on the margin; ligules 216. carneddorum strongly pilose-tipped 181. Leaves with stellate hairs on both surfaces; involucral bracts with dense stellate hairs; ligules glabrous-tipped 182. 182. Involucral bracts up to 13 × 1.5 mm, linear-lanceolate 233. cillense 182. Involucral bracts up to 11 × 1.3 mm, narrow linear-lanceolate, the inner almost filamentous 234. lasiophyllum 183. Leaves with obvious teeth other than denticulations or 184. just 1–2 swollen ones 183. Leaves entire, denticulate or with the occasional 220. shallow tooth 185. 184. Ligules obviously hairy at tips

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184. Ligules glabrous or with a very occasional hair at tips 187. 185. Involucral bracts not exceeding 11 mm 248. apiculatidens 186. 185. Involucral bracts up to 15 mm 186. Leaves glaucous not blotched, elliptical, lanceolate or oblong; involucral bracts with few stellate hairs 207. hypophalacrum 186. Leaves bluish-green, often faintly blotched, ovate, elliptical or lanceolate; involucral bracts with 243. eucallum numerous stellate hairs 188. 187. Involucral bracts 14 mm or more 201. 187. Involucral bracts not more than 12 mm 189. 188. Involucral bracts with few stellate hairs 188. Involucral bracts with numerous to dense stellate hairs 192. 189. At least some of the leaves oblong, oblong-lanceolate 211. vagense or oblong-elliptical 190. 189. Leaves ovate, lanceolate or subrotund 190. Teeth of leaves more or less regular and mostly pointed 254. britannicoides at apex 190. Teeth of leaves irregular and at least some of them mammiform with a rounded apex and longer than the 191. rest 191. Inner involucral bracts filamentous 258. subbritannicum 191. Inner involucral bracts narrow but not filamentous 261. eustomon 193. 192. Leaves spotted and blotched 196. 192. Leaves not spotted or blotched 194. 193. At least some leaves ovate or subrotund 195. 193. Leaves lanceolate, elliptical or oblong-elliptical 194. Involucral bracts with few glandular hairs 219. tricolorans 194. Involucral bracts with numerous glandular hairs 255. britanniciforme 195. Leaves bluish-green, lanceolate-elliptical, 214. fucatifolium oblong-elliptical or elliptical 195. Leaves pale to medium green, lanceolate or elliptical 225. arvonense 196. Leaves with large, unequal, rounded, mammiform teeth 256. vagicola 196. Leaves subentire, with a few shallow teeth or sharply 197. dentate, but not with rounded mammiform teeth 197. Inner basal leaves up to 6 cm wide, ovate or 219. tricolorans elliptic-ovate 197. Inner basal leaves up to 3 cm wide, lanceolate, 198. oblong-lanceolate, oblong or ovate, rarely elliptical 198. All leaves sharply dentate, gradually narrowed at base 223. promontoriale 198. Leaves subentire, denticulate or with mammiform 199. teeth, abruptly contracted or subtruncate at base 199. Leaves dull green; inner involucral bracts more or less 220. subcyaneum acute, but not filamentous 200. 199. Inner involucral bracts filamentous 200. Petioles long; capitula 40–50 mm in diameter 221. decolor 200. Petioles short; capitula 30–40 mm in diameter 250. dicella 202. 201. Involucral bracts with few glandular hairs 210. 201. Involucral bracts with numerous glandular hairs 203. 202. Involucral bracts with few stellate hairs 202. Involucral bracts with numerous to dense stellate hairs 204. 204. angustatiforme 203. Leaves cuneate at base

251. sarcophylloides 203. Leaves rounded or truncate at base 204. Involucral bracts very narrow, 0.8–1.0 mm wide; leaves 222. inaequilaterum very irregularly dentate 204. Involucral bracts 1.0–1.5 mm wide; leaves more 205. regularly dentate 206. 205. Leaves cuneate at base; teeth sharp 205. Leaves rounded or truncate at base, teeth sinuate208. dentate or aquiline-mammiform 206. Leaves 5–11 × 2–6 cm, at least some broadly ovate 200. veterascens 206. Leaves 2.5–9.0 × 1–4, lanceolate to oblong or elliptical 207. 207. Peduncles without or with an occasional glandular hair 206. siluriense 207. Peduncles with few to numerous glandular hairs 223. promontoriale 208. Involucral bracts 6–13 × 1.0–1.2 mm, narrow 250. dicella linear-lanceolate, the inner filamentous 208. Involucral bracts 4–13 × 1.0–1.5 mm, linear-lanceolate 209. 209. Leaves 1.0–2.5 cm wide; involucral bracts erect in bud 220. subcyaneum 209. Leaves 1.5–4.0 cm wide; involucral bracts spreading 224. cyathis outwards at the tip in bud 211. 210. Involucral bracts with few stellate hairs 210. Involucral bracts with numerous to dense stellate hairs 215. 211. vagense 211. Leaves glaucous, deeply incise-dentate 211. Leaves green or glaucous, dentate but not 212. incise-dentate 212. Leaves mainly oblong, oblong-lanceolate or 259. stenolepiforme oblong-elliptical 213. 212. Leaves mostly ovate, elliptical or lanceolate 237. bettyhillense 213. Involucral bracts acute at apex 214. 213. Involucral bracts obtuse at apex 202. boswellii 214. Leaves bluish-green often tinted purplish 232. nitidum 214. Leaves bright green 215. Leaves mostly oblong or oblong-lanceolate, deeply 212. cambricum laciniate-dentate 215. Leaves mostly ovate or elliptical, dentate but not 216. laciniate-dentate 216. Leaves pale to medium rather bright green, speckled 225. arvonense with brownish-purple 217. 216. Leaves bluish-green, sometimes spotted 217. Leaves ovate, lanceolate or elliptical, cuneate at base 218. 217. Leaves broadly ovate or elliptical, rounded or truncate 219. at base 218. Involucral bracts 1.0–1.5 mm wide, linear-lanceolate, 218. pseudoleyi gradually narrowed to a subobtuse apex 218. Involucral bracts 0.8–1.0 mm wide, narrow linear-lanceolate, gradually narrowed to an acute apex 229. leyi 253. britannicum 219. Leaves 3–11 × 2–7 cm 257. naviense 219. Leaves 2–5 × 1.5–3.0 cm 221. 220. Involucral bracts with few stellate hairs 220. Involucral bracts with numerous to dense stellate hairs 232. 222. 221. Involucral bracts with few glandular hairs 223. 221. Involucral bracts with numerous glandular hairs 222. Leaves lanceolate, oblong or narrowly elliptical 205. angustatum

39. Hieracium 222. Leaves broadly elliptical, lanceolate or ovate 209. argentiforme 224. 223. Leaves spotted and blotched 225. 223. Leaves not spotted and blotched 213. amydrostictum 224. Leaves cuneate at base 244. saxorum 224. Leaves rounded or subtruncate at base 226. 225. Involucral bracts obtuse at apex 228. 225. Involucral bracts mostly acute at apex 208. argentatum 226. Leaves glaucous 227. 226. Leaves bright green 227. Leaves 3–6 cm wide, very broadly elliptical or broadly 196. hesperium ovate 227. Leaves 1.2–3.5 cm wide, elliptical, ovate or lanceolate 232. nitidum 229. 228. Leaves mostly cuneate at base 230. 228. Leaves rounded or subtruncate at base 229. Leaves linear, linear-lanceolate or narrowly elliptical 210. argenteum 229. Leaves elliptical to broadly elliptical, ovate or 245. dalense lanceolate 242. ebudicum 230. Leaves ovate to elliptical or subrotund 230. Leaves oblong, lanceolate-oblong, elliptical or 231. narrowly lanceolate 226. sublasiophyllum 231. Leaves 2–11 × 1.5–4.0 cm 228. peroblongum 231. Leaves 2–8 × 1.0–2.0 cm 233. 232. Involucral bracts with numerous glandular hairs 239. 232. Involucral bracts with few glandular hairs 239. hypochaeroides 233. Leaves much spotted and blotched 234. 233. Leaves unspotted or with an occasional spot 234. Involucral bracts with few simple eglandular hairs 249. breconense 234. Involucral bracts with numerous simple eglandular 235. hairs 236. 235. Ligule tips glabrous 237. 235. Ligule tips hairy 198. holophyllum 236. Leaves mostly cuneate-based 199. caledonicum 236. Leaves mostly subtruncate-based 237. Leaves glabrous or nearly so on upper surface 227. crinellum 238. 237. Leaves hairy on upper surface 238. Leaves 4.5–11.0 × 2.5–5.5 cm, ovate, elliptical or 197. dentulum subrotund 238. Leaves 2.5–8.0 × 1.5–3.5 cm, elliptical, ovate, 241. riddelsdellii ovate-lanceolate to elliptic-oblong 240. 239. Leaves spotted and blotched 242. 239. Leaves not spotted or blotched 215. sommerfeltii 240. Involucral bracts acute at apex 241. 240. Involucral bracts obtuse at apex 241. Leaves 2–6 × 1–2 cm, bluish-green, lanceolate or 217. basicrinum elliptical 241. Leaves 4–14 × 2.5–6.0 cm, medium yellowish-green, 238. stictum ovate, elliptical or lanceolate 243. 242. Involucral bracts obtuse at apex 246. 242. Involucral bracts acute at apex 243. Leaves broadly triangular-ovate or subrotund; ligules 203. leyanum usually poorly developed 243. Not with characteristic shape of last; ligules well 244. developed

231

244. Leaves 6–13 × 3–6 cm, cuneate at base, the cauline usually 2, large; involucral bracts 1.2–1.5 mm wide 195. subrubicundum 244. Leaves 3–11 × 2–5 cm, some rounded at base, the cauline 0–2, small; involucral bracts 1.5–1.7 mm 245. wide 198. holophyllum 245. Leaves cuneate at base 245. At least some leaves rounded at base 201. rubicundiforme 247. 246. Leaves obviously hairy on upper surface 248. 246. Leaves glabrous or nearly so on upper surface 235. brigantum 247. Involucral bracts up to 15 mm 236. schmidtii 247. Involucral bracts less than 12 mm 240. repandulare 248. Ligules very hairy at tip 249. 248. Ligules glabrous at tip 249. Leaves pale green, usually narrowly elliptical to 231. carenorum elliptical 250. 249. Leaves bluish-green, broader, at least some ovate 250. Stem with numerous glandular hairs in the upper part 246. subplanifolium 250. Stem with very few or no glandular hairs in the upper 250. dicella part

Section Vulgata and Sections Tridentata and Alpestria with few leaves 251. 251. 252. 252. 253. 253. 254. 254. 255. 255. 256. 256. 257. 257. 258. 258. 259. 259. 260. 260. 261. 261. 262. 262. 262a. 262a.

252. Cauline leaves more than 5 289. Cauline leaves 5 or less 253. Involucral bracts with few or no glandular hairs 263. Involucral bracts with numerous glandular hairs 254. Cauline leaves more or less amplexicaul at base 257. Cauline leaves not amplexicaul at base 68. solum Involucral bracts 11–13 mm 255. Involucral bracts 9–11 mm 56. subtruncatum Styles yellow 256. Styles discoloured 52. mirandum Leaves cuneate or rounded at base 65. difficile Leaves gradually narrowed at base 258. Involucral bracts with numerous stellate hairs 260. Involucral bracts with no or few stellate hairs 45. sparsifrons Styles yellow 259. Styles discoloured Leaves not spotted 263(3). vulgatum forma pseudosubramosum Leaves lightly spotted brownish-purple 267. caesionigrescens 36. charitodon Leaves laciniate with slender teeth Leaves subentire or dentate, but not laciniate with 261. slender teeth Involucral bracts with no or occasional simple 40. lissolepium eglandular hairs Involucral bracts with numerous simple eglandular 262. hairs Leaves 4–11 cm; involucral bracts more or less acute 37. gothicoides at apex Leaves 7–14 cm; involucral bracts more or less obtuse 262a. at apex Involucral bracts 1.5–2.0 mm wide 34. placerophylloides Involucral bracts 0.8–1.2 mm wide 41. subintegrifolium

232

1 4 8 . A S T E R AC E A E

263. Involucral bracts with numerous simple eglandular 264. hairs 263. Involucral bracts with no or few simple eglandular 274. hairs 265. 264. Cauline leaves more or less amplexicaul 269. 264. Cauline leaves not amplexicaul 58. pugsleyi 265. Styles yellow 266. 265. Styles discoloured 267. 266. Involucral bracts with dense stellate hairs 268. 266. Involucral bracts with few or no stellate hairs 50. carpathicum 267. Involucral bracts 10–13 mm 51. dewari 267. Involucral bracts 9–10 mm 268. Leaves sharply and irregularly dentate with the teeth 63. dovrense ascending-mammiform 64. australius 268. Leaves subentire or sharply denticulate 269. Involucral bracts more or less acute at apex 263(3). vulgatum forma pseudosubramosum 270. 269. Involucral bracts obtuse at apex 282. perscissum 270. Leaves deeply incise-dentate or lobate 271. 270. Leaves entire to dentate but not incise-dentate 271. Leaves 3–9 × 1–4 cm, elliptical, ovate or subrotund; 32. scullyi styles discoloured 271. Leaves 3–8 × 1.0–3.0 cm, elliptical, elliptic-lanceolate, elliptic-oblong, oblong or lanceolate; styles more or 272. less yellow 272. Involucral bracts 1.5–2.0 mm wide; receptacle pits 39. uiginskyense fimbriate-dentate 272. Involucral bracts 1.0–1.5 mm wide; receptacle pits 273. subulate-dentate or serrate-dentate 273. Leaves usually spotted and blotched, the petiole short; 42. sparsifolium achenes 3.0–3.5 mm 273. Leaves never spotted, the petiole long; achenes 46. hibernicum 3.5–4.0 mm 274. Involucral bracts with numerous to dense stellate hairs 275. 281. 274. Involucral bracts without or with few stellate hairs 276. 275. Involucral bracts obtuse at apex 278. 275. Involucral bracts acute at apex 44. linguans 276. Cauline leaves not amplexicaul 277. 276. Cauline leaves more or less amplexicaul 49. borreri 277. Leaves 2.5–6.0 cm wide 61. hethlandiae 277. Leaves 0.5–2.0 cm wide 278. Many leaves 3–4 times as long as broad 263(3). vulgatum forma pseudosubramosum 279. 278. Leaves not more than 3 times as long as wide 279. Leaves denticulate to shortly dentate; involucral bracts 310. argillaceum up to 13 mm 279. Leaves dentate to incise-dentate; involucral bracts less 280. than 11 mm 280. Basal leaves 6–15 × 2–6 cm, mostly oblong-elliptical, sometimes lanceolate to ovate or oblong-lanceolate, dentate to incise-dentate, cusped and up to 15 mm; the upper cauline deeply incise-dentate with narrow, 307. acuminatum curved teeth up to 20 mm 280. Basal leaves 4–12 × 2–5 cm, lanceolate to ovate or elliptic-ovate, more or less dentate, the teeth often cusped and up to 6 mm; the cauline lanceolate to linear, dentate to incise-dentate with teeth narrowly 308. consociatum mammiform and up to 10 mm 282. 281. Cauline leaves more or less amplexicaul

285. 281. Cauline leaves not amplexicaul 282. Leaves more or less glabrous on upper surface 62. praethulense 283. 282. Leaves hairy on upper surface 283. Hairs on upper surface of leaves bulbous-based 69. breve 284. 283. Hairs on upper surface of leaves not bulbous-based 284. Basal leaves present at flowering; capitula 25–35 mm 59. spenceanum in diameter; styles discoloured 284. Basal leaves absent at flowering; capitula 35–50 mm in 60. attenuatifolium diameter; styles yellow 285. Involucral bracts 1.5–2.0 mm wide 31. aphyllopodioides 286. 285. Involucral bracts 0.8–1.5 mm wide 286. Glandular hairs on involucral bracts slender and 287. spidery 286. Glandular hairs on involucral bracts short and stiff or 288. minute 287. Involucral bracts 1.0–1.5 mm wide, linear-lanceolate, 33. scabrisetum incumbent in bud 287. Involucral bracts 1.0–1.2 mm, narrowly linear-oblong, 302. subminutidens more or less porrect in bud 288. Leaves bluish-green, all more or less acute at apex 43. nidense 288. Leaves yellowish-green, at least the lower obtuse at 317. festinum apex 290. 289. Cauline leaves 2–5 358. 289. Cauline leaves 0–1, rarely 2 290. Involucral bracts with numerous simple eglandular 291. hairs 290. Involucral bracts with few or no simple eglandular hairs 323. 291. Involucral bracts with numerous to dense stellate hairs 292. 307. 291. Involucral bracts with few or no stellate hairs 293. 292. Involucral bracts with numerous glandular hairs 295. 292. Involucral bracts with few or no glandular hairs 293. Leaves coarsely and sharply dentate; ligules 276. fulvocaesium orange-yellow and very short 294. 293. Leaves dentate; ligules yellow and normal length 269. peccense 294. Involucral bracts acute at apex 284. lepidulum 294. Involucral bracts obtuse at apex 296. 295. Involucral bracts obtuse at apex 299. 295. Involucral bracts acute at apex 296. Involucral bracts with dense simple eglandular hairs up 264. cravoniense to 2 mm 296. Involucral bracts with less dense, shorter simple 297. eglandular hairs 267. caesionigrescens 297. Leaves lightly spotted 298. 297. Leaves not spotted 273. aequiserratum 298. Leaves equally sinuate-serrate 298. Leaves irregularly and often deeply serrate 279. rhomboides 299. Leaves with numerous stellate hairs beneath and 265. coniops sometimes also above 300. 299. Leaves with few or no stellate hairs 300. Leaves pale green; involucral bracts dull pale green 266. acroleucum 300. Leaves dark green; involucral bracts dark or 301. blackish-green 301. Leaves with numerous stiff hairs on upper surface, 108. lingulatum entire; involucral bracts blackish-green

39. Hieracium 301. Leaves glabrous or with few soft hairs on upper 302. surface; involucral bracts dark or greyish-green 302. Involucral bracts 1.0–1.5 mm, dark green 280. subramosum 303. 302. Involucral bracts 1.0–1.2 mm, greyish-green 303. Basal leaves few, 2.5–5.0 cm wide; cauline 2–4, the lower as large as basal with spreading mammiform 263(5). vulgatum forma sejunctum teeth 303. Basal leaves 1.0–3.0 cm side; cauline 2–8, often 304. smaller than basal, the teeth often sharper 304. Cauline leaves 4–8 263(3). vulgatum forma pseudosubramosum 305. 304. Cauline leaves 2–4 305. Basal leaves few, 4–10 × 2.0–2.5 cm 263(1). vulgatum forma vulgatum 306. 305. Basal leaves numerous forming a spreading rosette 306. Leaves irregularly serrate-dentate; involucral bracts up 263(2). vulgatum forma subfasciculare to 12 mm 306. Leaves mammiforme-dentate; involucral bracts up to 263(4). vulgatum forma subravusculum 10 mm 308. 307. Involucral bracts with numerous glandular hairs 316. 307. Involucral bracts with few glandular hairs 309. 308. Involucral bracts acute at apex 312. 308. Involucral bracts obtuse at apex 310. 309. Leaves sinuate-dentate or incise-dentate 311. 309. Leaves denticulate with an occasional large tooth 277. amnicola 310. Leaves densely hairy on upper surface 278. stenophyes 310. Leaves with few hairs on upper surface 311. Leaves 3.5–6.5 × 1.5–3.5 cm, incise-dentate towards the cuneate base; involucral bracts up to 9 × 0.8–1.0 285. subviolascens mm 311. Leaves 5–12 × 3–6 cm, sinuate-dentate with shallow mammiform teeth, rounded or truncate at base; involucral bracts up to 11 × 1.0–1.2 mm 295. subamplifolium 313. 312. Leaves denticulate to shallowly dentate 314. 312. Leaves incise-dentate with mammiform teeth 271. robertsii 313. Leaves glabrous on upper surface 313. Leaves more or less hairy on upper surface 305. megapodium 74. perscitum 314. Leaves 0.5–4.0 × 0.8–2.5 cm 315. 314. Leaves 5–12 × 2–5 cm 315. Leaves with teeth long and narrow; involucral bracts 0.8–1.2 mm wide, narrow linear-lanceolate 282. perscissum 315. Leaves irregularly-dentate, the teeth fairly large and mammiform; involucral bracts 1.0–1.5 wide, 283. subirriguum linear-lanceolate 317. 316. Leaves faintly spotted and blotched 318. 316. Leaves not spotted or blotched 66. amaurostictum 317. Leaves entire or denticulate 317. Leaves denticulate to dentate, the teeth sometimes large 268. pollichiae 262. triviale 318. Involucral bracts acute at apex 319. 318. Involucral bracts obtuse at apex 319. Peduncles with numerous glandular hairs; involucral 274. vinifolium bracts 0.8–1.0 mm wide 319. Peduncles with few or no glandular hairs; involucral 320. bracts 1.0–1.7 mm wide

233

321. 320. At least some of the leaves acute at apex 322. 320. Leaves more or less obtuse at apex 321. Leaves greyish-green, narrowly to broadly elliptical or 270. caesiopilosum oblong-elliptical; styles discoloured 321. Leaves dull medium green, often suffused reddish, 275. rubiginosum lanceolate or ovate; styles yellow 322. Leaves glabrous or nearly so on upper surface 272. kintyricum 286. surrejanum 322. Leaves hairy on upper surface 324. 323. Involucral bracts with numerous stellate hairs 335. 323. Involucral bracts with few or no stellate hairs 325. 324. Leaves spotted and blotched with brownish-purple 324. Leaves not spotted and blotched but sometimes washed 328. with purple 326. 325. Involucral bracts obtuse at apex 327. 325. Involucral bracts acute at apex 326. Peduncles without or with a rare glandular hair; styles 292. asperatum discoloured 326. Peduncles with numerous glandular hairs; styles yellow 293. fictum 327. Glandular hairs of involucral bracts and peduncles very 291. naevuliferum short and not very numerous 327. Glandular hairs of involucral bracts and peduncles 294. spilophaeum larger (short) and more numerous 329. 328. Involucral bracts obtuse at apex 330. 328. Involucral bracts acute at apex 329. Leaves 4–11 × 2–6 cm; glands on involucral bracts 309. radyrense short to medium 329. Leaves 4–8 × 2–4 cm; glands on involucral bracts very 314. latebrosum short to short 330. Leaves with small sharp teeth, or denticulate, or entire 331. 332. 330. Leaves with teeth more or less mammiform 313. aviicola 331. Leaves serrate-dentate 316. lortetiae 331. Leaves entire to denticulate 332. Leaves incise-dentate, the teeth up to 15 mm, cusped 307. acuminatum and sometimes free on the petiole 332. Leaves dentate, but teeth not long, though sometimes 333. cusped 312. nemophilum 333. Leaves 2.5–6.0 × 1.5–2.0 cm 334. 333. Leaves 5–12 × 2–5 cm 334. Leaves mainly ovate or elliptical-ovate; involucral 308. consociatum bracts up to 10 mm 334. Leaves mainly elliptical or oblong-elliptical; involucral 310. argillaceum bracts up to 13 mm 336. 335. At least the uppermost leaves semiamplexicaul 338. 335. Leaves not amplexicaul 336. Involucral bracts with numerous glandular hairs 70. zetlandicum 337. 336. Involucral bracts with few glandular hairs 67. gratum 337. Involucral bracts 9–11 × 1.5 mm 68. solum 337. Involucral bracts 11–13 × 1.5–3.0 mm 339. 338. Involucral bracts obtuse at apex 349. 338. Involucral bracts acute at apex 339. Glandular hairs of involucral bracts short and very 340. short, without medium glandular hairs 339. At least some of the glandular hairs of the involucral 344. bracts medium in length

234

1 4 8 . A S T E R AC E A E

298. diaphanum 340. Leaves 1.5–2.5 cm wide 341. 340. Leaves 2–4 cm wide 341. At least some of the leaves ovate, the basal in a rosette and the cauline rapidly getting smaller upwards 297. submutabile

357. Leaves usually pale to medium green, denticulate to dentate; involucral bracts olive green; styles often 318. cheriense yellow 357. Leaves medium green often suffused purple, mostly 319. rectulum entire or denticulate; styles discoloured

341. Leaves oblong, elliptic-oblong or lanceolate-oblong, the basal few and the cauline large and only slowly 342. decreasing in size 296. mammidens 342. Leaf teeth large and mammiform 343. 342. Leaf teeth small and not mammiform

358. Involucral bracts with no or few simple eglandular hairs

343. Leaves long attenuate at the base with long petioles, 305. megapodium undulate-dentate; styles yellow 343. Leaves cuneate at base to a short petiole, more sharply 317. festinum toothed; styles discoloured 281. praesigne 344. Leaves glabrous on upper surface 345. 344. Leaves more or less hairy on upper surface 345. Involucral bracts with solid black glandular hairs 315. pulchrius 345. Involucral bracts with thin, spidery, black glandular 346. hairs 290. inquinatum 346. Leaves entire to denticulate 347. 346. Leaves dentate 300. lepidiceps 347. Leaves mainly ovate 347. Leaves mainly lanceolate, elliptical, oblong-elliptical, 348. oblong or elliptical-lanceolate 348. Leaves denticulate to incise-dentate, the teeth narrow; peduncles with numerous glandular hairs 302. subminutidens 348. Leaves denticulate to dentate with small, sharp to large 304. diaphanoides mammiform teeth 349. Some of the glandular hairs on the involucral bracts 350. medium in length 349. Glandular hairs on involucral bracts all short or very 353. short 350. Leaves narrowly elliptical, oblong-elliptical or 301. scanicum ovate-oblong, incise 351. 350. Leaves not like that 351. Leaves often suffused reddish; glands of involucral 287. erubescens bracts thin but stiff 351. Leaves often green; glands of involucral bracts thin and 352. spidery 352. Leaves 2–7 × 1–3 cm, subentire or with a few sharp 299. daedalolepioides teeth 352. Leaves 3–10 × 1.5–4.5 cm, dentate to deeply 303. anglorum incise-dentate 353. Leaves glabrous on upper surface or with hairs only 289. glanduliceps near the margin 354. 353. Leaves hairy on upper surface 288. commixtum 354. Leaves spotted, 4–5 × 1.0–1.5 cm 355. 354. Leaves not spotted, 2–11 × 1–5 cm 355. Leaves 5–11 × 2–5 cm, cauline leaves as large as basal, 311. chlorophyllum subentire to shortly dentate 356. 355. Leaves not with this combination of characters 356. Basal leaves narrowly elliptical or ovate, subentire or remotely denticulate; lowest cauline leaf low on the 306. lagganense stem, the upper small or bract-like 357. 356. Not so

Sections Hieracium and Subalpina 359. 358. Involucral bracts with numerous simple eglandular 413. hairs 359. Involucral bracts with numerous to dense stellate 360. hairs 386. 359. Involucral bracts with few or no stellate hairs 361. 360. Involucral bracts obtuse at apex 369. 360. Involucral bracts acute at apex 361. Inflorescence aggregated like an umbel 320. aggregatum 362. 361. Inflorescence not so 363. 362. Leaves glabrous or nearly so on upper surface 366. 362. Leaves hairy on upper surface 380. candelabrae 363. Leaves with mammiform teeth 364. 363. Leaves entire to dentate but teeth not mammiform 364. Leaves with dense stellate hairs beneath and 152. hyparcticoides sometimes also on surface 365. 364. Leaves with occasional or no stellate hairs 395. cardiophyllum 365. Leaves cordate or truncate at base 410. camptopetalum 365. Leaves cuneate or rounded at base 370. cuneifrons 366. Leaves cuneate at base 366. Leaves mainly rounded, truncate or subcordate at base 367. 367. Leaves 2–9 × 1–4 cm; ligules hairy at tip at least when 404. pruinale young 368. 367. Leaves 2–14 × 2–7 cm; ligules glabrous 368. Leaves 2–14 cm, at least some oblong-elliptical, the 386. stenstroemii teeth large and mammiform 368. Leaves 3–10 cm, ovate or elliptical and at least one usually large and subrotund, the teeth mammiform or 388. dowardense sharply mammiform 370. 369. Longest involucral bracts more than 11 mm 375. 369. Longest involucral bracts not more than 11 mm 370. Leaves hairy on upper surface; ligules with hairs at 371. apex 370. Leaves glabrous or nearly so on upper surface; ligules 372. glabrous at apex 371. Basal leaves mostly obtuse; involucral bracts 381. subcrassum blackish-green, incumbent in bud 371. Basal leaves mostly acute at apex; involucral bracts 382. cinderella rather pale green, porrect in bud 373. 372. Styles yellow 374. 372. Styles discoloured 373. Leaves usually spotted and blotched, 1–6 × 1–3 cm 136. lakelandicum 373. Leaves never spotted, 2–11 × 1.5–7.0 cm 137. chrysolorum 355. prolixum 374. Glands of involucral bracts 0.2–1.0 mm 374. Glands of involucral bracts 0.2–1.5 mm 380. candelabrae 375. At least the cauline leaves with numerous to dense 376. stellate hairs beneath 377. 375. Leaves without or with occasional stellate hairs 385. kentii 376. Leaves hairy on upper surface

39. Hieracium 376. Leaves glabrous or nearly so on upper surface 412. asteridiophyllum 377. Leaves with large mammiform or narrowly 378. mammiform teeth 377. Leaves not with mammiform teeth or rarely with small 380. mammiform teeth 378. Leaves spotted and blotched; styles discoloured 390. monstrosum 379. 378. Leaves not spotted or blotched; styles yellow 379. Leaves lanceolate, ovate or ovate-oblong, teeth 375. adelphicum mammiform 379. Leaves more elongate, teeth narrower mammiform and 384. aterrimum more pointed 381. 380. Leaves subentire or denticulate 383. 380. Leaves shallowly to fairly deeply dentate 381. Glandular hairs of involucral bracts very short, up to 407. hjeltii 0.6 mm 382. 381. Glandular hairs of involucral bracts up to 1 mm 398. microspilum 382. Leaves rounded or cordate at base 382. Leaves subtruncate or shortly cuneate at base 411. snowdoniense 383. Leaves glabrous or nearly so on upper surface 408. patale 384. 383. Leaves hairy on upper surface 391. medium 384. Leaves serrate with sharp teeth 385. 384. Leaves dentate with more or less blunt teeth 394. sublepistoides 385. Leaves ovate, lanceolate or elliptical 397. sylvivagum 385. Leaves mostly broadly triangular-ovate 386. Involucral bracts usually with an occasional or few 387. simple eglandular hairs 393. 386. Involucral bracts without simple eglandular hairs 388. 387. At least some of the leaves subtruncate at base 390. 387. Leaves cuneate or abruptly contracted at base 123. nigrifactum 388. Leaves not spotted 389. 388. Leaves spotted and blotched 135. clovense 389. Leaves 1.5–4.0 × 1.5–2.5 cm 369. liljeholmii 389. Leaves 5–9 × 2–4 cm 390. Leaves irregularly dentate or incise-dentate 128. oxyodus 391. 390. Leaves subentire to shallowly dentate 124. gracilifolium 391. Leaves markedly attenuate at base 392. 391. Leaves cuneate or abruptly contracted at base 392. Leaves 4–9 × 2–4 cm with shallow, mammiform teeth 125. atriglandulosum 392. Leaves 1.5–5.0 × 1.5–3.0 cm, denticulate to sharply 371. uistense dentate 394. 393. Leaves with large mammiform teeth 401. 393. Leaves entire or with small teeth 395. 394. Leaves ovate or elliptic-ovate 394. At least the inner leaves oblong, elliptic-oblong or 396. lanceolate-oblong 395. Involucral bracts 4–12 × 1.0–1.2 mm; styles 374. sylvularum discoloured 395. Involucral bracts 4–10 × 0.8–1.0 mm; styles yellow 383. koehleri when fresh 396. Leaves small, 1.5–8.0 × 1.0–3.5 cm, teeth small 405. gentile 397. 396. Leaves large, 3–16 × 2.5–7.0 cm, teeth large

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397. Leaves with teeth small at top getting longer towards 398. the base 397. Leaves with teeth large and very unequal mostly 399. throughout 398. Glandular hairs of involucral bracts 0.2–0.7 mm 376. firmiramum 398. Glandular hairs of involucral bracts 0.2–1.1 mm 377. onychodontum 399. Leaves very large, 9–16 × 4–7 cm, teeth very large 379. seriflorum 400. 399. Leaves smaller 3–11 × 2.5–4.0 cm 373. grandidens 400. Leaf teeth numerous and close together 378. quadridentatum 400. Leaf teeth large and few 402. 401. Styles discoloured 408. 401. Styles yellow 403. 402. At least some leaves oblong or oblong-lanceolate 405. 402. None of the leaves oblong or oblong-lanceolate 403. Glandular hairs on involucral bracts 0.2–1.5 mm 393. subaequialtum 404. 403. Glandular hairs on involucral bracts 0.2–1.0 mm 372. ronasii 404. Glands on involucral bracts 0.2–0.5 mm 399. oblongum 404. Glands on involucral bracts 0.3–1.0 mm 405. Leaves more or less hairy on upper surface 406. itunense 406. 405. Leaves more or less glabrous on upper surface 406. Involucral bracts 1.5–2.0 mm wide, with glandular 409. pollinarium hairs 0.2–0.5 mm 406. Involucral bracts 0.8–1.2 mm wide, with glandular 407. hairs 0.4–1.0 mm 407. Involucral bracts greyish-green, with dark glandular 400. integratum hairs 407. Involucral bracts almost black with black glandular 401. pellucidum hairs 396. severiceps 408. Leaves sharply serrate-dentate 409. 408. Leaves not sharply serrate-denate 409. Leaves 2.5–6.0 × 1.5–2.5 cm; glandular hairs of 392. hortense involucral bracts 0.2–0.5 mm 409. Leaves 2–14 × 2–5 cm; glandular hairs of involucral 410. bracts 0.2–1.7 mm 389. glevense 410. Involucral bracts up to 13 mm 411. 410. Involucral bracts not more than 11 mm 387. elevatum 411. Leaves mostly cuneate at base 412. 411. Leaves rounded or truncate at base 412. Involucral bracts narrow linear-lanceolate, 0.8–1.0 mm 402. exotericum wide 412. Involucral bracts linear-lanceolate, 1.0–1.2 mm wide 403. neosparsum 414. 413. Involucral bracts with few glandular hairs 444. 413. Involucral bracts with numerous glandular hairs 414. Involucral bracts with numerous to dense stellate hairs 415. particularly on the margins 426. 414. Involucral bracts with no or few stellate hairs 416. 415. Involucral bracts acute at apex 421. 415. Involucral bracts obtuse at apex 416. Leaves hairy on upper surface; involucral bracts 328. inspissatum 5–14 × 1.5–1.7 mm 416. Leaves glabrous or nearly so on upper surface; 417. involucral bracts 4–12 × 1.0–1.5 mm

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1 4 8 . A S T E R AC E A E 418. Tips of ligules very shortly hairy 420. Tips of ligules glabrous 142. oenophyllum Styles yellow 419. Styles discoloured Peduncles with few to numerous simple eglandular 140. petrocharis hairs and few glandular hairs

436. 437. 437. 438.

419. Peduncles with numerous glandular hairs and 144. glandulidens occasional simple eglandular hairs 420. Peduncles with a few short to medium simple eglandular hairs and occasionally a short glandular hair 150. arranense

439. 439. 440. 440. 441. 441. 442. 442.

417. 417. 418. 418. 419.

420. Peduncles with numerous short glandular hairs and 337. silvaticoides rarely a simple eglandular hair 422. 421. Leaves hairy on upper surface 424. 421. Leaves glabrous or nearly so on upper surface 338. breadalbanense 422. Styles yellow 423. 422. Styles discoloured 328. inspissatum 423. Involucral bracts 5–14 × 1.5–1.7 mm 331. oistophyllum 423. Involucral bracts 3.0–8.5 × 1.0–1.2 424. Leaves oblanceolate, oblong-oblanceolate or narrowly 118. anfractiforme elliptical 425. 424. At least some leaves ovate or elliptical 425. Leaves denticulate to incise-dentate, the teeth mostly mammiform or narrowly mammiform 321. caesiomurorum 425. Leaves regularly sinuate-denticulate or sinuate-dentate 341. crebridens 427. 426. Involucral bracts acute at apex 434. 426. Involucral bracts obtuse at apex 330. neocoracinum 427. Ligules glabrous at tip 428. 427. Ligules hairy at tip 138. aristidens 428. Leaves with sharply spinulose teeth 429. 428. Leaves with large teeth but not spinulose 430. 429. Involucral bracts 5–14 × 1.0–1.5 mm 432. 429. Involucral bracts 4–10 × 0.6–1.5 mm 430. Leaves denticulate or with an occasional tooth 151. thalassinum 431. 430. Leaves deeply dentate 431. Inner basal leaves mostly oblong or oblong-elliptical, with jaggedly elongate-mammiform teeth, sometimes 109. cremnanthes extending on petiole 431. Inner basal leaves narrowly to broadly elliptical, lanceolate or ovate-lanceolate, with narrow irregular 115. molybdochroum teeth 357. caesitium 432. Involucral bracts 4–8 × 0.6–1.0 mm 433. 432. Involucral bracts 4–10 × 0.8–1.5 mm 433. Peduncles with few simple eglandular hairs and few glandular hairs; involucral bracts blackish-green 134. diversidens 433. Peduncles with numerous simple eglandular hairs and few glandular hairs; involucral bracts greyish-green 329. subhirtum 435. 434. Leaves glabrous on upper surface 437. 434. Leaves hairy on upper surface 111. callistophyllum 435. Involucral bracts up to 13 mm 436. 435. Involucral bracts not more than 11 mm 323. armadalense 436. Styles discoloured

438.

443. 443. 444. 444. 445. 445. 446. 446. 447. 447. 448. 448. 449. 449. 450. 450. 451. 451. 452. 452. 453. 453. 454. 454. 455. 455. 456. 456. 457. 457. 458. 458. 459. 459.

335. uisticola Styles pure yellow 438. Involucral bracts up to 15 mm 439. Involucral bracts not more than 12 mm Leaves 2.5–7.0 × 1–3 cm, abruptly contracted or 112. hastiforme subtruncate at base Leaves 3–10 × 2–3 cm, rounded to cuneate at base 117. marshallii 440. At least some leaves subtruncate at base 441. Leaves cuneate to attenuate at base 139. dasythrix Involucral bracts not more than 10 mm 333. orithales Involucral bracts up to 12 mm 361. protentum Leaves with long cusped teeth below 442. Leaves not with cusped teeth below 278. stenophyes Leaves with few hairs on upper surface Leaves with numerous, subrigid hairs on upper surface 443. Leaves 6–12 × 2.5–4.5 cm, mostly elliptical, or 324. clivicola ovate-elliptical Leaves 3–8 × 2–4 cm, elliptical, ovate, lanceolate or 325. orcadense oblanceolate 445. Involucral bracts with few or no stellate hairs Involucral bracts with numerous to dense stellate hairs 468. 446. Leaves usually spotted and blotched 450. Leaves not spotted and blotched Involucral bracts not more than 10 mm 365. scotostictum 447. Involucral bracts more than 10 mm 448. Leaves hairy on upper surface 449. Leaves glabrous or nearly so on upper surface 332. lintonii Leaves with large teeth 334. maculoides Leaves with small teeth 353. varifolium Leaves glabrous on upper surface 368. gougetanum Leaves hairy on upper surface Leaves with numerous to dense stellate hairs on both 153. pseudanglicoides surfaces 451. Leaves without stellate hairs or with a few beneath 452. Styles yellow 455. Styles discoloured Leaves glabrous on upper surface; involucral bracts 354. anguinum more than 11 mm Leaves hairy on upper surface; involucral bracts less 453. than 11 mm 133. raveniorum Leaves with small sharp teeth 454. Leaves subentire or with blunt teeth Inner leaves elliptical, ovate or lanceolate 352. pauculidens Inner leaves elliptical to oblong-elliptical 362. micracladium 456. Involucral bracts obtuse at apex 460. Involucral bracts acute at apex Leaves dentate with mammiform teeth 132. melanochloricephalum 457. Leaves subentire to denticulate or with sharp teeth 347. mariae Leaves rounded or subtruncate at base 458. Leaves cuneate to attenuate at base 147. fissuricola Involucral bracts 12 mm or less 459. Involucral bracts more than 12 mm 113. isabellae Leaves hairy on upper surface 131. ochthophilum Leaves glabrous on upper surface

39. Hieracium 460. 460. 461. 461. 462. 462. 463. 463. 464. 464. 465. 465. 466. 466. 467. 467.

468. 468. 469. 469. 470. 470. 471. 471. 472. 472. 473. 473. 474. 474. 475. 475. 476. 476. 477. 477. 478. 478. 479. 479. 480. 480. 481. 481. 482.

461. Leaves abruptly contracted or subtruncate at base 463. Leaves cuneate to attenuate at base 342. crebridentiforme Leaves hairy on upper surface 462. Leaves glabrous on upper surface 145. sannoxense Ligules with short hairs at tip 356. mucronellum Ligules glabrous at tip Leaves deeply dentate with mammiform teeth 110. sinuans 464. Leaves entire to dentate but not deeply dentate Leaves sharply serrate; peduncles with numerous 126. dissimile glandular hairs and no eglandular hairs Leaves not sharply serrate; peduncles with simple 465. eglandular hairs as well as glandular hairs 120. centripetale Leaves mainly oblong-lanceolate 466. Leaves mainly ovate or elliptical 130. westii Capitula small and narrow 467. Capitula large and broad Glandular hairs of involucral bracts and peduncles 116. laetificum strong and black and obvious Glandular hairs of involucral bracts weaker and slender, and more obscured by eglandular hairs 141. pseudanglicum 469. Involucral bracts obtuse at apex 483. Involucral bracts acute at apex 470. Leaves obviously hairy on upper surface 475. Leaves glabrous or nearly so on upper surface 471. At least some of the leaves cuneate at base 473. Leaves rounded, subtruncate or cordate at base Leaves narrowly elliptical, oblong-lanceolate or 327. dipteroides lanceolate 472. Leaves broader 122. kingshousense Leaves 4–12 × 2.5–4.0 cm 146. insulare Leaves 2–6 × 1.5–2.0 cm Leaves 2.5–14.0 × 4.0–8.5 cm, often subcordate at 339. auratiflorum base; styles yellow Leaves 2–11 × 1.5–6.0 cm, never cordate at base; 474. styles discoloured Leaves regularly sinuate-dentate with apiculate teeth, 336. sinuolatum not spotted Leaves subentire, denticulate or shallowly dentate, 358. piligerum often spotted 476. Leaves with large and irregular teeth 477. Leaves with small teeth 127. cuspidens Leaves cuneate at base 322. irregularidens Leaves subtruncate at base 129. vennicontium Leaves cuneate at base 478. Leaves rounded or subtruncate at base 149. ovaliforme Leaves 1.5–5.0 × 1–3 cm 479. At least some leaves larger At least some leaves oblong-elliptical or oblong-ovate 480. 481. Leaves all broader, mostly ovate or elliptical 345. pictorum Peduncles with scattered glandular hairs Peduncles with numerous glandular hairs 349. sanguineum Leaves triangular-ovate; involucral bracts up to 14 mm 340. triangularifolium Leaves elliptical or ovate; involucral bracts not more 482. than 12 mm Involucral bracts gradually narrowed at apex 351. pachyphylloides

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482. Involucral bracts abruptly narrowed at apex 364. discophyllum 484. 483. Leaves glabrous or nearly so on upper surface 490. 483. Leaves obviously hairy on upper surface 484. Involucral bracts not more than 9 mm 363. subprasinifolium 485. 484. Involucral bracts more than 9 mm 485. Involucral bracts very narrow, 0.8–1.0 mm wide 343. pseudosarcophyllum 486. 485. Involucral bracts more than 1.0 mm wide 486. Leaves broadly elliptical or broadly ovate with large 359. pollinarioides mammiform teeth 486. Leaves narrower, some usually oblong or 487. oblong-elliptical 488. 487. Involucral bracts more than 11 mm 489. 487. Involucral bracts 11 mm or less 488. Leaves with mammiform or aquiline-mammiform 121. breacense teeth, cuneate at base 488. Leaves subentire to sharply dentate, subtruncate at base 143. cumbriense 326. subtenue 489. Leaves entire or denticulate 346. elongatifolium 489. Leaves with some large teeth 490. Leaves with numerous to dense stellate hairs beneath 344. cymbifolium and sometimes also on upper surface 491. 490. Leaves without or with an occasional stellate hair 492. 491. Leaves cuneate or attenuate at base 494. 491. Leaves rounded or subtruncate at base 492. Inner leaves narrowly elliptical, oblong or 119. vorlichense oblong-lanceolate and attenuate at base 493. 492. Inner leaves broader and cuneate at base 114. senescens 493. Involucral bracts more than 14 mm 327. dipteroides 493. Involucral bracts 10 mm or less 494. Leaves regularly sinuate-dentate, the teeth apiculate 336. sinuolatum 494. Leaves entire to mammiform-dentate, but not 495. sinuate-dentate with apiculate teeth 148. eustales 495. Leaves 3–6 × 1.0–1.5 cm 496. 495. Leaves 2–15 × 1.0–5.5 cm 366. baliophyllum 496. Ligules glabrous at apex 497. 496. Ligules hairy at apex 498. 497. Styles yellow 499. 497. Styles discoloured 348. rivale 498. Leaves regularly denticulate or dentate 498. Leaves with large irregular mammiform teeth at the 367. zygophorum base 360. duriceps 499. Capitula narrow and subcylindrical 500. 499. Capitula broad and round 348. rivale 500. Leaves denticulate or shallowly dentate 500. Leaves with some large mammiform teeth 350. boreoanglicum

Section 1. Sabauda (Fr.) Arv.–Touv. Taxon Sabauda Fr.; Series Sabauda (Fr.) P. D. Sell & C. West comb. inval.; Subsection Hirsuta Scheele, non Section Hirsuta Rouy; Subsection Subsabauda Gremli Aphyllopodous, rarely pseudophyllopodous perennial herbs with branched stocks. Stems 30–120 cm, usually roughly hairy, often becoming scaberulous. Leaves 10–70,

238

1 4 8 . A S T E R AC E A E

more or less heterophyllous, gradually decreasing in size upwards, or some leaves grouped in a false rosette, or sometimes becoming abruptly smaller, the lower gradually narrowed at base and sometimes shortly petiolate, not amplexicaul, usually hairy or scabrous, nervation not prominent. Inflorescence a raceform panicle, or paniculate-corymbose; peduncles bracteolate, rarely with glandular hairs. Capitula medium to rather large. Involucral bracts incumbent in bud, appressed or the outer lax, rarely with more or less recurved tips, 4–14 × (0.8–)1.0–1.5 mm, dark and usually more or less concolorous, linear-lanceolate, obtuse at apex, at most with only a few stellate hairs. Ligules glabrous. Styles usually discoloured, rarely yellow. Receptacle pits with margins markedly fibrillous-dentate. Achenes purplish- or blackish-brown. Flowers (6–)7–10. The species of this section differ from those of Section Foliosa in their usually more hairy stems, non-amplexicaul leaves which are often more hairy and less conspicuously veined, often with more or less concolorous involucral bracts which often have characteristic glandular hairs and microglands, usually discoloured styles and markedly fibrillous-dentate margins to the receptacle pits. An easily recognised group widespread in the lowlands of Great Britain on heaths, in open woods and railway and roadside banks and waste places and extending into the uplands by streams, rocky slopes and cliff ledges and screes. In Ireland one species is frequent in the south-east and another has been recorded once further north. The section is widespread in western and central Europe extending to southern Russia and Turkey. Secondary growth after they have been cut or grazed makes the species difficult to key out, but they can still be recognised once the species become familiar. 1. H. vagum Jord. Glabrous-headed Hawkweed H. sabaudum var. latifolium Gaudin; H. boreale var. vagum (Jord.) Gren. & Godr.; H. boreale subsp. virgultorum microgen. vagum (Jord.) Sudre; H. boreale subsp. vagum (Jord.) Rouy; H. sabaudum subsp. vagum (Jord.) Schinz & R. Keller; H. boreale auct.; H. sublactucaceum auct.; H. eminens auct.; H. subrectum auct. Aphyllopodous perennial herb with a branched stock. Stem 30–100 cm, pale yellowish-green, sometimes suffused brownish-purple, erect, mostly robust but sometimes slender, striate, with numerous, sometimes dense below, medium to long, pale simple eglandular hairs throughout, often becoming glabrous or scabrous in the upper half. Leaves 10–40(–70), usually gradually decreasing in size upwards, sometimes with a group in a false rosette towards the base, usually deep green, sometimes paler on the upper surface, paler beneath, sometimes suffused reddish-purple, the lamina of the lower 6–10(–14) × 2.5–4.0(–5.0) cm, elliptical to ovate, obtuse-mucronulate to more or less acute at apex, remotely denticulate or dentate and gradually narrowed below to a short winged petiole or sessile, the lamina of the median mostly ovate or elliptic-ovate, sometimes elliptical or rarely lanceolate, acute at apex, remotely denticulate to dentate or dentate-serrate, sometimes the teeth curved, and cuneate to rounded at the sessile base, the lamina of the upper ovate or lanceolate, acute at apex, entire or

with a few, small teeth and rounded at the sessile base, all glabrous or nearly so, or with medium or long, pale simple eglandular hairs beneath, particularly on the midrib and margin or scaberulous, the upper sometimes with scattered stellate hairs beneath. Inflorescence with numerous capitula in a large corymbose panicle; peduncles straight and erect, with numerous to dense stellate hairs, sometimes a few, medium, pale simple eglandular hairs and occasional linear bracts. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.3 mm, blackish-green, more or less concolorous or inner with slightly paler margins, all rather broadly linear-lanceolate, obtuse at apex, characteristically glabrous, but sometimes with a few stellate and/or medium, pale simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured or yellow. Receptacle pits with margins fibrillous-dentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 7–10. Apomictic. 2n = 27. In its typical widespread form H. vagum is a tall plant with mostly ovate, sparsely denticulate or shortly dentate leaves which gradually decrease in size up the stem, with many capitula, glabrous involucral bracts and discoloured styles. In upland areas of Wales there is a tendency for them to be shorter, have fewer more dentate leaves and fewer capitula with a group of leaves forming a pseudorosette towards the base (H. calvatum), but such plants can be matched in most of the rest of the range of the species. Yellow-styled plants seem to show no other distinctive differences. When H. vagum grows with H. sabaudum intermediate plants are sometimes found, but both species and intermediates are apogamous triploids. A diploid H. vagum is known in Slovakia. (1) Forma vagum H. sublactucaceum var. radnoricum Druce & Zahn; H. boreale var. calvatum F. Hanb.; H. calvatum (F. Hanb.) Pugsley; H. sabaudum subsp. nemorivagum var. sparsicilium forma calvatum (F. Hanb.) Zahn. Styles discoloured. (2) Forma croceostylum (Pugsley) P. D. Sell H. croceostylum Pugsley; H. subquercetorum Pugsley Styles yellow. Native. Open woods, stream banks, rocky places, dunes, cliff ledges and heaths where it is native, and banks, waysides, coppices, hedgerows, railway banks, wall-tops, quarries, waste ground and debris where it is probably introduced. It goes up to at least 350 m in Wales. A common species of central and northern England and Wales with a few outlying localities elsewhere in Great Britain. It occurs throughout a large part of Continental Europe. The main variant is forma vagum. Forma croceostylum is fairly frequent in Wales and occurs also in Yorkshire. 2. H. rigens Jord. Rigid Hawkweed H. boreale subsp. virgultorum var. rigens (Jord.) Sudre; H. sabaudum subsp. virgultorum var. rigens (Jord.) Schinz & R. Keller; H. sabaudum subsp. rigens (Jord.) Zahn Aphyllopodous perennial herb with a branched stock. Stem 50–100 cm, pale yellowish-green, sometimes tinted purplish, erect, usually slender, rarely robust, striate, flexuous,

39. Hieracium with numerous to dense, long, pale simple eglandular hairs near the base, with fewer, shorter ones upwards, or scaberulous throughout, or becoming glabrous upwards. Leaves 20–50, rarely aggregated near the base, gradually decreasing in size upwards, dark green on upper surface, paler beneath, sometimes tinted purplish, firm, the lamina of the lower 10–15 × 2–5 cm, lanceolate, long acute or acuminate at apex, sharply serrate or serrulate with fine, ascending teeth and long attenuate at base but scarcely petiolate, the lamina of the median 5–10 × 1.5–3.0 cm, lanceolate to ovate, long acute or acuminate at apex, finely, sharply and unequally serrate or in large plants sometimes serratedentate with larger teeth, in cultivation the teeth become even larger and the leaf is shortly cuneate to rounded at the sessile base, the lamina of the upper 1–4 × 0.4–2 cm, ovate-lanceolate or linear, long acute at apex, more or less serrulate and with a rounded sessile base, all glabrous or nearly so on the upper surface, with few to numerous, short to medium, pale simple eglandular hairs on the lower surface and margin or scaberulous, sometimes with a few stellate hairs on the lower surface of the upper. Inflorescence with 10–40(–100) capitula in a narrow panicle or small corymb, sometimes with a long lower branch; peduncles straight, with dense stellate hairs, without simple eglandular or glandular hairs or with very occasional simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, dark olive or blackish-green, the inner sometimes with slightly paler margins, all linear-lanceolate, obtuse at apex, with numerous, short, slender, unequal, dark glandular hairs and sometimes an occasional, short or medium pale simple eglandular hair. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins fibrillous-dentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 8–10. Apomictic. Native. Locally common on sandy heaths, open woodland, railway banks and grassy places in south-east England, where it is probably native. In a few outlying localities in central England and solitary records from Cumberland, the Lothians, Forfarshire and Kincardineshire, where it may be introduced. Pyrenees and central Europe. 3. H. salticola (Sudre) P. D. Sell & C. West Bluish-leaved Hawkweed H. virgultorum microgen. dispatulum var. salticola Sudre; H. sublactucaceum auct. Aphyllopodous perennial herb with a branched stock. Stem 50–120 cm, pale bluish-green, sometimes tinted purplish below, erect, slender to robust, sometimes flexuous, striate, with medium to long, pale simple eglandular hairs throughout, sometimes numerous, sometimes sparse or sometimes scaberulous especially in the upper part, and sometimes a few stellate hairs above. Leaves 20–40, sometimes gradually decreasing in size upwards, sometimes forming a pseudorosette near the base, sometimes abruptly becoming smaller halfway up and then remaining more or less the same size, slightly bluish medium green on upper surface, paler and more bluish beneath, sometimes faintly blotched or the lower tinted reddish, the lamina of the lower 6– 16 × 1–3 cm, narrowly elliptical or lanceolate, acute at apex, serrate with regular, spinulose teeth and narrowed

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at base to a short, winged petiole or sessile, the lamina of the median when gradually decreasing in size similar to lower but slightly smaller and more shortly narrowed to a rounded base, the lamina of the upper or all those in the upper part when abruptly smaller in size 1–6 × 0.4–2.5 cm, lanceolate or ovate, acute at apex, sharply serrate or the uppermost entire and cuneate or rounded at the sessile base, all glabrous or nearly so, or with medium, pale simple eglandular hairs along the midrib beneath or on the margin, or scaberulous. Inflorescence a panicle or small corymb; peduncles straight and erect, with numerous to dense stellate hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.3 mm, blackish-green, more or less concolorous, broadly linearlanceolate, obtuse at apex, glabrous or with a line of minute glandular hairs down the centre, rarely with a solitary simple eglandular hair. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins fibrillous-dentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 8–10. Apomictic. Records from Aberdeen Links and by the River Dee near Kingcausie in Aberdeenshire, dunes in Kincardineshire and heaths and commons in the south-east could be of native plants. Widely scattered records elsewhere in Great Britain, particularly in the Birmingham area, on railway banks, roadand lanesides, ballast pits, chalk quarries and coppice margins are probably introductions. It is a widespread species in France and Germany. 4. H. prominentidens P. D. Sell Large-toothed Hawkweed H. dispalatum var. macrodon Sudre, non H. macrodon N¨ageli & Peter; H. boreale var. lacerum Wimm. & Grab., non H. lacerum Reut. ex Fr. Aphyllopodous perennial herb with a branched stock. Stem 70–130 cm, pale yellowish-green, sometimes flushed purplish towards the base, robust, striate, with numerous, short and medium, pale simple eglandular hairs, and numerous stellate hairs above. Leaves 25–50, medium green on upper surface, paler and slightly bluish beneath, gradually decreasing in size upwards, the lamina of the lower 7–13 × 3–4 cm, lanceolate or oblong-lanceolate, more or less acute at apex, deeply and irregularly laciniate-dentate, the teeth up to 20 mm, narrow, acute and curved upwards and gradually narrowed to a short winged petiole, the lamina of the median lanceolate or ovate, acute at apex and toothed like the basal, the lamina of the upper ovate, lanceolate or linear-lanceolate, acute at apex, entire to sharply dentate and sessile, all glabrous or nearly so with scaberulous margins. Inflorescence with 20–30 capitula, elongatepaniculate, with short lower branches; peduncles with dense stellate hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.8– 1.2 mm, blackish-green, linear-lanceolate, obtuse at apex, glabrous or with a few, minute glandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with subulate-dentate margins. Achenes 3.0–3.5 mm, blackishbrown. Flowers 8–9. Apomictic. Native. Recorded from Barnes Common in Surrey, Northwood in Middlesex, where it grows with H. salticola,

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and Montgomeryshire. France, Germany, Switzerland and Austria. 5. H. virgultorum Jord. Long-leaved Hawkweed H. boreale subsp. virgultorum (Jord.) Sudre; H. sabaudum subsp. virgultorum (Jord.) Schinz & Keller Aphyllopodous, perennial herb with a branched stock. Stem 50–100 cm, pale yellowish-green, erect, slender, striate, flexuous, with numerous to dense, medium to long, pale, rather stiff, minutely bulbous-based simple eglandular hairs throughout, often fewer and shorter above and sometimes becoming nearly glabrous or scaberulous above. Leaves 20– 40, gradually decreasing in size upwards, bright green on upper surface, paler beneath, the lamina of the lower 8–15 × 1.0–2.5 cm, narrowly elliptical or narrowly lanceolate, acute at apex, denticulate or serrulate, the teeth rather distant and long attenuate at base to a winged petiole, the lamina of the median similar but slightly shorter and sometimes sessile, the lamina of the upper mostly lanceolate, acute at apex, entire to denticulate or serrulate and more or less cuneate to a sessile base, all glabrous or nearly so on the upper surface, with medium to long, pale simple eglandular hairs scattered on the lower surface, particularly the midrib, and on the margins and often becoming scaberulous. Inflorescence with 10–50 capitula, in a panicle, sometimes with long lower branches; peduncles straight and erect, slender, with numerous to dense stellate hairs and sometimes with a few short or medium, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.0–1.3 mm, dull or dark olive green, concolorous or the inner with slightly paler margins, all broadly linear-lanceolate, obtuse at apex, with numerous slender, unequal, dark glandular hairs and microglands, often with some, slightly longer, pale simple eglandular hairs, without stellate hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins fibrillous-dentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 8–10. Apomictic. Probably native. Open woodland and grassy banks, very local in south-east England; widespread and frequent as as an introduction on roadsides and railway banks in south-central Scotland with isolated records further north. Widespread in Continental Europe from Spain and France to Russia. 6. H. salicetorum Sudre Willow-leaved Hawkweed H. scabiosum var. salicetorum (Sudre) Sudre; H. sabaudum subsp. scabiosum var. salicetorum (Sudre) Zahn Aphyllopodous perennial herb with a branched stock. Stem 60–100 cm, pale yellowish-green, suffused purplish near the base, more or less robust, striate, with dense, long, pale simple eglandular hairs, sometimes becoming scabrous particularly above, and with stellate and minute glandular hairs in the upper part. Leaves 15–22, medium green on upper surface, paler beneath, gradually decreasing in size upwards, the lamina of the lower 8–15 × 1.5–3.0 cm, longlanceolate, oblong-lanceolate or narrowly elliptic-oblong, gradually narrowed to a more or less acute apex, subentire or with a few small teeth and gradually narrowed at the base, the lamina of the median similar but smaller and more acute at apex, the lamina of the upper lanceolate, acute at apex,

entire or with a small tooth and cuneate at base, all glabrous or nearly so on upper surface, with short to medium, pale simple eglandular hairs beneath, particularly on the midrib, hairy or scabrous on the margin. Inflorescence with 10–20 capitula, narrow in outline, with a corymb at the top and short branches lower down; peduncles with numerous stellate hairs, few simple eglandular hairs and minute glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–11 × 1.0–1.5 mm, pale olive green, the inner with paler margins, all linearlanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous, short, dark glandular hairs and with a few stellate hairs towards the base. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 7–8. Apomictic. This species is exactly like H. virgultorum in appearance, but has paler involucral bracts which are clothed with numerous simple eglandular as well as glandular hairs. Possibly native. Recorded only from Leigh Hill in Surrey, from the Lleyn Peninsula in Caernarvonshire and a roadside at Monteviot in Roxburghshire. France and Switzerland. 7. H. argutifolium Pugsley Sharp-toothed Hawkweed Aphyllopodous, perennial herb with a branched stock. Stem 50–90 cm, pale yellowish-green, sometimes slightly suffused purplish, slender to robust, erect, markedly striate, with short to long, pale, rough, wavy simple eglandular hairs especially below, often becoming scaberulous in the upper half and sometimes throughout. Leaves 15–30, deep green on upper surface, paler beneath, sometimes slightly tinted purplish, rather crowded on the stem and gradually decreasing in size upwards, the lamina of the lower 8–15 × 2– 5 cm, mainly broadly to narrowly elliptical, equally narrowed at the acute apex and sessile base and broadest in the middle, sometimes elliptic-lanceolate or almost rhombiclanceolate and with fine, very sharp or spinulose ascending teeth sometimes interspersed with serrulations, the lamina of the median slightly shorter but still large and broad and if anything more markedly toothed, the lamina of the upper ovate or lanceolate, long acute or acuminate at apex, sharply toothed or subentire, shortly narrowed or rounded at the sessile base, with short to long, pale, rather stiff, sometimes bulbous-based hairs on both surfaces and the margins, or only on the veins beneath, or only scaberulous. Inflorescence with 6–50 capitula usually in a narrow corymbose panicle, with long lower branches; peduncles suberect, with numerous to dense stellate hairs and few to numerous, short to long, pale simple eglandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–14 × 1.2–1.5 mm, blackish-green, almost concolorous, broadly linear-lanceolate, obtuse at apex, with numerous, unequal, fine dark glandular hairs and microglands and few to fairly numerous, short to long, pale, often darkbased simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins fibrillousdentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 6–9. Apomictic. This species can be distinguished from the variable H. sabaudum by its elliptical leaves equally tapered at both

39. Hieracium ends and broadest in the middle, with fine sharp or spinulose teeth. It seems to start flowering earlier than other species of the Section Sabauda. Probably introduced as its distribution seems to be associated with railways, roadsides and waste places, but it is not known outside Great Britain. Bewdley in Worcestershire in 1858; railway bank between Fearn and Tain, Ross-shire in 1891; hedgebank near Torpantau Station, Breconshire in 1892 and 1893; hedge Rhayader, Radnorshire in 1892; colliery mounds, Neath, Glamorganshire in 1906; rough bank Selham, Sussex in 1907; roadside near Doddington, Lincolnshire in 1907; Leigh-on-Sea, Essex in 1910; roadside near Abriachan, Inverness-shire in 1966; railway line Borthwick Bank, Midlothian in 1968; and shaded piles of cinders and ballast, Whittlesey Station, Cambridgeshire in 1970. 8. H. sabaudum L. Autumn Hawkweed Aphyllopodous perennial herb with a branched stock. Stem (30–)50–100(–120) cm, yellowish-green, often more or less tinted reddish, sometimes deep rose-purple, slender to robust, erect, striate, sometimes flexuous, variable in hairiness, sometimes with medium to long, numerous, wavy, pale, often bulbous-based simple eglandular hairs throughout, sometimes with them mainly in the lower half, sometimes partly or wholly scabrous. Leaves 20–40(–70), sometimes gradually reduced in size upwards, sometimes abruptly reduced in size part of the way up the stem, sometimes with leaves aggregated near the base or halfway up the stem to form a false rosette, medium to dark dull green on upper surface, paler beneath, sometimes the lower tinted reddish, occasionally the whole leaf suffused rosepurple, the lamina of the lower 5–15 × 1.5–3.0(–4.0) cm, mostly lanceolate, ovate-lanceolate or oblong-lanceolate, and widest below the middle, sometimes narrowly elliptical or oblong-elliptical, mostly acute but sometimes obtuse at apex, subentire to denticulate or with short ascending teeth, narrowed at base and sessile or with a short, winged petiole, the lamina of the median ovate-lanceolate or lanceolate, acute at apex, usually a little more toothed than lower and shortly contracted or rounded at base, the lamina of the upper ovate or lanceolate, acute at apex, subentire or denticulate and mostly rounded at base, all mostly glabrous on the upper surface, with short to long, pale simple eglandular hairs beneath, especially on the midrib and margins, sometimes only scaberulous. Inflorescence with (3–)20– 45(–60) capitula, in a corymbose panicle, often with long lower branches; peduncles suberect or curved, with numerous to dense stellate hairs, pale simple eglandular hairs few to numerous or absent. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–13 × 1.0–1.5 mm, blackish-green, nearly concolorous or with slightly paler margins, all broadly linear-lanceolate, more or less obtuse at apex, with numerous to dense, short, unequal, slender, dark glandular hairs and microglands, and often few to numerous, pale, short to long simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins fibrillous-dentate. Achenes 3–4 mm, purplish-black. Flowers 7–10. Apomictic. 2n = 27. H. sabaudum is most variable and seems to be subject to roadside cutting and grazing more than any other leafy

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species. In habit it can have the leaves gradually decreasing in size up the stem or grouped in a false rosette towards the base or halfway up with the remaining leaves becoming abruptly smaller. The leaves are usually narrower in outline than in H. vagum, but broad-leaved variants do occur. The amount of simple eglandular hairs on the peduncles and involucral bracts varies greatly and the two extremes have been called forms. When the two forms grow together the hairiness makes them look very different. A variant occurs having the stems and leaves deep rosy purple, but it has no other distinguishing features and grows with the ordinary green variant. Poorly grown and secondary growth plants can be very dwarf and have few leaves and capitula. (1) Forma sabaudum H. sabaudum subsp. sedunense var. perpropinquum Zahn; H. perpropinquum (Zahn) Druce; H. boreale subsp. dumosum microgen. propinquum Sudre; H. boreale subsp. obliquum var. propinquum (Sudre) Rouy; H. bladonii var. brunkeri Pugsley; H. boreale var. hervieri auct. Upper part of peduncles and involucral bracts with numerous simple eglandular hairs. 2n = 27 univalents in pollen mother cell (R. A. Finch). (2) Forma bladonii (Pugsley) P. D. Sell H. heterophyllum Bladon, non Retz.; H. bladonii Pugsley; H. obliquum Jord.; H. boreale subsp. obliquum (Jord.) Sudre; H. sabaudum subsp. obliquum (Jord.) Schinz & R. Keller Upper part of peduncles and involucral bracts without or with an occasional simple eglandular hair. Native. In the lowlands it is a species of open woodland, heaths and shady hedgerows particularly on sandy (often acid) soils, and is probably introduced where it occurs on roadside and railway banks, quarries and old walls. In upland areas it occurs by streams and on rocky (particularly shady) banks, cliff ledges and scree. Throughout Great Britain and in the Isle of Man, common in the south-east of England and in Wales, scattered records in Scotland; southeast Ireland. The most common leafy species in the lowlands and extending up to 500 m in Wales. Occurs throughout much of western and central Europe. Forma sabaudum is the most common variant, although forma bladonii occurs here and there in the range of the species, but has not been seen for Ireland. Care should be taken in looking for the glandular hairs and microglands on the involucral bracts of forma bladonii to separate it from H. vagum. 9. H. eminentiforme Pugsley Exmoor Hawkweed H. eminens auct.; H. occitanum auct. Aphyllopodous perennial herb with a branched stock. Stem (30–)50–110 cm, pale yellowish-green, sometimes suffused brownish-red, more or less robust, erect, striate, with numerous to dense, pale, wavy, long simple eglandular hairs in the lower part becoming fewer and shorter upwards, often becoming more or less glabrous or scaberulous in the upper half. Leaves 20–60, bright green on upper surface, paler beneath, rather crowded on the stem but widespread on the branches, decreasing in size upwards, the lamina of the lower 6–13 × 2–4 cm, narrowly to broadly elliptical, ovate

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or lanceolate, acute at apex, with a few, small, spreading, sharp and sometimes curved teeth and narrowed below to a short, winged petiole or sessile, the lamina of the median elliptical, ovate or lanceolate, acute at apex, with a few small or sometimes larger teeth and shortly cuneate or rounded at the sessile base, the lamina of the upper ovate or lanceolate, acute at apex, entire or sparsely denticulate and rounded at the sessile base, all glabrous or nearly so, or with an occasional, pale, medium to long simple eglandular hair on the upper surface and scattered ones on the lower surface, often becoming scaberulous. Inflorescence with 10–30(–80) capitula in a panicle, the branches long and open; peduncles long, with numerous to dense stellate hairs and without or with few to numerous, pale, medium to long simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–14 × 1.2– 1.5 mm, blackish-green, nearly concolorous, broadly linearlanceolate, obtuse at apex, the tips of the outer spreading, with numerous, very short to medium, slender, unequal, dark glandular hairs and usually few to numerous, medium to long, pale, dark-based simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins fibrillous-dentate. Achenes 3.5–4.0 mm, purplishblack. Flowers 8–10. Apomictic. It is questionable whether H. eminentiforme is a distinct species from the variable H. sabaudum. It is best recognised from that species by its less hairy habit, its broader leaves none of which are elongated as in that species, its more open inflorescence with long branches, larger capitula and longer, wider involucral bracts which have more glandular hairs. Some plants are much nearer H. sabaudum than others. It retains its characters in cultivation. Native. Rocky slopes, by streams, rocky open woodland and on roadsides and embankments. Along the north coastal region of Exmoor from Culbone in Somerset to Pickwell Downs, Woolacombe, Devonshire and both sides of the Afon Teifi between Cardigan and Cilgerran in Pembrokeshire and Cardiganshire. Endemic. 10. H. dumosum Jord. Yellow-glandular Hawkweed H. boreale var. dumosum (Jord.) Gren. & Godr.; H. sabaudum subsp. dumosum (Jord.) Sudre Aphyllopodous perennial herb with a branched stock. Stem 30–80 cm, pale yellowish-green, sometimes slightly purplish below, slender to robust, sometimes slightly flexuous, markedly striate, with long, wavy, pale, numerous to dense, shaggy, often bulbous-based simple eglandular hairs throughout or becoming scaberulous in the upper part, and few to numerous stellate hairs and an occasional minute, yellowish glandular hair in the upper part. Leaves 10–40, gradually decreasing in size upwards, rather pale yellowishgreen on upper surface, paler beneath, the lamina of the lower 8–15 × 2.5–4.0 cm, lanceolate or elliptic-lanceolate, acute at apex, with a few, small, spreading teeth and narrowed at base to a short, winged petiole, the lamina of the median lanceolate or ovate, acute at apex, with sharp, ascending, sometimes spinulose teeth, and narrowed or rounded at the sessile base, the lamina of the upper lanceolate or ovate, acute at apex, subentire to sharply toothed and rounded at the sessile base, all usually with the upper surface

more or less glabrous or with an occasional simple eglandular hair and with numerous, pale, short to long, sometimes bulbous-based simple eglandular hairs beneath and on the margin, sometimes becoming scaberulous. Inflorescence with 4–30 capitula, paniculate-corymbose,with long lower branches; peduncles rather short, suberect, with numerous to dense stellate hairs, numerous to dense, long, pale, wavy simple eglandular hairs and a few minute glandular hairs. Capitula 35–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.2–1.5 mm, olive green, most, but especially the inner with broad pale margins, all linear-lanceolate, obtuse at apex, with numerous to rather dense, medium and long, pale, sometimes dark-based simple eglandular hairs intermixed with numerous, medium and short, very unequal, slender, yellowish to rather dark glandular hairs and some stellate hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins fibrillousdentate. Achenes 3.5–4.0 mm, blackish-red. Flowers 7–9. Apomictic. Presumably introduced. Known only from sandy ground near the National Trust car park, Crantock, near Newquay, Cornwall, Hazeley Heath in Hampshire, near Llandysul in Cardiganshire and walls of the Botanic Garden, Glasgow. It is widespread in central and western Europe. Section 2. Hieracioides Dumort. Hieracioides Moench, non Fabr.; Section Aphyllopoda Froel. ex W. D. J. Koch; Subsection Leiocephala Froel.; Section Accipitrina W. D. J. Koch; Series Accipitrina (W. D. J. Koch) Fr.; Taxon Umbellata Fr.; Section Umbellata (Fr.) Gremli; Series Umbellata (Fr.) P. D. Sell & C. West comb. inval. Aphyllopodous perennial herbs with a branched stock. Leaves numerous, linear to oblong or elliptic-oblong, more rarely elliptical or ovate, subentire or with distinct teeth, the teeth usually small, rarely large, more or less narrowed at base, with recurved, scaberulous margins, not reticulateveined beneath. Inflorescence of few to numerous capitula, usually more or less umbellate or with an umbellate top to the panicle. Involucral bracts incumbent in bud, 4–12 × 1.2–1.4 mm, linear-lanceolate, obtuse at apex, the outer squarrose with more or less reflexed tips, glabrous or nearly so. Ligules yellow, glabrous. Styles yellow, or rarely slightly discoloured. Receptacle pits with margins setaceous-dentate to dentate-fimbriate. Achenes blackishbrown. Flowers 7–9. An easily recognised group by its leaves having recurved, scaberulous margins, the inflorescence usually more or less umbellate, the involucral bracts squarrose and more or less reflexed at the tips and nearly glabrous and the receptacle pits setaceous-dentate to fimbriate-dentate. 11. H. umbellatum L. Umbellate Hawkweed Aphyllopodous perennial herb with a branched stock. Stem 10–100 cm, pale yellowish-green, often reddish or brownish-purple towards the base, sometimes throughout, slender to robust, striate, usually rigid, sometimes flexuous, with few to numerous, medium to long, pale simple eglandular hairs below, sometimes a few above, sometimes scaberulous throughout, sometimes with few to numerous

39. Hieracium stellate hairs above. Leaves flaccid to rigid, numerous, pale, bright or dark green on upper surface, paler beneath but without reticulate venation, very variable in shape and texture but intermediates between all variants, usually gradually decreasing in size upwards, sometimes clustered below and abruptly becoming smaller about halfway up the stem, the lamina of the lower 4–15 × 0.5–4.0 cm, linear, oblong or oblong-elliptical, rarely broadly elliptical, obtuse-mucronate to acute, entire to long-dentate, the margins recurved, gradually narrowed at base and sessile or with a short, winged petiole, the lamina of the median linear, narrowly elliptical, oblong, lanceolate or ovate, obtuse to more or less acute at apex, entire to dentate, to long dentate, sometimes the teeth narrow and curved, the margins recurved and cuneate or attenuate at base, the lamina of the upper linear or lanceolate, acute at apex, entire or dentate and cuneate at base, all usually scaberulous beneath and on the margins and glabrous on the upper surface, sometimes with few to numerous short to medium, pale simple eglandular hairs beneath, especially on the midrib, sometimes also stellate hairs beneath. Inflorescence with 1–60 capitula, usually cymose-umbellate, usually with short lower branches, sometimes with a long panicle with the top umbellate, sometimes in very dwarf plants or secondary growth with a single capitulum; peduncles more or less erect, slender, with dense stellate hairs, rarely with an occasional, medium, pale simple eglandular hair, sometimes slightly scabridulous. Capitula 30–35 mm in diameter, rounded to truncate at base. Involucral bracts incumbent in bud, 4–12 × 1.2–1.4 mm, olive or blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, the outer lax to squarrose, usually with reflexed tips, usually glabrous, rarely with a few stellate hairs or an occasional medium, pale simple eglandular hair or short, dark glandular hair. Ligules deep yellow, glabrous. Styles usually yellow, rarely slightly discoloured. Receptacle pits with margins irregularly dentate to setaceous-dentate or fimbriate-dentate. Achenes 3–4 mm, blackish-brown. Flowers 7–9. Sexual or apomictic. 2n = 18 (sexual) bivalents in pollen mother cell; 27 (apomictic) univalents in pollen mother cell. All our plants in this section are best included in one species, which contains both sexual diploids and apomictic triploids. No morphological characters have been found which distinguish the two cytotypes. The variation is therefore treated as though the plants are all sexual. Intermediates occur between all the taxa. Plants are often attacked by a gall-wasp, Aulucidea hieracii (Bouch´e), which causes large swellings on the stems. (a) Subsp. umbellatum Leaves dark green on upper surface, more or less rigid, narrowly linear to oblong, elliptic-oblong, oblong-lanceolate or lanceolate, at least the upper usually acute at apex, when leaves are broad usually markedly toothed. Outer involucral bracts recurved. (i) Var. dunense Reyn. H. umbellatum var. litorale Lindb. Stem 15–30 cm. Leaves up to 1.0 cm wide, linear, more or less crowded, especially towards the base, remotely toothed. Capitula few.

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(ii) Var. umbellatum H. umbellatum var. linariifolium Wallr.; H. umbellatum var. filifolium Fr. Stem (20–)30–80(–100) cm. Leaves up to 1.0 cm wide, linear, sometimes crowded, entire or with small, remote teeth up to 4 mm. Capitula few to numerous. (iii) Var. coronopifolium Bernh. ex Hornem. Stem 30–80(–100) cm. Leaves up to 2 cm wide, linear to oblong, with distant, long, curved teeth up to 10 mm. Capitula few to many. (iv) Var. commune Fr. H. umbellatum var. paniculatum Cariot; H. umbellatum var. serum Sudre. Stem 30–100 cm. Leaves up to 2.5 cm wide, oblong, elliptical-oblong or lanceolate-oblong, with 2–3, short teeth on each side. Capitula many, often in an elongate panicle. (b) Subsp. ogwenii (E. F. Linton) W. R. Linton H. ogwenii E. F. Linton; H. corymbosum forma subumbellatum F. N. Williams; H. umbellatum var. ogwenii (E. F. Linton) F. Hanb.; H. umbellatum var. paniculatum auct. Leaves up to 2.5 cm wide, bright green on upper surface, rather stiff, narrowly elliptical or lanceolate, long acute at apex, regularly dentate, narrowed at base. Outer involucral bracts not recurved. (c) Subsp. bichlorophyllum (Druce & Zahn) P. D. Sell & C. West H. laevigatum subsp. bichlorophyllum Druce & Zahn; H. bichlorophyllum (Druce & Zahn) Pugsley; H. umbellatum var. monticola auct. Stem 25–50(–70) cm. Leaves rather pale green, flaccid, broadly oblong, elliptic-oblong to lanceolate, obtuse to acute at apex, entire or sparsely denticulate. Outer involucral bracts recurved. (v) Var. curtum E. F. Linton H. bichlorophyllum var. curtum (E. F. Linton) Pugsley; H. umbellatum var. littoreum auct. Stem 10–30 cm, slender. Leaves up to 1.5 cm wide, narrowly elliptical, obtuse at apex, entire or remotely denticulate, often clustered near the base and becoming abruptly smaller halfway up the stem. (vi) Var. sarniense P. D. Sell Stem 10–30 cm, robust and fleshy. Leaves thick and fleshy, narrowly to broadly elliptical, obtuse at apex, the lower up to about 5 cm wide, clustered near the base, becoming smaller upwards. (vii) Var. taylori Bab. H. umbellatum var. monticola auct. Stem 25–50(–70) cm. Leaves broadly oblong, ellipticoblong or lanceolate, getting gradually smaller up the stem, many more than 3 times as long as wide, often obtuse at apex. (viii) Var. latifolium (W. R. Linton) P. D. Sell H. umbellatum forma latifolium W. R. Linton

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Stem 25–50(–70) cm. Leaves up to 4 cm wide, elliptical, broadly elliptical, ovate or lanceolate, most less than 3 times as long as wide, often acute at apex. Native. Heaths, wood margins, hedgebanks, scrub, dunes and shingle in the lowlands, rough pasture, stream banks and open woodlands in the uplands, sometimes introduced on roadside and railway banks. Throughout Great Britain and Ireland, but scarce in many areas and possibly most common in Wales. Europe; northern Asia; North America. Circumpolar Wide-temperate element. Probably the most widespread species of Hieracium. Subsp. umbellatum occurs throughout the range of the species as probably does var. umbellatum. Var. dunense is a plant of dunes and shingle and some Welsh upland plants seem indistinguishable from it. It is also found elsewhere on the north-west coast of Europe. Var. coronopifolium is scattered through Great Britain and Ireland, often growing with var. umbellatum. It is widespread in Europe and northern Asia and is recorded for North America. Var. commune is of scattered occurrence in Great Britain and is recorded for Ireland. It is widespread in Europe, Asia and North America. Subsp. ogwenii is endemic to river valleys in Caernarvonshire. Subsp. bichlorophyllum is a plant of south-west Britain, Wales and the border counties, Ireland and the Channel Islands and is endemic. The usual plant is var. taylori. Var. curtum occurs on the west and south coasts of Wales and the Channel Islands and is the subsp. bichlorophyllum equivalent of var. dunense. Var. sarniense is endemic to the north cliffs of Jersey. Var. latifolium is known only from woods near Kingston in Devonshire. Section 3. Foliosa (Fr.) Arv.-Touv. Taxon Foliosa Fr.; Series Foliosa (Fr.) P. D. Sell & C. West comb. inval. Aphyllopodous perennial herbs with a branched stock. Leaves numerous, the lamina linear-elliptical, linearoblong, lanceolate, oblong-elliptical, broadly to narrowly elliptical, oblanceolate or oblong-lanceolate, subentire to shortly dentate, markedly reticulate-veined beneath and at least the median and upper more or less amplexicaul. Inflorescence of few to numerous capitula, condensed or open. Involucral bracts incumbent in bud, 3–13 × (1.0–)1.2–1.6 mm, linear-lanceolate, more or less obtuse at apex, more or less hairy. Ligules glabrous. Styles discoloured or yellow. Receptacle pits sharply dentate, sometimes with setaceouspointed or fimbriate teeth. Achenes purplish or blackishbrown. Flowers 7–10. Recognised by its markedly reticulate-veined, semiamplexicaul leaves. The species of the Section Foliosa are mainly plants of upland Great Britain and Ireland found near mountain streams, and are thus plants of the north and west. Central Europe and from Scandinavia eastwards into Russia and Siberia and westwards to Iceland and Greenland. 12. H. maritimum (F. Hanb.) F. Hanb. Maritime Hawkweed H. crocatum var. maritimum F. Hanb.; H. umbellatum subsp. maritimum (F. Hanb.) Zahn; H. inuloides subsp.

strictum forma angustifolium Dahlst.; H. maritimum var. pauciceps Pugsley; H. obesifolium Pugsley; H. umbellatum var. pauciflorum auct. Aphyllopodous perennial herb with a branched stock. Stem 30–70(–90) cm, pale yellowish-green, often suffused with reddish-purple, slender to very robust, striate, sometimes flexuous, with numerous, medium to long, pale simple eglandular hairs below, becoming asperous-scabridulous above, sometimes throughout, sometimes almost glabrous throughout, usually with numerous stellate hairs in the upper part. Leaves 12–35, crowded at least below, gradually becoming smaller up the stem or abruptly smaller halfway up, dark green on upper surface, sometimes flushed reddishpurple, very thick and fleshy, very variable in width, the lamina of the lower 4–10 × 0.5–2.5 cm, narrowly linearelliptical, linear-oblong, lanceolate or narrowly elliptical, rounded-obtuse to acute at apex, entire, remotely denticulate or rarely with a few small teeth, narrowed at base, sessile or subpetiolate and semiamplexicaul; the lamina of the median similar but often more acute at apex, the lamina of the upper linear or lanceolate, acute at apex, entire, slightly rounded and semiamplexicaul at base, all glabrous on the upper surface, with short to medium, rather stiff, pale hairs on the lower surface and asperous-scaberulous on the slightly thickened margins and prominent midrib beneath. Inflorescence with (1–)2–12(–25) capitula, paniculatecorymbose, sometimes with a long lower branch; peduncles erecto-patent, with numerous to dense stellate hairs and asperous with minute projections. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all broadly linear-lanceolate, obtuse at apex, with a few stellate hairs and a few, solitary, dark glandular hairs, without simple eglandular hairs. Ligules yellow, glabrous. Styles deep yellow. Receptacle pits with sharply dentate to ciliate-dentate margins. Achenes 3.5–4.0 mm, blackish-brown. Flowers 7–8. Apomictic. A variable species in width and dentation of leaves and size of capitula, but the large colony at Carrick River, Co. Donegal shows all the variation found elsewhere and continues to do so during many years of cultivation. H. maritimum bears a superficial resemblance to H. umbellatum but can be distinguished by its semiamplexicaul leaves, very broad involucral bracts which have a few glandular hairs and by not usually forming an umbellate inflorescence. Native. Shingle and sand by lochs, grassy sea cliffs, rock ledges and sides of mountain streams. Scattered localities in Kirkcudbrightshire, Ayrshire, Perthshire, Invernessshire, Argyllshire, Kintyre, Isle of Mull, Outer Hebrides, Sutherland, Caithness, Orkney and Shetland Islands and Co. Galway, Co. Mayo, Co. Clare and Co. Donegal in Ireland. Endemic. 13. H. pycnotrichum (W. R. Linton) Roffey Dense-leaved Hawkweed H. corymbosum var. pycnotrichum W. R. Linton; H. inuloides subsp. pycnotrichum (W. R. Linton) Zahn. Aphyllopodous perennial herb with a branched stock. Stem 40–90 cm, pale yellowish-green, sometimes suffused

39. Hieracium brownish-purple below, robust, striate, with medium to long, pale simple eglandular hairs in the lower part or glabrous or slightly scabridulous throughout. Leaves 15–40, more or less crowded, gradually decreasing in size upwards, medium green on upper surface, paler beneath with prominent veins, thickened at margin; the lamina of the lower 5–13 × 1.5–2.0 cm, narrowly elliptical to oblong-elliptical or oblong, obtuse to acute at apex, subentire or with a few remote teeth, attenuate at base and often petiolate; the lamina of the median narrowly elliptic-oblong or lanceolate, long acute at apex, entire or denticulate, narrowed to a semiamplexicaul base; the lamina of the upper lanceolate or linear-lanceolate, more or less acute at apex, more or less entire, rounded and semiamplexicaul at base, all with numerous, short to long, pale simple eglandular hairs on both surfaces and the margins, or nearly glabrous above, or scaberulous particularly on the margin, sometimes with stellate hairs on the underside of the upper leaves. Inflorescence with 2–25 capitula, compactly corymbose, sometimes with lower branches; peduncles rather spreading, slender, with numerous to dense stellate hairs, few, medium, pale simple eglandular hairs and sometimes a few, very small glandular hairs. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 3–12 × 1.0–1.3 mm, dull olive green, the inner with broad pale margins, all linear-oblong, obtuse at apex, with numerous very short and short, dark glandular hairs and microglands, without stellate or simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins incisedentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Only known from near the Gayle Beck at Hawes in Yorkshire, where it was collected in 1888 and 1889, and by the River Wear at Bishop Auckland in Co. Durham in 1871. It is also found in Scandinavia. 14. H. bakeranum Pugsley Baker’s Hawkweed H. crocatum auct.; H. angustum auct.; H. strictum var. angustum auct. Aphyllopodous perennial herb with a branched stock. Stem 50–90 cm, pale yellowish-green, sometimes tinted purplish towards the base, slender, glabrous or nearly so throughout, or scaberulous, or with some medium to long, pale simple eglandular hairs towards the base. Leaves 15–30 (–40), gradually decreasing in size upwards, rather pale yellowish-green on upper surface, paler beneath; the lamina of the lower 7–10 × 1–2 cm, oblong-lanceolate or narrowly elliptical-oblong, mostly obtuse at apex, subentire or remotely denticulate and long attenuate below to short, winged petioles, the lamina of the median similar to lower but sometimes with remote teeth and a sessile, semiamplexicaul base, the lamina of the upper linear-lanceolate, lanceolate or oblong, long acute at apex and with a rounded, semiamplexicaul base, all with soft, pale, few to numerous, medium to long, simple eglandular hairs on both surfaces and margins, or nearly glabrous especially on upper, or scaberulous especially on the margins. Inflorescence with 3–10(–12) capitula, paniculate-corymbose; peduncles more or less erect and slender, with scattered to dense stellate hairs and rarely a solitary simple eglandular hair.

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Capitula 30–35 mm in diameter, rounded or narrowed at the base. Involucral bracts incumbent in bud, 4–11 × 1.0– 1.2 mm, blackish-green, the inner with broad pale margins, all linear-lanceolate to oblong-lanceolate, obtuse to subacute at apex, with numerous, very short and short, unequal, dark glandular hairs and microglands and occasional short or medium, pale, dark-based simple eglandular hairs, without stellate hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.5–4.0 mm, dark purplish-brown. Flowers 7–9. Apomictic. Native. Known only from the banks of the River Tees in the region of Winch Bridge, Upper Teesdale. Endemic. Named after John Gilbert Baker (1834–1920). 15. H. subumbellatiforme (Zahn) Roffey Slender Hawkweed H. aestivum subsp. subumbellatiforme Zahn; H. corymbosum var. umbellatiforme W. R. Linton; H. strictum var. amplidentatum F. Hanb.; H. inuloides subsp. tridentatifolium var. amplidentatum (F. Hanb.) Zahn; H. pseudamplidentatum Pugsley; H. pseudamplidentatum var. stenophyllum Pugsley; H. strictum var. angustum auct.; H. aestivum subsp. angustum auct.; H. angustum auct.; H. corymbosum var. salicifolium auct. Aphyllopodous perennial herb with a branched stock. Stem 30–100 cm, pale yellowish-green, often tinted brownishpurple in the lower half, slender to robust, striate, sometimes flexuous, more or less glabrous, or with a few, medium to long, pale simple eglandular hairs in the upper half, or scaberulous. Leaves 20–40, gradually decreasing in size upwards, rather pale green on upper surface, paler and slightly glaucous beneath with prominent veins and thickened margins; the lamina of the lower (6–)10–15 × 1.5–2.5 cm, oblong or narrowly elliptic-oblong, obtuse-mucronate to acute at apex, subentire to denticulate and attenuate at base to long or short petioles; the lamina of the median often crowded, narrowly elliptical or lanceolate, long acute at apex, subentire, denticulate or sparingly sinuate-dentate and shortly narrowed or rounded at the sessile, semiamplexicaul base, the lamina of the upper lanceolate or linear-lanceolate, long acute at apex, entire and rounded at the sessile, semiamplexicaul base; all with stellate hairs on both surfaces or only beneath, the lower often with some pale, simple eglandular hairs particularly on the veins beneath, sometimes all leaves scaberulous particularly on the margins and veins beneath. Inflorescence with 6–20 capitula, broadly paniculate-corymbose, often with lower branches; peduncles slender, often curved, with few to dense stellate hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts 4–11 × 1.0–1.4 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with very short and short, dark, slender, unequal glandular hairs down the median line and a few scattered stellate hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins more or less fimbriate-dentate. Achenes 3.5–4.0 mm, dark purplish-brown. Flowers 7–9. Apomictic. Native. Streamsides and river banks. Widespread in central Scotland. Endemic.

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16. H. strictiforme (Zahn) Roffey Strict Hawkweed H. inuloides subsp. strictiforme Zahn; H. opsianthum (Dahlst. ex F. Hanb.) Roffey; H. strictum var. opsianthum Dahlst. ex F. Hanb.; H. inuloides subsp. opsianthum (Dahlst. ex F. Hanb) Zahn; H. lycopifolium var. norvegicum Lindb.; H. strictum auct.; H. listerae Pugsley; H. crocatum auct. Aphyllopodous perennial herb with a branched stock. Stem 30–90 cm, pale yellowish-green, often suffused brownishpurple in the lower half, more or less robust, striate, with few to numerous, medium to long, pale simple eglandular hairs, mainly in the lower half, or scaberulous, with few to numerous stellate hairs towards the inflorescence. Leaves 15–30(–40), gradually decreasing in size upwards, pale to rather deep green on upper surface, paler and reticulateveined beneath, thickened on the margin, the lamina of the lower 5–13 × 1.0–3.5 cm, narrowly oblong to narrowly elliptical or oblanceolate to obovate-oblanceolate, obtusemucronate to acute at apex, subentire or denticulate, sometimes with 1–2 large teeth and gradually narrowed at base to a short, winged, semiamplexicaul petiole; the lamina of the median oblong, narrowly elliptical or elliptic-lanceolate, obtuse-mucronate to acute at apex, denticulate to shortly dentate and mostly rounded or rarely shortly narrowed at the sessile, semiamplexicaul base, the lamina of the upper oblong, lanceolate or linear-lanceolate, rarely ovate, acute at apex, entire or denticulate or rarely dentate, all with short to medium, occasionally long, pale, sometimes rather stiff simple eglandular hairs on both surfaces and the margins, the upper particularly sometimes scaberulous, with numerous stellate hairs on the lower surface of at least the upper, and sometimes on lower leaves also. Inflorescence with (2–)6–15(–30) capitula, mostly paniculate-corymbose, sometimes with long lower branches; peduncles suberect and slender, with numerous to dense stellate hairs and sometimes few to rarely numerous, short or medium, pale simple eglandular hairs. Capitula 30–35 mm in diameter, truncate at base. Involucral bracts 4–13 × 1.2–1.5 mm, blackishgreen, the inner with broad pale margins, all broadly linearlanceolate, obtuse at apex, with numerous very unequal, very short and short, dark glandular hairs intermixed with few to numerous, short to medium, pale, sometimes darkbased simple eglandular hairs and sometimes with few, scattered stellate hairs. Ligules pale yellow, glabrous, or rarely with a few, very short simple eglandular hairs at the tips. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.5–4.0 mm, dark purplish or blackishbrown. Flowers 7–8. Apomictic. This variable species can be recognised by its oblong to narrowly elliptical leaves, involucral bracts with numerous, unequal glandular hairs mixed with longer simple eglandular hairs and discoloured styles. The two extremes of leaf shape, oblong to narrowly elliptical, especially if there is only one type in a colony, make the plants look very different, hence the division by H. W. Pugsley (1948) into three species, but the bulk of the large collection of specimens in CGE fall in the middle of the range and a series sometimes occurs in a single colony. There is also a slight difference in capitulum size and number of simple eglandular hairs on

the involucral bracts, but these characters do not fit in with those of the leaves, and it seems best to leave one variable species. Native. Mainly rocks and grassy banks by streams, also on cliff bases and sandhills up to 800 m, and possibly introduced on roadside and railway banks. Common in central and north Scotland, a few scattered localities in southern Scotland and northern England, and north-east and southeast Ireland. It also occurs in Norway. 17. H. reticulatiforme P. D. Sell Reticulate-leaved Hawkweed H. reticulatum var. angustifolium Pugsley, non H. angustifolium Hoppe; H. reticulatum auct. Aphyllopodous perennial herb with a branched stock. Stem 30–70(–100) cm, pale yellowish-green, often flushed purplish below, slender to robust, striate, glabrous, scaberulous or with a few, pale simple eglandular hairs here and there and a few stellate hairs above. Leaves 15–40, yellowishgreen on upper surface, paler, slightly glaucous, sometimes tinted reddish and reticulate-veined beneath; the lamina of the lower 8–14 × 1.5–2.0(–2.5) cm, lanceolate, oblonglanceolate, narrowly elliptical or oblong-elliptical, acute at apex, entire or with few, small teeth, attenuate at base and sessile or with a short winged petiole, the lamina of the middle and upper gradually decreasing in size, oblong, oblong-lanceolate or linear-lanceolate, gradually narrowed to an acute apex, subentire or with a few, small, sharp teeth and cuneate or rounded at the sessile, semiamplexicaul base, all glabrous or with short to medium, pale simple eglandular hairs on the upper surface particularly near the margin, with numerous, short to long, pale simple eglandular hairs beneath and on the margin, especially the midrib, or scabrous on the margin, sometimes with stellate hairs beneath. Inflorescence with few to numerous capitula, paniculate-corymbose; peduncles rather long, with few to numerous stellate hairs and few to numerous, short to medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.2–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous unequal, short and very short, dark glandular hairs and a few, short and medium, pale simple eglandular hairs, with a few stellate hairs near the base. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, blackishbrown. Flowers 7–8. Apomictic. H. reticulatum Lindeb. is a slender plant with narrow, more strongly toothed leaves and smaller capitula which is found in Scandinavia and does not occur in Britain. H. reticulatiforme is distinguished from H. reayense by its longer, narrower, entire or little toothed leaves. Native. Streamsides and grassy banks. Scattered over central and north Scotland and more rarely in south Scotland. Endemic. 18. H. reayense (Pugsley) P. D. Sell Reay Hawkweed H. reticulatum var. reayense Pugsley; H. reticulatum auct. Aphyllopodous perennial herb with a branched stock. Stem 30–70 cm, pale green, often suffused brownish-purple,

39. Hieracium slender to robust, sometimes flexuous, striate, often nearly glabrous in lower half or with a few, short to medium, pale simple eglandular hairs throughout, with few to numerous stellate hairs in the upper part. Leaves 15–35, medium green on upper surface, paler beneath, gradually decreasing in size upwards, the lamina of the lower 5–10 × 1.0–2.5(–3.0) cm, narrowly elliptical or oblong-elliptical, obtuse-mucronate to subacute at apex, with several, small, sharp teeth on each side and narrowed to a sessile or shortly petiolate base, the lamina of the remainder becoming more drawn out and acute at apex and shortly narrowed to a rounded, semiamplexicaul base, the uppermost lanceolate, and with a few, sharp teeth on each side, all usually glabrous on upper surface although sometimes with a few, long pale hairs on the lower, sometimes glabrous, sometimes scabrous or sometimes with medium to long, pale simple eglandular hairs on the lower surface, especially the midrib, with numerous medium to long, pale simple eglandular hairs on the margin or sometimes scaberulous, sometimes with stellate hairs on the under surface of the upper. Inflorescence with few to numerous capitula, laxly paniculatecorymbose; peduncles with few to numerous stellate hairs and few to numerous, medium to long, pale simple eglandular hairs. Capitula 24–40 mm in diameter, with broad, rounded bases. Involucral bracts incumbent in bud, 4– 11 × 1.4–1.6 mm, dark olive green, linear-lanceolate to oblong-lanceolate, obtuse at apex, with numerous, short, dark glandular hairs and microglands and an occasional, medium, pale simple eglandular hair and a few stellate hairs towards the base. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Streamsides and rocky and grassy places. Widespread in central and northern Scotland. Endemic. 19. H. melanoglochin (E. F. Linton) P. D. Sell Dark-headed Hawkweed H. corymbosum var. melanoglochin E. F. Linton; H. reticulatum var. melanoglochin (E. F. Linton) Pugsley; H. reticulatum auct. Aphyllopodous perennial herb with a branched stock. Stem 40–70(–120) cm, pale yellowish-green, often suffused brownish-purple, usually more or less robust, usually scaberulous, sometimes with a few, medium or long, pale simple eglandular hairs, with few to numerous stellate hairs in the upper part. Leaves 15–40, yellowish-green on upper surface, paler and reticulate-veined beneath, gradually decreasing in size up the stem, the lamina of the lower 8–12 × 1.5–3.5 cm, narrowly elliptical to elliptical, gradually narrowed above to an obtuse to acute tip, more or less dentate, gradually narrowed at base and sometimes with a short, winged petiole, the lamina of the median and upper oblong, oblong-lanceolate, or lanceolate, the upper sometimes ovate, long drawn out to an acute tip, strongly serratedentate, the teeth narrow, very sharp and up to 10 mm and narrowed to a cuneate or rounded, semiamplexicaul base, all glabrous on the upper surface, or a least the lower with short to long, pale, often subrigid simple eglandular hairs, with numerous medium to long, pale simple eglandu-

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lar hairs beneath (especially on the midrib) and margin, but sometimes only scaberulous on the upper ones, sometimes with stellate hairs on the lower surface. Inflorescence with numerous capitula, usually widely and laxly paniculatecorymbose; peduncles long, with numerous to dense stellate hairs, few to numerous, short to long, pale simple eglandular hairs and few to rather numerous, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linearlanceolate, obtuse at apex, with numerous, short, unequal, fine, dark glandular hairs, usually a few to fairly numerous, short to medium, pale simple eglandular hairs and a few stellate hairs towards the base. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, blackish-brown. Flowers 7–8. Apomictic. H. melanoglochin is closely allied to H. reayense, but can be distinguished by its much longer teeth on the middle and upper cauline leaves and the few to numerous glandular hairs on the peduncles. Native. Usually on stream banks, rarely on low cliffs. Recorded from Perthshire, Aberdeenshire, Inverness-shire, Ross-shire and Sutherland. Endemic. 20. H. drummondii Pugsley Drummond’s Hawkweed Aphyllopodous perennial herb with a branched stock. Stem 40–100 cm, pale yellowish-green, often brownishred towards the base, usually robust, striate, with numerous, short to long, pale simple eglandular hairs throughout or absent from the lower part, or scabrous, with few to numerous stellate hairs in upper part. Leaves (20–)30–45, crowded except in the lower part of the stem, pale to dark green on upper surface, paler and reticulate-veined beneath, gradually decreasing in size upward, the lamina of the lower 6–8 × 2–3(–4) cm, elliptical, ellipticaloblong, oblanceolate or lanceolate, obtuse-mucronate to acute at apex, subentire to remotely undulate-denticulate and attenuate to short-winged petioles, the lamina of the median lanceolate or elliptic-lanceolate, acute at apex, denticulate to coarsely and unequally denticulate-dentate and sessile with rounded, semiamplexicaul base, the lamina of the upper ovate-lanceolate or lanceolate, acute at apex, irregularly dentate, with a sessile, amplexicaul base, all with short to medium, pale simple eglandular hairs on both surfaces or the upper becoming glabrous or scaberulous and at least the upper with numerous to dense stellate hairs on both surfaces. Inflorescence with 5–35 capitula, compactly paniculate-corymbose; peduncles slender, crowded, with numerous to dense stellate hairs and rarely a solitary, short or medium, pale simple eglandular hair. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts 4–12 × 1.0–1.3 mm, dark olive green, the inner with paler margins, all linear-lanceolate, obtuse to subacute at apex, with numerous, unequal, very short to medium, fine, dark glandular hairs and dense microglands intermixed with a few, medium, dark-based simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins subulate-dentate and more or less

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fimbriate. Achenes 3.0–3.5 mm, dark purplish-brown. Flowers 7–9. Apomictic. H. drummondii stands between H. subcrocatum and H. strictiforme from which it differs in the characteristic clothing of its involucral bracts and its stellately hairy leaves. Native. Grassy banks and streamsides. In a few localities in Kirkcudbrightshire, West Lothian, Argyllshire, Kintyre, Perthshire and Forfarshire. Endemic. Named after Thomas Drummond (c. 1793–1835). 21. H. subcrocatum (E. F. Linton) Roffey Dark-styled Hawkweed H. strictum var. subcrocatum E. F. Linton; H. aestivum subsp. subcrocatum (E. F. Linton) Zahn; H. bartonii Pugsley; H. neocorymbosum Pugsley; H. neocorymbosum var. crocatiforme Pugsley; H. corymbosum auct.; H. corymbosum var. salicifolium auct. Aphyllopodous perennial herb with a branched stock. Stem 35–100 cm, pale yellowish-green, sometimes suffused brownish-purple, especially below, slender to robust, striate, with few to numerous, short to long, pale simple eglandular hairs especially in the lower half, often becoming glabrous or scaberulous in the upper half and sometimes throughout, with few stellate hairs in the upper part. Leaves (5–)15–30, deep, dull green on upper surface, paler and reticulate-veined beneath, sometimes suffused purplish, the lamina of the lower 6–12 × 1.5–3.0 cm, oblong-lanceolate, oblong-oblanceolate, elliptical or oblong-elliptical, obtusemucronate to acute, subentire to undulate-denticulate more rarely shortly dentate, and attenuate to a sessile semiamplexicaul base or sometimes with a short winged petiole, the lamina of the upper gradually decreasing in size upwards, ovate-lanceolate, lanceolate, ovate or oblonglanceolate, acute at apex, subentire to denticulate or more rarely dentate and cuneate or rounded at the sessile, more or less amplexicaul base, all glabrous or becoming glabrous, or the lower with short to medium, pale simple eglandular hairs on the lower surface and on the margins or scaberulous, sometimes with stellate hairs on one or both surfaces. Inflorescence with few to numerous capitula, paniculatecorymbose, sometimes with a long, lower branch; peduncles with numerous stellate hairs and sometimes with occasional, short to medium, pale, dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, rounded to subtruncate at base. Involucral bracts 4–12 × 1.2–1.5 mm, olive to dark green, linear-lanceolate, more or less obtuse at apex, with few to numerous, very short or short, unequal, dark glandular hairs, sometimes with a few stellate hairs, rarely with an occasional simple eglandular hair. Ligules yellow, glabrous. Styles darkly discoloured. Receptacle pits with margins subulate-dentate, sometimes setaceous. Achenes 3.0–4.0 mm, dark purplish-brown. Flowers 7–9. Apomictic. Varies in the width and toothing of the leaves and the amount of glandular hairs on the involucral bracts. Several taxa have been described, but the characters occur in numerous different combinations. H. subcrocatum as here understood is characterised by its more or less amplexicaul leaves, few to numerous glandular hairs on its involucral bracts and dark styles. Both H. strictiforme and H. drummondii, which

have dark styles, have simple eglandular hairs mixed with the glandular ones on the involucral bracts. Native. Mainly on grassy slopes or rocks by streams, occasionally in open woodland and on hedgebanks up to 700 m. Common in upland areas of Wales, the Derbyshire dales, northern England and Scotland and local in eastern Ireland. Probably endemic. 22. H. latobrigorum (Zahn) Roffey Yellow-styled Hawkweed H. inuloides subsp. latobrigorum Zahn; H. johnstonii Dahlst.; H. aestivum subsp. hemitrachys Druce & Zahn; H. polyphyllum Dahlst., non Willd.; H. subpolyphyllum Pugsley; H. salicifolium auct.; H. auratum auct.; H. rigidum auct.; H. striatum Tausch var. pseudauratum auct.; H. inuloides subsp. latobrigorum var. pseudauratiforme Zahn; H. latobrigorum forma angustifolium Pugsley Aphyllopodous perennial herb with a branched stock. Stem 30–90(–100) cm, yellowish-green, often reddish towards the base, sometimes flexuous, striate, with few, medium to long, pale simple eglandular hairs in the lower half and numerous stellate hairs in the upper part, or glabrous or scaberulous throughout. Leaves 12–25(–30), dull yellowish-green on upper surface, paler and more or less conspicuously veined beneath, gradually decreasing in size upwards, the lamina of the lower 6–10 × 2–3(–4) cm, oblanceolate, oblong, lanceolate or elliptical, obtuse to acute at apex, subentire to shortly dentate and attenuate at base to a sessile or short winged petiole, the lamina of the median elliptical, rhombic-lanceolate, oblong or oblonglanceolate, more or less acute at apex, denticulate to irregularly but shortly dentate and cuneate or rounded at the sessile, semiamplexicaul base, all scaberulous on the lower surface and margins or with short to medium pale simple eglandular hairs, usually without but sometimes with a few simple eglandular hairs on the upper surface, with numerous stellate hairs on both or only the lower surface of at least the upper leaves, sometimes the simple hairs on the lower leaves bulbous-based. Inflorescence with few to numerous capitula, compactly paniculate-corymbose with long, spreading branches; peduncles with dense stellate hairs and sometimes an occasional, short or medium, pale simple eglandular hair. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts 4–12 × 1.3–1.5 mm, dark olive green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with few stellate hairs near the base, numerous, unequal, short and medium, dark glandular hairs and microglands and sometimes a few, medium, dark simple eglandular hairs. Ligules yellow, glabrous. Styles pure yellow. Receptacle pits with margins setaceous-subulate-dentate. Achenes 3–4 mm, dark purplish-brown. Flowers 7–9. Apomictic. This species varies in the width and dentation of its leaves, but is easily distinguished by its glandular-hairy involucral bracts and pure yellow styles. Native. Grassy banks and rocks by streams, grassy sea-cliffs and upland pastures. Common in Scotland and in the Orkney Islands, scarce in northern England south to Staffordshire and one locality in north Wales; scarce in eastern Ireland. Central Europe.

39. Hieracium 23. H. tavense (Ley) Ley Tawe Hawkweed H. rigidum var. tavense Ley; H. laevigatum subsp. tavense (Ley) Zahn; H. protractum auct. Aphyllopodous perennial herb with a branched stock. Stem 50–90 cm, pale yellowish-green, sometimes tinted purplish at base, rather slender, sometimes flexuous, striate, glabrous or with a few, medium to long, pale simple eglandular hairs especially towards the base. Leaves 15–30, dull rather greyish-green on upper surface, paler and reticulate-veined beneath, gradually decreasing in size up the stem; the lamina of the lower 6–10 × 1.5–2.5 cm, narrowly elliptical to elliptic-lanceolate or lanceolate, acute at apex, remotely denticulate or with a few, small unequal teeth and attenuate at base to long or short, winged, semiamplexicaul petioles, the lamina of the median similar, but more frequently lanceolate, long acute or acuminate at apex, denticulate or sometimes with a few spinulose teeth and cuneate to rounded at the sessile, semiamplexicaul base, the lamina of the upper lanceolate or linear-lanceolate, long acute at apex, entire to denticulate and more or less rounded at the sessile, semiamplexicaul base, all sometimes glabrous or nearly so, or with a few, short and medium, pale simple eglandular hairs beneath, particularly on the midrib, or scaberulous on the midrib and margin, usually with numerous stellate hairs on the under surface of the upper leaves. Inflorescence with 3–14 capitula, paniculate-corymbose, often with a long lower branch; peduncles rather long and slender, more or less erect, with numerous to dense stellate hairs and sometimes an occasional very short, dark glandular hair and a short or medium, pale, dark-based simple eglandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–10 × 1.2–1.5 mm, olive green, the inner with broad scarious margins, all linearlanceolate or linear-oblong, obtuse at apex, with numerous short, dark glandular hairs and microglands, sometimes a few stellate hairs about the base, without simple eglandular hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins setaceous-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. This distinct species has only been collected four times, in 1899, 1902, 1957 and 1999. Native. Known only from rocks by a waterfall, Nant-yllyn, at the head of the Upper Tawe Valley, Breconshire. Endemic. Section 4. Tridentata (Fr.) Arv.-Touv. Taxon Tridentata Fr.; Series Tridentata (Fr.) P. D. Sell & C. West comb. inval.; Section Rigida (Lindeb.) Dahlst.; Hieracium vulgatella F. N. Williams. Aphyllopodous, pseudophyllopodous or hypophyllopodous perennial herbs with branched stocks. Stems 20–110 cm, hairy or becoming scaberulous. Leaves 4–30, more or less heterophyllous, gradually decreasing in size upwards, or some leaves grouped in a false rosette, or sometimes abruptly decreasing in size, not or hardly amplexicaul, margins thickened or revolute and shortly hairy or scaberulous. Inflorescence a raceform cyme or paniculate-corymbose; peduncles with stellate and simple eglandular hairs, but only sometimes with glandular hairs. Capitula medium to rather

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large. Involucral bracts 3–13 × 0.8–2.0 mm, incumbent in bud, appressed, dark, the inner usually with paler margins, all obtuse or subacute at apex, usually with only a few stellate hairs. Ligules glabrous. Styles yellow to discoloured. Receptacle pits with margins more or less subulate-dentate, rarely fimbriate-dentate. Achenes blackish-brown. Flowers (6–)7–9. The species of this section connect the Section Vulgata with the Sections Foliosa and Hieracioides. From the Section Foliosa they differ in their non-amplexicaul leaves and from the Section Hieracioides usually in their fewer broader leaves. They are mostly species of upland grassy banks and streamsides, more rarely on cliffs, and in heathy areas in the lowlands. They are widespread in Wales, Scotland, Ireland and northern England, and local in eastern and southern England. They range over most of Europe, northern Asia and North America. 24. H. ornatilorum P. D. Sell & C. West Large-flowered Hawkweed H. sparsifolium var. grandescens auct.; H. rigidum var. friesii auct. Aphyllopodous perennial herb with a branched stock. Stem 40–110 cm, pale yellowish-green, often suffused purplish in the lower half, more or less slender, often flexuous, striate, with sparse, short, medium or long, pale simple eglandular hairs and few stellate hairs. Leaves 10–17, gradually decreasing in size upwards, dark green on upper surface, paler beneath, the lamina of the lower and median 6–10 × 2– 3 cm, elliptical, ovate or lanceolate, narrowed to an obtuse or subacute apex, dentate, with long, narrow teeth, cuneate at base and sessile or shortly petiolate, the lamina of the upper ovate or lanceolate, acute at apex, sharply dentate or serrate-dentate, rounded at base and sessile, all glabrous or nearly so on the upper surface, with few to fairly numerous, short to medium, pale simple eglandular hairs beneath and on the margin. Inflorescence with 2–12 capitula, paniculatecorymbose, with slender, divaricate branches; peduncles with dense stellate hairs, few, short or medium, pale simple eglandular hairs, rarely with a short, dark glandular hair. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–13 × 1.0–1.3 mm, dark olive green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an obtuse apex, with few, short or medium, dark simple eglandular hairs and few, short dark glandular hairs. Ligules deep yellow, glabrous. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Grassy places and streamsides. Endemic to Yorkshire and Lancashire where it was collected at Silverdale, the Scarborough area and Wharfedale by J. Cryer between 1908 and 1914. 25. H. eboracense Pugsley Northern Hawkweed H. tridentatum var. decipiens Ley; H. tridentatum auct. Aphyllopodous, rarely hypophyllopodous perennial herb with a branched stock. Stem 50–100 cm, pale yellowishgreen, often suffused purplish, more or less robust, striate, with numerous, medium to long, pale, bulbous-based simple eglandular hairs, but often becoming scaberulous,

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with few stellate hairs and numerous minute glandular hairs above. Leaves 10–30, bright to deep green on upper surface, paler beneath, the lower sometimes tinted reddish, gradually decreasing in size upwards and often crowded, the lamina of the lower 5–9 × 2–3 cm, ovate to ovate-lanceolate, narrowed at apex, but obtuse-mucronate, denticulate to dentate and narrowed below to short petioles, the lamina of the median mostly ovate, acute at apex, usually with more or less numerous, sharp, unequal, ascending teeth, some very narrow, and interspersed with denticulations, attenuate below and usually sessile, the lamina of the upper ovate-lanceolate or lanceolate, gradually narrowed to an acute apex, usually with several, narrow teeth, all glabrous or with short, pale simple eglandular hairs on the upper surface, with short to medium, pale simple eglandular hairs beneath particularly on the midrib and the upper often with some stellate hairs, the margins with short simple eglandular hairs or scaberulous. Inflorescence with 2–50 capitula, paniculate-corymbose, in large plants often elongate and often with lower branches from the upper leaf axils; peduncles with numerous to dense stellate hairs and often an occasional simple eglandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.8–1.2 mm, olive green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an obtuse apex, with more or less numerous, short and medium, pale, dark-based simple eglandular hairs and numerous short, dark glandular hairs and microglands. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins dentate-fimbriate. Achenes 2.5–3.0 mm, blackish-brown. Flowers 7–9. Apomictic. H. eboracense is closely allied to H. calcaricola and H. trichocaulon, but differs from both in its broader, more dense leaves, more floriferous panicle and its fimbriate receptacle pits. Native. Open woodland, grassy places and heaths. It is a fairly frequent plant of south Wales, and south-eastern and northern England, with a few outlying localities in central England. Endemic. 26. H. cantianum F. Hanb. Kent Hawkweed H. laevigatum subsp. cantianum (F. Hanb.) Zahn Aphyllopodous or slightly pseudophyllopodous perennial herb with a branched stock. Stem 20–90 cm, pale yellowishgreen, often slightly purplish at base, more or less robust, striate, with dense, long, pale, wavy simple eglandular hairs below becoming less numerous above and often absent, with numerous stellate hairs in the upper part. Leaves 10– 20(–35), yellowish-green on upper surface, paler beneath and the lower sometimes tinted reddish, the lamina of the lower 4–9 × 2–4 cm, sometimes slightly aggregated, elliptical, oblong-elliptical or ovate, obtuse-mucronate to acute, denticulate to dentate, the teeth narrowly mammiform, narrowed below to short hairy petioles, the lamina of the remainder gradually decreasing in size upwards, elliptical, ovate or lanceolate, mostly acute at apex, irregularly dentate, the teeth often narrowly mammiform, sometimes even narrower and ascending, long to short cuneate at base and sessile, often with stellate hairs on both surfaces especially the upper, the lower with long, pale simple eglandular

hairs on the upper surface, the upper often without them, all with short to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with (5–)10–25(–35) capitula, narrowly paniculate-corymbose, with slender, erectspreading branches; peduncles with dense stellate hairs and occasional simple eglandular or glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–9 × 1.0–1.5 mm, olive green, the inner with paler margins, all narrowly linear-lanceolate, narrowed to an obtuse apex, with numerous, pale, dark-based, short to medium simple eglandular hairs, few to numerous short, dark glandular hairs and some microglands, without stellate hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 2.5–2.7 mm, blackish-brown. Flowers 6–8. 2n = 27, mostly univalents at meiosis (R. A. Finch). Native. Rides and roadsides in wooded areas. Kent, Surrey and Sussex. Endemic. 27. H. acamptum P. D. Sell & C. West Witley Hawkweed H. cantianum var. subrigidum E. F. Linton ex F. Hanb. Aphyllopodous or pseudophyllopodous perennial herb with a branched stock. Stem 40–60 cm, pale yellowish-green, often suffused purplish, slender but strict, striate, with some stellate hairs throughout, but numerous to dense in the upper part, dense, long, pale simple eglandular hairs below becoming sparse in the upper part and without glandular hairs. Leaves 10–16, dull green on upper surface, paler beneath and frequently suffused purplish, the lower often aggregated, the median and upper gradually decreasing in size, the lamina of the lower 5–10 × 2–3 cm, lanceolate, lanceolate-oblong, elliptical or oblong-elliptical, narrowed to an obtuse-mucronate apex, subentire, undulate-dentate or with several mammiform teeth and narrowed at base to a short, winged petiole, the lamina of the median and upper lanceolate or oblong-lanceolate, gradually narrowed at apex but the tip more or less obtuse, dentate and with several mammiform or narrowly aquiline-mammiform teeth on each side, the median cuneate based, the upper sometimes rounded; all glabrous or nearly so on upper surface or the lowest sometimes with a few short simple eglandular hairs, with few to numerous, short to medium simple eglandular hairs on the lower surface and numerous medium to long simple eglandular hairs on the margin and midrib beneath. Inflorescence of 15–25 capitula with small corymbs on long branches and usually a long lower branch; peduncles with dense stellate hairs, sometimes an occasional, pale simple eglandular hair, without glandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–9 × 1.0–1.3 mm, olive green, inner very pale, all linear-lanceolate, obtuse at apex, with numerous stellate hairs towards the base, numerous, medium, pale dark-based simple eglandular hairs and few, short, yellowish glandular hairs. Ligules yellow, with a few, very short simple eglandular hairs at apex. Styles yellowish. Receptacle pits with the margins shortly dentate. Achenes 2.5–3.0 mm, blackish-red. Flowers 6–8. Apomictic. Native. Known only from Church Lane, Witley in Surrey where it seems to have been last recorded in 1896. Endemic.

39. Hieracium 28. H. trichocaulon (Dahlst.) Johanss. Hairy-stemmed Hawkweed H. rigidum subsp. trichocaulon Dahlst.; H. laevigatum subsp. affine var. trichocaulon (Dahlst.) F. N. Williams; H. rigidum var. trichocaulon (Dahlst.) W. R. Linton; H. laevigatum subsp. trichocaulon (Dahlst.) Zahn; H. tridentatum auct. Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 30–90 cm, pale yellowish-green, often suffused reddish-purple throughout much of its length, more or less robust, striate, with long, pale simple eglandular hairs in the lower half but often becoming scabrous, with few stellate hairs and numerous, minute glandular hairs in the upper part. Leaves 8–20, gradually becoming smaller upwards, deep green on upper surface, the lower often flushed rosy purple, the basal not usually present at anthesis, the lamina 6–9 × 2.5–3.0 cm, elliptical or ovate, obtusemucronate to acute at apex, coarsely dentate, the teeth aquiline-mammiform, and cuneate at base to a short petiole, the lamina of the lower cauline 7–11 × 2–3 cm, narrowly to broadly elliptical, oblong-elliptical or ovate-lanceolate, subobtuse to acute at apex, denticulate to irregularly dentate, the teeth sometimes narrowly aquiline-mammiform and becoming very large in late-flowering plants, and narrowed at the base to short petioles, the lamina of the median and upper cauline lanceolate or oblong-lanceolate, gradually narrowed to an acute apex, denticulate or dentate and cuneate at the sessile base, all glabrous or with a few simple eglandular hairs on the upper surface, with few to fairly numerous, short to medium, pale simple eglandular hairs beneath and often with stellate hairs on the under surface of the upper, the margins with short simple eglandular hairs or scaberulous. Inflorescence with 2–20(–30) capitula, paniculate-corymbose, with suberect branches and often a long lower branch; peduncles with numerous to dense stellate hairs and numerous minute glandular hairs hidden amongst them, rarely with an occasional simple eglandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–10 × 1.0–1.5 mm, olive green, the inner with paler margins, all linearlanceolate, obtuse to subacute at apex, with few to numerous, short to medium, pale simple eglandular hairs, numerous, short and minute, dark glandular hairs and few or no stellate hairs. Ligules yellow, glabrous. Styles yellowish to discoloured. Receptacle pits with margins subulate-dentate. Achenes 2.5–3.0 mm, blackish-brown. Flowers 6–7(–9). Apomictic. Native. Sandy heaths, open woodland, shady roadsides and railway banks. Fairly common in southern England with a few outlying localities further north in England and Wales. Scandinavia. 29. H. calcaricola (F. Hanb.) Roffey Toothed Hawkweed H. rigidum var. calcaricola F. Hanb.; H. laevigatum subsp. calcaricola (F. Hanb.) Zahn; H. tridentatum forma glandulosiceps (Zahn) Pugsley; H. laevigatum subsp. tridentatum var. genuinum subvar. glandulosiceps Zahn; H. tridentatum var. polyphyllum (Zahn) Pugsley; H. laevigatum subsp. tridentatum var. polyphyllum Zahn; H. tridentatum auct.

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Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 25–100 cm, pale yellowish-green, sometimes brownish-purple in the lower half, slender to robust, sometimes flexuous, striate, with numerous to dense, short to long, pale simple eglandular hairs, often becoming scabrous, with few to numerous stellate hairs in the upper part. Leaves 8–30, fairly widely spaced and slowly decreasing in size upwards, bright or pale green on upper surface, paler beneath, sometimes tinted reddish, the lamina of the lower 3–10 × 1–2 cm, narrowly elliptical or lanceolate, gradually narrowed to a more or less acute apex, sharply dentate, often interspersed with denticulations and gradually narrowed at base to a short petiole, the lamina of the median similar, but very sharply dentate and sessile, the upper lanceolate or linear-lanceolate, sharply acute at apex, sharply dentate and shortly narrowed at the sessile base, all glabrous or with a few short or medium simple eglandular hairs on upper surface, with more or less numerous, short to medium simple eglandular hairs beneath and on the margin or scaberulous on the margin, sometimes with numerous stellate hairs beneath. Inflorescence with 4–20(–50) capitula, usually paniculate-corymbose, rarely with a lower branch; peduncles with dense stellate hairs and few to numerous, short to medium, pale simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.5 mm, dark olive green, the inner with paler margins, all linearlanceolate, obtuse to subacute at apex, usually with numerous, short and medium, pale, dark-based simple eglandular hairs and always numerous short, dark glandular hairs and microglands, sometimes with stellate hairs along the margin and at the base. Ligules yellow, glabrous. Styles yellow to discoloured. Receptacle pits with margins subulatedentate and sometimes slightly fimbriate. Achenes 2.5–3.0 mm, blackish-brown. Flowers 7–8. Apomictic. 2n = 27, mostly univalents at meiosis (R. A. Finch). This species includes most of the narrow-leaved plants which were included in British H. tridentatum. True H. tridentatum Fr. is a slender, short plant with eglandular involucral bracts which does not seem to occur in our area. Native. Grassy places and heathland. Scattered records in England and Wales; introduced in waste ground in Edinburgh. Probably widespread in Europe. 30. H. stewartii (F. Hanb.) Roffey Stewart’s Hawkweed H. friesii var. stewartii F. Hanb.; H. gothicum var. stewartii (F. Hanb.) F. Hanb.; H. rigidum var. stewartii (F. Hanb.) W. R. Linton; H. stewartii var. praegeri Pugsley; H. laevigatum subsp. stewartii (F. Hanb.) Zahn; H. ardaricum Pugsley; H. donegalense Pugsley; H. hartianum Pugsley; H. backhouseanum auct. Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 25–100 cm, pale yellowish-green, often suffused reddish-purple in the lower half, slender to robust, sometimes flexuous, striate, with few to dense, medium to long, pale simple eglandular hairs below becoming fewer or scaberulous upwards, with few to fairly numerous stellate hairs above. Leaves 10–20, gradually becoming smaller upwards, dull deep green on upper surface,

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paler and bluish beneath, the lower often suffused reddishpurple and sometimes aggregated, the lamina of the lower 6–11 × 2–5 cm, elliptical, broadly elliptical, lanceolate or ovate, obtuse-mucronate to acute, subentire to denticulate or dentate and cuneate or attenuate to a short petiole, the median and upper elliptical, ovate or lanceolate or linearlanceolate, more or less acute at apex, more or less dentate, sometimes with long, curved teeth and narrowed or rounded at the sessile base, all glabrous on the upper surface or the lower with a few, short, rigid simple eglandular hairs, with few to numerous, short to medium, pale simple eglandular hairs beneath and on the margin, or scaberulous, sometimes with stellate hairs on the lower surface of the upper. Inflorescence with 1–20 capitula, paniculatecorymbose, with long, spreading branches; peduncles with few to numerous stellate hairs and sometimes an occasional minute glandular or short simple eglandular hair. Capitula 35–45 mm in diameter, rounded or subtruncate at base. Involucral bracts incumbent in bud, 5–12 × 1.0–1.6 mm, dark olive green, the inner with paler margins, all broad linear-lanceolate, obtuse at apex, with more or less numerous short and very short dark glandular hairs, few to numerous, short to medium pale simple eglandular hairs, and more or less numerous stellate hairs towards the base. Ligules yellow, glabrous. Styles usually more or less yellow, sometimes discoloured. Receptacle pits with margins subulatedentate. Achenes 3–4 mm, blackish-brown. Flowers 7–8. Apomictic. As variable within localities as between localities, both in the wild and in cultivation, in the dentation of the leaves and the hairiness of the peduncles and involucral bracts. The short plants with fewer capitula occur on rocks in and by the streams, the taller plants are on the banks. Native. Rocky and grassy sides of streams and open woodland. Co. Donegal, Co. Down and Co. Derry in Ireland, Stirlingshire, Perthshire, Forfarshire, Aberdeenshire and Nairn in Scotland and Breconshire and Caernarvonshire in Wales. Named after Samuel Alexander Stewart (1826–1910). 31. H. aphyllopodioides F. N. Williams Broad-leaved Hawkweed H. gothicum var. latifolium Backh.; H. friesii var. latifolium (Backh. fil.) E. F. & W. R. Linton; H. gothicum forma latifolium (Backh. fil.) W. R. Linton; H. laevigatum subsp. backhouseanum Zahn; H. backhouseanum (Zahn) Roffey nom. illegit. Aphyllopodous or pseudophyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowishgreen, often suffused reddish, usually more or less robust, sometimes slender, striate, erect, with medium to long, pale, bulbous-based hairs throughout, sometimes becoming scaberulous above. Leaves 6–14(–20), gradually decreasing upwards or in a false rosette then abruptly decreasing upwards, dull medium green on upper surface, paler beneath, the lamina of the lower 4–8 × 2.0–3.5 cm, ovate, lanceolate or elliptical, obtuse-mucronate to more or less acute at apex, entire or minutely denticulate and cuneate or rounded at base to a short petiole, the lamina of the median lanceolate or ovate, acute at apex, subentire or sparsely

denticulate, cuneate at base and sessile or with a short winged petiole, the lamina of the upper lanceolate or linear, acute at apex, more or less entire and sessile; all glabrous or nearly so on upper surface, with numerous short to long, pale simple eglandular hairs beneath and on the margin and particularly on the midrib. Inflorescence with 6–12 capitula, paniculate-corymbose, with rather long, erecto-patent branches; peduncles with few to numerous stellate hairs and sometimes an occasional simple eglandular hair. Capitula 30–40 mm in diameter, large, broad and truncate-based. Involucral bracts incumbent in bud, 5–13 × 1.5–2.0 mm, dull blackish-green, the inner with paler margins, all broad linear-lanceolate, obtuse at apex, with scattered, minute glandular hairs and an occasional simple eglandular hair. Ligules pale yellow, glabrous. Styles yellow. Receptacle pits with margins fimbriate-dentate. Achenes 2.5–3.0 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Heathy hillocks and grassy cliffs. Clova Mountains, Forfarshire, near Uig in Skye and the coast of Sutherland and Caithness. Endemic. 32. H. scullyi E. F. Linton Scully’s Hawkweed H. carpathicum subsp. scullyi (E. F. Linton) Zahn Pseudophyllopodous perennial herb with a branched stock. Stem 30–60(–75) cm, pale yellowish-green, sometimes purplish at the base, slender to robust, flexuous, striate, with numerous to dense, long, pale, wavy, bulbous-based simple eglandular hairs below becoming sparse upwards, with minute glandular hairs and sparse stellate hairs in the upper part. Leaves 6–15(–30 cult.), deep green on upper surface, paler beneath and with prominent veins, the lower and basal usually aggregated, the lamina 3–9 × 1–4 cm, elliptical or sometimes nearly subrotund, rounded-mucronulate to shortly acute at apex, entire or denticulate (dentate in cultivation), and narrowed to short, winged petioles, the lamina of the remaining cauline gradually decreasing in size upwards, elliptical or ovate, more or less acute at apex, entire, denticulate or with a few large teeth, the lower cuneate at base, the upper rounded and sessile at base, all often glabrous on upper surface but sometimes the lower with short, rigid simple eglandular hairs, with numerous short to medium, pale simple eglandular hairs beneath and on the margins and a few stellate hairs beneath on the upper ones. Inflorescence with 2–12(–20 cult.) capitula, paniculate-corymbose, with erect-spreading branches; peduncles with stellate and minute glandular hairs and very occasional simple eglandular hairs. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts 10– 12 × 1.5–2.0 mm, blackish-green and slightly paler on the margins, all broadly linear-lanceolate, more or less obtuse at apex, with numerous microglands, fewer longer glandular hairs, a few, short or medium simple eglandular hairs, and a few stellate hairs at the base. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Known only from rocks by the Roughty River in Co. Kerry, Ireland. Endemic. Named after Reginald William Scully (1858–1935).

39. Hieracium 33. H. scabrisetum (Zahn) Roffey Scabrous Hawkweed H. laevigatum subsp. scabrisetum Zahn; H. tridentatum var. setigerum Ley; H. rigidum var. rubefactum E. F. Linton; H. rubefactum (E. F. Linton) Roffey; H. melanocephalum Lindeb., non Tausch; H. septentrionale Arv.-Touv., non Norrl.; H. laevigatum subsp. boreophilum Zahn; H. boreophilum (Zahn) Roffey; H. cambricogothicum var. glandulosum Pugsley; H. fragilicaule Pugsley; H. fragilicaule forma subhirsutum Pugsley; H. backhouseanum var. radnoricum Pugsley; H. pseudacrifolium Pugsley; H. rhayaderense Pugsley; H. septentrionale var. amphibolum auct.; H. angustatum subsp. amphiboloides auct.; H. rigidum var. pullatum auct.; H. tridentatum var. acrifolium auct. Aphyllopodous, pseudophyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 20–70 cm, pale yellowish-green, sometimes purplish in lower half, slender to more or less robust, striate, with numerous to dense, medium to long, pale, bulbous-based simple eglandular hairs in lower part, becoming fewer upwards and sometimes part or all of the stem becoming scaberulous, with more or less numerous minute glandular hairs and numerous stellate hairs in the upper part. Leaves 4–20, gradually decreasing in size upwards, sometimes aggregated near the base, dull green on upper surface, paler beneath, rarely tinted reddish, the lamina of the lower narrowly elliptical, ellipticlanceolate, elliptic-oblong or lanceolate, obtuse-mucronate to acute at apex, subentire, denticulate or remotely dentate, and narrowed at base to a very short petiole or sessile, the lamina of the upper lanceolate, oblong-lanceolate or linearlanceolate, acute at apex, entire to sharply dentate, cuneate or rounded at base and sessile, all glabrous on upper surface or the lower with few to numerous, bulbous-based, short, stiff simple eglandular hairs, with short to long, numerous, pale simple eglandular hairs beneath and on the margin, sometimes with stellate hairs beneath or on both surfaces. Inflorescence with 1–20(–30) capitula, paniculatecorymbose, in large plants apical part sometimes subumbellate, in a tight corymb or with erect-spreading branches; peduncles with numerous to dense stellate hairs, numerous minute glandular hairs and sometimes a few larger ones, and few to numerous, short to medium, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–10 × 1.0–1.5 mm, dark olive to blackish-green, linear-lanceolate, obtuse to subacute at apex, with more or less numerous, very unequal, short to medium, slender dark glandular hairs and very few to fairly numerous, short to medium simple eglandular hairs, without stellate hairs. Ligules yellow, glabrous. Styles yellowish to discoloured. Receptacle pits with margins sharply dentate. Achenes 2.5–3.5 mm, blackish-brown. Flowers 6–8. Apomictic. Variable mainly in size and number of leaves and capitula. Plants growing on rocks in streams are short with few leaves and few capitula (H. fragilicaule), plants in better soil in vegetation are much taller with up to 20 leaves and capitula (H. scabrisetum) while those on the river bank are intermediate (H. pseudacrifolium). All forms can be recognised by the very unequal, slender glandular hairs of the involucral bracts.

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Native. Grassy and rocky places, particularly by streams. Widespread in Wales, with an adjacent locality in Herefordshire, with outlying localities at Chapel-en-le-Frith in Derbyshire, Glen Devonshire in Perthshire and between Kingussie and Kincraig in Inverness-shire. Norway. 34. H. placerophylloides Pugsley Purplish-leaved Hawkweed H. sparsifolium var. placerophyllum auct., non H. sparsifolium var. placerophyllum Dahlst. ex W. R. Linton; H. levigatum subsp. placerophyllum auct., non (Dahlst. ex W. R. Linton) Zahn; H. placerophyllum auct., non (Dahlst. ex W. R. Linton) Roffey; H. sparsifolium var. grandescens auct., non Dahlst. ex W. R. Linton; H. levigatum subsp. grandescens auct., non (Dahlst. ex W. R. Linton) Zahn; H. grandescens auct., non (Dahlst. ex W. R. Linton) Roffey Aphyllopodous or pseudophyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale yellowish-green, often suffused deep brownish-purple, slender to robust, flexuous, striate, with numerous to dense, long, pale, wavy simple eglandular hairs below, decreasing in number upwards and often absent in the upper part, and with few to numerous stellate hairs and numerous minute glandular hairs in the upper part. Leaves 6–12(–15) rather deep dull green on upper surface, paler beneath, often suffused purplish, the lamina of the lower 8–11 × 2–3 cm, lanceolate, oblonglanceolate to narrowly elliptical, narrowed to an obtusemucronate or subacute apex, subentire, shallowly undulatedentate or with a few, sharp, narrow teeth and narrowed below to slender petioles; the lamina of the median smaller lanceolate or oblong-lanceolate, long drawn out to an obtuse or acute apex, dentate, the teeth sometimes long, curved and narrow, and cuneate to a sessile or shortly petioled base, the lamina of the upper narrowly lanceolate, acute or acuminate, entire or denticulate or dentate near the base, sessile, all glabrous or with scattered, short or medium, pale simple eglandular hairs on the upper surface, with few to numerous, medium to long, pale simple eglandular hairs beneath especially on the midrib, with few to numerous, short to long, pale simple eglandular hairs on the margin or scaberulous with the bases of broken hairs. Inflorescence of (2–)5–16 capitula, usually paniculate-corymbose, sometimes with long lower branches; peduncles with numerous to dense stellate hairs, few to numerous, short to medium, pale, dark-based simple eglandular hairs and numerous minute glandular hairs often difficult to see when stellate hairs are dense. Capitula 30–45 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.5–2.0 mm, blackish-green, the inner with paler margins, all linearlanceolate, obtuse at apex, with few stellate hairs near the base, numerous, short to medium, pale, dark-based simple eglandular hairs intermixed with few, short, dark glandular hairs. Ligules yellow, glabrous. Styles yellowish or slightly discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 7–8. Apomictic. The original publication of both H. sparsifolium var. placerophyllum and var. grandescens includes both Scandinavian and British plants. In both cases the varieties are typified by H. Dahlstedt’s Scandinavian specimens. The British

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plants which have been placed under both varieties are not the Scandinavian plants and both belong to H. placerophylloides. Native. Rocky streamsides, cliff ledges and among boulders, apparently more or less confined to the limestone. Fairly common in northern England and more local in Wales. Endemic. 35. H. substrigosum (Zahn) Roffey Long-haired Hawkweed H. rigidum var. strigosum Ley; H. sparsifolium var. strigosum (Ley) W. R. Linton; H. laevigatum subsp. substrigosum Zahn; H. friesii var. hirsutum F. Hanb.; H. rigidum var. longiciliatum (F. Hanb.) W. R. Linton; H. laevigatum subsp. longiciliatum (F. Hanb.) Zahn; H. longiciliatum (F. Hanb.) Roffey; H. lapponicum auct.; H. sparsifolium var. longiciliatum (F. Hanb.) W. R. Linton Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 30–100 cm, pale yellowish-green, often suffused reddish-purple in the lower half, slender to robust, often flexuous, striate, with dense, long, pale simple eglandular hairs sometimes becoming scaberulous with wear. Leaves 6–15, rather pale yellowish-green on upper surface, paler beneath, the basal when present with lamina 5–8 × 2.0–2.5 cm, elliptical, more or less obtuse at apex, denticulate to shortly dentate, cuneate at base and shortly petioled; the lamina of the lower and median 5–11 × 1–3 cm, elliptical, oblong-lanceolate or lanceolate, mostly acute at apex, with a few coarse, unequal teeth, sometimes only denticulate, and sessile or shortly petiolate; the lamina of the upper linear-lanceolate or linear, acute at apex, usually with a few teeth but sometimes laciniate and sessile, all with few to numerous, short to long, pale, bulbous-based hairs on both surfaces and the margins. Inflorescence with 3–20 capitula, with long branches and acladium forming a loose paniculate corymb; peduncles with numerous stellate hairs and numerous, short and medium, pale, dark-based simple eglandular hairs. Capitula 25–35 mm in diameter, rounded or subtruncate at base. Involucral bracts incumbent in bud 4–11 × 1.0–1.5 mm, dark greyish or blackish-green, linearlanceolate, obtuse at apex, with numerous to dense, short to long, pale, dark-based simple eglandular hairs and a few, short, dark glandular hairs near the base, without stellate hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Rocky and grassy places, usually by streams. Fairly frequent in Wales, particularly in Breconshire. Plants from by the River Clunie at Braemar, Aberdeenshire on which H. longiciliatum was based seem indistinguishable and other rather poor specimens from Perthshire seem to be the same species. Endemic. 36. H. charitodon P. D. Sell Slender-toothed Hawkweed Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 35–100 cm, pale yellowish-green, often purplish at the base, slender to robust, with dense, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs in the upper part. Leaves pale to medium green on upper surface,

paler beneath, 5–15, gradually decreasing in size upwards; basal and lower cauline with lamina 6–11 × 1.5–4.0 cm, narrowly elliptical, oblong-elliptical or lanceolate-oblong, gradually narrowed to an obtuse-mucronate or acute apex, dentate, with mammiform to rather long, narrow teeth, attenuate at base to a winged petiole up to 4 cm; median cauline with lamina 4–9 × 1.5–4.0 cm, linear-elliptical or oblong-elliptical, gradually narrowed to a more or less acute apex, dentate, with long narrow teeth up to 14 mm, gradually attenuate at base, sessile or with a short petiole; upper cauline with lamina linear or linear-lanceolate, long-acute at apex, entire or with narrow teeth, sessile; all glabrous or with some very short hairs on upper surface, with numerous, short to medium, pale simple eglandular hairs beneath and on the margin. Inflorescence with 1–21 capitula, paniculatecorymbose, often with long lower branches; peduncles slender and erect, with numerous, short to medium, pale, sometimes dark-based simple eglandular hairs and dense stellate hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.2–1.5 mm, olive to blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous to dense, short to medium, pale, often dark-based simple eglandular hairs, few, short, dark glandular hairs and a few stellate hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins fibrillous-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. Riverside rocks. Upper Tawe Valley, Cwm Tarell and Mellte Glen in Breconshire. Endemic. When describing H. rigidum var. strigosum (i.e. H. substrigosum) A. Ley mentioned a plant gathered at Glyn Tarell in 1883, which he included in his new variety but mentioned a number of differences. The specimens from Glyn Tarell in his herbarium are this species. The description of his new variety fits the other specimens he listed and from them the type of H. rigidum var. strigosum has been chosen. The long narrow teeth of H. charitodon become even more marked in cultivation. 37. H. gothicoides Pugsley Broad-headed Hawkweed H. gothicum auct.; H. gothicum var. basifolium auct.; H. gothiciforme auct.; H. rigidum var. longiciliatum auct. Aphyllopodous or pseudophyllopodous perennial herb with a branched stock. Stem 30–80 cm, pale yellowish-green, sometimes purplish near the very base, slender to robust, striate, with numerous to dense, medium to long, pale simple eglandular hairs below, becoming less numerous and often absent upwards, with stellate and minute glandular hairs above. Leaves 5–16, rather pale yellowish-green on upper surface, paler and the lowest sometimes tinted reddish beneath, the lower often slightly aggregated, the lamina of the lower 4–11 × 1.5–2.5 cm, narrowly elliptical, ellipticoblong or lanceolate, narrowed to an obtuse-mucronate or more or less acute point, usually sharply denticulate, sometimes with larger teeth and narrowed below to a short, winged petiole, the lamina of the upper gradually decreasing in size upwards or suddenly decreasing in size halfway up, lanceolate, oblong-lanceolate or the uppermost linear, usually acute at apex and denticulate or serrate-dentate, the uppermost sometimes entire or nearly so, all glabrous

39. Hieracium or with occasional simple eglandular hairs on upper surface, with few to numerous, short to medium, pale simple eglandular hairs beneath and on the margin, often with stellate hairs beneath or on both surfaces. Inflorescence with 2–10(–25) capitula, paniculate-corymbose, narrowly compact, with slender, suberect branches; peduncles with dense stellate hairs hiding minute glandular hairs, sometimes with an occasional simple eglandular hair. Capitula 30– 40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.5 mm, blackish-green, the innermost with pale margins, all linear-lanceolate, more or less acute at apex, with more or less numerous, short to medium, pale, dark-based simple eglandular hairs, without glandular or stellate hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Grassy and rocky places. A frequent species of central Scotland with isolated records in Westmorland, Kirkcudbrightshire, Wigtownshire and the Shetland Islands. Endemic. 38. H. cambricogothicum Pugsley Llanfairfechan Hawkweed Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 20–50(–100 cult.) cm, pale yellowishgreen, often suffused with brownish-purple, more or less robust, striate, flexuous but rigid, with dense, medium to long, pale, wavy simple eglandular hairs near the base which become steadily less numerous upwards until they are almost absent in the upper part, with numerous stellate hairs in the upper part, without glandular hairs. Leaves 10–15(–25), deep green on upper surface, paler and sometimes suffused purplish beneath, the lower sometimes aggregated, the lamina of the lower 5–7 × 2–3 cm, elliptical to ovate-lanceolate, more or less obtuse-mucronate at apex, irregularly serrate-denticulate with small mucronulate teeth and sessile with a shortly cuneate to rounded base, the lamina of the upper gradually becoming smaller, elliptical or lanceolate, obtuse to acute at apex, with a few, narrow mucronulate teeth and sessile, all glabrous or with short to medium simple eglandular hairs and stellate hairs on the upper leaves beneath, and numerous medium to long simple eglandular hairs on the margin. Inflorescence with 5– 16(–40 cult.) capitula, narrowly paniculate with long lower branches, the upper part often subumbellate; peduncles slender, with dense stellate hairs, and glandular or simple eglandular hairs. Capitula 20–25 mm in diameter, rounded to subtruncate at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.3 mm, blackish-green, at least the inner with a paler margin, all linear-lanceolate, obtuse at apex, with a few, very short glandular hairs along the median line, without simple eglandular or stellate hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, blackish. Flowers 7–8. Apomictic. Native. This species occurs in two widely separated areas. The type material comes from roadside cliffs at Llanfairfechan in Caernarvonshire and is almost certainly native. In Kent it is well established on some old ragstone walls and

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waste ground above Riverhead by Sevenoaks, where it is almost certainly introduced. Endemic. 39. H. uiginskyense Pugsley Uig Hawkweed H. trinitatis Pugsley; H. gothicum auct.; H. laevigatum subsp. subgracilipes auct. Aphyllopodous or pseudophyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, often flushed reddish-purple towards the base, slender to more or less robust, flexuous, striate, with numerous to dense, medium to long, pale, bulbous-based simple eglandular hairs particularly in the lower half, sometimes nearly glabrous above, with scattered stellate hairs, and minute, yellowish glandular hairs above. Leaves 5–14, quickly becoming small upwards, rather pale yellowishgreen on upper surface, paler and sometimes flushed purplish beneath, the lamina of the lower 4–8 × 1.5–3.0 cm, elliptical, elliptic-lanceolate, elliptic-oblong or lanceolate, obtuse-mucronate at apex, subentire to sparsely dentate, cuneate to attenuate at base and with short to medium, long-hairy petioles, the lamina of the median and upper lanceolate, acute at apex, denticulate to dentate, cuneate at base and sessile, all glabrous or with a few simple eglandular hairs on the upper surface, with few to numerous, medium to long, pale simple eglandular hairs especially on the midrib and few to numerous stellate hairs beneath, often scaberulous on the margin. Inflorescence with 2–5(–10) capitula, narrowly and compactly corymbose; peduncles with dense stellate hairs, sometimes with a few, medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–11 × 1.5–2.0 mm, blackish-green, the inner with paler margins, all broad linear-lanceolate, obtuse at apex, with numerous, short to medium, pale simple eglandular hairs and dense dark microglands, stellate hairs mainly near the base. Ligules yellow, glabrous. Styles yellow. Receptacle pits with fimbriate-dentate margins. Achenes 2.5–3.0 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Grassy and rocky banks of streams, cliffs and grassy slopes. Breconshire, Radnorshire, Carmarthenshire and Merionethshire in Wales, the Tarn Beck above Seathwaite in Cumbria, central and western Scotland and Co. Donegal and Co. Antrim in Ireland. Endemic. 40. H. lissolepium Roffey Hairless-bracted Hawkweed H. lineatum Almq. ex Stenstr¨om, non Arv.-Touv.; H. rigidum subsp. lineatum Dahlst.; H. laevigatum subsp. lissolepium Zahn nom. illegit.; H. calviceps Pugsley; H. calviceps var. robustum Pugsley; H. gothicum auct.; H. longiramosum Pugsley Aphyllopodous or pseudophyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale yellowish-green, often suffused reddish-purple, slender to more or less robust, erect, striate, strict, with a few, scattered stellate and occasional simple eglandular hairs or more numerous in robust plants. Leaves 6–15, rather pale yellowishgreen, sometimes slightly suffused reddish, either gradually decreasing in size upwards or abruptly decreasing in size halfway up the stem, sometimes aggregated below to form a pseudorosette, the lamina of the lower 5–10 × 1.5–2.5 cm,

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narrowly elliptical to lanceolate, more or less acute at apex, usually subentire to denticulate, rarely shortly dentate, and gradually narrowed at base to a short petiole, the lamina of the median lanceolate or ovate-lanceolate, acute or acuminate at apex, denticulate or shortly serrulate, and shortly narrowed or rounded at the sessile base, the lamina of the upper lanceolate, linear or bract-like, acute at apex, entire to denticulate and sessile, with few to numerous stellate hairs on both surfaces and a few, short, pale simple eglandular hairs, the margins very shortly ciliate. Inflorescence with 1–5(–18) capitula, in a compact corymb; peduncles with numerous stellate hairs, rarely with a simple eglandular hair. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linearlanceolate, gradually narrowed to an obtuse or subacute apex, glabrous or with occasional stellate or simple eglandular hairs. Ligules yellow, glabrous. Styles yellow, rarely discoloured. Receptacle pits with the margins dentatefimbriate. Achenes 2.5–3.0 mm, blackish-brown. Flowers 6–8. Apomictic. Native. Rocky streamsides, grassy slopes and rocky outcrops. The headquarters of this species is the northern Pennines, the Lake District and southern Scotland, but there are records for north and south Wales and isolated specimens from the north-east end of Loch Tummel in Perthshire and the Shetland Islands. It occurs also in Scandinavia and central Europe. The Shetland plants have dark styles, but no other character can be found to distinguish them. 41. H. subintegrifolium Pugsley Glenridding Hawkweed Aphyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often suffused purplish below, slender to robust, striate, with numerous medium to long, pale simple eglandular hairs at least below, becoming scaberulous above and with numerous, small stellate hairs above. Leaves 5–10(–16), dark green on upper surface, paler beneath with conspicuous venation, the lamina of the lower 7–14 × 1.5–2.5 cm, lanceolate, linear-lanceolate or oblonglanceolate, more or less acute at apex, subentire, remotely denticulate or with an occasional larger tooth, long attenuate at base to a short or medium petiole; the lamina of the upper lanceolate, long-acute or acuminate at apex, entire, denticulate or with a few larger teeth and cuneate to a sessile base; all with numerous, short to medium, pale, bulbousbase hairs on both surfaces or more or less glabrous on the upper surface. Inflorescence with 3–12 (numerous in cult.) capitula, paniculate-corymbose, with long, spreading branches; peduncles with numerous to dense stellate hairs and few, short to medium simple eglandular hairs, without glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 0.8–1.2 mm, blackish-green, the inner with paler margins, all linearlanceolate, more or less obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short glandular hairs and numerous microglands, without stellate hairs. Ligules glabrous. Styles more or less yellow to slightly discoloured. Receptacle pits with margins

incise-dentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 7–8. Apomictic. Native. Discovered by H. W. Pugsley in limited quantity on grassy banks in Glenridding in Westmorland in 1927 and seen again in the same locality by J. E. Raven in 1953. It has not been found since despite much searching. Endemic. 42. H. sparsifolium Lindeb. Sparse-leaved Hawkweed H. stictophyllum Dahlst.; H. laevigatum subsp. sparsifolium (Lindeb.) Zahn; H. laevigatum subsp. stictophyllum (Dahlst.) Zahn; H. laevigatum subsp. phyllopodioides var. gothicum forma centonata F. N. Williams; H. rigidum var. serpentinum F. Hanb.; H. stictophyllum var. serpentinum (F. Hanb.) W. R. Linton; H. laevigatum subsp. stictophyllum var. serpentinum (F. Hanb.) Zahn; H. stictophyllum var. concolor Pugsley; H. pseudoprotractum Pugsley, non Notø; H. protractum var. shetlandicum Dahlst.; H. faeroense Dahlst.; H. epileucoides Dahlst.; H. norvegicum var. confertum auct.; H. protractum auct. Aphyllopodous, pseudophyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 20–80 cm, pale yellowish-green, often suffused with reddish-purple, slender to robust, striate, usually with numerous, medium to long, pale simple eglandular hairs in the lower half, glabrous or becoming so above and sometimes becoming so below. Leaves 4–18, dull or bluish green, usually richly blotched and spotted brownish-purple on upper surface, paler and flushed reddish-purple beneath, some colonies without markings, the lamina of the lower 3–11 × 1–3 cm, narrowly elliptical, oblong-elliptical or spathulate, obtusemucronate to subacute at apex, entire or with an occasional small tooth and narrowed below to a short petiole, the lamina of the remainder gradually or abruptly decreasing in size upwards, narrowly oblong-elliptical, oblong, linear or linear-lanceolate, more or less acute at apex, entire or with sparse, short teeth, sessile and sometimes slightly semiamplexicaul, all glabrous on upper surface, with short to medium, pale simple eglandular hairs beneath and on the margin and sometimes some stellate hairs. Inflorescence with 2–20 capitula, paniculate-corymbose, with slender, erect-spreading branches; peduncles with numerous to dense stellate hairs and sometimes few to numerous, short to medium, pale simple eglandular hairs and an occasional short glandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.5 mm, olive or blackish-green, linearlanceolate, obtuse to subacute at apex, with few stellate hairs, more or less numerous, short to medium, pale, darkbased simple eglandular hairs and numerous, short, dark glandular hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 7–8. Apomictic. Most plants of H. sparsifolium can be easily recognised by their narrow, subentire, spotted and blotched leaves. Some plants occur without the spotted and blotched leaves. They may form distinct colonies or be mixed with the blotched plants. Some Welsh plants, var. serpentinum, have broader leaves and larger teeth but are mixed with normal plants. Shetland and Faeroes plants tend to have more hairy

39. Hieracium peduncles, but similar plants are found elsewhere, particularly in Wales. In cultivation all the variants get larger with longer, more pointed leaves and have no simple eglandular hairs on the peduncles. Native. Grassy banks, streamsides, rocks, cliff ledges and slate debris. Fairly common in north and south Wales, northern England and Scotland and in a few localities in western Ireland. Iceland, Norway and Faeroes. 43. H. nidense (F. Hanb.) Roffey Nidd Hawkweed H. rigidum var. nidense F. Hanb.; H. laevigatum subsp. nidense (F. Hanb.) Zahn Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 35–90 cm, pale yellowish-green, often purplish below, slender to more or less robust, flexuous or rigid, striate, with numerous, medium to long, pale simple eglandular hairs near the base, becoming few or absent upwards and with few stellate hairs in the upper part. Leaves 8–15, pale to rather bluish-green on upper surface, paler and more bluish beneath, sometimes flushed purplish, the lamina of the lower 5–12(–15 cult.) × 1.0–2.5(–5.0 cult.) cm, lanceolate or oblong-lanceolate, acute at apex, dentateserrate, with a few, unequal, sharp, narrow teeth and narrowed at base to a short, winged petiole, the lamina of the upper gradually smaller, linear-lanceolate, long-acute, with a few, sharp teeth, cuneate at base and sessile, with short, stiff, pale simple eglandular hairs on both surfaces and the margins or the upper more or less glabrous on the upper surface, the upper sometimes also with stellate hairs on the lower surface. Inflorescence with 2–9(–20 cult.) capitula, narrowly paniculate-corymbose, with long, slender branches; peduncles with dense stellate hairs, an occasional, medium, pale simple eglandular hair and rarely a solitary glandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts more or less porrect, 5–12 × 0.8–1.5 mm, dark olive green, at least the inner with paler margins, all narrowly linear-lanceolate, obtuse at apex, with a few stellate hairs, numerous short glandular hairs and microglands and few, short to medium simple eglandular hairs. Ligules glabrous. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, blackish-brown. Flowers 6–8. Apomictic. Native. Riverside rocks and cliff ledges. Mellte, Hepste, Taf-fechan, Haffes and Tawe Glens in Breconshire, the Wye in Radnorshire and Llyn-y-fan-fechan in Carmarthenshire. Endemic. 44. H. linguans (Zahn) Roffey Tongue-leaved Hawkweed H. sparsifolium var. lingua Ley; H. laevigatum subsp. linguans Zahn Aphyllopodous perennial herb or basal leaves withering at the time of flowering. Stem 30–60 cm, slender, flexuous, often purplish towards the base, with scattered short and medium, pale simple eglandular hairs particularly towards the base and numerous, minute, pale glandular hairs throughout, stellate hairs few or absent. Leaves medium, slightly bluish-green on upper surface, paler and often tinted purplish beneath, the basal with lamina 30–80 × 15–30 mm, oblanceolate to narrowly obovate, rounded-mucronulate at

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apex, denticulate or with an occasional tooth and cuneate at base, the cauline 5–10, 1.5–13.0 × 0.2–4.0 cm, middle leaves largest, with lamina oblong-lanceolate, narrowly elliptical, linear-lanceolate or lanceolate, gradually narrowed towards a long rounded-obtuse or acute apex, asymmetrically dentate with a few, aquiline-mammiform teeth or denticulations and cuneate to a sessile base, the lower sometimes with a short petiole, with short, stiff, medium simple eglandular hairs on the basal leaves, glabrous or nearly so on upper surface of cauline leaves, with scattered very short, pale simple eglandular hairs or bases of hairs on the lower surface and margin. Inflorescence with 2–8 capitula, in a narrow cyme, peduncles medium to long, straight, suberect, with dense stellate and pale minute glandular hairs mixed with an occasional medium, pale simple eglandular hair. Capitula 30–55 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 8–12 × 1.0–1.7 mm, greyish-green, the inner with paler margins, all broad linear-lanceolate, more or less obtuse at apex, with dense stellate hairs, numerous short, dark glandular hairs and few to numerous, short and medium simple eglandular hairs. Ligules irregularly dentate, yellow, glabrous. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 0.5–4.0 mm, dark. Flowers 6–7. Apomictic. Native. Known only from rocks by a waterfall in Haffes Glen, Breconshire. Endemic. 45. H. sparsifrons P. D. Sell & C. West Kerry Hawkweed H. sparsifolium var. oligodon E. F. Linton; H. oligodon (E. F. Linton) Pugsley, non N¨aegeli & Peter; H. subintegrum auct. Aphyllopodous perennial herb or basal leaves withered at the time of flowering. Stem 20–55 cm, robust, often purplish towards the base, with medium to long, pale simple eglandular hairs, frequent below, becoming sparse upwards, scattered stellate hairs throughout and minute glandular hairs in the upper part. Leaves dull dark green on upper surface, paler and slightly bluish beneath, the basal few to numerous, the lamina 4–6 × 1–2 cm, narrowly elliptical, acute at apex, with a few denticulations or small teeth, gradually attenuate below to a short petiole, the cauline 4–12, the lamina 1–12 × 0.2–2.0 cm, gradually decreasing in size upwards, the lower oblong-elliptical to narrowly oblong-lanceolate, the upper lanceolate to linear, mostly acute at apex, subentire, denticulate or with few, irregular teeth, narrowed to a sessile base, glabrous or nearly so above, with scattered, short or medium, pale simple eglandular hairs or hair bases beneath and on the margin and sometimes a few stellate hairs on the upper leaves. Inflorescence with 1–7(–10) capitula, laxly and narrowly paniculate; peduncles medium to long, suberect, with numerous stellate and pale, minute glandular hairs and occasionally a few, medium, pale simple eglandular hairs. Capitula 22–35 mm in diameter, narrowed at base. Involucral bracts incumbent in bud, 9–11 × 1.0– 1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse to subacute at apex, with numerous stellate hairs particularly towards the base, scattered short or medium, dark-based simple eglandular hairs and few, short, dark glandular hairs. Ligules yellow, glabrous. Styles

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yellow. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, dark. Flowers 7–8. Apomictic. Native. Known only from the rocky banks of the Clydagh and Roughty Rivers in Co. Kerry in the south of Ireland. Endemic. 46. H. hibernicum F. Hanb. Irish Hawkweed H. norvegicum subsp. hibernicum (F. Hanb.) Zahn; H. hibernicum var. vennicniorum Pugsley Aphyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, sometimes suffused purplish below, rather slender, flexuous, striate, nearly glabrous, or with few to fairly numerous, pale simple eglandular hairs below, and fairly numerous stellate hairs above. Leaves 7–10, bluish-green on upper surface, paler beneath, the lamina of the lower 7–8 × 2–3 cm, oblong, obtusemucronulate at apex, entire or remotely denticulate and gradually narrowed below to a more or less long, winged, semiamplexicaul petiole, the lamina of the median long (–15 cm), narrowly lanceolate or linear-lanceolate, acute at apex, remotely denticulate or with 2–3 sharp teeth on each side and more or less attenuate below, the lamina of the upper linear-lanceolate, decreasing in size to bracts, acute at apex, entire and sessile, glabrous or with a few, short pale simple eglandular hairs and stellate hairs beneath, margins thickened and scaberulous. Inflorescence usually with 1–3(–6) capitula but in var. vennicniorum up to 22; peduncles with numerous stellate hairs, without glandular hairs, without or with few, short, pale simple eglandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 6–12 × 1.0–1.5 mm, dark green, linear-lanceolate, obtuse at apex, with a few stellate hairs on the margins, few to numerous, short, dark glandular hairs and few to numerous, medium, dark simple eglandular hairs. Ligules yellow, glabrous. Styles yellowish. Receptacle pits with serrate-dentate margins. Achenes 3.5–4.0 mm, blackish. Flowers 7–8. Apomictic. Native. Rare in rocky places in the Mourne Mountains, Co. Down and Moynalt, near Laghy, Co. Donegal. Endemic. Section 5. Prenanthoidea W. D. J. Koch Series Prenanthoidea (W. D. J. Koch) P. D. Sell & C. West comb. inval.; Section Euprenanthoidea Gremli nom. illegit. Aphyllopodous, rarely pseudophyllopodous perennial herbs with branched stock. Stems 30–100 cm, glabrous or nearly so below, more or less hairy above. Leaves 6–30, more or less heterophyllous, gradually decreasing in size upwards, the lower narrowed at base and shortly petiolate, the upper sessile and more or less amplexicaul, nervation reticulate. Inflorescence paniculate-corymbose; peduncles with numerous glandular hairs. Capitula small and narrow. Involucral bracts 3–10 × 0.8–1.5 mm, incumbent in bud, appearing to be almost in 2 rows, dark linear-lanceolate to lanceolate, obtuse at apex, stellate hairs on margin, very glandular-hairy. Ligules hairy at tips. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes pale brown. Flowers 7–9.

This distinct section is recognised by its numerous more or less amplexicaul cauline leaves and numerous, small, narrow capitula clothed with glandular hairs. In Great Britain the species are found on rocky banks, grassy places and by streams, but in Europe and western Asia the plants are mainly inhabitants of mountain woods. 47. H. prenanthoides Vill. Rough-leaved Hawkweed H. prenanthoides subsp. prenanthoides; H. prenanthoides subsp. strictissimum auct.; H. subelatum auct. Aphyllopodous perennial herb with a branched stock. Stem 40–100 cm, pale yellowish-green, sometimes purplish in the lower half, slender to robust, striate, more or less glabrous below, with few to fairly numerous, short to medium, pale dark-based simple eglandular hairs and sometimes a few, very short, dark glandular hairs in the upper part. Leaves 12–30, dull medium or yellowish-green on upper surface, paler and rather greyish-green with reticulate venation beneath, the lamina of the lower 6–11 × 1.5–3.0 cm, narrowly to broadly elliptical, oblong-spathulate or oblanceolate, obtuse-mucronulate at apex, entire or remotely denticulate, gradually narrowed below to a short, broadly winged petiole, the lamina of the remaining cauline gradually decreasing in size upwards, lanceolate, narrowly elliptical, oblong-lanceolate and ovate, mostly acute at apex, entire to denticulate, sometimes with a few larger teeth, cordate-semiamplexicaul at base, all glabrous or nearly so on both surfaces, or with scattered to numerous, short, pale simple eglandular hairs on one or both surfaces and with numerous short to medium simple eglandular hairs on the margin and midrib beneath. Inflorescence with few to many capitula, paniculate-corymbose, sometimes with a long, lower branch; peduncles with dense stellate hairs, numerous, equal or unequal, very short and short, slender, dark glandular hairs and sometimes an occasional dark or dark-based simple eglandular hair. Capitula 15–20 mm in diameter, with a rounded base. Involucral bracts incumbent in bud, 3–10 × 0.8–1.3 mm, dark olive green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with few to numerous stellate hairs on the margin, numerous more or less equal or unequal, dark or yellowish, short glandular hairs and sometimes an occasional, dark, medium simple eglandular hair. Ligules yellow, with very short simple eglandular hairs at the apex. Styles discoloured. Receptacle pits with subulate-dentate margins. Achenes 3.5–4.0 mm, pale brown. Flowers 7–9. Apomictic. Variable in leaf toothing and hairiness and colour and size of glandular hairs on the involucral bracts. Native. Rocky banks, grassy places and streamsides. South Wales, Derbyshire, northern England, south and central Scotland, north to Braemar with an isolated record on boulder clay at Red Bay, Co. Antrim. In England and Wales it is usually on limestone. French and Swiss Alps, and Scandinavia. 48. H. lanceolatum Vill. Braemar Hawkweed H. prenanthoides subsp. lanceolatum (Vill.) Zahn; H. prenanthoides subsp. strictissimum auct. Aphyllopodous perennial herb with a branched stock. Stem 40–70 cm, pale yellowish-green, sometimes suffused

39. Hieracium brownish-purple, slender to robust, rigid, striate, with few, medium to long, pale, dark-based hairs in the lower part, but more numerous above, and with few to numerous stellate hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath and reticulate-veined, the lamina of the lower 7–10 × 2–4 cm, oblanceolate-oblong, elliptical or ovate, more or less obtuse at apex, and entire or remotely denticulate, the remainder gradually decreasing in size, lanceolate, ovate or elliptical, acute at apex, all sessile and auriculate-amplexicaul, glabrous or nearly so on the surfaces, with numerous short and medium, pale, rather stiff simple eglandular hairs on the margins. Inflorescence with numerous capitula, compactly paniculate-corymbose, sometimes with lower branches; peduncles mostly short, with dense stellate hairs, numerous, unequal, short and very short, black glandular hairs and numerous, pale, dark-based, short and medium simple eglandular hairs. Capitula 15–20 mm in diameter, small, with a rounded base. Involucral bracts incumbent in bud, 3–10 × 1.0–1.5 mm, blackish, the inner with paler margins, all lanceolate, obtuse at apex, with numerous stellate hairs on the margin, with numerous, short, black glandular hairs and few to numerous, dark-based simple eglandular hairs. Ligules yellow, with very short simple eglandular hairs at apex. Styles blackish. Receptacle pits with subulate-dentate margins. Achenes 3.5–4.0 mm, pale brown. Flowers 7–9. Apomictic. Native. This species was recorded by K. H. Zahn (1921–2) from Braemar. A series of specimens collected from the banks of the River Clunie at Braemar in Aberdeenshire seem to fit this species which differs from H. prenanthoides by its more rigid habit, numerous simple eglandular hairs in the inflorescence and almost black, broader involucral bracts. However, typical H. prenanthoides also grows at the same locality and the situation needs examining in the field. H. lanceolatum occurs widely in the mountains of much of Continental Europe. Zahn’s record for H. prenanthoides subsp. strictissimum (Froel.) Zahn for Braemar is probably also this species. The only specimens seen from Fortingal in Perthshire, the other locality given by Zahn for H. prenanthoides subsp. strictissimum, are typical H. prenanthoides. 49. H. borreri Syme Borrer’s Hawkweed H. denticulatum auct.; H. juranum subsp. pseudelatum auct. Aphyllopodous perennial herb or rarely with basal leaves withered at the time of flowering, with a branched stock. Stem 30–100 cm, pale yellowish-green, sometimes slightly reddish towards the base, slender to robust, striate, with few, medium to long, pale simple eglandular hairs scattered throughout and a few, short, slender glandular hairs in the upper part. Leaves bright bluish-green on upper surface, paler and sometimes tinted purplish beneath, with reticulate venation, the lamina of the basal 4–9 × 2.5–3.5 cm, lanceolate to narrowly elliptical or oblong-elliptical, rounded-obtuse-mucronulate to acute at apex, subentire to denticulate, cuneate at base and with the petioles up to 8 cm, which have pale, medium to long simple eglandular hairs, cauline 6–12(–17) the lamina of the lower 7–15 × 3–6 cm, gradually becoming smaller up the stem,

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elliptic, lanceolate or ovate, obtuse-mucronulate to acute, and entire to denticulate, the lower narrowed to a short petiole, the upper sessile and amplexicaul, the uppermost sometimes cordate-based, the lowermost sometimes with a winged, semiamplexicaul petiole, all with scattered, short to medium, pale simple eglandular hairs on both surfaces and numerous medium to long ones on the margins. Inflorescence with many capitula paniculate-corymbose; peduncles rather short, slender, suberect, with dense stellate hairs and numerous, subequal, short, dark glandular hairs, without simple eglandular hairs. Capitula 15–20 mm in diameter, small and narrow with a rounded base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, almost in 2 rows, olive green, most with paler margins, all linear-oblong, obtuse at apex, with dense stellate hairs along the margins, numerous, short, subequal, dark glandular hairs and an occasional, dark, medium simple eglandular hair. Ligules yellow, with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.5–4.0 mm, pale brown. Flowers 7–9. Apomictic. This species has a strange history. It is known only from cultivated plants which were said to have been originally collected in Harehead Wood, near Selkirk. It has never been re-found in this locality and is not known elsewhere. Described after William Borrer (1781–1862). Section 6. Alpestria (Fr.) Arv.-Touv. Taxon Alpestria Fr.; Subsection Alpestria (Fr.) Arv.-Touv.; Section Subaphyllopoda Arv.-Touv.; Subsection Stenocephala F. N. Williams; Subsection Pachycephala F. N. Williams Aphyllopodous perennial herbs or with basal leaves withered at the time of flowering or rarely phyllopodous. Cauline leaves usually fairly numerous, 2–19, rarely thickened at the margins, at least the upper more or less rounded at the base and semiamplexicaul. Inflorescence cymosepaniculate. Capitula usually few, dark, with broad, more or less obtuse involucral bracts, which are usually almost without stellate hairs. Achenes dark. Flowering in July or August. The species of this artificial group have a mixed origin. Some of them appear to be dwarf forms of the Section Foliosa which have been selected to survive in the exposed situations in which they occur, some even strongly approaching that group in cultivation. Others would appear to have originated from hybridisation between members of the Section Foliosa and other groups which almost certainly include the Sections Tridentata, Vulgata and Oreadea. The Section Alpestria is characterised by having few or no basal leaves, more or less amplexicaul cauline leaves and few capitula with broad, obtuse involucral bracts. It is distinguished from the Section Prenanthoidea by the darker achenes and usually fewer leaves and from the Section Foliosa by the fewer cauline leaves, longer internodes and by the leaves having less thickened margins. H. carpathicum, dewari and mirandum are related to those species occurring in the mountains of central Europe, while the remainder are most nearly related to those of the Faeroes, Iceland and Scandinavia. Important accounts dealing with

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the Section are Sell & West (1965), Scott & Palmer (1987) and Scott & Palmer (1995). 50. H. carpathicum Besser Perth Hawkweed H. dovrense var. spectabile E. S. Marshall; H. perthense F. N. Williams; H. carpathicum subsp. perthense (F. N. Williams) Zahn Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem up to 90 cm, pale yellowish-green, sometimes purplish at the very base, flexuous, striate, with long, pale simple eglandular hairs throughout but less dense above, the upper sometimes dark-based, with numerous stellate hairs above, without glandular hairs. Leaves 6–17, gradually decreasing in size upwards, 2–9(–11 cult.) × 1.0– 3.5(–4.0) cm, dark green on upper surface, paler beneath with a prominent midrib; the lamina of the lowermost broadly elliptical or obovate, rounded-obtuse-mucronulate at apex, entire or remotely and minutely denticulate and attenuate below to short, broadly winged petioles; the lamina of the lower and median broadly elliptical, elliptical or ovate-lanceolate, obtuse to acute at apex, denticulate to finely, sharply and remotely dentate, sessile and semiamplexicaul; the lamina of the upper ovate-lanceolate or ovate, acute to long-acute at apex, denticulate or finely and sharply dentate and with a rounded, sessile, semiamplexicaul base; all glabrous or nearly glabrous on upper surface, with medium, pale simple eglandular hairs on the lower surface and margins, the upper sometimes with scattered stellate hairs on the lower surface. Inflorescence with 3–12(–20 cult.) capitula, narrowly and compactly cymose, the acladium with a medium to long peduncle; peduncles medium to long, suberect, densely stellately hairy, usually with a few, short to medium, pale, dark-based simple eglandular hairs, rarely with a very short, dark glandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 10–13 × 1.0–1.3 mm, blackishgreen, the inner with paler margins, all linear-lanceolate, obtuse at apex, stellately hairy, densely so on the margins, sometimes with a slight tuft at the apex, with dense, short to medium, pale, dark-based simple eglandular hairs and numerous, very short to short, dark, slender glandular hairs. Ligules yellow, with dense, very short hairs at apex. Styles discoloured. Receptacle pits with margins fimbriatedentate. Achenes 4.0–4.5 mm, dark brown. Flowers 7–8. Apomictic. 2n = 36. It seems unlikely that the plants of the Tatra Mountains are exactly the same as those found in Scotland, but no characters can be found to distinguish them. Native. Known only from two localities on grassy riversides in Glen Shee, Perth at 300–400 m. Also in a few localities in the Tatra Mountains of east-central Europe. 51. H. dewari Syme Dewar’s Hawkweed H. carpathicum subsp. dewari (Syme) Zahn Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem up to 110 cm, yellowish-green, occasionally purplish below, flexuous, more or less robust, with long, pale simple eglandular hairs throughout, less densely so above, the upper sometimes dark-based, with scattered stellate hairs above, usually without glandular hairs, but

occasionally with a very short glandular hair near the inflorescence. Leaves 6–20, all except the lowermost widely spaced, gradually decreasing in size upwards; the lamina 3–15 × 0.5–4.5 cm, medium green on upper surface, paler beneath with a prominent midrib, the lowermost broadly elliptical or obovate, obtuse-mucronulate to acute at apex, entire or remotely denticulate and attenuate to a winged petiole, the lamina of the lower and median similar to the lowermost but never obovate and the petioles are either short or absent, the lamina of the upper ovate or ovate-lanceolate, acute to long-acute at apex, entire or denticulate and with a rounded, sessile, semiamplexicaul base; all with the indumentum variable, usually very hairy on both surfaces and the margins with pale, subrigid, medium simple eglandular hairs, but sometimes the hairs are fewer or nearly absent. Inflorescence with up to 17(–35 cult.) capitula, narrowly and compactly cymose, the acladium with a short to medium peduncle; peduncles medium, suberect, with dense stellate hairs, with more or less numerous, dark-based, short simple eglandular hairs and more or less numerous, very short, dark, fine glandular hairs, the numbers of simple eglandular and glandular hairs, and the proportions of one to the other, varying greatly. Capitula 20–25 mm in diameter, rounded below. Involucral bracts incumbent in bud, 9–10 × about 1.0 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse or rarely subacute at apex, with scattered stellate hairs towards the base and on the margins and with a tuft at the apex, with numerous short or very short, dark glandular hairs and few to many, short to medium, darkbased simple eglandular hairs. Ligules yellow, with very short simple eglandular hairs at the apex. Styles discoloured. Receptacle pits with the margins fimbriate-dentate. Achenes 3.5–4.0 mm, dark brown. Flowers 7–8. Apomictic. Native. Banks of streams and rocky ledges up to 730 m and mainly on schist. Mountains of west-central and central Scotland. Endemic. Named after Andrew Dewar (c. 1792– 1870). 52. H. mirandum P. D. Sell & C. West Remote Hawkweed Hypophyllopodous perennial herb with a branched stock. Stem up to 45(–70 cult.) cm, pale yellowish-green, purplish below, flexuous, slender, with pale, long simple eglandular hairs throughout, in the upper part fewer and dark-based, without stellate or glandular hairs. Leaves with lamina 2–6(–8 cult.) × 0.5–3.0(–4.0 cult.) cm, medium green on upper surface, caesious beneath; basal with lamina elliptical or ovate, more or less acute at apex, denticulate, attenuate to a medium, narrowly winged petiole, with scattered, pale, medium simple eglandular hairs on both surfaces and the margins; cauline 6–9(–15 cult.), the lamina ovate or lanceolate, subacute to acute at apex, denticulate or with a few shallow teeth, the lower cuneate at base to a short petiole, the upper rounded, sessile and amplexicaul at base, glabrous on upper surface and with a few, pale, medium simple eglandular hairs beneath and on the margin. Inflorescence with 2–5(–23 cult.) capitula, compact, the acladium with a medium peduncle; peduncles suberect, medium, with dense stellate hairs and numerous, dark-based, medium simple eglandular hairs, without

39. Hieracium glandular hairs. Capitula 20–25 mm in diameter, rounded below. Involucral bracts incumbent in bud, 9–10 × 1.5– 2.0 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with occasional stellate hairs mainly at the base, with few, very short to short, dark glandular hairs and a few, short, dark-based simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins dentate. Achenes 3.5–4.0 mm, blackish-brown. Flowers 7–8. Apomictic. 2n = 27. Native. Near Well House, Stean at 255–300 m and on banks of streams at Ribbleshead in Yorkshire, Masson Hill, Bonsall in Derbyshire and near Seathwaite in Westmorland; in all localities rare. Endemic. 53. H. vinicaule P. D. Sell & C. West Wine-stemmed Hawkweed H. crocatum var. vinaceum Beeby; H. strictum var. humilius Beeby; H. congestum forma vinaceum (Beeby) Roffey; H. platylepium auct.; H. vinaceum (Beeby) Pugsley, non Johanss. & Sam. Aphyllopodous perennial herb with a branched stock. Stem up to 70 cm, pale yellowish-green, purple below, sometimes throughout, flexuous, slender to fairly robust, usually with scattered, pale, medium simple eglandular hairs throughout, with few stellate hairs especially above, sometimes with a few, very short, dark glandular hairs near the inflorescence. Leaves all cauline, 10–19(–30 cult.), 2–12(–14 cult.) × 0.5–2.5 cm, medium or dark green on upper surface, paler beneath, slowly decreasing in size upwards; lamina of the lower narrowly elliptical to elliptic-oblong, more or less acute at apex, obscurely denticulate or with an occasional shallow tooth, attenuate below to short or medium, narrowly winged petioles, and with sparse, pale, short to medium simple eglandular hairs on both surfaces, the margins and the petioles; lamina of the median and upper lanceolate, linearoblong or narrowly elliptical, subacute to acute at apex, subentire to shallowly and remotely denticulate or occasionally dentate, abruptly or gradually narrowed below to a semiamplexicaul base, clothed like the lower except that the simple eglandular hairs are sometimes absent on the upper surface and stellate hairs are frequent beneath and rare above, and the margins are sometimes only scaberulous. Inflorescence with 2–42(–350 cult.) capitula, narrowly and laxly cymose, with long, slender, suberect branches exceeding the medium peduncled acladium; peduncles medium to long, suberect, with scattered stellate hairs and with darkbased, short to medium simple eglandular hairs and very short, dark glandular hairs in varying proportions, one or other type of hair sometimes absent. Capitula 20–25 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 9.0–10.5 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, sometimes narrowed above but always obtuse at apex, with few, scattered stellate hairs, with more or less numerous, very short to short, dark glandular hairs and usually a few, medium, darkbased simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 4.0–4.5 mm, blackish-brown. Flowers 7–9. Apomictic. 2n = 27.

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The most variable of the Shetland species of the Series Alpestria in size, leaf colour and leaf denticulation, each colony being slightly different from every other. Native. Grassy or rocky streamsides, sea-banks, meadows and pastures, sometimes on crags, holms in lochs and on grassy roadsides. Occasional in North Mainland and rare in Central and West Mainland in the Shetland Islands. Endemic. 54. H. northroense Pugsley North Roe Hawkweed H. crocatum var. congestum Beeby; H. congestum (Beeby) Roffey, non Freyn Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 17–35(–45) cm, pale yellowish-green, sometimes purplish at base, slender to fairly robust, flexuous, with numerous, pale, medium simple eglandular hairs at the base, usually less numerous above or scaberulous, with scattered stellate hairs, without glandular hairs. Leaves 1.5–9.0 × 0.5–2.0 cm, pale to medium green, often reddishpurple on the margins and veins; basal with lamina elliptical, obtuse at apex, regularly denticulate and attenuate to a short, winged petiole; cauline (7–)9–15, suddenly decreasing in size about halfway up the stem, the lamina of the lower lanceolate to broadly elliptical, obtuse to acute at apex, with numerous, fine, closely placed denticulations and narrowed to a semiamplexicaul base, the lamina of the upper linearlanceolate to lanceolate (to ovate-lanceolate in cultivation), subacute to acute at apex, denticulate as the lower and sessile, rounded and semiamplexicaul at the base, all with short to medium, pale simple eglandular hairs beneath and on the margins, the lower sometimes having them above; all with numerous stellate hairs on the lower surface, the upper with stellate hairs on both surfaces. Inflorescence with 2–9(–20 cult.) capitula, cymose, laxly so in cultivation, the acladium with a medium peduncle; peduncles medium to long, suberect, with scattered stellate hairs, with a few, very short, dark glandular hairs and occasionally a few, pale, medium simple eglandular hairs. Capitula 25–30 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 9–11 × 1.5–2.0 mm, green with a dark centre, lanceolate, obtuse at apex, with scattered stellate hairs, with a few, very short or short, dark glandular hairs and sometimes a few, pale, medium simple eglandular hairs in a row down the centre and at the base. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with the margins shortly dentate. Achenes about 3.5 mm, dark brown. Flowers 7–8. Apomictic. 2n = 27. H. northroense is very closely allied to H. vinicaule, but differs in its more dwarf, compact habit, more numerous denticulations on the leaves which are less drawn out at the apex and in its paler less densely clothed involucral bracts. Native. Heathy pastures and rocky sea-cliffs. Known only from below Windy Scord, Voe of Snarraness in West Mainland, and a hay meadow below Burravoe, North Mainland in the Shetland Islands. Endemic. 55. H. klingrahoolense Walter Scott & R. C. Palmer Klingrahool Hawkweed Usually aphyllopodous perennial herb with a branched stock. Stem 20–70 cm, pale yellowish-green, often suffused

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with brownish-purple, flexuous, striate, glabrous or nearly so or with a few stellate or minute hairs. Leaves medium yellowish-green on upper surface, paler beneath, sometimes tinted purple; basal usually absent at flowering but sometimes with lamina small, elliptical, obtuse at apex and cuneate at base; cauline 10–19, rapidly decreasing in size in the upper half of the stem, the lower with lamina 4–11 × 1–2 cm, narrowly linear-elliptical or oblong, narrowed to a usually acute apex, subentire or with minute teeth and gradually narrowed at base and more or less amplexicaul, the upper small and linear or linear-lanceolate, acute at apex, entire and amplexicaul; all glabrous or with a few, minute hairs, especially on the margins. Inflorescence with 1–10 capitula; peduncles with numerous stellate hairs, usually without obvious simple eglandular or glandular hairs, but with very minute glandular hairs. Capitula 25–30 mm in diameter, rounded or subtruncate at base. Involucral bracts incumbent in bud, 5–11 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with few to numerous, short to medium, dark glandular hairs, sometimes an occasional simple eglandular hair, and a few stellate hairs towards the base. Ligules yellow, glabrous at tip. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.5–4.0 mm, deep red. Flowers 7–8. Apomictic. Native. Dry rocky pasture, steep rocks, ravines and rocky banks. Mouth of the Burn of Crookadale, Kelo Wick, Lunna Ness, between Brough and Eswick, South Nesting and Burn of Skelladale near Brae in Mainland; North Burn south of West Yell and Loch of Lumbister in Yell; and Muckle Roe, all in the Shetland Islands. Endemic. The name is derived from the Norse description of the first place it was found and means ‘the place where the wild rose or bramble grows’; the rose is still there today. 56. H. subtruncatum Beeby Mainland Hawkweed H. friesii auct.; H. truncatum auct.; H. rigidum var. friesii auct.; H. carpathicum subsp. truncatum auct.; H. laevigatum subsp. friesii auct.; H. subtruncatum var. glussburnense Pugsley Aphyllopodous perennial herb with a branched stock. Stem up to 75 cm, pale yellowish-green, usually purplish below, sometimes throughout, more or less robust, flexuous, usually with pale, medium to long simple eglandular hairs throughout, those above dark-based, often densely so below, sometimes only sparingly so or scaberulous, with scattered stellate hairs throughout, sometimes more or less dense near the inflorescence, without glandular hairs. Leaves all cauline, 8–20, 2–11 × 0.5–2.5 cm, gradually decreasing in size upwards, pale to medium green on upper surface, paler and often suffused with purple beneath; lamina of the lowest broadly elliptical or oblong, obtuse at apex, subentire or denticulate, narrowed (sometimes abruptly) to a sessile base or occasionally shortly petiolate and with numerous, pale, subsetiform, bulbous-based, medium simple eglandular hairs on both surfaces and the margins; lamina of the median lanceolate to oblong, subacute at apex, denticulate or with a few, shallow teeth intermixed, contracted to a rounded, semiamplexicaul base and clothed as lower except that the hairs are fewer, the upper surface is

sometimes glabrous and the lower surface sometimes has scattered stellate hairs; lamina of the upper lanceolate to ovate-lanceolate, often gradually narrowed from the middle to an acute apex, denticulate, sometimes with a single shallow tooth, with a broad, rounded and semiamplexicaul base and clothed like the median. Inflorescence with 1–13 capitula, narrowly and laxly cymose the acladium with a long peduncle; peduncles long and slender, suberect, with few to numerous stellate hairs, few to numerous, usually dark-based, medium simple eglandular hairs, sometimes only the dark bases remaining and occasionally a few, very short, dark glandular hairs. Capitula 20–30 mm in diameter, rounded below. Involucral bracts incumbent in bud, 9–11 × about 1.5 mm, blackish-green, the inner with paler margins and often reddish-tipped, all linearlanceolate, obtuse at apex, without stellate hairs, usually with a few, very short, dark glandular hairs and solitary, dark, very short simple eglandular hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with the margins shortly dentate. Achenes 3.5–4.0 mm, blackish-brown. Flowers 7–8. Apomictic. Variable in leaf colouring, leaf width and stem and peduncle hairiness. Var. glussburnense differs only in having fewer and shorter simple eglandular hairs on the stem and peduncles. Native. Rocky or grassy streambanks and crags, rarely on sea-banks or in meadows and pastures, mainly on granite and diorite, very rarely on limestone. Occasional in North Mainland, very local or rare in South, Central and West Mainland, Shetland Islands. Endemic. 57. H. dilectum P. D. Sell & C. West Purple-tinted Hawkweed Aphyllopodous perennial herb with a branched stock. Stem up to 55 cm, pale green often suffused purple, slender, flexuous, rarely glabrous, usually with a few (rarely more numerous), pale, medium simple eglandular hairs especially at the nodes, with scattered stellate hairs and a few, very short glandular hairs above. Leaves all cauline, 9–16(–20), 2– 10 × 0.3–2.0 cm, pale green on upper surface, paler and usually suffused with purple, especially beneath; lamina of the lower narrowly elliptical or sometimes lanceolate, obtusemucronulate to subacute at apex, regularly and remotely denticulate and attenuate below to short, broad-winged, semiamplexicaul petioles; lamina of the median narrowly elliptical or sometimes linear, acute at apex, remotely denticulate with an occasional small, sharp dentation and shortly narrowed to a cuneate or rounded, semiamplexicaul, sessile base; lamina of the upper long-lanceolate, long-acute at apex, remotely denticulate or entire and sessile with a rounded, semiamplexicaul base; all glabrous or with few to many, pale, medium simple eglandular hairs on the midrib beneath, the median and upper often with scattered stellate hairs on both surfaces. Inflorescence with 1–6 capitula, loosely cymose, the acladium medium to long; peduncles suberect, with scattered stellate hairs, sometimes scaberulous and occasionally with a solitary, pale, medium simple eglandular hair. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 9–11 × 1.2 – 1.5 mm, blackish-green, the median and inner with paler

39. Hieracium margins, all linear-lanceolate, obtuse at apex, with scattered stellate hairs, scattered, short and very short, dark simple eglandular hairs, and scattered very short or short, dark glandular hairs. Ligules yellow, glabrous. Styles yellow to discoloured. Receptacle pits with margins fimbriatedentate. Achenes 4.0–4.5 mm, dark brown. Flowers 7–8. Apomictic. Native. Rocky or grassy streamsides and crags. Known only from one site in Central Mainland, two in West Mainland and one in North Mainland, Shetland Islands. Endemic. 58. H. pugsleyi P. D. Sell & C. West Pugsley’s Hawkweed Aphyllopodous perennial herb with a branched stock. Stem up to 60 cm, pale green, often suffused purple, slender, flexuous, with dense, long, pale simple eglandular hairs throughout, those of the upper half with dark bases, with few, scattered stellate hairs, sometimes with a few, very short glandular hairs. Leaves all cauline, 8–12(–15), 1– 11 × 0.5–3.0 cm, pale green on upper surface, often suffused with purple especially on the lower surface; lamina of the lower elliptic-oblong to oblong, obtuse-mucronate to subacute at apex, subentire to minutely denticulate and attenuate at base to a short, winged petiole; lamina of the median elliptic-oblong to oblong, more or less acute at apex, subentire to regularly denticulate and rounded at the sessile, semiamplexicaul base; lamina of the upper lanceolate to ovate-lanceolate, acute at apex, subentire and with a rounded sessile base; all with pale, medium simple eglandular hairs on both surfaces and the margins, the uppermost with very few stellate hairs beneath. Inflorescence with 3–9(–22) capitula, cymose, sometimes with long, leafy branches in the leaf axils, the acladium with a short to medium peduncle, with slender, suberect branches; peduncles short to medium, sometimes geminate, with scattered stellate hairs, numerous medium, dark-based simple eglandular hairs and a few, very short glandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 9–10 × 1.5–1.7 mm, blackish-green, sometimes reddish at the tip, the inner with paler margins, all linear-lanceolate, obtuse at apex, with very few stellate hairs, numerous, medium, dark-based simple eglandular hairs and few or many, very short, dark glandular hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with the margins shortly dentate. Achenes 3.0–3.5 mm, dark brown. Flowers 7–9. Apomictic. Native. Grassy or sometimes rocky streamsides and seacliffs, very rarely on roadside banks. Burn of Laxdale, Cunningsburgh in South Mainland, Burn of Weisdale in Central Mainland, Tumblin in West Mainland, Laxo Burn in North Mainland, and two sites in Yell (near Otterswick and at Whale Firth) in the Shetland Islands. Endemic. Named after Herbert William Pugsley (1868–1947). 59. H. spenceanum Walter Scott & R. C. Palmer Spence’s Hawkweed Phyllopodous perennial herb with a branched stock. Stem (10–)20–40(–80) cm, pale yellowish-green, usually suffused reddish throughout, but sometimes reddish only at

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base and green above, rarely green throughout, robust, with stellate hairs few below and numerous above, with fairly numerous simple eglandular hairs throughout and especially below, without glandular hairs. Leaves 5–9 × 1.0– 2.5 cm, deep green on upper surface, paler and suffused with purple beneath and sometimes also above, and with reddish margins; basal (0–)3–5(–7), at least some nearly always present at flowering and fruiting and often forming a well-developed rosette, the lamina subrotund, broadly elliptical or ovate-elliptical, broadly rounded and often minutely apiculate at apex, subentire or finely denticulate, and more or less rapidly contracted below to a winged petiole, with few stellate hairs and few simple eglandular hairs beneath, but more numerous on the midrib and with more or less numerous simple eglandular hairs on the upper surface; cauline 3–7, often concentrated in the lower part of the stem, the lamina 2.5–3.5 times longer than broad, that of the lowest typically elliptical, acutely apiculate at apex, distinctly and more or less regularly denticulate, sometimes with a few, shallow teeth and narrowed to a winged petiole, the lamina of the median and upper ovate-elliptical, toothed as lowest, sessile and semiamplexicaul, all the cauline with scattered stellate hairs on both surfaces and numerous simple eglandular hairs beneath especially on the midrib, the upper often becoming glabrous. Inflorescence with (1–)2– 6(–24) capitula, compactly cymose, sometimes with long branches from the upper leaf axils, the acladium with a short peduncle; peduncles with numerous stellate hairs, and scattered, pale, dark-based simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 10–11 × about 2 mm, dark green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with few stellate hairs, few, pale, dark-based simple eglandular hairs and numerous, very unequal, dark glandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with the margins fimbriate-dentate. Achenes about 4.5 mm, dark brown. Flowers 7–8. Apomictic. Native. Crags, low rocky sea-banks, streamsides and rocky pastures. From Norby to Bousta, both sides of West Burra Firth, Ward of Scollan, between The Runie and Maw Loch, and below Brindister, all in West Mainland, Shetland Islands. Endemic. Named after Prof. D. H. N. Spence (1925–1985). 60. H. attenuatifolium P. D. Sell & C. West Laxo Burn Hawkweed Aphyllopodous perennial herb with a branched stock. Stem up to 90 cm, flexuous, pale yellowish-green, often more or less flushed purple, with long, pale simple eglandular hairs above, these often becoming deciduous, their bases scaberulous, and with a few stellate hairs, without glandular hairs. Leaves all cauline, 6–15, 2–10 × 0.5–3.0 cm, deep or pale green on upper surface, paler beneath and often suffused with purple; lamina of the lowest broadly elliptical, rounded-obtuse-mucronulate at apex, subentire or remotely denticulate and cuneate or attenuate at base to winged petioles; lamina of the lower elliptical, sometimes broadly so, obtuse-mucronate to subacute at apex, distantly denticulate or occasionally with 1 or more sharp teeth and gradually narrowed below to broad, winged petioles; lamina of the

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median narrow to broadly elliptical, more or less acute at apex, distantly denticulate or rarely with a few, sharp, shallow teeth and narrowed or abruptly contracted to a sessile, semiamplexicaul base; lamina of the upper lanceolate to linear-lanceolate, acute at apex and subentire, or bract-like; lower and median with medium, pale simple eglandular hairs on both surfaces and the margins, the median occasionally with a few stellate hairs beneath, the upper with or without an occasional simple eglandular hair, and with scattered stellate hairs on both surfaces, the margins and veins puberulous. Inflorescence with 2–7(–13 cult.) capitula, loosely cymose, the acladium with a long peduncle; peduncles mostly long, with scattered stellate hairs, usually with numerous, medium, dark-based simple eglandular hairs and an occasional very short to short, dark glandular hair. Capitula 35–50 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 10–12 × 1.5–2.0 mm, the outer blackish-green, very short and triangular-ovate, the median with paler margins and lanceolate, the inner similar to the median but paler and rather narrower, all obtuse at apex, with very few stellate hairs, numerous, unequal, very short to medium, dark glandular hairs and a few, medium, dark simple eglandular hairs. Ligules pale yellow, glabrous. Styles yellow, darkening when dry. Receptacle pits with margins shortly dentate. Achenes 4.5–5.0 mm, dark brown. Flowers 7–8. Apomictic. Native. Grassy banks of the Laxo Burn, North Mainland in the Shetland Islands. Endemic. 61. H. hethlandiae (F. Hanb.) Pugsley Cliva Hill Hawkweed H. dovrense var. hethlandiae F. Hanb.; H. dovrense subsp. humidorum var. hethlandiae (F. Hanb.) Beeby; H. aestivum subsp. crocatum subvar. hethlandiae (F. Hanb.) Zahn; H. crocatum forma hethlandiae (F. Hanb.) Roffey; H. crocatum var. hethlandiae (F. Hanb.) Druce Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem up to 40 cm, pale yellowish-green, purplish below, often throughout, slender, flexuous, with long, pale simple eglandular hairs only at the very base and then often only sparingly, and more or less numerous stellate hairs especially above, without glandular hairs. Leaves all cauline, 5–9, 1–8 × 0.5–2.0 cm, caesious medium green on upper surface, paler beneath and often purple-tinted especially beneath and on the margins, the upper rapidly decreasing in size upwards; lamina of the lowest broadly elliptical, more or less obtuse at apex, subentire, attenuate to short, winged, semiamplexicaul petioles, glabrous or with pale, medium, subrigid, bulbous-based simple eglandular hairs above and on the margins, and the petioles with medium, pale simple eglandular hairs; lamina of the lower narrowly elliptical, gradually narrowed at both ends, acute at apex, denticulate, with short, winged petioles, or sessile and semiamplexicaul, and glabrous or with pale, medium simple eglandular hairs on both surfaces and the margins; lamina of the median as lower, but sometimes broader and more rounded at the base, with a long-acute apex, sessile, glabrous or nearly so on both sides and the margins puberulous or with a few, short simple eglandular hairs; lamina of the upper gradually narrowed to an acute, apiculate tip from

a broad, rounded, semiamplexicaul base, glabrous except for a few stellate hairs below. Inflorescence with 2–4 capitula, compactly cymose, acladium with a medium peduncle; peduncles medium, suberect, slender, stellately hairy, usually without simple eglandular hairs, without glandular hairs. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 10.5–12.0 × 1.5–1.7 mm, blackish-green, the inner with paler margins, all linearlanceolate, obtuse at apex, with stellate hairs scattered over the whole surface, very short to medium, dark glandular hairs and some short to medium, dark simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with the margins long-dentate. Achenes 3–5 mm, blackish-brown. Flowers 7–8. Apomictic. H. hethlandiae has no near allies among the British species, but it is related to the Scandinavian H. humidorum Almq. Native. Recently exterminated in its only certain station. Formerly on steep, grassy, heathery rock-faces at Cliva Hill, Mavis Grind, Shetland Islands. Also recorded for Ronas Voe, but the specimen may be mislabelled. The species survives in cultivation. An unsuccessful attempt was made to introduce it at Hurda Field and a more recent similar introduction at the Burn of Skelladale shows a little more promise. 62. H. praethulense Pugsley Thule Hawkweed H. auratum var. thulense F. Hanb.; H. crocatum var. thulense (F. Hanb.) Druce Hypophyllopodous or aphyllopodous perennial herb with a branched stock, or with basal leaves withering at the time of flowering. Stem up to 60 cm, usually rather slender but some large plants robust, flexuous, markedly striate, pale yellowish-green, often purplish below, sometimes throughout, with long, pale simple eglandular hairs throughout, but decreasing in density upwards, sometimes without simple eglandular hairs except at the very base, with scattered stellate hairs in the upper half, without glandular hairs. Leaves 1.5–9.0(11.0 cult.) × 0.5–3.0(–4.0 cult.) cm, pale green on upper surface, paler beneath and often suffused with purple; basal few or absent, the lamina usually broadly elliptical and sometimes almost subrotund, rounded or obtuse at apex, remotely denticulate, narrowed (sometimes abruptly) to a winged petiole, with scattered, medium simple eglandular hairs or glabrous above and with long simple eglandular hairs beneath and on the margins; cauline 7–12(–16) (in cultivation with numerous other leaves on the branches), the lower variable, the lamina elliptical, ovate or oblonglanceolate, obtuse-mucronulate to acute at apex, remotely denticulate, attenuate to a winged petiole or abruptly narrowed to a sessile, semiamplexicaul base and clothed as basal, the lamina of the median elliptical or lanceolate, acute at apex, remotely denticulate, with a rounded or subcordate, semiamplexicaul base, glabrous above, with few simple eglandular hairs beneath, and the margins with simple eglandular hairs or scaberulous, the lamina of the upper ovate-lanceolate to lanceolate, gradually narrowed to an acute apex, entire or with an occasional denticulation, rounded or subcordate and semiamplexicaul at the base, glabrous above, with few simple eglandular and stellate

39. Hieracium hairs beneath and the margins ciliate or scaberulous. Inflorescence with 2–9(–100 cult.) capitula, compactly cymose (in cultivation cymose-corymbose), with medium acladium; peduncles slender, suberect, medium to long, sparingly stellately hairy and sometimes with a few, pale, short simple eglandular hairs, without glandular hairs. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 9–10 × (1.0–)1.5–2.0 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with scattered stellate hairs, few to many, very short to short glandular hairs and occasionally a few, short simple eglandular hairs intermixed. Ligules yellow, glabrous. Styles yellow, sometimes discoloured when dry. Receptacle pits with margins shortly and irregularly dentate. Achenes 4.0–4.5 mm, dark reddish brown. Flowers 7–8. Apomictic. In cultivation, H. praethulense often becomes a very large plant with many capitula and then has a superficial resemblance to H. latobrigorum of the Series Foliosa. It is closely allied to the Icelandic species H. phrixoclonum Omang and H. halfdanii Osk. Native. Steep, rocky sea-banks, crags, and holms in lochs on granite and diorite. From just north of Mavis Grind to the holms in Swabie Water, particularly by the north side of Ronas Voe and at Swabie Water where it is frequent and rarely locally abundant, North Mainland, Shetland Islands. Endemic. 63. H. dovrense Fr. Dovre Hawkweed H. carpathicum subsp. dovrense (Fr.) Zahn Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem to 50 cm, pale green, slightly purpletinted below, flexuous, fairly robust, with long, pale simple eglandular hairs throughout, sometimes sparsely, sometimes densely, sometimes the hairs deciduous, sometimes with a few stellate hairs above, with an occasional, very small glandular hair. Leaves 2–10 × 0.8–3.5 cm, pale green on upper surface, paler beneath; basal few, the lamina elliptical or subrotund, obtuse-mucronulate at apex, remotely denticulate to coarsely dentate, attenuate to winged petioles which are broadened below, midrib very prominent, nearly glabrous above, with a few, pale, medium simple eglandular hairs below, margins and petioles with medium, pale simple eglandular hairs; cauline 4–8, slowly decreasing in size upwards, the lamina of the lower and median elliptical, obtuse-mucronulate to acute at apex and denticulate to sharply and irregularly dentate with the teeth ascendingmammiform, the lower attenuate below to a winged semiamplexicaul petiole, the median to a sessile or abruptly contracted semiamplexicaul base, with clothing as basal, the lamina of the upper lanceolate, acute to acuminate at apex, denticulate or shallowly dentate, cuneate to a sessile, semiamplexicaul base, and glabrous except on the midrib below. Inflorescence usually with 3–9(–12) capitula, rather compactly cymose, the acladium geminate; peduncles medium, suberect, slender, stellately hairy, otherwise variable in clothing with few to numerous, short to medium, dark-based simple eglandular and very short, black glandular hairs. Capitula 20–25 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 9–11 × 2.0– 2.5 mm, blackish-green, the inner with paler margins, all

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linear-lanceolate, obtuse at apex, sparingly stellately hairy with a tuft at the apex, with many, short to medium, dark-based simple eglandular and very short, black glandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins long-dentate. Achenes 2.0– 2.5 mm and dark brown in Scandinavian plants. Flowers 7–8. Apomictic. 2n = 36. Native. Cliff ledges from 130 to 330 m. Local on Hornblende Gneiss on north-facing cliffs of Ben Loyal, in a gorge near Rhiconich, burnside at Foinaven and at Creag na h’Uidhe all in Sutherland. It is common in the Dovre district of Norway. 64. H. australius (Beeby) Pugsley Unst Hawkweed H. dovrense subsp. demissum var. australius Beeby; H. demissum var. australius (Beeby) Druce; H. polycomum auct.; H. polycomatum auct. Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem up to 65 cm, pale green, purplish below, usually robust, markedly striate, usually with long, pale simple eglandular hairs below, sometimes throughout, the hairs often deciduous above, with a few stellate hairs just below the inflorescence, without glandular hairs. Leaves 2– 10(–11) × 0.5–3.0(–4.0) cm, pale to dark green on upper surface, sometimes with purplish spots; basal soon drying up and dropping off, the lamina elliptical to broadly elliptical, obtuse-mucronate to acute at apex, denticulate, narrowed below to winged petioles, with pale, medium, subrigid, bulbous-based hairs on both surfaces, the margins and petioles; cauline 5–12(–numerous cult.), gradually decreasing in size upwards, the lamina of the lower more or less elliptical, obtuse-mucronate to acute at apex, subentire or sharply denticulate, attenuate to a winged petiole and clothed as basal, the lamina of the median very variable, elliptical, lanceolate or oblong-lanceolate (sometimes curved in the shape of a crescent), acute at apex, sometimes gradually narrowed from the middle, usually with sharp denticulations, gradually narrowed to a semiamplexicaul base, hair clothing as on lower, but sparser and some plants with abortive branches growing in the axils of the leaves, the lamina of the upper ovate-lanceolate, gradually narrowed from a rounded base to an acute apex, denticulate or sometimes with a few, sharp teeth, usually semiamplexicaul and nearly glabrous above otherwise as lower, the uppermost sometimes with scattered stellate hairs beneath. Inflorescence with 2–10(–80 cult.) capitula, compactly cymose, in cultivation laxer and cymose-corymbose, the medium acladium subtended by the branches; peduncles suberect, stellately hairy, with an occasional short, darkbased simple eglandular hair. Capitula 20–35 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 9.0–10.5 × 1.5–2.0 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with a few stellate hairs near the base, many, very short, pale and dark glandular hairs and few or many, usually short, rarely medium, pale, dark-based simple eglandular hairs, sometimes without simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins long-dentate. Achenes 3.0–3.5 mm, dark brown. Flowers 7–8. Apomictic.

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In cultivation this species gets very large with numerous capitula and leaves and looks like a hybrid between a species of the Section Tridentata and a species of the Section Foliosa. Native. Rocky banks by lochs, steep grassy sea-banks and granite crags. Locally common in a restricted area about the Loch of Cliff in northernmost Unst, rare by the Wick of Tresta, Fetlar, and occasional on both sides of Ronas Voe, North Mainland in the Shetland Islands. Endemic. 65. H. difficile P. D. Sell & C. West Okraquoy Hawkweed Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem up to 70 cm, pale green, flushed purplish especially below, flexuous, rather robust, with long, wavy, pale, sometimes dark-based simple eglandular hairs throughout, but only scattered above, and scattered stellate hairs throughout, without glandular hairs. Leaves 2.5– 11.0 × 0.7–3.0 cm, medium green on upper surface, paler and sometimes purplish tinged beneath; basal with lamina broadly elliptical, obtuse-mucronulate at apex, subentire, attenuate to a short, winged petiole, with dense, medium, pale, subrigid simple eglandular hairs on both surfaces and the margins; cauline 4–8, widely spaced and gradually decreasing in size upwards, sometimes with abortive branches in the axils of the leaves, the lamina of the lower elliptical, acute to long-acute at apex, remotely and shallowly denticulate, gradually narrowed to a broad, sessile, semiamplexicaul base or with a winged, semiamplexicaul petiole, and hairs as on basal but fewer in number on the upper surface, the lamina of the upper lanceolate, gradually tapered from a rounded, sessile, semiamplexicaul base to a long-acute apex, with scattered, medium simple eglandular hairs and sometimes with stellate hairs on both surfaces. Inflorescence with 3–12 capitula, narrowly and compactly cymose, the acladium with a medium peduncle; peduncles suberect, with stellate hairs, sometimes with occasional, short, dark-based simple eglandular hairs and a very short dark glandular hair. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 9–10 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse to subacute at apex, with scattered stellate hairs at the base, scattered, short and/or very short, dark glandular hairs and an occasional, short, dark-based simple eglandular hair. Ligules yellow, glabrous. Styles discoloured. Achenes about 4 mm, dark brown. Flowers 7–8. Apomictic. Very variable in hair clothing. Native. Banks of stream in a limestone ravine. Known only from near the Bay of Okraquoy in South Mainland in the Shetland Islands. Endemic. 66. H. amaurostictum Walter Scott & R. C. Palmer Semblister Hawkweed Phyllopodous or aphyllopodous perennial herb with a branched stock. Stem 20–45 cm, pale yellowish-green, often suffused with brownish-purple, slender, flexuous, striate, with numerous, often dense, long, pale, wavy simple eglandular hairs below, becoming slightly less numerous upwards, and with numerous stellate hairs and rarely a

solitary, short, dark glandular hair in the upper part. Leaves medium yellowish-green and faintly blotched and spotted brownish-purple on upper surface, pale and often suffused purple beneath; basal few or absent at flowering, the lamina 2–7 × 2.0–2.5 cm, narrowly elliptical to oblong-elliptical, acute at apex, entire or with minute denticulations, cuneate to attenuate at base, with its petiole up to 4 cm, often purplish and with dense, long, simple eglandular hairs; cauline 3–6, 1.5–7.0 × 0.2–2.5 cm, gradually becoming smaller upwards, the lower like the basal and petiolate, the median elliptical or lanceolate, acute at apex, entire or denticulate, cuneate to a sessile base, the upper linear-lanceolate or linear, acute at apex, entire and sessile, or bract-like; all with numerous, long, pale simple eglandular hairs on both surfaces and on the margins. Inflorescence with 1–3 capitula; peduncles with dense stellate hairs, numerous short and medium, dark-based simple eglandular hairs and a few, short, dark glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 6–12 × 1.3–1.6 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse or subacute at apex, with numerous short and medium, dark-based simple eglandular hairs, few, short, dark glandular hairs and stellate hairs almost completely absent. Ligules yellow, glabrous or with few very short hairs at apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 4–5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Frequent over a small area of low sea-banks near the north end of the Loch of Semblister, West Mainland, Shetland Islands. Endemic. 67. H. gratum P. D. Sell & C. West Handsome Hawkweed H. pulchellum auct.; H. demissum var. pulchelliforme W. R. Linton; H. dovrense subsp. demissum var. pulchelliforme (W. R. Linton) Beeby; H. demissum subsp. demissum var. pulchelliforme (W. R. Linton) Zahn; H. pulchelliforme (W. R. Linton) Pugsley, non Dahlst. ex Omang Phyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 15–35(–40) cm, pale green, usually purple below, slender, flexuous, markedly striate, with long, pale, wavy simple eglandular hairs below, becoming nearly glabrous upwards, but usually with some simple eglandular hairs at the nodes, with scattered stellate hairs throughout and without glandular hairs. Leaves 1.5–7.5 × 3.0–3.5 cm, pale green on upper surface, paler and often suffused purplish beneath and sometimes tinged purple above; basal few, soon dying off, the lamina narrowly to broadly elliptical or ovate-lanceolate, obtuse-mucronulate to subacute at apex, with up to 4 sharp, patent denticulations on each side and more or less attenuate to a winged petiole; cauline 2–4(–7), rapidly decreasing in size upwards, the lamina of the lower elliptical to lanceolate, obtuse-mucronulate to acute at apex, toothed as basal and narrowed to a winged, semiamplexicaul petiole, the lamina of the median ovate-lanceolate or lanceolate, obtuse to acute at apex, the apex sometimes tapering from near the middle, abruptly contracted to a narrow, sessile, semiamplexicaul base and toothed as lower, the lamina of the upper lanceolate or ovate-lanceolate, acute at apex,

39. Hieracium which sometimes tapers from the middle, and entire or with up to 3 denticulations of each side; all with long, pale, subrigid simple eglandular hairs on both surfaces and on the margins, the upper cauline sometimes only sparingly so, and sometimes sparsely hairy beneath. Inflorescence with 1–6 capitula, compactly cymose, sometimes with 1–2 branches from the upper leaf axils; peduncles usually long, slender and suberect, sometimes with several bracts, with stellate hairs, with or without a very occasional, short simple eglandular hair or very short glandular hair. Capitula 20–25 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 9–11 × about 1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse or occasionally acute at apex, with a few stellate hairs near the base, usually sparingly covered with short to medium, pale, dark-based simple eglandular hairs and slender, very short, dark glandular hairs, sometimes, however, the simple hairs are numerous and sometimes nearly absent. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with the margins fimbriate-dentate. Achenes 2.5–3.0 mm, blackishbrown. Flowers 7–8. Apomictic. H. gratum shows no close affinity with any British species, but is closely allied to the Icelandic H. elegantiforme Dahlst. Native. Steep grassy sea-cliffs and rocky banks by lochs, mainly on quartzite. Restricted to an area of northernmost Unst (about the Loch of Cliff and nearby Burra Firth), and recently found on sea-cliffs at Whale Firth in the neighbouring island of Yell, both in the Shetland Islands. Endemic. 68. H. solum P. D. Sell & C. West Solitary Hawkweed Pseudophyllopodous perennial herb with a branched stock. Stem up to 50 cm, pale green, often flushed purplish, flexuous, slender, with a few, medium to long, pale, sometimes dark-based simple eglandular hairs at the very base, with scattered stellate hairs above, without glandular hairs. Leaves 2.5–9.0(–12.0 cult.) × 0.3–2.5(–4.5 cult.) cm, caesious on upper surface, paler beneath and sometimes purpletinged; basal with lamina broadly elliptical, narrowly elliptical or oblong, obtuse at apex, subentire and cuneate to a sessile, semiamplexicaul base; cauline 3–8, often with abortive branches in the axil of the leaves, the lamina lanceolate, gradually tapering from a rounded, sessile, semiamplexicaul base to a long-acute apex, subentire; all with scattered, pale, medium to long simple eglandular hairs on the margins and midrib, rarely beneath, sometimes without simple eglandular hairs, sometimes with puberulous margins, sometimes with scattered stellate hairs on one or both surfaces. Inflorescence with 3–7(–11) capitula, compactly cymose, often with branches from the axils of the upper cauline leaves, the acladium with short or medium peduncles; peduncles suberect, medium to long, with stellate hairs and occasionally with a solitary, very short simple eglandular or glandular hair. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 11–13 × 1.5–2.0 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with a few, scattered stellate hairs, scattered dark, medium simple eglandular hairs and scattered very short, black glandular hairs. Ligules yellow, glabrous. Styles discoloured.

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Receptacle pits with margins shortly dentate. Achenes 3.0– 3.5 mm, blackish-red. Flowers 7–8. Apomictic. This species has no near allies and is isolated geographically. Native. Dry, schistose rock-ledges between 310–500 m. Cliffs of Loch an Dughaill, An Stuchd and Cruach Lusach, in Kintyre. Endemic. 69. H. breve Beeby Rare Hawkweed Aphyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 15–33(–40 cult.), pale green, robust or slender, markedly striate, with dense, long simple eglandular hairs especially below, those of the upper part dark-based, with a few stellate hairs above, without glandular hairs. Leaves 12–55 × 5–35 mm, deep green on upper surface, becoming copper-coloured with age; basal usually few, or occasionally up to 9, the lamina more or less broadly elliptical to elliptic-rhomboidal, obtusemucronulate to acute at apex, remotely denticulate or occasionally with a few, shallow, acute, patent teeth, abruptly contracted to a rounded or cuneate base and the petioles broadly winged; cauline 4–11(–13 cult.), the lamina of the lower ovate-lanceolate to ovate, rarely ovate-rhomboidal, obtuse to acute at apex, denticulate or with 2–3, short, acute, patent teeth on each side, cuneate-based and subpetiolate or sessile, the lamina of the upper smaller, ovatelanceolate, acute at apex, toothed as lower, and the base rounded and semiamplexicaul; all with straight, medium, bulbous-based, subrigid simple eglandular hairs on both surfaces and the margins (in cultivated specimens sometimes very few), and on the petioles where they are similar but longer. Inflorescence with 2–6(–13 cult.) capitula, compactly cymose (in cultivated specimens the branches more elongated), the acladium with a medium peduncle; peduncles medium, suberect, with few stellate hairs, fairly numerous, short, patent simple eglandular hairs and occasionally a few, very short glandular hairs. Capitula about 20 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 8–10 × 1.7–2.0 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, rarely with a few stellate hairs at base, with numerous, very short glandular hairs, sometimes with a few, short simple eglandular hairs. Ligules yellow, glabrous. Styles discoloured. Receptacle pits with margins long-dentate. Achenes 3.0– 3.5 mm, blackish-brown. Flowers 7–8. Apomictic. No closely allied species to H. breve are known. Native. Exceedingly rare on granite crags on the north side of Ronas Voe, North Mainland in the Shetland Islands. Endemic. 70. H. zetlandicum Beeby Shetland Hawkweed H. demissum subsp. zetlandicum (Beeby) Zahn Phyllopodous or rarely hypophyllopodous perennial herb with a branched stock. Stem up to 40(–50) cm, pale green often tinted purplish, flexuous, slender, with numerous, medium to long, soft simple eglandular hairs at the base, which become gradually fewer upwards, the uppermost sometimes dark-based, usually stellately hairy throughout, but sometimes sparingly so or not at all near the base, some plants with an occasional, very short, blackish glandular

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hair near the inflorescence. Leaves deep green on upper surface, often suffused with purple; basal 3–6, variable, the lamina usually broadly elliptical, obtuse-mucronate to acute at apex, denticulate to shallowly but sharply serrate and narrowed below, or ovate, obtuse-mucronulate at apex, denticulate and cuneate-based, or in large specimens nearly rhomboidal, acute at apex, irregularly serrate with the teeth interspersed with denticulations and cuneate-based, with numerous, unequal, short to long, subrigid simple eglandular hairs on both surfaces and the margins, the petioles winged and with few to many, long simple eglandular hairs; cauline 2–4(–6 cult.), the lamina of the lower elliptical to elliptic-lanceolate or sometimes ovate, acute (occasionally obtuse-mucronate) at apex, denticulate to sharply serrate and cuneate-based with a winged petiole, the lamina of the median ovate to ovatelanceolate, acute to acuminate at apex, denticulate (sometimes finely serrate), cuneate- or rounded-based and semiamplexicaul, the lamina of the upper linear-lanceolate, linear or bract-like, all the cauline glabrous or with a few, medium simple eglandular hairs above, medium simple eglandular hairs beneath and on the margins and sometimes with sparse stellate hairs on one or both surfaces. Inflorescence with 1–7(–30 cult.), often geminate capitula, compactly cymose; peduncles short to medium, suberect, with numerous stellate hairs, numerous, short, black glandular hairs and a few, medium, usually dark-based simple eglandular hairs. Capitula 20–25 mm in diameter, cylindrical in bud, rounded at base. Involucral bracts incumbent in bud, 8–9(–10 cult.) × 1–2 mm, linear-lanceolate, the inner pale green, sometimes with a darker centre, more or less acute and sometimes abruptly narrowed near the apex, the outer blackish-green, shorter and less acute at apex, all with scattered stellate hairs, numerous black, unequal, very short to short glandular hairs and sometimes an occasional, pale, short, dark-based simple eglandular hair. Ligules deep yellow, glabrous. Styles yellow, becoming discoloured when dry. Receptacle pits with the margins sharply dentate. Achenes 3.5–4.0 mm, blackish-brown. Flowers 6–8. Apomictic. 2n = 27. This very distinct species has no near allies in the British Isles, but is closely allied to the Faeroese H. hartzianum Dahlst. and H. ostenfeldii Dahlst. and to the Icelandic H. arrostocephalum Omang. In pastures it is a small, neat plant often only up to 20 cm, but in sheltered sites is much taller. Native. Dry heaths and pastures, rocky, heathery streamsides and grassy sea-banks, mainly on gneiss. Formerly common in a restricted area near North Roe, extending from Burgo Taing northwards to Benigarth in North Mainland but now much diminished through agricultural pressures and grazing; recently detected at a number of new coastal sites in the North Roe and Ronas Voe areas of North Mainland. Also recorded from the Burn of Lunklet in Central Mainland, on rocky geos below Windy Scord, and by the west side of Snarra Ness (both in West Mainland), all in the Shetland Islands. Endemic. Section 7. Amplexicaulia (Griseb.) Scheele Taxon Amplexicaulia Griseb.; Taxon Balsamea Arv.-Touv.; Subsection Balsamea (Arv.-Touv.) Rouy

Plants with a very hairy apex to the rhizome and a basal rosette and stems, leaves and involucres with dense glandular hairs. Cauline leaves 3–6, with additional small ones on branches, amplexicaul. Capitula large, with narrow, more or less acute involucral bracts. Receptacle pits shortly dentate and fibrillous-ciliate. Achenes dark. Flowering from June to August. Natives of the mountains of central and south Europe and Morocco; widely introduced elsewhere. The species of this group are easily recognised by their strong glandular clothing throughout and amplexicaul cauline leaves. 71. H. speluncarum Arv.-Touv. Cave Hawkweed H. amplexicaule var. subhirsutum Arv.-Touv.; H. amplexicaule subsp. speluncarum (Arv.-Touv.) Zahn Phyllopodous perennial herb with a branched, hairy stock. Stem 20–85 cm, yellowish-green, sometimes flushed purplish, markedly striate, robust, with dense, long, pale simple eglandular hairs below, slowly petering out upwards, with few, unequal, short to medium, brownish-yellow glandular hairs below, becoming numerous to dense upwards, with few stellate hairs above. Leaves yellowish-green to rather dark green, sometimes slightly spotted or blotched on upper surface, paler beneath, the basal 7–19 × 2–8 cm, in a rosette, narrowly to broadly elliptical or oblong-elliptical, more or less acute at apex, regularly serrate or serratedentate, with aquiline-mammiform teeth, attenuate at base to a long, winged petiole, the cauline 3–8 with additional small ones on the branches, gradually decreasing in size upwards, 1.5–20.0 × 0.5–8.0 cm, the lower broadly oblong, acute at apex, serrate-dentate, the upper broadly ovate, acute to acuminate at apex, entire or nearly so, the lower shortly narrowed, the upper rounded or subtruncate at the amplexicaul base, all with numerous to dense, short to long, pale simple eglandular hairs on the margins, petioles and midvein beneath and fewer on the lower surface, with numerous, short, yellowish-brown glandular hairs on both surfaces and the margins. Inflorescence with 4–40 capitula, paniculatecorymbose, with long, robust, leafy branches, acladium with a long peduncle; peduncles suberect, short to long, with numerous to dense stellate hairs and numerous to dense, short to medium, yellowish-brown glandular hairs, without simple eglandular hairs. Capitula 30–50 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 7–16 × 1.5–2.0 mm, greyish-green, the inner with paler margins, all oblong-lanceolate, long drawn out to an acute apex, with dense short to long, unequal, yellowishbrown glandular hairs and numerous stellate hairs, without simple eglandular hairs. Ligules medium yellow, with dense simple eglandular and glandular hairs at the apex and on part of the surface. Styles yellow, discoloured when dry. Receptacle pits with the margins shortly dentate and fibrillous-ciliate. Achenes 3.5–4.0 mm, blackish. Flowers 6–8. Apomictic. Introduced. Garden escape established on walls and long known in several localities. Mells, Somerset, and Richmond Hill, Vivian Road and Cobblestone Mews, Bristol in Gloucestershire, Ham House in Surrey, Hitcham Abbey Farm in Norfolk, Colwyn Bay in Denbighshire, Darwen in Lancashire and Kenmore in Perthshire. The Kenmore plant

39. Hieracium was originally identified as H. pulmonarioides. H. speluncarum is native of the Alps and mountains of Corsica. 72. H. amplexicaule L. Sticky Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale yellowish-green, robust, markedly striate, with dense, unequal, short to long, yellowish glandular hairs throughout and numerous stellate hairs in the upper part, without simple eglandular hairs. Leaves yellowishgreen on upper surface, paler beneath, the basal numerous, in a rosette, 3–22 × 1–6 cm, oblong, elliptical, oblanceolate-oblong, obovate-oblong, obtuse-mucronate at apex, undulate-denticulate, gradually narrowed at base to a short to rather long, winged petiole, the cauline 4–6 plus 1–4 on the branches, gradually decreasing in size upwards, 2–17 × 0.7–7 cm, broadly elliptical, oblong-ovate, oblongobovate, oblong-oblanceolate or ovate, obtuse-mucronate to acute at apex, undulate-denticulate to subentire, gradually narrowed or rounded at the amplexicaul base, with numerous to dense brownish-yellow glandular hairs on both surfaces and the margins, without simple eglandular or stellate hairs. Inflorescence with 3–40 capitula, racemosecorymbose, usually with long, robust, leafy branches, the acladium with a rather long peduncle; peduncles short to long, with dense stellate hairs and dense, unequal, short to long, brownish-yellow glandular hairs, without simple eglandular hairs. Capitula 20–40 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 5– 14 × 1.2–1.5 mm, dark or greyish green, the inner with paler margins, the outer rather lax, broadly linear-lanceolate, gradually narrowed to an acute apex, with dense, unequal, short to medium glandular hairs, numerous stellate hairs and with a tuft at the apex, without simple eglandular hairs. Ligules medium yellow, with dense glandular hairs at the apex and on part of the surface. Styles yellow. Receptacle pits with margins shortly dentate and fibrillous-ciliate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. 2n = 36. Introduced. A garden escape found on walls and in waste places. Known for over 200 years on walls round the Botanic Garden and in stonework along the canal at the back of Magdalene College, Oxford. It occurred for many years on the walls of Cleish Castle in Fifeshire and was recorded in Upper Yoredale in Yorkshire. Native of mountain rocks in central and south Europe and Morocco; naturalised on walls elsewhere. 73. H. pulmonarioides Vill. Wall Hawkweed H. amplexicaule subsp. pulmonarioides (Vill.) Zahn; H. amplexicaule var. pulmonarioides (Vill.) Gaudin; H. speluncarum var. lanceolatum Arv.-Touv.; H. pulmonarium Sm. nom. illegit. Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, robust, markedly striate, with long, pale simple eglandular hairs mixed with numerous, unequal, short to medium, yellowish-brown to dark glandular hairs, the glandular hairs numerous to dense in the upper part, the simple eglandular hairs slowly petering out and dense stellate hairs above. Leaves dark or bluish green on upper surface, paler beneath, the basal 4–11 × 1.5–4.0 cm, in a rosette, obovate, oblanceolate,

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ovate, elliptical or oblong, rounded-mucronulate at apex, serrate-dentate, the teeth narrowly aquiline-mammiform, shortly narrowed at base to a medium, winged petiole, the cauline 3–5, 2–10 × 1–5 cm, gradually decreasing in size upwards, oblong, lanceolate or ovate, rounded-obtuse to the amplexicaul base or the lower sometimes shortly petiolate, all with numerous, short to long, pale simple eglandular hairs mixed with shorter dark glandular hairs on both surfaces and the margins. Inflorescence with 1–20 capitula, racemose-corymbose, with long, robust branches, sometimes from near the base, the acladium rather long; peduncles straight or arcuate, medium to long, with dense stellate hairs and dense, short to medium, dark glandular hairs. Capitula 25–40 mm in diameter, broadly rounded at base. Involucral bracts 5–13 × 1.5–1.8 mm, linear-lanceolate, gradually narrowed to an acute apex, with numerous stellate hairs especially on the margins, numerous glandular and without simple eglandular hairs. Ligules medium yellow, with dense glandular hairs on the apex and part of the surface. Styles yellow, discoloured when dry. Receptacle pits with margins shortly dentate, and fibrillous-ciliate. Flowers 6–8. Apomictic. 2n = 27, 36. Introduced. A garden escape. On walls surrounding Nottingham Castle and a nearby house where it is probably extinct, on a railway viaduct at Saltburn in Yorkshire and Stretford in Lancashire and at Gayle and Upper Yoredale in Yorkshire. Native of the Alps and Pyrenees. Section 8. Alpina (Griseb.) Gremli Taxon Alpina Griseb.; Subsection Eualpina F. N. Williams Phyllopodous perennial herbs with a branched stock. Stems usually rather short (up to 35(–75) cm), often with numerous simple eglandular hairs. Leaves pale to deep green, rarely bluish-green, usually with few to numerous, very short or short glandular hairs particularly visible on the margins, the basal usually more or less numerous and often in a rosette, the cauline mostly small and bract-like with the lowest one larger and usually narrow. Capitula 25–60 mm in diameter, usually solitary, sometimes 2–few, usually dark. Involucral bracts incumbent in bud, usually narrow, the outer lax, the inner rarely tightly appressed, often with numerous to dense, long simple eglandular hairs, usually obscuring the glandular hairs and stellate hairs usually absent. Ligules usually hairy on the back and tips; when hairs are said to project beyond the apex it means beyond the margins of the apical lobes. Receptacle pits with margins usually shortly dentate. Flowers 7–8. Mainly on cliffs, boulders, stony slopes and rocky sides of streams on mountains and usually above 550 m, but lower in the extreme north of Scotland. Chiefly in Scotland with two species in the English Lake District and one in North Wales. Absent from Ireland. Mountains of Europe and Arctic and Subarctic regions of Europe and Asia. The descriptions in this Section have been made by David Tennant from a combination of living plants collected in the field and cultivated by him and of herbarium specimens. This Section is the most difficult in the genus in which to define species, many of its populations occurring in isolated localities, while in other places as many as ten species occur together. The descriptions are very long and include, in particular, ligule characters for which information is

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not available for species in the other Sections. When the outer ligules are straight the top of the open capitulum will be more or less flat or broadly obconical, when they are upturned it will be more or less concave. Measurement of the diameter of the open capitulum has been made from living plants while in other Sections it has been made from herbarium sheets and is thus less exact. This more detailed description can be made in this group because the Section Alpina is well defined and once seen will not be forgotten. In the field many of the species have a distinctive range of involucre and leaf shapes, although difficult to define in words. Only H. perscitum and perhaps H. hanburyi and to a lesser extent H. completum, H. backhousei and H. subgracilentipes might be mistaken for a species of the Section Subalpina. All except one of the species are either triploid or tetraploid, the exception being a pentaploid, the first ever recorded in the genus as here defined. The species must have originated when there was a sexual diploid and an apomictic tetraploid which had fertile pollen and the apomictic genes were carried into the triploids. It would seem that one parent would have had a large hairy capitulum and one glandular hairs on the leaves. The dwarf habit, often small stem leaves and solitary capitulum, typical of mountain plants, must have been brought about by selection while the plants were still sexual, although some species may have been created as apomictic sports. The triploids may have originated from crosses between more than two species or may be segregates from the same cross. A detailed account of the chromosomes in this Section is given by C. A. Stace et al. (1995). 74. H. perscitum P. D. Sell & C. West Neat Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–27 cm, usually slender, often flexuous, tinted with pale violet below, with numerous to dense, slender, wavy, medium to very long, white, mainly dark-based simple eglandular hairs towards the base and scattered or sparse, very slender, very wavy, short to long ones in the upper part, with scattered, pale, very short glandular hairs in the lower part, and fairly numerous, very short to short, dark ones in the upper part, with scattered stellate hairs in the lower part which become more numerous, but not dense, in the upper part. Leaves rather pale to rather bright medium green or deeper, slightly bluish-green, faintly glossy or matt, usually tinted or marbled with dirty, dull violet on both surfaces and the margins, or paler on the lower surface; basal few to several, the outer with lamina 5–40 × 8–25 mm, subrotund, broadly ovate or broadly elliptical, broadly rounded-obtuse at apex, entire or denticulate or sometimes shallowly sinuate-dentate and cuneate-truncate to rounded at base, the inner with lamina 15–50 × 6–20 mm, ovate, ovate-lanceolate or elliptical, sometimes slightly undulate, rounded-obtuse, subacute or acute and frequently apiculate at apex, somewhat rigid but not or only slightly coriaceous, remotely denticulate or irregularly serrate-dentate, with narrow to broad, often ascending and reddish-violet-coloured, mammiform teeth and cuneate or occasionally attenuate at base, all with scattered to rather dense, slender, soft or subrigid, wavy, short, medium and occasionally long, white

simple eglandular hairs with pale or dark bulbous bases, often uniformly distributed on the upper surface of the inner leaves and becoming dense and very wavy on the margins, with rather fewer on the lower surface which become more sparse on the outer leaves and with a few very short, yellowish glandular hairs on the margins and both surfaces, the petioles usually rather short, slender, deeply tinted reddishviolet at the base and clothed with numerous or occasionally dense, medium to long simple eglandular hairs; cauline (0–)1–3 (–5), the lowest rigidly patent, frequently undulate and positioned one-third to halfway up the stem, with lamina 10–40(–70) × 2–15(–20) mm, usually lanceolate, occasionally narrowly ovate, often resembling the innermost basal, usually gradually and very sharply acute, acuminate or more or less cuspidate at apex, denticulate or irregularly serrate with a few, sharp or narrowly mammiform teeth, cuneate or attenuate at the often sessile, occasionally petiolate and often semiamplexicaul base, sparsely clothed with simple eglandular hairs on the upper surface but densely so on the margin and lower surface with black-based or pale-based ones, and the lower surface also with several stellate hairs, the second lowest semi-patent, sometimes with a rudimentary bud in its axil, linear or linear-lanceolate, small, entire and sessile or bract-like and glabrous on the upper surface but densely clothed with stellate hairs on the lower surface, the remainder if present extremely small, appressed or semipatent, filamentous, acute at apex and sometimes with a rudimentary bud in the axil. Capitula 1–2, 23–42 mm in diameter, the involucre greenish-black or blackish in appearance, narrowly campanulate, rounded or subtruncate at the base; peduncles when present clothed as the upper stem. Involucral bracts incumbent in bud, blackish-green, the inner with slightly paler margins, inner 10–13(–14) × 1.2–1.7 mm, closely appressed, rather broadly linearlanceolate, mainly obtuse with some subacute at apex, the outer shorter and more or less appressed; all with numerous, medium to long (–2.0 mm), slender, somewhat greyish, black-based simple eglandular hairs, numerous, mainly very short and short (–1.0 mm), dark glandular hairs few of which are obscured by the simple eglandular hairs and few or no stellate hairs but the inner with a tuft of short hairs at the apex. Ligules bright medium yellow, the outer up to 3.5 mm wide, more or less discrete, straight or slightly upturned and deeply lobed or laciniate-lobed at apex, with few or no pale simple eglandular hairs on the lower surface and few or no extremely short ones projecting beyond the apex. Styles dark greenish-yellow-grey. Receptacle pits with margins sometimes fimbriate-dentate. Achenes 3.9–4.7 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Rocky knolls and stream banks from 530 to 760 m on Torridonian Sandstone and Moine Schist. On the An Teallach range, Ben More Coigach, Braemore Forest and Fannich Forest hills in Ross-shire and Canisp and Carn Dearg in Sutherland. Endemic. 75. H. hanburyi Pugsley Hanbury’s Hawkweed H. chrysanthum Backh. fil., non Ledeb.; H. nigrescens subsp. chrysanthum Zahn Phyllopodous perennial herb with a branched stock. Stem 7–28(–35) cm, pale green, usually reddish-violet tinted

39. Hieracium at base, usually slender, very rarely somewhat robust, frequently flexuous, the lower part with few or scattered, medium to long, often dark-based, slender, wavy simple eglandular hairs, very few or no very short, pale or dark glandular hairs and scattered or sparse stellate hairs, the upper part with many to fairly numerous, mainly medium, occasionally rather long, slender, wavy, white or more often greyish simple eglandular hairs with long black bases, or occasionally none, very numerous, very short and short to medium, unequal, blackish glandular hairs and moderately to densely clothed with stellate hairs. Leaves pale to rather deep, usually bright, glossy, occasionally dull green, slightly paler beneath, frequently tinted with reddish-violet; basal numerous, sometimes in a neat rosette, the outer with lamina 7–50 × 5–35 mm, variable, subrotund, broadly ovate or obovate, occasionally undulate, rounded-obtuse, often mucronate, occasionally retuse at apex, denticulate or serrate-dentate with shallowly mammiform teeth, very occasionally sublaciniate or entire and broadly cuneate to shortly narrowed and occasionally more or less truncate at the base, the inner with lamina 10–100 × 4–35 mm, sometimes rigid, not or only slightly coriaceous, variable, narrowly to broadly elliptical, lanceolate, ovate, obovate or subrotund, obtuse to acute and mucronate, apiculate or occasionally cuspidate and slightly twisted at apex, subentire, denticulate or deeply incise-dentate and very irregularly dentate or serrate, with often large, broad or narrowly mammiform to spinulose, sometimes cusped and occasionally a few, retrorse teeth with long, rounded-obtuse-apiculate apices mainly on the lower two-thirds of the margin and which are interspersed with denticulations, occasionally with a few disjunct teeth descending on to the petiole and cuneate, shortly narrowed or attenuate at base, the petioles rather long, rarely short, usually slender, only occasionally winged and frequently reddish-violet tinted at base, the indumentum of all the basal very variable with numerous, medium or long simple eglandular hairs on the margin, which often become numerous to dense and long and wavy at the base, the majority of the leaves usually more or less uniformly clothed on the upper surface with scattered to numerous, short, medium and long, slender, wavy or apically curved, subrigid, white simple eglandular hairs with mainly pale, sometimes discoloured, but not black, bulbous bases, the outermost leaves only sparsely clothed or with none except near the margins, the innermost usually with numerous, rather long ones, the lower surface with scattered to numerous, short to medium, very slender, very wavy, sometimes appressed ones which are often denser on the midrib and many, mainly medium ones on the margins, with scattered, very short yellowish glandular hairs on both surfaces and the margins and sometimes with a few stellate hairs on the lower surface; cauline 0–3 (–4), the lowest suberect or semipatent, occasionally undulate, sometimes rigid, with lamina 10–55 × 2–20 mm, linear, linear-oblanceolate or oblanceolate, sometimes like the innermost basal, usually very acute, often acuminate or cuspidate at apex, entire or sometimes with several, irregular, acute teeth, attenuate to an often very distinct petiole, semiamplexicaul at base, usually nearly glabrous on the upper surface, moderately clothed on the margin and lower surface with often dark-based simple eglandular hairs and several to

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numerous stellate hairs, the remainder small or bract-like, acute at apex, suberect, the uppermost filamentous. Capitula 1–4(–5), 30–50(–55) mm in diameter, occasionally with additional stems arising from the base each bearing a capitulum, often with a rudimentary bud in the axil of a cauline leaf, the involucre usually blackish-grey or black, seldom greenish in appearance, campanulate, sometimes broadly campanulate, occasionally narrow and cylindricalcampanulate, very rounded at base and often slightly constricted above the base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, with paler margins, the inner 12–16 × 0.8– 1.5 mm, closely appressed, narrowly linear-lanceolate, straight or curved and attenuate to a finely or abruptly acute or acuminate apex, the outer very few, much shorter, semilax, suberect, narrow and subacute or obtuse at apex, all with few to numerous, mainly short and medium, sometimes rather long (up to 2.5 mm), slightly wavy, usually greyish, occasionally white simple eglandular hairs with long, black bases, or occasionally none, very numerous to dense, unequal, very short, short and medium, and sometimes several longish (up to 2.0 mm), black glandular hairs which are not obscured by the simple eglandular hairs or only partially obscured, usually without stellate hairs but the inner usually with a very distinct tuft of short hairs at the apex. Ligules bright medium to deep yellow, the outer up to 4.5 mm wide, very unequal, the longest very discrete, upturned or often involute towards the lobed or deeply lobed apex, usually glabrous or with some very short, pale, appressed simple eglandular hairs on the lower surface and with some extremely short, rarely a few very short ones (–0.2 mm) projecting beyond the apex. Styles pure yellow or slightly discoloured (olive) yellow, rarely more deeply discoloured, the stigmas usually yellow. Receptacle pits with margins shortly dentate. Achenes 3.5–4.5 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. This plant varies in almost all parts including size, shape and toothing of the leaves, size of capitula and proportion of glandular hairs to simple eglandular hairs. The variation seems to have no geographical or ecological significance. The three following forms are those that are most distinct. (1) Forma pusillum P. D. Sell & D. J. Tennant Plant often small. Stem with numerous glandular hairs and more or less numerous, medium to longish simple eglandular hairs in the uppermost part. Leaves often spreading, with lamina often ovate or subrotund and subentire or with small teeth, sometimes elliptical with larger teeth, rounded, cuneate-truncate or cuneate at base with short petioles. Involucral bracts abruptly acute to obtuseacuminate at apex with numerous simple eglandular and glandular hairs. Styles medium discoloured (olive-grey). 2n = 36. (2) Forma hanburyi H. chrysanthum var. microcephalum Backh. fil.; H. hanburyi var. microcephalum (Backh. fil.) Pugsley; H. hanburyi var. humile Pugsley Plant variable, often large. Stem with numerous glandular hairs and more or less numerous, medium to rather long simple eglandular hairs in the upper part. Leaves usually suberect, the majority with lamina usually elliptical,

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frequently with large teeth, cuneate to attenuate at base into medium to long petioles. Involucral bracts narrowly to broadly acute or acuminate at apex with more or less numerous simple eglandular hairs and numerous glandular hairs. Styles pure yellow to discoloured (olive-yellow). 2n = 36. (3) Forma atraticeps (Pugsley) P. D. Sell & D. J. Tennant H. hanburyi var. atraticeps Pugsley; H. atraticeps (Pugsley) P. D. Sell & C. West Plant variable, often large. Stem with numerous glandular hairs but very few or no simple eglandular hairs in upper part. Leaves usually suberect, the majority with lamina usually elliptical, often with large teeth, cuneate to attenuate at base into medium to long petioles. Involucral bracts narrowly to broadly acute or acuminate at apex, with dense unequal glandular hairs and very few or no simple eglandular hairs. Styles pure yellow to discoloured (olive-yellow). 2n = 36. Native. Cliff ledges, rocks, rocky knolls, grassy and heather slopes and by rocky streams between 550 and 1060 m on granite, quartz-biotite mica-schist and other formations. Widespread and not infrequent on the mountains of the Scottish Highlands, but not recorded from Sutherland. Endemic. The common widespread variant is forma hanburyi. Forma atraticeps is rare but occurs throughout most of the range of forma hanburyi. Forma pusillum is rare and known only from a few places in the western glens of Forfarshire. Named after Frederick Janson Hanbury (1851–1938). 76. H. kennethii P. D. Sell & D. J. Tennant Kenneth’s Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–24 cm, slender or medium, sometimes flexuous, occasionally tinted with reddish-violet at base, with very few to numerous, often slender, medium to long (–3 mm), wavy, dark-based, white simple eglandular hairs, scattered, very short, yellowish glandular hairs and very few or scattered stellate hairs in the lower part, with several to numerous, medium and long (–2 mm), white or greyish, black-based simple eglandular hairs, numerous to very numerous, very short to medium (–1.0 mm), dark glandular hairs and numerous to dense stellate hairs in the upper part. Leaves medium to rather deep bright green, glossy on both surfaces, slightly paler on lower surface; basal few to numerous, with the lamina of the outer 5–35 × 4–30 mm, subrotund, broadly ovate or obovate, often asymmetrical, broadly rounded-obtuse or sometimes retuse or apiculate at apex, denticulate or more or less sinuate-dentate, with several very irregular teeth and sometimes with a few, retrorse, mammiform or occasionally narrowly mammiform teeth and broadly cuneate to truncate at base, the lamina of the inner 6–45 × 4–25 mm, somewhat coriaceous, broadly ovate to elliptical, broadly acute and apiculate to obtuse at apex, sparsely denticulate to irregularly serrate-dentate, with medium to narrow, patent, mammiform teeth and broadly cuneate to narrowly cuneate, seldom attenuate at base, all with numerous to dense, slender, medium and long (–2 mm), subrigid, white, pale-based simple eglandular hairs more or less uniformly

distributed on the upper surface and margin but the outer often with few about the midrib, with several, very short, yellowish glandular hairs on the surface and margins, sometimes with a few stellate hairs on either surface about the midrib, the petioles short to rather long with dense, long, white simple eglandular hairs; cauline 1–5, the lowest semipatent, with lamina 3–20 × 2–10 mm, oblanceolate, occasionally narrowly elliptical and sometimes almost at the base of the stem and resembling the innermost basal, acute or obtuse, sometimes acuminate or cuspidate at apex, entire or serrate-dentate with several, irregular, sharp teeth, attenuate or very narrowly cuneate at base into a usually narrowish petiole, with few, pale simple eglandular hairs more or less uniformly scattered on both surfaces and the margins, becoming dense on the petiole, those on the the lower surface with blackish bases and occasionally with some stellate hairs mainly on the midrib, the remainder small, entire and more or less bract-like. Capitula usually solitary, sometimes 2, 30–48 mm in diameter, the involucre greyishgreen to greenish-black in appearance, campanulate but slightly narrowed at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green or green, the inner 11–15 × 1.2–1.8 mm, unequal, closely appressed, linear-lanceolate, and mainly abruptly acute, subacute or sometimes obtuse at apex, the outer shorter, erect and semiappressed, all with numerous, slender, wavy, medium or long (–3 mm), white or greyish, black-based simple eglandular hairs and numerous, short and very short, blackish glandular hairs largely obscured by the simple eglandular hairs, without stellate hairs but the inner with a pronounced tuft of short hairs at the apex. Ligules uniformly rather deep, occasionally medium yellow, the outer 3.5–4.0 mm wide, slightly discrete, almost straight to slightly upturned, and more or less deeply lobed at apex, almost straight to slightly upturned, with very few to several, very short, pale simple eglandular hairs on the lower surface and none to several extremely short or very short ones projecting beyond the apex. Styles more or less uniformly dark greenish-grey. Receptacle pits with margins dentate. Achenes 3.9–4.5 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Occurs rather locally on rocky knolls and steep, grassy slopes from 450 to 700 m on Torridonian Sandstone on An Teallach, on north-eastern and north-western sides of the An Teallach range, Dundonnell in Rossshire and on Cnoc na Creige, near Glen Coul in Sutherland. Endemic. Named after Archibald Graham Kenneth (1915–89). 77. H. completum P. D. Sell & C. West Complete Hawkweed H. curvatum auct.; H. nigrescens auct.; H. nigrescens var. curvatum auct.; H. pseudocurvatum auct. Phyllopodous perennial herb with a branched stock. Stem 13–30 cm, often robust, sometimes woody at base and usually thickened beneath the involucre, striate, usually reddish-violet tinted at base and sometimes flecked on the ridges in the lower part, with few to many, pale, dark-based, short to long simple eglandular hairs, usually with numerous long ones near the base and becoming shorter with longer blackish bases in the upper part, with few to numerous, very

39. Hieracium short and short, pale and dark glandular hairs and some stellate hairs, both stellate and glandular hairs becoming more numerous in the upper part. Leaves pale to medium, bright green, usually rather glossy or rarely matt, usually blotched or marbled with dull brownish-violet on the whole upper surface, often paler on lower surface with midrib tinted reddish-violet; basal usually numerous, occasionally coriaceous, the outer with lamina 5–45 × 3–35 mm, subrotund, ovate, elliptical or rarely obovate, rounded-obtuse to obtuse and often mucronulate at apex, denticulate, mainly broadly cuneate or sometimes truncate at base, the inner with lamina 10–70(–100) × 4–30 mm, mainly ovate, some broadly elliptical or broadly lanceolate, broadly acute or subacute, occasionally obtuse and usually mucronate or apiculate at the sometimes twisted apex, more or less regularly serratedentate with evenly spaced, acute, narrowly mammiform teeth or sometimes only denticulate, and cuneate, rounded or attenuate at the base, the majority with few to numerous, white, subrigid, short, medium and longish, wavy or apically curved, rather slender, usually pale-based simple eglandular hairs more or less uniformly distributed on the upper surface and margin, numerous, softer, wavy, short to medium ones on the lower surface and a few, very short, yellowish glandular hairs on the surfaces and margins, the petioles short or long, usually reddish-violet tinted especially at the base and with few to numerous, medium to long, sometimes appressed simple eglandular hairs; cauline 1–3(–4), the lowest with lamina 10–60 × 2–23 mm, frequently resembling the inner basal, lanceolate, elliptical or narrowly ovate, acute-mucronate or acuminate at apex, denticulate or irregularly serrate-dentate, with a few, narrow, acute teeth and often cuneate at base and distinctly petiolate, the remainder erect, linear and acute at apex or more or less bract-like, the uppermost filamentous and all with numerous simple eglandular hairs, some very short, yellowish glandular hairs and often some stellate hairs on the lower surface. Capitula usually solitary, but not infrequently 2–3, 35–50 mm in diameter, the involucre blackish or dark greenishgrey in appearance, usually broadly campanulate, markedly constricted in the middle; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, greenish-black, the inner with indistinct greenish margins; inner 11–15(–18) × 1.2–1.8(–2.0) mm, unequal, markedly appressed, more or less broadly linear-lanceolate, abruptly acute to subacute at apex; outer shorter, erect and more or less obtuse at apex; all with rather dense, slender, medium to long (up to 3 mm), white to greyish simple eglandular hairs with long, thickened black bases and numerous, very short and short, yellowish and dark glandular hairs which are only partially obscured by the simple eglandular hairs, without stellate hairs but the inner with a tuft of short branched and simple hairs at the apex. Ligules bright and rather deep, occasionally medium yellow, the outer rather broad (up to 4.0(–4.5) mm wide), slightly discrete, upturned, markedly ascending and deeply lobed at the apices, with none or few, very short to short, pale simple eglandular hair on the lower surface and several extremely short ones projecting beyond the apex. Styles uniformly very dark greyish. Receptacle pits with margins shortly dentate. Achenes 3.6–4.5 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36.

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Plants vary in size, degree of leaf toothing and number of capitula. This species has been long mistaken for H. curvatum Elfstrand and H. pseudocurvatum. Native. Rock-ledges, dry gullies and streambanks on both schist and granite between 600 and 900 m. Recorded on the Cairnwell in Perthshire; in several places in and around the heads of Glen Clova and Glen Isla in Forfarshire; in Glen Callater, on Lochnagar and at Dubh Loch in Aberdeenshire; on Creag an Dail Bheag (Little Craigandail) and in Coire Etchachan in Aberdeenshire and on many of the mountains bordering both sides of the Lairig Ghru Pass in Aberdeenshire and Inverness-shire and in Glen Einich and Coire Garbhlach in Inverness-shire, all in the Cairngorm mountains and on the adjacent Monadhliath Hills in Invernessshire. Endemic. 78. H. probum P. D. Sell & C. West Excellent Hawkweed Phyllopodous perennial herb with a branched stock. Stems 13–35 cm, slender to robust, sometimes slender above and robust below, often flexuous and distinctly striate, very sparsely clothed with simple eglandular, glandular and stellate hairs in the lower part and with a few to many, short to rather long, slender, whitish simple eglandular hairs with long, black, rather swollen bases, but rarely with very many long ones, many to numerous, very short to short (–0.7 mm), blackish glandular hairs and occasionally with few, usually numerous to dense stellate hairs in the upper part. Leaves rather pale to deep, rather drab green, barely glossy or matt, occasionally with a few brownish-violet markings on the upper surface, paler beneath and occasionally with reddish-violet markings; basal numerous, the lamina of outer 10–30 × 7–25 mm, ovate, obovate to subrotund or elliptical, sometimes undulate, rounded-obtuse, sometimes apiculate and occasionally retuse at apex, entire or denticulate and attenuate, cuneate or subtruncate at base, the lamina of inner 35–70 × 13–25 mm, obovate, oblanceolate, oblong-elliptical or elliptical, undulate, often obtuse, sometimes subacute or broadly acute, usually apiculate and slightly twisted at apex, somewhat coriaceous, often rigid, incise-dentate or serrate-dentate, with irregular, broadly to narrowly mammiform or aquiline-mammiform teeth which occasionally descend onto the petiole, or denticulate and cuneate or attenuate at base, all with few to many, medium to long (–2 mm), subrigid, wavy, white simple eglandular hairs with rather pale, bulbous bases on the upper surface or none except at or near the margins, few to many shorter ones more or less uniformly distributed on the lower surface and on the margins, and a few, very short, fine glandular hairs on the surfaces and the margins, the petioles rather slender and sometimes long (up to 70 mm) to occasionally broadly winged, reddish-violet at base and with few to many, medium to rather long, sometimes appressed, pale simple eglandular hairs on the margin or none; cauline 1–3(–5), the lowest erect and appressed or spreading, with lamina 10–50 × 2–12(–20) mm, sometimes undulate, linear-elliptical or linear-oblanceolate, acute, abruptly acute or obtuse and often twisted at apex, subentire or dentate with a few teeth, shortly narrowed or attenuate at base, more or less petiolate and often winged, with many, pale, medium simple eglandular hairs, often confined towards

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and at the margins on the upper surface with many medium, often black-based ones and sometimes with a few stellate hairs on the lower surface, the remainder erect, linear and bract-like or filamentous, acute at apex. Capitula usually solitary, sometimes 2(–5), or with additional stems from the basal rosettes bearing capitula, 35–45 mm in diameter, the involucre dark greenish-grey or greenish-black in appearance, campanulate or broadly campanulate, round but often slightly narrowed into peduncle at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, the inner with indistinct paler margins, inner (13–)15–17 × 1.0–1.7 mm, unequal, appressed, linear-lanceolate, mainly abruptly acute, subacute or obtuse and acuminate, sometimes incurved at apex, the innermost often narrower and acute; the outer shorter, appressed or rather lax; all with numerous, white to somewhat greyish, slender, medium to long (–3 mm) simple eglandular hairs with long, blackish, thickened bases and several to numerous, obvious very short to short (–0.7 mm) dark glandular hairs and without stellate hairs, but with a distinct tuft of branched and simple hairs at the apex. Ligules rather dull, deep yellow, dense centrally, the outer up to 4.0 mm wide, moderately to markedly discrete, slightly to distinctly upturned and deeply lobed or occasionally not developed or rolling inwards at apex, eglandular hairs on the apical part of the lower surface and very few to several, extremely short (to very short), ones projecting beyond the apex. Styles dark greenish-grey, the stigma only marginally paler. Receptacle pits with margins shortly dentate. Achenes 3.2– 4.2 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Local on rocks and ledges between 550 and 915 m, on hard mica-schist and granite. On Creagan Liatha, Stob Binnein, An Caisteal and Cruach Ardrain all near Crianlarich in Perthshire, and Coire Ardair, Coire Moy and Coire nan Gall around Creag Meagaih, Aonach Mor and Aonach Beag in Inverness-shire and Bidean nam Bian in Glen Coe in Argyllshire. Endemic. 79. H. pensum P. D. Sell & C. West Valuable Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–30 cm, medium to robust, often flexuous, sometimes reddish-tinted at base, with numerous to dense, slender, medium to long, white, blackish-based simple eglandular hairs, few, very short, mainly pale glandular hairs and rather sparse stellate hairs in the lower part, several to many, slender, long, usually somewhat greyish, black-based simple eglandular hairs, numerous, very short and short (–1.0 mm), dark glandular hairs and dense stellate hairs in the upper part. Leaves pale or medium green, with a faint, bluishgrey surface bloom, only slightly glossy or matt, sometimes lightly tinted with reddish-violet on the margins and near the apex, with conspicuous pale veins, the lamina of outer 10– 50 × 8–27 mm, subrotund, obovate or elliptical, roundedobtuse at apex, sharply denticulate or with a few, shallow to narrowly mammiform teeth and cuneate at the base, the lamina of inner 20–50(–80) × 8–30 mm, oblanceolate to obovate or elliptical, occasionally ovate, flat or slightly channelled, occasionally slightly undulate, obtuse and distinctly mucronate to subacute and cuspidate at the sometimes

twisted apex, serrate with irregularly ascending or retrorse, narrowly mammiform, sometimes cusped teeth and cuneate or attenuate at base, all with numerous to dense, medium to very long, slender, wavy, subrigid, pale-based simple eglandular hairs on the margins, the majority of the leaves with no simple eglandular hairs on the upper surface except the innermost which are sometimes uniformly clothed, and the lower surface moderately clothed, either uniformly or more densely on the midrib with very wavy, short to long, sometimes dark-based ones, with some very short, yellowish glandular hairs on the lower surface and margins; petioles medium to long, slender or winged, mainly rather densely clothed with long, wavy, patent or retrorse simple eglandular hairs; cauline (2–)3–5(–7), the lowest sometimes near the base of the stem and more or less spreading, the lamina 10– 45(–60 cult.) × 2–20 mm, sometimes undulate, oblanceolate or narrowly elliptical, narrowly acute at apex, denticulate or with some, small, narrow teeth, attenuate at base and petiolate, densely clothed with long, dark-based simple eglandular hairs on the margins, becoming very dense, very long and wavy at the base, with few or none on the upper surface and few stellate hairs on the midrib on both surfaces and the margins or none, the remainder narrowly elliptical or linear and entire or the uppermost bract-like. Capitula usually solitary, sometimes with a rudimentary bud in the axil of a cauline leaf, occasionally with 2 or more capitula or with a second stem arising from the basal rosette bearing a capitulum, 35–58 mm in diameter, the involucre large, dark greenish-grey or greenish-black in appearance, campanulate, broadly rounded at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, greenish-black; inner 12–20 × 1.1–1.8 mm, somewhat lax, slightly curved, linear-lanceolate and mostly sharply acute at apex; outer lax, slightly shorter and abruptly acute at apex; all densely clothed with long (to 3.5 mm), white to somewhat greyish, black-based simple eglandular hairs, numerous very short to short and some medium (up to 1.0 mm), dark glandular hairs only partially obscured by the simple eglandular hairs, without stellate hairs, but the inner with a tuft of short hairs at the apex. Ligules rather pale to medium yellow, the outer up to 4.7 mm wide, slightly discrete, almost straight and deeply and often irregularly lobed at the apex, with a few, very short and short pale simple eglandular hairs on the lower surface, or none, and few to several very short, and sometimes a few, short ones (–0.4 mm) projecting beyond the apex. Styles uniformly dark greenish-grey. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Cliff ledges and large rocks and infrequently rock detritus on quartz-biotite mica-schist and weathered microgranite between 520 and 915 m. On Beinn Dearg, Seana Bhraigh, Fannich Forest and Braemore Forest hills in Rossshire. Endemic. 80. H. calenduliflorum Backh. fil. Marigold Hawkweed H. halleri var. calenduliflorum (Backh. fil.) F. N. Williams; H. nigrescens subsp. calenduliflorum (Backh. fil.) Zahn Phyllopodous perennial herb with a branched stock. Stem 10–26(–32) cm, medium to robust, sometimes flexuous,

39. Hieracium usually only faintly reddish-violet tinted at the extreme base, with medium and long, thick, wavy, white to somewhat greyish simple eglandular hairs with thickened black bases throughout, often dense and very long towards the base, shorter and greyer in the upper part with long blackish bases, with very short to short, dark glandular hairs, becoming numerous in the upper part, and with scattered stellate hairs becoming dense in the upper part. Leaves rather pale to medium bluish-green and pruinose, rarely deeper green, matt or only faintly glossy; basal few to numerous, the lamina of outer 15–60 × 10–37 mm, subrotund, obovate or broadly elliptical, broadly rounded-obtuse at the apex, sometimes retuse, subentire, denticulate, or with a few teeth like those of the inner and attenuate, cuneate or shortly narrowed at the base, the lamina of inner 25–90 × 15–35 mm, sometimes rather rigid but not coriaceous, mostly broadly elliptical, obovate-spathulate or narrowly obovate to oblanceolate, rounded-obtuse to subacute, frequently mucronulate to apiculate, occasionally cuspidate at the slightly twisted apex, subentire, denticulate or more frequently serrate-dentate with large irregular, sometimes incise, unevenly spaced, narrowly mammiform, cuspedmammiform, slender aquiline-mammiform or occasionally spinulose teeth which are often rounded-obtuse-apiculate at their apices, most frequently confined to the lower part of the margin and sometimes descending onto the petiole, occasionally sinuate-dentate with very irregular, broad, rather obtuse teeth and cuneate to attenuate at the base, the majority usually uniformly clothed with numerous, white, bulbous-based simple eglandular hairs on both surfaces and the margin, which are very dense, thick, wavy, very long, patent and black-based on the innermost leaves, especially in eastern populations, sometimes shorter and less dense in western populations with black or pale-based, subrigid, long or medium ones, the outermost often with few except on the margin, all with some very short yellowish glandular hairs on both surfaces and on the margin, the petioles usually more or less broadly winged, rather short to long, usually only faintly reddish-violet tinted at the base, those of the inner leaves at least with dense, very long, coarse, wavy, white, blackish-based simple eglandular hairs on the margin which are often less dense and shorter in the western populations; cauline 2–3(–5), the lamina of lowest 20–50 × 1–16 mm, linear-oblanceolate to oblanceolate, acute or subacute and apiculate, occasionally acuminate at apex, entire or denticulate, sometimes with a few, larger teeth like those of the inner basal, very rarely markedly dentate in eastern populations, subpetiolate, frequently decurrent and attenuate at base, the remainder more or less linear, bract-like, entire, suberect and acute at apex, all the cauline often sparsely clothed with simple eglandular hairs on the upper surface but with numerous longish ones on the margin. Capitula 1– 2(–3), 40–60 mm in diameter in eastern populations, 30–50 mm in western populations, not infrequently with additional stems arising from the basal rosette each bearing a capitulum, the involucre broadly campanulate, broadly rounded or rounded at base, very dark grey in appearance; peduncles when present clothed like the stem. Involucral bracts incumbent in bud, greenish-black, inner 12–18 × 1.0–1.8 mm in eastern populations, 9–15 × 1.3–2.1 mm in western pop-

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ulations, numerous, appressed to somewhat lax, narrowly to broadly linear-lanceolate, subacute, sharply acute or acuminate at apex; outer shorter, semilax, erect, narrower than the inner in eastern populations or frequently broader in western populations, more or less acute at apex; densely clothed with long, wavy, whitish or greyish simple eglandular hairs with long, blackish, thickened bases, many very short to short (–0.6 mm), blackish glandular hairs with few to several obvious ones which are unobscured by the simple eglandular hairs, without stellate hairs but the inner with some very short, branched and simple hairs at the apex. Ligules bright, pale yellow, occasionally almost medium yellow, the outer broad (up to 5 mm), outermost long and very discrete (eastern populations) or shortly and moderately discrete (western populations), upturned and sometimes ascending at the deeply lobed apex, with several to numerous, very short and short or medium, sometimes rather appressed, pale and often discoloured simple eglandular hairs on the lower surface, usually with several to numerous, very short and often a few short ones projecting beyond the apex. Styles dark grey; stigmas not or only slightly paler. Receptacle pits with margins shortly dentate. Achenes 3.7–5.2 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Eastern and western populations tend to differ from one another in size, length of hairs, size and dentation of cauline leaves and length and width of involucral bracts and ligule arrangement. The eastern populations show very little variation, but the western populations show considerable variation making it difficult to distinguish a separate taxon. The most densely hairy plants of the western populations strongly approach H. notabile, but can be distinguished by their less attenuate basal leaves, slightly broader-based involucre and much darker, longer achenes. They all appear to be tetraploid. Native. Cliff ledges, rocky streambanks and amongst short Calluna between 520 and 980 m. The eastern populations occur on granite on Lochnagar and at Dubh Loch in Aberdeenshire and locally in the Clova Mountains of Forfarshire and are recorded near Glen Shee in Perthshire. Western populations occur on granite and gneiss on the Ben Nevis range and the hills of Glen Spean in Invernessshire and around Glen Coe and Glen Etive in Argyllshire. Endemic. 81. H. notabile P. D. Sell & C. West Remarkable Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–26 cm, yellowish-green, sometimes reddish-violet tinted near the base, medium to somewhat robust, often striate and flexuous, with numerous, short, medium and very long, rather thick, wavy, blackish-based simple eglandular hairs, becoming dense in the lower part with sometimes semiappressed, often retrorse, very long white ones, and usually with numerous to dense, very long, whitish ones in the upper part, with few, very short, pale and dark glandular hairs in the lower part becoming mainly short and more numerous above, and with few stellate hairs towards the base becoming numerous to dense above. Leaves rather pale to medium, faintly bluish-green and pruinose,

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matt or only faintly glossy on upper surface, occasionally reddish-violet tinted; basal few to numerous, the lamina of outer 6–55 × 5–30 mm, obovate, broadly elliptical, narrowly obovate or occasionally subrotund, rounded-obtuse and often mucronate at apex and usually subentire and cuneate or attenuate at base, the lamina of inner 3–75 × 3– 25 mm, often rather rigid, but not coriaceous, sometimes undulate, narrowly to broadly elliptical or broadly oblanceolate, obtuse and mucronate to apiculate or subacute at the sometimes slightly twisted apex, either subentire, denticulate, dentate or serrate-dentate, with several, large, irregular, shallowly mammiform or narrowly aquiline-mammiform teeth, often rounded-obtuse-apiculate at their apices, or with cusped teeth and often very attenuate at base, all with numerous medium to very long (up to 5 mm), often subrigid, white, blackish-based simple eglandular hairs uniformly distributed on both surfaces of the majority of the leaves and on the margins and few to many, very short, yellowish glandular hairs on both surfaces and the margins, the petioles short to long, frequently winged, with several to numerous, long, white, dark-based simple eglandular hairs on the margin; cauline 2–3(–4), the lowest rigidly semipatent, the lamina 40–70 × 8–20 mm, oblanceolate, acute or subacute at apex, often undulate and serrate-dentate on the margin with a few, narrow, mammiform teeth, often similar to the innermost basal, sessile and sometimes slightly amplexicaul at base, the second lowest not infrequently also rather large, often semipatent, the lamina 20–55 × 4–13 mm, narrowly oblanceolate, subacute to acute at apex and petiolate to decurrent and semiamplexicaul at base, the remainder smaller, more or less bract-like, suberect to semipatent, all clothed with very long, dense, wavy simple eglandular hairs, becoming very long at the bases of the leaves. Capitulum solitary, 30–45(–50) mm in diameter, the involucre campanulate, rounded but slightly narrowed at base, silvery grey in appearance. Involucral bracts incumbent in bud, blackish-green, inner 11–16 × 1–2 mm, somewhat lax, usually unequal, broadly linear-lanceolate or lanceolatesubulate and attenuate-acute or occasionally acuminate at apex; outer 7–10 × 1.0–1.8 mm, rather lax, erect or occasionally recurved, unequal, lanceolate, acute to subacute at apex; densely clothed with medium and long (up to 5 mm), very wavy, whitish simple eglandular hairs with long, thickened blackish bases more or less completely or only partially obscuring the few to numerous, very short, dark glandular hairs, without stellate hairs, but the inner often with a tuft of very short hairs at the apex. Ligules bright, pale to occasionally medium yellow, the outer broad (up to 5 mm), numerous, slightly discrete, straight, occasionally slightly upturned or very slightly recurved and deeply lobed at apex, with few to numerous, pale, or often slightly greyish, very short to medium simple eglandular hairs on the lower surface and several to numerous very short and short ones projecting beyond the apex. Styles dark greenish-grey, stigmas only marginally paler. Receptacle pits with margins shortly dentate. Achenes (2.5–)3– 4 mm, reddish-brown, never purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Cliff ledges on mica-schist between 800 and 950 m. Originally found on the east side of Ben More

in Perthshire in 1910, where it has never been re-found. Now known only from the Coire Ardair, and Moy Corrie on Creag Meagaidh and Coire nan Gall, all on the north side of Loch Laggan, Inverness-shire, where it is not infrequent. Endemic. 82. H. subgracilentipes (Zahn) Roffey Borrowdale Hawkweed H. chrysanthum var. gracilentiforme F. Hanb.; H. nigrescens subsp. subgracilentipes Zahn Phyllopodous perennial herb with a branched stock. Stem 12–34 cm, pale green, often reddish-violet towards the base, medium in thickness, sometimes flexuous, often striate, the lower part usually with few, short to medium simple eglandular hairs, sparse, extremely short to short, pale to discoloured glandular hairs and scattered stellate hairs, the upper part with few to numerous, short to medium and often some long, slender, wavy, usually rather greyish, blackbased simple eglandular hairs, many very short to short (–medium) dark glandular hairs which become numerous to dense above, and numerous to dense stellate hairs. Leaves drab, pale to somewhat deepish green, often with reddishviolet tints, more or less matt on the upper surface, paler on lower surface and minutely granulate in appearance; basal numerous, the lamina of outer 10–50 × 5–25 mm, subrotund, broadly elliptical, obovate or obovate-lanceolate, sometimes undulate, broadly obtuse and often mucronulate or apiculate at apex, entire, denticulate or occasionally remotely dentate and cuneate or shortly narrowed at base, the lamina of inner 15–70(–110) × 3–22(–28) mm, narrowly elliptical or elliptical to oblanceolate, rigid, semicoriaceous, frequently undulate and flat to markedly channelled, obtuse-apiculate to acute and sometimes acuminate at the often slightly twisted apex, denticulate or more frequently serrate-dentate with irregular, ascending, mammiform or aquiline-mammiform teeth which often have long, obtuse-apiculate apices and are mainly confined to the lower two-thirds of the margin and attenuate or sometimes shortly narrowed at the base, very sparsely or moderately clothed on the upper surface with mainly short to medium, sometimes long, wavy, subrigid, white simple eglandular hairs with pale or sometimes discoloured or blackish bases, usually with fewer or none about the midrib, the outer leaves often with none or with many short to medium ones on the margins, and with scattered ones on the lower surface which are often denser on the midrib, with scattered, very short, yellowish glandular hairs on both surfaces and on the margins and sometimes with a few stellate hairs, the petioles slender or sometimes winged, short to medium, occasionally long, frequently reddish-violet tinted, clothed on the margin with few to numerous, medium to long, wavy simple eglandular hairs which occasionally become dense at the base and with many, very short, yellowish glandular hairs; cauline (1–)2–3, the lowest patent or suberect, the lamina 20–35 × 2–10 mm, occasionally slightly undulate, linear-elliptical to linear-oblanceolate, usually finely acute or acuminate at the apex, entire or denticulate and attenuate at base to an obscure petiole, the remainder linear, bract-like and acute at the apex, the uppermost filamentous. Capitula solitary or sometimes 2 or more, sometimes with

39. Hieracium a rudimentary bud in the axil of a cauline leaf, 33–48 mm in diameter, the involucre dark greenish-grey in appearance, campanulate and rounded at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, the inner with paler margins; inner unequal, 12–19 × 0.8–1.9 mm, more or less appressed, linear or narrowly linear-lanceolate and very gradually and finely acute, acuminate or rather abruptly acute at apex; outer short, acute or acuminate at apex; all with dense, often somewhat greyish, medium to long, wavy, slender simple eglandular hairs with long black bases, numerous, mainly short, dark glandular hairs, which are only partially obscured by the simple eglandular hairs and without stellate hairs, but the inner with a tuft of short hairs at the apex. Ligules rather pale to almost medium yellow, the outer up to 4 mm wide, slightly discrete at the rather shallowly but narrowly lobed apices, with very few to many, short to medium and often numerous very short, pale or slightly discoloured simple eglandular hairs on the lower surface and very few to several, extremely short or very short and occasionally a few short ones projecting beyond the apex. Styles markedly discoloured, greenish-grey. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Rock and cliff ledges on basic Borrowdale Rock in Cumbria between 550 and 840 m. On Catstye Cam, Dollywagon Pike, Keppel Cove, Striding Edge and the head of Glencoynedale in the Helvellyn Range, Rampsgill Head, Kidsty Pike and Caudale Crags in the High Street Range, and Pillar in Ennerdale. Recorded on Glaramara at the head of Borrowdale. Endemic. 83. H. optimum P. D. Sell & C. West Splendid Hawkweed Phyllopodous perennial herb with a branched stock. Stem 15–30 cm, striate, often reddish-violet at base, with scattered to numerous, short to long, whitish, dark-based simple eglandular hairs, scattered, very short, yellowish glandular hairs and a few stellate hairs in the lower part, with scattered to numerous, usually somewhat greyish, short to long, black-based simple eglandular hairs, numerous, very short and some short, yellowish and dark glandular hairs and numerous stellate hairs in the upper part. Leaves rather drab, rather pale to medium, bluish-grey-green, matt, often partially tinted or suffused with dull reddish-violet; basal few to numerous, the lamina of outer 5–60 × 5–25 mm, subrotund or broadly ovate, obtuse-mucronate at apex, entire, denticulate or with a few, shallow teeth and more or less cuneate at base, the lamina of inner 25–70 × 5–25 mm, mostly broadly to narrowly elliptical, occasionally oblanceolate or obovate, obtuse-mucronate or subacute at apex, entire, denticulate or with a few, shallow, acute teeth and attenuate at base, the petioles sometimes long, and with numerous, white, medium to long simple eglandular hairs; cauline 1–2(–3), the lowest often rather patent, the lamina 10–45 × 4–11 (–20) mm, oblanceolate to lanceolate, obtuse to acute at apex, entire, often undulate, attenuate at base and shortly petiolate, the remainder more or less bract-like, all with scattered, white, pale or dark-based, short to medium simple eglandular and very short glandular hairs and with very

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numerous stellate hairs especially on the lower surface, the innermost basal often with very numerous, long, blackbased simple eglandular hairs on the lower surface and margins. Capitula usually solitary, sometimes 2, and often with a rudimentary bud in the axil of a cauline leaf, 30– 45 mm in diameter, the involucre campanulate, broadly rounded at base and greyish-black in appearance; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, inner 15–17 × 1.5–2.0 mm, semiappressed, rather broadly linear-lanceolate, more or less tapering to a sharply or abruptly acute apex; outer short; blackish-green, densely clothed with long (up to 5 mm), wavy, white or more usually greyish, black-based simple eglandular hairs and numerous, very short and short, blackish-stalked, yellow-headed glandular hairs which are more or less completely obscured by the simple eglandular hairs, without stellate hairs but the inner with numerous, short, branched hairs at the apex; outer short. Ligules bright, rather pale to medium yellow, the outer rather narrow (up to 3.5 mm), discrete, shallowly upturned or straight and irregularly lobed at apex, with few to many, very short or short, pale simple eglandular hairs on the lower surface and several, very short to short ones (–1.0 mm) projecting beyond the apex. Styles dark olive-grey. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 7–8. Apomictic. Native. A local and very rare, distinct species occurring on rock ledges on Dalradian Mica-schist from 700 to 820 m. Meall Buidhe and Beinn a’ Chreachain in Argyllshire. Endemic. 84. H. grovesii Pugsley Groves’s Hawkweed Phyllopodous perennial herb with a branched stock. Stem 12–30 cm, pale green, reddish-violet tinged towards the base, slender to slightly robust, often flexuous, striate, the lower part with scattered short to medium, pale-based simple eglandular hairs, very short, fine, yellowish glandular hairs and some stellate hairs, the upper part with numerous, slender, short to medium and sometimes several long (up to 2 mm), pale, black-based simple eglandular hairs, several to many, very short and a few short, dark glandular hairs and dense stellate hairs. Leaves pale to somewhat deepish green, mostly matt on the upper surface, or the innermost leaves more or less glossy, frequently with dark reddishviolet markings or tints, especially on the margin, both surfaces minutely granulate in appearance; basal numerous, the lamina of outer 8–55 × 4–22 mm, broadly elliptical, obovate or oblong-elliptical, rounded-obtuse-mucronate, occasionally slightly retuse at apex, entire or denticulate and cuneate or shortly narrowed at base, the lamina of inner 15– 80 × 5–20 mm, rigid and coriaceous, sometimes undulate, mainly narrowly elliptical, often with some oblanceolate to linear-elliptical and obtuse to acute, sometimes apiculate and often twisted at the apex, occasionally subentire or denticulate but usually rather sparsely or irregularly dentate or serrate-dentate with mammiform, narrowly mammiform or cusped and obtuse-apiculate teeth which occasionally descend onto the petiole and attenuate or sometimes shortly narrowed at base, all with the upper surface with few or no simple eglandular hairs which when present are

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confined to or near the margins occasionally with scattered, short to medium, slender, white, pale-based ones especially on the innermost leaves, the margins usually with many medium, slender, wavy simple eglandular hairs and scattered or very few, short to medium, pale-based ones on the lower surface, few to numerous, very short, yellowish glandular hairs on both surfaces and the margins and numerous to dense stellate hairs uniformly distributed on the lower surface with some on the margin and sometimes a few on the upper surface, the petioles medium to long, slender or occasionally winged, reddish-violet tinted, with numerous, medium, slender, wavy simple eglandular hairs, those near the base long, and numerous stellate hairs; cauline (0–)1–2(–3), usually all small, linear or bract-like and entire, upper filamentous, the lowest with lamina very rarely narrowly oblanceolate, petiolate and slightly dentate. Capitula usually solitary, occasionally 2, 30–50 mm in diameter, the involucre greenish-grey or dark grey in appearance, campanulate, rounded to broadly rounded at base and somewhat constricted in the middle, becoming very broad at base when in fruit; peduncles when present not long and clothed like the upper stem. Involucral bracts incumbent in bud, dark green, the inner much paler; inner 11–16 × 0.8– 1.8 mm, very appressed, narrowly linear-lanceolate, mainly gradually and finely acute at apex; outer short, narrow and more or less appressed; all with numerous to dense, short to long (up to 3 mm), slender, wavy, white or greyish, black-based simple eglandular hairs, numerous, very short (–short), fine, mostly yellowish but sometimes dark glandular hairs which are completely or very largely obscured by the simple eglandular hairs, and few, scattered stellate hairs, the inner with a tuft of very short hairs at the apex. Ligules deep, bright yellow, the outer 3.2–4.0 mm wide, sometimes rather few and often markedly discrete at the apices, very upturned and often ascending at the shallowly to rather deeply lobed apex, with few to numerous, very short to short, pale or slightly discoloured, sometimes rather appressed simple eglandular hairs on the lower surface and a few very short (–0.4 mm) ones or occasionally none projecting beyond the apex. Styles pure yellow. Receptacle pits with margins shortly dentate. Achenes 4.0–4.5(–5.0) mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. In the wild H. grovesii varies little except in the degree of dentation of the basal leaves. In cultivation the leaves become larger and more dentate. Native. A local and rare species occurring in very small numbers in crevices of rock slabs, on rock-ledges and stable scree at the base of cliffs on granite from 610 to 1030 m. Confined to the Cairngorms at Loch Etchachan, Coire Etchachan, Derry Cairngorm and Beinn a’ Bhuird in Aberdeenshire, Coire Raibairt above Loch Avon in Banffshire and the west side of Glen Einich, east side of Cairn Gorm and Creag an Leth-choin above the Lairig Ghru in Inverness-shire. Endemic. Named after Henry Groves (1835–91). 85. H. larigense (Pugsley) P. D. Sell & C. West Lairig Hawkweed H. globosiflorum var. larigense Pugsley Phyllopodous perennial herb with a branched stock. Stem 10–25 cm, pale green, often reddish-violet tinted at base,

slender to somewhat robust, sometimes flexuous, the lower part moderately clothed with medium to long, slender, wavy simple eglandular hairs, very few, very short glandular hairs and numerous stellate hairs, the upper part with numerous to dense, medium and long (up to 5 mm), slender, wavy, often somewhat greyish, black-based simple eglandular hairs, several to numerous, very short to short, pale and dark glandular hairs and numerous to dense stellate hairs. Leaves medium to rather deep, occasionally pale green and glossy above, paler and less glossy beneath, frequently reddish-violet tinted on the midrib of both surfaces; basal numerous (very numerous in cultivation), the lamina of outer 7–45 × 5–23 mm, broadly elliptical or obovate, broadly rounded-obtuse to obtuse-mucronulate at apex, entire, denticulate or sometimes with a few mammiform teeth and cuneate or shortly narrowed at base, the lamina of inner 15–80 × 3–20 mm, narrowly to broadly elliptical, oblong-elliptical, oblanceolate or lanceolate, rather rigid, semicoriaceous, often channelled, sometimes strongly so (cymbiform), frequently undulate or twisted, mainly acute or subacute and apiculate at the often downwardly twisted apex, subentire to very irregularly dentate or serrate-dentate, mainly on the lower half of the margin and often descending onto the petiole in larger plants, with often ascending, narrowly mammiform or long, acute, cusped teeth which have long, rounded-obtuse-apiculate apices and mainly attenuate or sometimes narrowly cuneate at base, all usually without simple eglandular hairs on the upper surface, or at least about the midrib, the innermost sometimes with scattered ones, but the margins with numerous to dense, medium or frequently long, rather coarse, subrigid, wavy or straight, white simple eglandular hairs with pale or slightly discoloured bases, which are usually directed inwards, especially near the apex of the leaf, the lower surface with uniformly scattered, shorter, softer ones, or sometimes denser on the midrib, and with many very short, yellowish glandular hairs on both surfaces and the margins, the petioles slender or winged, shortish to long, with long (up to 4 mm), dense simple eglandular hairs on the margin and especially about the base, and with a reddish-violet tinted midrib; cauline 0–2(–5), frequently all small and narrow, the lowest typically up to 20 × 2(–6 cult.) mm, linear or bract-like, sessile, rarely with a few, acute teeth and acute at apex, the remainder bract-like, acute at apex and more or less appressed, the uppermost filamentous. Capitula solitary, occasionally 2, sometimes with an additional stem from the base bearing a capitulum, often with a rudimentary bud in the axil of a cauline leaf, 30–48 mm in diameter, the involucre dark greenish-grey or greyish-black in appearance, broadly campanulate to campanulate and broadly rounded at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackishgreen, the inner with paler margins; inner 11–16 × 1.0– 1.5(–1.8) mm, appressed, linear-lanceolate, acute at apex or many finely attenuate-acute or narrowly acuminate, the innermost narrow; outer shorter, loosely appressed, erect; all with numerous to dense, medium to long, wavy, often more or less greyish, black-based simple eglandular hairs, numerous very short to short, mainly dark glandular hairs, largely or only partially obscured by the simple eglandular hairs, without stellate hairs but the inner with a tuft of very

39. Hieracium short hairs at the apex. Ligules bright medium yellow, the outer 3.0–3.9 mm wide, numerous but more or less discrete, almost straight to very slightly recurved (flower opening almost flat) and deeply lobed at the apex with narrow lobes, with very few to several, extremely short and very short, pale or sometimes slightly discoloured simple eglandular hairs on the lower surface and projecting beyond the apex. Styles more or less uniformly olive-grey, dirty greenish-yellow or occasionally yellow (but with discoloured scales present). Receptacle pits with margins shortly dentate. Achenes 3.0– 4.0 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 27 or 36. Native. Local and rather rare on cliff ledges and in crevices of sloping rock slabs on coarse-grained granite and sometimes descending on to the turf and scree below, between 730 and 870 m. Cairngorm Mountains, on Creag an Leth-choin in the Lairig Ghru Pass in Inverness-shire and in An Garbh Choire, Coire an t-Saighdeir, Glen Geusachan and on Devil’s Point, all on Cairn Toul in Aberdeenshire. Both chromosome counts come from plants collected in the Lairig Ghru, but one is probably incorrect because of mixed material. Endemic. 86. H. deargicola P. D. Sell & D. J. Tennant Beinn Dearg Hawkweed Phyllopodous perennial herb with a branched stock. Stem 8–23(–28) cm, pale green, medium to rather slender, with scattered to numerous, short, medium and long, wavy or straight, pale, mainly dark-based simple eglandular hairs throughout, often becoming greyish with long black bases in the upper part, scattered, very short glandular hairs, becoming more numerous to rather dense, black, very short, short and medium (up to 1.0 mm) in upper part, and scattered or numerous stellate hairs becoming more numerous in upper part and partially obscuring the short glandular hairs throughout. Leaves rather pale to medium, sometimes rather drab green and slightly glossy on upper surface, slightly paler beneath, usually without reddish-violet tints, noticeably graniculate; basal more or less numerous, sometimes in a fairly neat rosette, thin or only very slightly coriaceous, slightly undulate, shallowly channelled or flat, the outer with lamina 5–35 × 4–25 mm, ovate, obovate or subrotund, more or less broadly rounded-obtuse with a few slightly retuse at apex, subentire, remotely denticulate or with a few unequal, broad, rather blunt teeth on the margin and mainly cuneate or cuneate-truncate at the base into a medium or rather long petiole, the inner with lamina 8–50 × 5–25 mm, mainly elliptical, with some narrowly elliptical, ovate or obovate, the innermost with lamina often oblanceolate, mainly more or less broadly obtuse and some subacute or acute and slightly twisted at the apex, subentire, denticulate or serrate-dentate on the margin, with shallow to narrowly mammiform, or broadly acute, more or less cusped, sometimes reddish-tipped teeth and mainly cuneate at base, the innermost almost attenuate at base into a medium to long petiole, the outer with lamina often subglabrous, with very numerous medium to rather long simple eglandular hairs on the margin and scattered to numerous, usually medium ones on the lower surface which are often denser on the midrib, and with several or few, very short, yellowish glandular hairs mainly on the margin, without stellate hairs, the

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inner with lamina with numerous (or scattered), rather rigid, wavy, medium and long, white, pale-based, simple eglandular hairs, uniformly distributed or sometimes confined towards the margin on the upper surface, the majority of the petioles with numerous, long, wavy, simple eglandular hairs and usually without reddish-violet coloration; cauline 1–4, the lowest rigidly suberect or somewhat patent, the lamina 10–40 × 2–12 mm, linear-elliptical, linear-lanceolate or oblanceolate, usually acute, very often with a long, fine acumen at the apex, subentire or sometimes slightly dentate and undulate on the margins and attenuate to a sessile base or with a shortish, often winged petiole, the next lowest linearoblanceolate, finely acuminate at apex, entire, or as lowest, the remainder suberect, rather long, very narrowly linear, finely acute bracts, all with few to many, medium to long simple eglandular hairs on the upper surface, very numerous, wavy, more or less black-based ones and scattered to many stellate hairs, or none, on the margins and lower surface. Capitula often solitary to 3 (4 in cult.), 30–45 mm in diameter, the involucre very dark olive-grey or olive-black in appearance, rather broadly campanulate, rounded or very slightly narrowed into the peduncle and usually constricted in the middle; peduncles when present somewhat dusky pale green, clothed as the upper stem. Involucral bracts incumbent in bud, blackish green, the inner unequal, the longest 11–15 × 1.0–1.6 mm, appressed or loosely appressed, narrowly lanceolate-subulate, sharply acute or acuminate or sharply acute with a few subacute, the innermost finely acuminate at the apices, the outer much shorter, lax, more or less erect, acute or subacute; with numerous to dense, very wavy, medium and long (to 4(–5) mm), whitish to silvery grey, black-based simple eglandular hairs which almost completely obscure the very short to short (–0.6 mm) blackish glandular hairs or sometimes with a few visible, without stellate hairs but with a matted tuft of short branched and simple hairs at the apex. Ligules medium (to deepish) yellow, rather dense centrally, the outer up to 3.5(–4.0) mm wide, fairly discrete, straight to shallowly upturned and with rather long, narrow lobes at the apex, with very few to several, short to medium, often slightly discoloured (pale golden brown) simple eglandular hairs on the lower surface, or none, and very few to many, very short to short ones on the lobes at the apex, hardly projecting beyond the margins of the lobes. Styles pure yellow. Receptacle pits with margins shortly dentate. Achenes 3.3–4.4 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Somewhat intermediate between H. kennethii and H. subglobosum, differing from the former in its less dentate leaves, more acute involucral bracts and yellow styles and from the latter by its more hairy leaves, fewer obvious glandular hairs on the involucral bracts and yellow styles. Native. Cliff ledges and slopes below the cliffs. Above 800 m on Beinn Dearg in Ross-shire. Endemic. 87. H. subglobosum P. D. Sell & C. West Subglobose Hawkweed Phyllopodous perennial herb with a branched stock. Stems 6–20(–30) cm, slender or medium, frequently flexuous, reddish-violet at base, with few to many, slender, wavy, short, medium and long, white to greyish simple eglandular hairs with thickened, blackish bases throughout, scattered

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pale and sometimes retrorse ones in the lower part, several to many, medium to long (–4 mm), patent, often greyish ones with long, black bases in the upper part, scattered, very short, yellowish glandular hairs in the lower part which become numerous to dense, blackish, short and medium (–1.0 mm) in the upper part, and with scattered stellate hairs below which become numerous to dense in the upper part. Leaves medium, often bright green, glossy on upper surface, paler on lower surface, frequently tinted with dull reddish-violet especially on upper side of teeth and margins and on the midrib beneath; basal rather few to numerous, often small and in a rather neat rosette, the lamina of outer 3–50 × 3–25 mm, subrotund, ovate or obovate, entire or denticulate, broadly rounded-obtuse at apex and cuneate at base, the lamina of inner 15–90 × 2–30 mm, rather thin to coriaceous, occasionally channelled, narrowly to broadly elliptical or narrowly obovate, acute to obtuse and often apiculate at the apex, subentire, denticulate, sometimes undulate, or irregularly serrate-dentate with incise, occasionally large and very irregular, patent, narrowly mammiform or aquiline-mammiform teeth, and cuneate to shortly attenuate at base, the inner leaves usually almost glabrous on the upper surface, but with few to numerous, medium to long, subrigid, white simple eglandular hairs, often with discoloured or black, bulbous bases, towards and at the margins, which become long and dense on the innermost leaves, the lower surface of the majority of leaves with scattered, softer ones which are dense on the midrib, with some very short, yellowish glandular hairs on the margins and few on both surfaces, and with few stellate hairs on either surface, the petioles short to long, slender, often tinted reddishviolet towards the base, sometimes with a few teeth and with scattered to dense, medium and long, subrigid, sometimes wavy simple eglandular hairs on the margin and especially numerous towards the base; cauline (0–)1–3(–5), the lowest semipatent to erect, the lamina 6–25 × 1–7 mm, linearelliptical, sharply attenuate-acute or very rarely obtuse at apex, usually entire, occasionally remotely denticulate or with a few, sharp teeth, attenuate at base, glabrous on upper surface, the margin and lower surface with medium, subrigid, often black-based simple eglandular hairs becoming long and often dense towards the base and with scattered stellate hairs, the remainder small, filamentous or bractlike, appressed and acute at apex. Capitula solitary, rarely 2, often small, 25–40 mm in diameter, the involucre campanulate to narrowly campanulate, shortly narrowed into the peduncle but rounded at the base and constricted in the middle, blackish or dark greenish-grey in appearance; peduncles when present clothed as the upper stem. Involucral bracts incumbent in bud, blackish green, some inner ones with inconspicuous greenish margins, inner 9–16 × 0.9–1.7 mm, unequal, often curved, rarely very numerous, rather closely appressed, linear-lanceolate, the innermost very narrow, attenuate to a very sharply acute or acuminate, rarely subacute or obtuse apex; all with moderate to dense, medium to rather long (–4 mm), patent, wavy, white or more frequently greyish simple eglandular hairs with very long, thickened, black bases, numerous, very short pale and short, blackish glandular hairs which are only partially obscured by the simple eglandular hairs, without stellate

hairs but with some, short-branched hairs at the apex; outer shortish, erect and acute at apex. Ligules bright, pale yellow, occasionally medium yellow, the outer up to 4 mm wide often markedly discrete, straight to shallowly upturned and, deeply and sometimes irregularly lobed at the apex, without hairs or sometimes with very few, short, pale simple eglandular hairs on the lower surface and with several extremely short and occasionally a few very short more or less greyish ones projecting beyond the apex. Styles more or less uniformly very dark grey or medium greenish-grey. Receptacle pits with margins shortly dentate. Achenes 3.5–4.2 mm, purplish-black. Flowers 7–8. 2n = 27. Native. Mainly on gentle rocky slopes or rocky knolls, occasionally on rock-ledges on Moine and other schists, (?)quartzite, microgranite and Torridonian Sandstone, usually between 550 and 850 m, but descending to 350 m just south of Cape Wrath. Near Loch Cluanie and in Glen Affric and Glen Cannich near the western border of east Invernessshire, on several hills in the northern part of west Invernessshire, locally on the Island of Skye in the Inner Hebrides, on numerous hills in west Ross-shire, and several in east Ross-shire including Seana Bhraigh and Beinn Dearg and in Glen Carron, on hills south and east of Loch More in east Sutherland and many hills throughout west Sutherland. Endemic. 88. H. globosiflorum Pugsley Round-headed Hawkweed H. globosum Backh. fil., non Desf.; H. nigrescens var. globosum Hook. fil.; H. nigrescens subsp. globosum Zahn Phyllopodous perennial herb with a branched stock. Stem 10–20(–30) cm, slender to medium, often flexuous, sometimes striate and reddish-violet towards the base, with few or scattered, medium to longish simple eglandular hairs, very short glandular hairs and scattered to numerous stellate hairs in the lower part, the upper part with many to numerous, short and medium, slender, greyish and sometimes several to many, long, very wavy, whitish, black-based simple eglandular hairs, many to very numerous, very short and short, dark glandular hairs and numerous, but often not very dense stellate hairs. Leaves medium to deepish drab green and often matt or barely glossy on the upper surface, occasionally paler, brighter green glossy, and paler beneath, often with reddish-violet tints and markings, especially on the midrib; basal numerous to very numerous, often with a rather neat rosette, the lamina of outer 8–40 × 5–20 mm, subrotund, ovate or broadly elliptical, sometimes undulate, rounded-obtuse, sometimes mucronate, occasionally slightly retuse at the often twisted apex, subentire, denticulate or occasionally with larger, acute teeth and more or less cuneate at the base, the lamina of inner 20–50(–80) × 6–20 mm, variable, mainly narrowly to broadly elliptical, sometimes ovate, obovate, ovate-lanceolate, all very rigid and coriaceous, sometimes undulate, often broadly twisted at the rounded-obtuse, subacute or acute, sometimes apiculate apex, denticulate, shallowly sinuate-dentate or irregularly to very irregularly dentate or serrate-dentate with mammiform or occasionally aquiline-mammiform teeth, or dentate with sharply acute teeth interspersed with denticulations and the teeth when present often becoming large towards the base

39. Hieracium of the margins, narrowly cuneate or attenuate at the base, the upper surface often more or less glabrous, with numerous, uniformly short or medium, slender, subrigid, white, mainly pale-based simple eglandular hairs on, and sometimes near, the margins and occasionally with scattered long ones on the upper surface especially on the innermost leaves, with many short to medium softer, wavy or curved ones, more or less uniformly distributed on the lower surface, the outer leaves often without simple eglandular hairs except on the margins which usually have shortish ones, and with several to numerous, very short, yellowish glandular hairs on both surfaces and the margins, the petioles mainly short (rarely up to 50 mm), pale violet-tinted at base and with numerous, short or medium simple eglandular hairs on the margin which become long and sometimes dense at the base; cauline (0–)1–3(–5), the lowest erect, the lamina 10–40 × 1–6 mm, occasionally undulate, linear or bract-like, occasionally linear-oblanceolate, very acute or finely acuminate at apex, entire or rarely with very few acute teeth, obscurely petiolate or more usually sessile, without simple eglandular hairs on the upper surface, but with numerous, shortish, sometimes dark-based ones on the margin and lower surface, becoming dense and longer at the base, with some very short glandular hairs on the margin and occasionally with a few, stellate hairs on the midrib of either surface, the remainder very small, bractlike, appressed and acute, the upper filamentous. Capitula usually solitary, occasionally 2–3, sometimes with a rudimentary bud in the axil of a cauline leaf, 30–45 mm in diameter, the involucre dark olive-grey or very occasionally blackish in appearance, almost spherical in bud, swollen in fruit but more or less cuneate at base and tapering towards the apex, whilst in flower distinctive in shape, campanulate and rounded at base, but shortly narrowed into the peduncle and markedly constricted in the middle; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, often numerous, blackish-green, some of the inner with inconspicuous greenish margins; inner 10–16 × 1.0–1.5 mm, unequal, very appressed, sometimes curved, linear-lanceolate (the innermost narrowly so) and gradually and finely acute or acuminate, or infrequently abruptly acute at apex; outer short, semilax, erect and acute at apex; all with numerous to dense, medium to longish (up to 2 mm), very wavy, slender, whitish or greyish, black-based simple eglandular hairs, numerous, very dark or yellow-headed, very short to short glandular hairs, only partially obscured by the simple eglandular hairs, without stellate hairs but the inner with a distinct tuft of very short hairs at the apex. Ligules bright medium yellow, the outer usually not very wide (up to 3.7 mm), densely arranged, alternating long and short, usually barely discrete, straight to shallowly upturned (capitulum opening almost flat), and deeply and narrowly lobed at apex, often without hairs, or with some extremely short, appressed simple eglandular hairs, or sometimes with a few, seldom with many, very short or short, pale or very slightly discoloured ones on the lower surface, and with few to numerous, extremely short to short ones projecting beyond the apex. Styles pure yellow, rarely with a greenish tinge; stigmas always yellow (sometimes darkening markedly when dried). Receptacle

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pits with margins sharply dentate. Achenes 3.2–4.5 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 27. Most frequently grows in open situations, but when occurring in sheltered places it is taller and more hairy. Native. Crevices in rock slabs, cliff ledges, rocky slopes and stream banks on coarse-grained granite and occasionally felspar, between 600 and 1,050 m. Confined to the Cairngorms where it is not infrequent on all the hills bordering the Lairig Ghru, towards the head of Strath Nethy, also Loch Avon and in Coire Etchachan, but more local and rather scarce in Glen Derry, Glen Einich, on Cairn Gorm, Beinn a’ Bhuird and Craig an Dail Bheag (Little Craigandal). It has been recorded for Norway and Lapland, but these records belong to other species, and it is almost certainly endemic. 89. H. graniticola W. R. Linton Granite Hawkweed H. gracilentum var. graniticola (W. R. Linton) W. R. Linton; H. nigrescens subsp. graniticola (W. R. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 13–25(–30) cm, pale green, reddish-violet tinted in lower part, slender to somewhat robust, sometimes flexuous, often striate, the lower part with a sparse to moderate clothing of medium and long, wavy, dark-based simple eglandular hairs, with few, very short, pale or dark glandular hairs and scattered stellate hairs, the upper part with very numerous, short to medium, grey and some whitish, long and wavy, black-based simple eglandular hairs, numerous to dense, rarely few, short dark glandular hairs and numerous to dense stellate hairs. Leaves drab, rather pale to deepish green, sometimes tinted with reddish-violet, matt or barely glossy, the lower surface paler, minutely granulate in appearance and with a very pronounced midrib; basal numerous, usually in a neat rosette, the lamina of outer 6–40 × 4–25 mm, subrotund, obovate or broadly elliptical, broadly obtuse or sometimes slightly retuse at apex, entire or shallowly sinuate-dentate with obtuse teeth and cuneate at base, the lamina of inner 10–50(–70) × 5–22 mm, coriaceous and frequently undulate or twisted, sometimes slightly channelled, mainly elliptical or oblanceolate, sometimes narrowly elliptical, oblong-elliptical or ovate, mainly obtuse-mucronate, sometimes subacute or occasionally acute at the apex, denticulate or more frequently very irregularly sinuate-dentate or serrulate-dentate with shallowly to narrowly mammiform and often long, cusped teeth with obtuse-apiculate apices, mainly on the lower half of the margins, becoming longer at the base and often descending onto the petiole and shortly or gradually narrowed at base, all with the upper surface with numerous, medium to long, wavy, coarse, subrigid, white simple eglandular hairs, with pale or only slightly discoloured bases, which are usually confined to near and on the margins, the innermost sometimes with uniformly scattered ones, the outer often with few, and with more or less uniformly scattered, softer, more slender ones, or none on the lower surface, with numerous, very short, yellowish glandular hairs on both surfaces and on the margin, and occasionally with a few, stellate hairs on the midrib above, the petioles usually shortish to medium, sometimes reddish-violet tinted, with

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numerous, long, somewhat coarse, wavy simple eglandular hairs on the margins which become dense at the base and dense throughout on the petioles of the innermost leaves and with short to medium, subrigid and slender ones on those of the outer leaves, and all petioles with many, very short, yellowish glandular hairs; cauline (1–)2–4(–6), the lamina of lowest 5–35 × 1–9 mm, linear, linear-lanceolate or bract-like, acute, acuminate or obtuse-mucronate at apex, usually entire, occasionally denticulate or with few, small teeth and usually sessile, the remainder very small, bractlike, appressed and acute at apex, the uppermost usually filamentous. Capitula 1(–2), or occasionally with a rudimentary bud in the axil of a cauline leaf, 30–40(–45) mm in diameter, the involucre medium greenish-grey to very dark grey in appearance, campanulate or more frequently broadly campanulate and often rounded at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, the innermost with pale green margins; inner unequal, 9–15 × 0.9–1.7 mm, more or less closely appressed, linear-lanceolate, the innermost narrower and linear, abruptly acute, acuminate or subacute at apex; outer shorter and erect; all with very dense, medium to long (up to 3 mm), slender, very wavy, somewhat appressed and interwoven, whitish to somewhat pale greyish, black-based simple eglandular hairs, numerous, very short to short, darker glandular hairs which are almost fully obscured by the simple eglandular hairs and without stellate hairs, the inner with a very pronounced tuft of very short hairs at the apex. Ligules bright medium, very occasionally rather deep yellow, the outer 3.0–3.6(–4.0) mm wide, often numerous, densely arranged and often barely discrete and more or less straight, occasionally marginally recurved (capitula opening flat), often deeply and narrowly lobed or sometimes involute or with the outer lobes broad and rounded and the inner all very small at apex, with very few to numerous, very short to short, pale simple eglandular hairs on the lower surface and many very short and a few short ones projecting beyond the apex. Styles discoloured yellow or pure yellow. Receptacle pits with margins shortly dentate. Achenes 3.4–4.5 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 27. This species shows little variation in nature or in cultivation. Native. Rare and very local on exposed rocky slopes at high altitude, amongst tufts of Calluna over stabilised fine granite scree or at the margin of unstable scree by crumbling coarse-grained granite rocks between 800 and 1,050 m. Confined to the Cairngorms in Coire Etchachan, above Loch Etchachan and on Derry Cairngorm in Aberdeenshire, the south side of Cairn Gorm above Loch Avon in Banffshire and on the east side of Cairn Gorm at the head of Strath Nethy in Inverness-shire. Endemic. 90. H. pseudocurvatum (Zahn) Pugsley Cairngorm Hawkweed H. nigrescens subsp. pseudocurvatum Zahn Phyllopodous perennial herb with a branched stock. Stem 7–15(–20) cm, pale green, often violet-tinted at base, rather slender, sometimes flexuous, the lower part sparsely to moderately clothed with medium to long (up to 4 mm), darkbased, wavy simple eglandular hairs, few extremely short

glandular hairs and scattered stellate hairs, the upper part with several to many short to medium, slender, greyish, black-based simple eglandular hairs, with very few longer ones (up to 2 mm), with numerous very short to short, very dark glandular hairs and many stellate hairs. Leaves medium to deep, rather bright or rather drab green, rarely paler, sometimes with reddish-violet tints, rather glossy to somewhat matt on upper surface; basal numerous, in a compact, somewhat prostrate rosette, the lamina 4–40 × 3–16 mm, the outer obovate to subrotund, undulate and twisted, broadly obtuse at the apex, entire, denticulate or irregularly dentate with shallowly mammiform teeth and attenuate or shortly narrowed into the petiole, the lamina of inner more or less broadly elliptical, broadly obtuse and apiculate or subacute at the twisted apex, and very undulate on the margin, irregularly serrate-dentate with several shallow to narrow, sometimes incise, mammiform teeth, or subentire and attenuate at base, all with few to many, slender, mostly short to medium with some long, blackish or pale-based simple eglandular hairs, or none except towards and at the margins, the lower surface with scattered more or less uniformly distributed, short to medium, slender, very wavy simple eglandular hairs or sometimes with none except on the margins which also have a few, very short, yellowish glandular hairs, the petioles short, occasionally winged, strongly tinted reddish-violet and with several to many, medium simple eglandular hairs; cauline 3–5(–6), the lowest usually near the base, rigidly semipatent, its lamina 10–30 × 4–6 mm, undulate, oblanceolate, subacute and sometimes twisted at apex, serrate-denticulate or subentire and attenuate at base but with a distinct petiole, the remainder all small, linear or filamentous, more or less appressed, entire and acute at apex. Capitulum solitary, 30–40(–50) mm in diameter, the involucre very dark greenish-grey or blackish in appearance, campanulate and rounded at base. Involucral bracts incumbent in bud, blackish-green, the inner much paler towards the margins; inner 9–13 × 1.0–1.6 mm, rather closely appressed, linear-lanceolate and gradually tapering to an acute, subacute or acuminate apex; outer appressed, much shorter and narrower than inner, more or less acute at apex; all only moderately or rather sparsely clothed with slender, mainly medium and short (up to 2 mm), seldom longer, somewhat greyish, blackbased simple eglandular hairs which are sometimes sparse or absent towards the margins and several to many, obvious, very short to short, blackish glandular hairs, without stellate hairs but the inner with a tuft of short hairs at the apices. Ligules bright, pale yellow, the inner very numerous, the outer rather narrow (up to 3.7 mm), numerous but slightly discrete, slightly upturned and often deeply lobed at the apex, with only a few very short, sometimes slightly greyish simple eglandular hairs on the lower surface and very few, extremely short, with occasionally a few very short ones (up to 0.2 mm), projecting beyond the apex, or glabrous. Styles uniformly very dark grey (blackish). Receptacle pits with margins shortly dentate. Achenes 3.5–4.2 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 27. Native. Cliff ledges on coarse-grained granite between 680 and 750 m. Local and very scarce; known only from Craig an Dail Bheag (Little Craigandal) and a single coire in Glen Derry in Aberdeenshire. Endemic. Most Scottish

39. Hieracium plants previously called H. curvatum or H. pseudocurvatum are referable to H. completum. 91. H. einichense P. D. Sell & D. J. Tennant Glen Einich Hawkweed Phyllopodous perennial herb with a branched stock. Stem 8–25 cm, medium or occasionally rather slender, with scattered to many, short and medium to longish, very wavy simple eglandular hairs throughout, becoming mainly dark grey and black-based in upper part, scattered very short and short glandular hairs which become numerous to dense, black and mainly short to medium in the upper part, and scattered to many stellate hairs throughout which become more or less dense in the upper part. Leaves deep green, almost matt and largely or completely suffused with dark, dull reddish-violet on the upper surface, paler, faintly glaucousgreen and with prominent, often purplish midrib beneath; basal few to numerous, in a fairly neat, slightly spreading rosette, rather rigid, somewhat coriaceous, the outer with lamina 7–40 × 4–25 mm, obovate, broadly elliptical or subrotund, broadly rounded-obtuse at apex, subentire, denticulate or with some small teeth on the margin and more or less cuneate or rounded at the base into an often shortish, slightly winged petiole, the inner with lamina 20–55 × 5–25 mm, mainly elliptical to oblong-elliptical, some obovate and occasionally a few ovate, the innermost often more or less oblanceolate, obtuse and often mucronate, many broadly rounded-obtuse, the innermost subacute and sometimes apiculate at apex, denticulate or with several rather regular, smallish, sharply acute, occasionally cusped or narrowly mammiform teeth towards the base, and cuneate to narrowly cuneate and the innermost more or less attenuate at base into a broadly winged to somewhat slender, short to rather long petiole, the inner with lamina with more or less numerous, subrigid, pale or dark violet based, whitish, mainly medium simple eglandular hairs on the upper surface, either moderately and uniformly distributed or more often confined to the area near the margin, sometimes becoming very numerous and longish on the innermost, the outer with lamina more or less subglabrous, or with very numerous, mainly medium, but some longer and some shorter, more or less rigid simple eglandular hairs on the margins, with scattered to very numerous ones on the lower surface, becoming rather denser on the midrib, the outer often subglabrous beneath and with many very short glandular hairs on the margins and some on both surfaces, and sometimes with several stellate hairs on the lower surface, the petioles sparsely to moderately clothed on the margin with wavy medium and long simple eglandular hairs; cauline (0–)3– 5(–6), the lowest suberect or rigidly semipatent, the lamina 10–50 × 1–9 mm, usually linear-oblanceolate, sometimes narrowly linear and bract-like, more or less acute at apex, entire, or more often denticulate or with several smallish, sharply acute teeth on the sometimes undulate margin, sessile and semiamplexicaul or sometimes petiolate and often with many stellate hairs on the lower surface, the remainder more or less linear, entire and the uppermost bractlike. Capitula usually solitary (–2), 38–48 mm in diameter, occasionally a second stem arising from the basal rosette bearing a capitulum (often in cultivation), the involucre blackish in appearance, campanulate, rounded at base

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and constricted in the middle; peduncles when present clothed as the upper stem. Involucral bracts incumbent in bud, blackish-green; the inner unequal, the longest (11.5–) 13–15 × 1.1–1.6 mm, rather appressed, linear-lanceolate, mainly abruptly acute, with some subacute-acuminate and narrowly obtuse, the innermost sharply acute at the apices, the outer shorter and more or less appressed; with many to numerous, mainly medium and some rather long (to 2.5 mm), wavy, usually greyish or very dark, black-based simple eglandular hairs, and rather dense, very short, short and medium (to 1.5 mm) black glandular hairs which are not significantly obscured by the simple eglandular hairs, the inner with a distinct tuft of short branched and simple hairs at the apex, the outer with some stellate hairs at the base. Ligules deepish (to more or less medium) yellow, rather dense centrally, the outer up to 3.6(–4.0) mm wide, discrete, shallowly upturned to very upturned at apex, with longish, narrow lobes, with very few to several, very short (to short), sometimes rather appressed simple eglandular hairs on the lower surface and usually some extremely shortly (0.1–0.2 mm) projecting beyond the apex, or more or less none. Styles medium to deeply discoloured, uniformly dark olivegrey. Achenes 3.2–3.8 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Only known and extremely scarce in one or two places on ledges of granite cliffs, at about 800 m, on the west side of Glen Einich in Inverness-shire. Endemic. 92. H. tenuifrons P. D. Sell & C. West Tenuous-leaved Hawkweed H. gracilentum auct. Phyllopodous perennial herb with a branched stock. Stem 7–24 cm, very slender to medium, seldom rather robust, sometimes flexuous, usually pale green throughout or sometimes reddish-violet at base, the lower part with rather thinly scattered to many, mainly medium to long, often irregularly twisted, whitish simple eglandular hairs, several to many, very short, pale or discoloured glandular hairs and scattered stellate hairs, the upper part with many to numerous, or occasionally scattered, medium and long, very wavy and frequently irregularly twisted, usually whitish, black-based simple eglandular hairs, numerous, mainly very short and short (up to 0.8 mm) unequal, pale and dark glandular hairs and scattered to dense stellate hairs. Leaves pale to rather deep, bright green and usually glossy on the upper surface, paler and sometimes slightly glossy beneath, usually without reddish-violet colour, often slightly graniculate in appearance; basal leaves numerous, the lamina of the outer 5–40 × 3–28 mm, subrotund, obovate or elliptical, broadly rounded-obtuse to subacute at apex, entire, denticulate or with several larger teeth on the margins, cuneate or shortly narrowed at base, the lamina of the inner 15–70 × 4–28 mm, often rather rigid, slightly coriaceous, undulate, channelled or flat and with prominent, raised veins beneath, variable in shape, obovate, obovate-spathulate or oblanceolate to broadly or narrowly elliptical, subacute to broadly rounded-obtuse, mucronate to somewhat emarginate and often twisted at apex, the innermost sometimes acute, the margins subentire, denticulate, or serrate-dentate with mainly narrowly mammiform, often unequal, sometimes cusped-mammiform teeth, narrowly cuneate or shortly

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narrowed to attenuate at base, often with many to very numerous, long, wavy, pale-based simple eglandular hairs on the upper surface, usually fewer or none about the midrib, sometimes uniformly distributed, or occasionally absent, with many medium to longish ones on the margins and few to numerous, pale-based, or sometimes black-based ones on the lower surface, with some very short, yellowish glandular hairs which are often rather numerous on the margins and occasionally a few stellate hairs on the midrib beneath; petioles rather short to long, slender or winged, without reddish-violet colour or occasionally tinted at the base, with several to very numerous, long, wavy simple eglandular hairs; cauline few, (0–)1–2(–3), the lowest semierect, sometimes undulate, with the lamina 5–45(–70) × 1–3 mm, usually linear-elliptical or linear-oblanceolate, usually broadly acute to obtuse at apex, entire or occasionally with a few small, acute teeth on the margin, narrowed at base into an often longish, slender petiole, usually with many simple eglandular hairs on the upper surface, at least near the margins, and usually with very numerous, long, often black-based ones on the lower surface and on the margins, the remainder much smaller and more or less linear, the uppermost rarely filamentous. Capitula solitary (rarely 2), but occasionally with an additional stem arising from the base bearing a capitulum, 25–40(–50) mm in diameter, the involucre medium olive-grey to silvery olive-grey in appearance, narrowly campanulate to campanulate, narrowly subtruncate or rather narrowly rounded at base. Involucral bracts incumbent in bud, blackish-green; inner (11.5–) 14–17 × 0.8–1.5 mm, appressed to loosely appressed, linear or linear-lanceolate, sometimes lanceolate-subulate, abruptly acute to subacute, often with some obtuse and acuminate at apex, the innermost often attenuate-acute; outer few, shorter, erect and slightly lax, obtuse or subacute at apex; with very numerous to dense, medium and long, very wavy, sometimes interwoven, white to pale silvery grey, black-based simple eglandular hairs, numerous very short and short, pale and dark glandular hairs, which are fully obscured by the simple eglandular hairs, or sometimes with a few obvious ones near the base, without stellate hairs but the inner with a very distinct tuft of short, pale hairs at the apex. Ligules medium yellow, often with a very faint greenish tinge, sometimes pale yellow, the outer 3.0–4.0(–4.3) mm wide, often rather few and very discrete, slightly to markedly upturned, often ascending at apex, with few to many, short and medium, pale or greyish-yellow simple eglandular hairs on the lower surface and several to many, very shortly to shortly (0.2–0.5 mm) projecting beyond the apex. Styles discoloured, but usually not very dark (olive-grey-yellow to pale olive-grey). Receptacle pits with margins shortly dentate. Achenes (3.5–)4.0–4.7 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Plants that were formerly called H. tenuifrons which occur in the central and eastern Scottish Highlands have now been recognised as a distinct species and named H. subtenuifrons P. D. Sell & D. J. Tennant. Native. Cliff ledges, crevices in rocks, stream banks and exposed rock detritus, usually between 650 and 960 m, but descending to 440 m in Glen Coe, on granite, porphyrite and mica-schist. On several mountains in the Ben Nevis Range

above the south-western side of Glen Spean, in Invernessshire, throughout the hills on the south side of Glen Coe and several mountains east of Bridge of Orchy, in Argyllshire, just extending into west and central Perthshire on Beinn Dubhchraig and Beinn Heasgarnich. Endemic. 93. H. subtenuifrons P. D. Sell & D. J. Tennant Highland Hawkweed H. tenuifrons auct. Phyllopodous perennial herb with a branched stock. Stem 10–25(–32) cm, rather slender to somewhat robust, sometimes striate, occasionally flexuous, usually reddish-violettinted at base, with scattered to many, mainly medium to long, very wavy, whitish simple eglandular hairs, usually rather few, very short, pale or discoloured glandular hairs, and scattered stellate hairs in lower part and with many to very numerous, seldom few, medium and long, almost straight to very wavy, occasionally irregularly twisted, whitish or frequently greyish, black-based simple eglandular hairs, several to many very short and some short (up to 0.7 mm), pale and dark glandular hairs, and scattered to very numerous stellate hairs in upper part. Leaves medium to deepish, bright or occasionally drab and very faintly bluishgreen, and all, or at least the innermost, rather glossy on the upper surface, rather paler and slightly glossy to subglaucous on the lower surface, usually tinged, often strongly suffused with reddish-violet, especially on the lower surface; basal numerous, the outer with lamina 8–45 × 3– 28 mm, obovate, elliptical or oblanceolate, usually obtuse to broadly rounded-obtuse and often twisted at apex, subentire, denticulate or with several larger teeth and cuneate to attenuate at base, the inner with lamina 12–70 × 4–30 mm, rigid, sometimes rather coriaceous, often wavy and somewhat channelled, usually narrowly to broadly elliptical, sometimes oblanceolate, mainly rather broadly acute to subacute and often mucronate or apiculate, sometimes obtuse and often markedly twisted at apex, usually dentate to serrate-dentate with shallowly to narrowly mammiform or occasionally cusped-mammiform teeth, occasionally subentire, cuneate to attenuate at base, all with several to many, medium to long, slender to rather thick, subrigid, wavy or apically curved, pale-based or occasionally black-based simple eglandular hairs on the upper surface, usually more numerous near the margin, often with none about the midrib, occasionally uniformly distributed or absent, sometimes very numerous and long on the innermost leaves, with numerous to very numerous, often long, straight or wavy ones on the margin, and with few to many, mainly medium to rather long, pale-based or black-based ones, or none, on the lower surface, with rather few to scattered, very short, yellowish glandular hairs which are most obvious on the margin and usually without stellate hairs, the petioles rather short to longish, slender to somewhat winged, usually strongly reddish-violet-tinted, at least near the base, and with scattered to very numerous, medium to long, sometimes very wavy simple eglandular hairs; cauline numerous, (3–)4–5(–9), the lowest semispreading, often undulate, the lamina 6–30(–40) × 0.5–7.0 mm, linear or linear-oblanceolate, sharply acute or acuminate to subacute at apex, entire, denticulate or with a few, small, acute

39. Hieracium or cusped teeth, sessile or petiolate, with few to several simple eglandular hairs on the upper surface and numerous, often black-based ones on the margin and lower surface, the remainder similar, smaller and semi-erect, the uppermost bract-like. Capitula solitary or few (rarely to 7), 35– 55 mm in diameter, the involucre medium olive-grey to olive-black in appearance, campanulate to broadly campanulate, broadly to narrowly rounded at base. Involucral bracts incumbent in bud, blackish-green; inner 12–16 × 1.0–1.6 mm, appressed or semiappressed, linear-lanceolate or often lanceolate-subulate, more or less acute or acuminate at apex; outer shorter, slightly lax, suberect, subacute or acuminate at apex; all with very numerous to dense, medium to long (up to 4(–5) mm), almost straight to very wavy, often more or less grey, pale-tipped, black-based simple eglandular hairs, numerous very short and some short, pale and dark glandular hairs which are nearly always fully obscured by the simple eglandular hairs, without stellate hairs, but the inner with some short, pale hairs forming an indistinct tuft at the apex. Ligules bright pale yellow, the outer up to 3.5–4.3 mm wide, slightly to moderately discrete in the apical part, straight to slightly curving upwards, without simple eglandular hairs on the lower surface, or with several (to many) mainly short, pale ones often with some grey- or brownishtinted ones, and with none at the apex or a few to several, very shortly (up to 0.2 mm) projecting beyond the apex. Styles medium-discoloured to dark (medium olive-grey to dark grey, stigmas sometimes slightly paler). Receptacle pits with margins shortly dentate. Achenes 3.0–4.1 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 27, 36. Plants that occur in the central Scottish Highlands are sometimes more robust, often have wider basal leaves than those from the eastern Scottish Highlands and are triploid (2n = 27), whereas the eastern plants are tetraploid (2n = 36). Another very local and scarce variant occurs in Glen Feshie in the Cairngorms and most resembles those in the central Highlands but differs in its involucre being constricted in the middle and having deeper yellow, more hairy ligules. It is 2n = 36. Native. Cliff ledges, rocks on stream banks and rocky slopes in the central Scottish Highlands between 580 and 870 m, mainly on mica-schist and in the eastern Highlands between 850 and 1,000 m on granite rocks. Local and uncommon in the central Highlands, on several mountains throughout most of Breadalbane, including Ben Lawers in central Perthshire and in Glen Taitneach and Glen Tilt in eastern Perthshire, with an isolated locality on Creag Meagaidh above Loch Laggan in Inverness-shire; in the eastern Highlands now apparently confined to the Cairngorms on Cairn Toul, Coire Etchachan and Glen Dee in Aberdeenshire, and in Glen Einich, Coire Garbhlach and the Lairig Ghru in Inverness-shire, with a single old record from Forfarshire. Endemic. 94. H. milesii P. D. Sell & C. West Miles’s Hawkweed H. gracilentum Backh. fil. (1856), non (Fr.) Backh. fil. Phyllopodous perennial herb with a branched stock. Stems single or paired, 8–20(–28) cm, slender to robust, often flexuous, somewhat reddish-violet at base, with few to numerous, slender, wavy, short, medium and long, white

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to rather greyish simple eglandular hairs with thickened, blackish bases throughout, moderately clothed with them in the lower part and with numerous, short to medium or occasionally long often greyish ones in the upper part, several to numerous, very short, pale and dark glandular hairs below which become numerous to dense and longer (–1.0(–1.5) mm) in the upper part and many stellate hairs throughout which become rather dense in the upper part. Leaves pale to rather deep, occasionally slightly bluishgreen, matt or glossy on upper surface, paler on lower surface, often tinted with reddish-violet; basal few to numerous, the lamina of outer 6–40 × 5–26 mm, ovate, obovate or broadly elliptical, sometimes asymmetrical, often undulate and twisted, broadly rounded-obtuse at apex, subentire, denticulate or serrate-dentate, with several shallow to narrowly mammiform teeth usually confined to lower two-thirds of margins and cuneate or shortly narrowed at base, the lamina of inner 15–75 × 3–30 mm, rigid and subcoriaceous, narrowly to broadly elliptical, broadly acute and often apiculate or occasionally obtuse at apex, subentire or irregularly sinuate-dentate or with large, narrow to broad, mammiform, sometimes cusped teeth often with a roundedapiculate apex and confined to lower two-thirds of margins and cuneate to attenuate or rarely subtruncate at base, all usually with short, medium and long, subrigid simple eglandular hairs with pale or discoloured, occasionally blackish, bulbous bases uniformly distributed on the upper surface or restricted towards and at the margins, with scattered, rather shorter, softer ones on the lower surface or none, some extremely short, yellowish glandular hairs on the surfaces and the margins and sometimes with a few stellate hairs on the margins, the petioles short to long, more or less slender, often strongly reddish-violet tinted at base, with few to rather dense, medium to long, pale, subrigid simple eglandular hairs; cauline (1–)2–4(–5), the lowest rigidly semi-erect to patent, the lamina of lowest 10–80(–90) × 2–18 mm, sometimes undulate, linear-lanceolate or narrowly elliptical, acute or rarely rounded-obtuse at apex, usually entire, attenuate at base and usually with a distinct, sometimes winged petiole, semiamplexicaul at base, with few or no, often darkish-based simple eglandular hairs on the upper surface, but with several short to medium ones and very short, pale glandular hairs and few to numerous stellate hairs on the margins, and occasionally beneath and on the midrib above, the second leaf from base sometimes similar to the lowest, rigidly semi-erect to patent and up to 45 × 4 mm, the rest smaller, linear and bract-like are more or less appressed. Capitula solitary, rarely 2(–3), occasionally with a second stem from the base bearing a capitulum, 25–40(– 50) mm in diameter, sometimes nodding, the involucre deep greenish-grey or greenish-black in appearance, campanulate or broadly campanulate, rounded at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green with paler margins; inner 10–17 × 1.1–1.8 mm, broadly linear-lanceolate, abruptly acute, often acuminate, sometimes subacute and occasionally twisted at apex, somewhat appressed or often rather lax; outer somewhat shorter, sometimes lax, erect and subacute at apex; rather densely clothed with medium to long (–3 mm), slender, wavy, very pale to dark grey,

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black-based simple eglandular hairs, with numerous, very short, short and medium, dark glandular hairs which are usually only slightly obscured or occasionally fully obscured by the simple eglandular hairs, without stellate hairs but the inner with a very distinct tuft of short, pale hairs at the apex. Ligules pale (to medium) yellow, the inner dense, the outer medium to rather broad up to 4.0(–4.5) mm wide, slightly discrete, straight or shallowly upturned and deeply lobed at apex with more or less slender lobes, with few to many, very short to short, pale and often with some rather greyish simple eglandular hairs on the lower surface, or none, and several to many extremely short, sometimes with a few very short to short ones projecting beyond the apex. Styles pure yellow to dirty greenish-yellow with yellow stigmas. Receptacle pits with margins shortly dentate. Achenes 3.2– 4.0 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Cliff ledges and rock crevices on granite, quartzporphyry and quartz-biotite mica-schist from 740 to 1,010 m. Local and scarce on Carn nan Sac and Glas Tulaichean near Glen Shee in Perthshire; above Loch Wharral, near Braedownie, at the head of Glen Fee and in Coire Sharragh (Corrie Sharrock), all in Glen Clova, at the heads of Caenlochan and Canness Glens in Glen Isla, and in Glen Prosen in Forfarshire; not uncommon in the north-east coire of Lochnagar and in several localities above Loch Kander in Glen Callater, but local and very scarce in the Cairngorms in the Lairig Ghru, in Glen Geusachan at the southern and eastern foot of Cairn Toul and recorded on Craig an Dail Bheag (Little Craigandal) in Aberdeenshire. Endemic. Named after Beverley Alan Miles (1937–70). 95. H. memorabile P. D. Sell & C. West Memorable Hawkweed H. backhousei auct.; H. subgracilentipes auct. Phyllopodous perennial herb with a branched stock. Stem 10–24(–32) cm, pale yellowish-green, usually reddishviolet tinted towards the base, slender to medium, sometimes flexuous, usually markedly striate, the lower part with several, medium to long, slender, wavy, white, darkbased simple eglandular hairs, sometimes very long (up to 6 mm) at the base, few to scattered, very short, yellowish glandular hairs, and sparse stellate hairs, the upper part with several to many, short to medium and a few, long, usually greyish, occasionally white, black-based simple eglandular hairs, very numerous to dense, very short and short and occasionally medium (up to 1.2 mm), usually very dark glandular hairs, which usually greatly outnumber the simple eglandular hairs, and with a moderate clothing of stellate hairs. Leaves usually bright medium to deepish green, glossy, often with reddish-violet tints, rather paler beneath; basal usually numerous, the lamina of outer 8–70 × 7–23 mm, subrotund, obovate or broadly elliptical, often undulate, broadly obtuse-mucronate and sometimes twisted at apex, entire or denticulate to incise-dentate with mammiform or cusped teeth and cuneate shortly narrowed at base into short petioles, the lamina of inner 30–100 × 15–20 mm, narrowly elliptical to elliptical, obtuse to acute and apiculate and sometimes slightly twisted at apex, rigid, somewhat to markedly coriaceous, entire to more usually serrate-dentate, with irregular, often incise,

narrowly mammiform or cusped, teeth which have a long, rounded-obtuse-apiculate apex, attenuate at base, all with the upper surface often moderately and uniformly clothed with rather slender, curved, subrigid, white, pale or sometimes discoloured at base, medium and long (up to 3 mm) simple eglandular hairs, occasionally with only scattered ones or none, with many, mainly medium ones on the margin and scattered, softer, more wavy ones on the lower surface, with several to many, very short, yellowish glandular hairs on both surfaces and the margin and sometimes a few stellate hairs on the midrib beneath, the petioles rather short to medium, slender to sometimes winged, strongly reddishviolet-tinted, with numerous, short to medium, occasionally long simple eglandular hairs on the margin, often becoming very long and wavy about the base and with many, very short, yellowish glandular hairs; cauline (0–)1–3(–5), the lowest semipatent, the lamina 10–80 × 1–10(–20) mm, often linear-oblanceolate, sometimes linear or bract-like, sharply acute or acuminate at apex, usually entire, occasionally remotely denticulate, rarely with larger teeth and attenuate at base into a narrow or winged, sometimes semiamplexicaul petioles, the remainder bract-like and suberect, the uppermost filamentous. Capitula 1–2(–4), frequently with a second stem from the base bearing a capitulum, occasionally with a rudimentary bud in the axil of a cauline leaf, 30–45 mm in diameter, the involucre blackish or dark greenish-grey in appearance, cylindrical or narrowly campanulate, rounded or more or less subtruncate at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, the inner with paler margins, inner 10–16 × 0.9–1.5 mm, rather appressed, linear-lanceolate to linear, the innermost very narrow, subacute, abruptly acute or acuminate at apex; outer suberect, shorter and acute at apex; all with numerous to dense, unequal, very wavy and somewhat interwoven, slender, medium to long (up to 2.5 mm), greyish or occasionally white, black-based simple eglandular hairs, numerous very short to short and medium (–1.5 mm) dark glandular hairs the majority of which are not obscured by the simple eglandular hairs and without stellate hairs but the inner with a distinct tuft of short hairs at the apex. Ligules deepish (to medium) yellow, frequently with a slight greenish tinge, the outer up to 4 mm wide, often markedly discrete, usually very upturned and, often ascending at the shallowly to deeply lobed or rounded apex, with few to numerous, extremely short, very short and short, pale or very slightly discoloured simple eglandular hairs on the lower surface, and several, extremely short, very short and sometimes short ones projecting beyond the apex. Styles olive-grey, rarely yellow. Receptacle pits with margins shortly dentate. Achenes (3.0–)3.8–5.0 mm, purplish-black. Flowers 7–8. Apomictic 2n = 36. Some plants differ from the rest in their broader involucre and occasionally less obvious glandular hairs on the involucre and sometimes yellow styles. Native. A local and uncommon species on cliff ledges, rocks, scree and the banks of streams from 670 to 1,065 m, most frequently on schist, quartz-biotite-feldspar gneiss, but also on granite. Recorded on Ben Lawers and Beinn Heasgarnich in Perthshire; Canness, Glen Isla in Forfarshire;

39. Hieracium Cairntoul, Beinn a’ Bhuird and the head of Glen Callater in Aberdeenshire; Loch Avon in Banffshire; Glen Einich, Glen Feshie, the Lairig Ghru, the Ben Nevis Range, Glen Spean and hills above Loch Laggan, in Inverness-shire; and Glen Coe, Glen Etive and the Glen Orchy hills in Argyllshire. Endemic. 96. H. insigne Backh. fil. Noble Hawkweed H. alpinum var. insigne (Backh. fil.) Bab.; H. melanocephalum var. insigne (Backh. fil.) Bab.; H. alpinum forma grande sensu F. N. Williams; H. alpinum subsp. nigrosetosum auct. Phyllopodous perennial herb with a branched stock. Stem 12–32 cm, pale green, sometimes reddish-violet tinted at base, slender to robust, usually striate in lower part, often slightly flexuous, the lower part with few to many, short, medium and long, wavy, dark-based, white simple eglandular hairs, scattered, very short, pale and dark glandular hairs and sparse stellate hairs, the upper part with many short to medium and sometimes several long (up to 3 mm), black-based simple eglandular hairs, several to numerous, very short, short and sometimes medium, dark glandular hairs and rather densely clothed with stellate hairs. Leaves rather pale to rather deep bright green, somewhat glossy on the upper surface, paler and glossy or slightly glossy on lower surface, often tinted or suffused with reddish-violet; basal several to numerous, the lamina of outer 6–60 × 3–28 mm, variable, subrotund, ovate, oblong or elliptical, broadly rounded-obtuse, sometimes retuse at apex, often undulate, entire, denticulate, with a few, large teeth or sinuate-dentate with broad, mammiform teeth, cuneate to broadly cuneate, subtruncate, or occasionally attenuate at base and petiolate, the lamina of inner 15–80 × 3–28 mm, rigid or slightly coriaceous, broadly to narrowly elliptical, oblanceolate or narrowly obovate, frequently somewhat asymmetrical, sometimes slightly channelled and undulate or twisted, broadly rounded-obtuse, occasionally subacute or the innermost acute and usually apiculate at the sometimes downwardly twisted apex, sometimes subentire or denticulate, or more usually with several, very irregular, more or less widely spaced, medium to very large, patent and forwardly directed teeth and sometimes a few retrorse, shallowly and narrowly mammiform or cusped teeth which have long, roundedobtuse-apiculate apices and which are usually confined to the lower two-thirds of the margin and sometimes descend onto the petiole, attenuate or narrowly cuneate at base, all with the upper surface rather sparsely to moderately and uniformly clothed with white, subrigid, apically curved or wavy, short, medium and long (up to 3 mm) simple eglandular hairs which have swollen, pale or discoloured bases, or occasionally with few or none except towards and on the margins, the innermost leaves usually uniformly and more densely clothed with long simple eglandular hairs, the lower surface with few to numerous simple eglandular hairs, often with discoloured or black bases, on the lower surface, with few to numerous, very short, yellowish glandular hairs on both surfaces and on the margin, and sometimes with a few, stellate hairs on the lower surface, the petioles rather long, rarely short, sometimes slightly winged, rather sparsely to densely clothed with medium and long simple eglandular

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hairs on the margins and usually tinted with reddish-violet at base; cauline (1–)2–4(–8), the lowest frequently rigid and semipatent, the lamina 11–90 × 2–15(–20) mm, linearelliptical or oblanceolate, sometimes like the innermost basal, often undulate and twisted, broadly acute or subacute and apiculate at apex, entire, denticulate or with some larger, broadly mammiform teeth, and gradually narrowed to an often sessile or semiamplexicaul base, sometimes with a winged petiole, the second lowest semipatent, and the lamina up to 70 × 14 mm in large plants, the remainder small, linear-elliptical or linear, bract-like, more or less erect and subacute at apex. Capitula usually solitary, occasionally 2–3, or with a rudimentary bud in the axil of a cauline leaf, 25–60(–65) mm in diameter, the involucre very dark grey or greenish-grey in appearance, broadly to narrowly campanulate and rounded to broadly rounded at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, the inner with slightly paler margins; inner 10–19 × 1.0–2.0 mm, unequal, semilax, to appressed, linear-lanceolate, gradually and sharply acute or acuminate and sometimes incurved at apex; outer shorter, semilax, erect, sometimes slightly recurved and acute to obtuse at apex; all with dense, short to long (up to 2.5 mm), wavy, slender white or somewhat greyish, blackbased simple eglandular hairs and numerous, very short and short, dark glandular hairs which are partially to almost completely obscured by the simple eglandular hairs and without stellate hairs, but the inner with a tuft of short hairs at the apex. Ligules medium (to palish), sometimes deepish yellow, the outer 3.5–5.0 mm wide, slightly discrete, upturned to straight and shallowly to very deeply lobed at apex, sometimes with very long (up to 5.0 mm) narrow lobes, sparsely to moderately clothed on the lower surface with very short and short, pale or slightly discoloured simple eglandular hairs, or rarely with none, and with several to many extremely short and very short ones projecting beyond the apex. Styles usually pure yellow, sometimes slightly dirty yellow, often darkening when dried. Receptacle pits with margins dentate. Achenes 3.0–4.5 mm, purplish-black. Flowers 7–8. Apomictic. (1) Forma insigne Capitulum solitary, 45–65 mm in diameter. Involucral bracts 12–19 × 1.0–2.0 mm, lax, very narrow at apex. 2n = 36. (2) Forma celsum P. D. Sell & D. J. Tennant Capitula 1–3, 25–50(–55) mm in diameter. Involucral bracts 10–16 × 1.0–1.6 mm, appressed, less narrow at apex. 2n = 27, 36. Native. A local and very scarce species found on rockledges and sometimes rocky slopes on granite, quartzporphyry (red feldspar) and possibly Moine Schist from 670 to 1,000 m. Forma insigne is known only from cliffs above Loch Kander (Loch Ceanndin) in Glen Callater in Aberdeenshire. Forma celsum occurs in Coire an Lochain Uaine, Cairn Toul in Aberdeenshire and in Coire Garbhlach in Glen Feshie, Creag an Leth-choin in the Lairig Ghru and several places on Beinn na Socaich, Glen Spean in Inverness-shire. The species is endemic. The 2n = 27 count

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is from a Beinn na Socaich plant; four counts from eastern localities are all 2n = 36. 97. H. backhousei F. Hanb. Backhouse’s Hawkweed H. nigrescens subsp. backhousei (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 8–28(–38) cm, slender to rather robust, markedly striate in lower part, reddish-violet tinted at base, the lower part with few to scattered, mainly short and medium, slender, wavy, dark-based, white simple eglandular hairs, few very short glandular hairs and scattered or sparse stellate hairs, the upper part with many to very numerous, short to medium and frequently several long, black-based, white or somewhat greyish simple eglandular hairs, scattered to numerous, very short and short dark glandular hairs and moderately clothed with stellate hairs. Leaves medium to rather deep, occasionally pale, drab green, matt or only faintly glossy, frequently tinted with dull reddish-violet, especially on the margins, distinctly paler and minutely granulate in appearance on the lower surface; basal numerous, the lamina of outer 8–50 × 5–27 mm, subrotund, obovate or broadly elliptical, broadly obtuse and usually mucronate at apex, entire, denticulate and sometimes with a few larger teeth and rounded to shortly narrowed at base, the lamina of inner 20–110 × 5–24 mm, usually elliptical to narrowly elliptical, lanceolate or oblanceolate, gradually narrowed to an obtuse or acute and apiculate sometimes slightly twisted apex, occasionally rather undulate, rigid and coriaceous, subentire to irregularly and remotely serrate-dentate, often with long, ascending, narrowly mammiform, mammiform or cusped, acute teeth with long, rounded-obtuse-apiculate apices and are usually confined to the lower two-thirds of the margin, and gradually or shortly narrowed at base, all with the upper surface sparsely clothed with subrigid, rather pale-based, white, medium and long simple eglandular hairs, or often with none except on the margins which have numerous short to medium ones, the lower surface sparsely clothed or with none except on the midrib and the innermost often more densely clothed with simple eglandular hairs, with scattered, very short, yellowish glandular hairs on both surfaces and on the margins and sometimes with a few, stellate hairs on the lower surface or margins, the petioles short to long, slender or more frequently winged, usually reddish-violet-tinted; cauline (2–)3–4(–8), the lowest rigidly semipatent and sometimes like the innermost basal, the lamina 20–60(–85 cult.) × 2–10(–2.2 cult.) mm, usually very narrowly elliptical, sharply acute or acuminate at apex, denticulate or more frequently with several, sharply acute, narrow, often spinulose teeth, seldom entire, often with a short but distinct petiole which is often winged and semiamplexicaul, usually with no simple eglandular hairs on the upper surface but with numerous short to medium ones on the margin and on the lower surface, and with scattered, very short, yellowish glandular hairs on both surfaces, the upper smaller, bract-like or filamentous, acute at apex and moderately clothed with stellate hairs, often densely so on the lower surface. Capitula solitary, or not infrequently 2–3(–8 in cult.), 30–54 mm in diameter, often with a rudimentary bud in the axil of a cauline leaf, the involucre very dark greenish-grey in appearance, campanulate, rounded

or subtruncate at base; peduncles when present often long and clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, with the innermost paler; inner 11–17 × 1.0–1.8 mm, unequal, appressed, linear-lanceolate or lanceolate-subulate, more or less acute or subacute at apex; outer short, semilax, erect, subacute at apex; all with dense white or somewhat greyish, black-based, wavy, somewhat interwoven, medium and long simple eglandular hairs, numerous very short and short, yellowish to dark glandular hairs which are usually almost fully obscured by the simple eglandular hairs, without stellate hairs, but the inner with a pronounced tuft of short hairs at the apex. Ligules bright rather pale (to medium) yellow, the outer rather broad (up to 4.5 mm wide), slightly discrete, straight to shallowly upturned and deeply lobed at apex with narrow lobes, with few to many pale or slightly discoloured, very short or short simple eglandular hairs on the lower surface and many extremely short and very short ones projecting beyond the apex. Styles pure yellow to slightly discoloured yellow. Receptacle pits with margins incise-dentate. Achenes 3.0– 4.2 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Rare and local in crevices and pockets on shelving granite slab-rock and boulders, usually by fast-running streams and waterfalls from 670 to 830 m. About the head of Glen South Esk, Clova in Forfarshire; near Dubh Loch, Glen Muick, and on Derry Cairngorm, and in Coire Etchachan, in Aberdeenshire. Endemic. Named after James Backhouse jun. (1825–90). 98. H. mundum P. D. Sell & C. West Elegant Hawkweed Phyllopodous perennial herb with a branched stock. Stem 7–24(–30) cm, pale yellowish-green, strongly tinted with reddish-violet at base, rather slender to somewhat robust, often flexuous, with few to numerous, slender, wavy, short, medium and long, white to somewhat greyish simple eglandular hairs with thickened, blackish bases throughout, often dense, very long, very wavy and sometimes retrorse ones in the lower part, becoming moderately clothed with medium to long, patent ones in the upper part, the lower part with very few, very short, fine glandular hairs becoming numerous to dense in the upper part, with numerous stellate hairs which often become dense in the upper part. Leaves somewhat drab medium to rather deep green, matt or only faintly glossy on upper surface, rather paler on lower surface which is usually strongly tinted or blotched with reddish-violet, especially on the midrib, margin and teeth; basal few to numerous, the lamina of outer 6–55 × 4–23 mm, mainly subrotund or elliptical, rounded-obtuse at apex, usually denticulate, sometimes dentate and cuneate at base, the lamina of inner 20–80 × 6–20 mm, lanceolate, oblanceolate or narrowly elliptical, often somewhat channelled and slightly undulate, slightly coriaceous and rather rigid, more or less obtuse or the innermost acute at apex, serrate-dentate with more or less regular or irregular, narrow, incise, often apiculate teeth and attenuate at base, all with numerous slender, subrigid, medium to long simple eglandular hairs, with pale or discoloured bulbous bases on the upper surface, the outer usually sparsely clothed, the innermost more or less densely clothed with uniformly distributed ones or the

39. Hieracium hairs restricted towards and at the margins, the lower surface with few to numerous, more or less uniformly distributed, short to medium, often black-based simple eglandular hairs, with some very short, yellowish glandular hairs and often with a few stellate hairs on both surfaces and the margins, the petioles rather short, rarely long, rigid, more or less winged, reddish-violet-tinted; cauline (2–)3– 6(–7), the lowest spreading, the lamina 10–75 × 1–15(–22) mm, often undulate, linear or linear-lanceolate, acute or subacute at apex, with long, narrow, reddish-violet-tinted, ascending teeth, or regularly spinulose-serrate, very seldom entire, often like the innermost basal, very robust plants often have up to 4 large, spreading cauline leaves situated almost at the base of the stem, all with very narrow teeth and attenuate at base into a winged or more or less amplexicaul short petiole, the remainder small, acute at apex and more or less bract-like. Capitula solitary or up to 3, 25–40(–47) mm in diameter, the involucre very dark greenish-grey or blackish-grey in appearance, campanulate, rounded at base; peduncles when present clothed like the stem. Involucral bracts incumbent in bud, blackish-green or tinted with red or brown; inner 11–15 × 1.0–1.5 mm, more or less appressed, linear-lanceolate, mostly subacute at apex or some obtuseapiculate and the innermost abruptly acute; outer much shorter, erect and obtuse at apex; all densely clothed with very short to long (up to 2.5 mm), slender, wavy, white or slightly dusky, black-based simple eglandular hairs, numerous very short to short (up to 0.8 mm), dark glandular hairs only partially obscured by the simple eglandular hairs and without stellate hairs but with a pronounced tuft of very short hairs at the apex. Ligules uniformly medium to rather deep yellow, the outer rather broad (up to 4.3 mm), appreciably discrete, slightly upturned and shallowly to deeply lobed at apex, with very few to many, very short to short, pale or slightly discoloured simple eglandular hairs on the lower surface, and several to many, very short and often a few short ones projecting beyond the apex. Styles more or less uniformly greenish-grey. Receptacle pits with margins shortly dentate. Achenes 3–4 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Rocky knolls, detritus and shelving rocks, less frequently on cliff ledges, on fine-grained quartz-biotite and other rocks between (425–)580 and 760 m. Fannich Forest hills, Braemore and the Beinn Dearg Range in Ross-shire; Carn Dearg, Foinaven, Ben Hope and near Glen Coul in Sutherland. Endemic. 99. H. marginatum P. D. Sell & C. West Spear-leaved Hawkweed H. globosiflorum var. lancifolium Pugsley, non H. lancifolium Vuk. Phyllopodous perennial herb with a branched stock. Stem 5–17(–25) cm, pale green, often reddish-violet-tinted towards the base, slender, sometimes slightly flexuous, often striate, with few to numerous, slender, usually short to medium simple eglandular hairs, few to several, very short glandular hairs and very few or scattered stellate hairs in the lower part, the upper part with few to very numerous, medium and longish (up to 2.5 mm), slender, wavy, whitish simple eglandular hairs with long, thickened,

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black bases, several to numerous, very short and short, dark or pale-tipped glandular hairs and scattered to numerous, but seldom dense stellate hairs. Leaves rather pale to medium, very seldom rather deep, drab green, sometimes slightly glossy and reddish-violet-tinted on the upper surface, paler beneath; basal more or less numerous, often in a fairly neat rosette, the lamina of outer 3–50 × 2–20 mm, mainly elliptical or obovate, sometimes oblong-elliptical or ovate, broadly rounded-obtuse or obtuse at apex, denticulate or shallowly serrate-dentate with mammiform teeth and cuneate at base, the lamina of inner 10–50(–70) × 4–20 mm, mainly narrowly elliptical to elliptical, sometimes oblanceolate to lanceolate, often somewhat channelled, sometimes cymbiform and undulate at the margins, rather coriaceous and rigid, with conspicuously impressed veins, acute or obtuse and sometimes mucronate or apiculate at apex, subentire, denticulate or occasionally with several patent or ascending, acute, long, narrowly mammiform teeth, and mostly attenuate, seldom cuneate at base, all with the upper surface moderately or densely clothed with mainly short to medium, occasionally long, slender, subrigid, wavy or curved, usually pale-based simple eglandular hairs, or the majority more frequently sparsely clothed, or with none and the lower surface and margins usually moderately and uniformly clothed with slender, wavy, short or medium ones, with scattered, very short, yellowish glandular hairs on the lower surface and margins and with several to many stellate hairs on the margin and sometimes a few on the midrib above; petioles usually short to occasionally rather long, slender or winged, with numerous, long simple eglandular hairs and often reddish-violet tinted at base; cauline 0–2(–3), the lowest more or less erect, the lamina 5–30 × 1–3 mm, linear or bract-like, acute at apex and attenuate at base and occasionally petiolate, the remainder bract-like, appressed, acute at apex and the uppermost filamentous. Capitula usually solitary, or rarely 2, and seldom with a rudimentary bud in the axil of a cauline leaf, 25–45 mm in diameter, the involucre silvery grey to dark greenishgrey in appearance, often rather small and narrow, cylindrical or narrowly campanulate, subtruncate or rounded at base, sometimes constricted in the middle; peduncles when present, long and clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, the inner with very pale margins; inner 9–17 × 0.8–1.5 mm, often unequal, very appressed, narrowly linear-lanceolate, the innermost often filamentous, mainly very gradually narrowed to a sharply acute apex, sometimes with a few subacute and occasionally obtuse ones present; outer rather lax, suberect, shorter and more or less acute at apex; all with numerous to dense, medium to long (up to 4 mm), wavy, slender, white or somewhat greyish, black-based simple eglandular hairs, numerous very short and short, dark or pale-tipped glandular hairs which are completely obscured by the simple eglandular hairs or seldom with very few obvious ones and without stellate hairs, the inner sometimes with a tuft of short hairs at apex. Ligules bright medium, occasionally rather pale yellow, the outer often rather broad (up to 4.5 mm), usually few and very discrete, straight or slightly upturned, sometimes ascending at the apex, which is deeply and irregularly lobed, or sometimes partially developed with the

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2 outermost lobes large and the remainder very small, with few to many, very short or short, pale or faintly discoloured simple eglandular hairs on the lower surface and several to numerous, occasionally few, very short ones or rarely none, projecting beyond the apex. Styles pure yellow or faintly discoloured (olive-yellow); stigmas pure yellow. Receptacle pits with margins shortly dentate. Achenes 4.0–4.5 mm, purplish-black. Flowers 7–8. Apomictic. (1) Forma marginatum Involucral bracts with dense simple eglandular hairs up to 2.5 mm. 2n = 36. (2) Forma chaetocephalum P. D. Sell & C. West Involucral bracts with dense simple eglandular hairs up to 4 mm. Sometimes hairs on stem, petioles and leaves more numerous than in forma marginatum. 2n = 36. A plant which is similar to H. marginatum but differs mainly in having few simple eglandular hairs and numerous, obvious, yellow black-based glandular hairs on its narrow, greenish involucre occurs locally on Torridonian Sandstone on An Teallach in Ross-shire. Native. Rock-ledges, exposed rocky knolls and screeslopes between 530 and 915 m on Moine Schist, quartzbiotite mica-schist and Torridonian Sandstone and in one locality on syenite, a form of granite. Probably on most of the hills in Ross-shire and not infrequent on the hills of west Sutherland, east and south of Loch More and south of Carn Dearg in east Sutherland and above Loch Laggan in Inverness-shire where it is scarce. Endemic. The two forms often occur in the same populations and occur throughout the greater part of the range of the species, but forma chaetocephalum occurs in smaller numbers and is the only form in Inverness-shire. 100. H. calvum P. D. Sell & D. J. Tennant Bald-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 13–20(–27) cm, sometimes 2 or more, slender or rather robust, often flexuous, sometimes reddish-violet-tinted at base, the lower part with several to numerous, medium and long, wavy, white, black-based simple eglandular hairs becoming denser and very long below, few to several, extremely short, yellowish glandular hairs and few or no stellate hairs, the upper part with numerous, medium and long (or in uppermost part very long), white or greyish simple eglandular hairs with thickened black bases, scattered very short and short dark glandular hairs and several to many but not dense stellate hairs. Leaves pale or medium, or rarely rather deep green, the inner usually glossy on the upper surface, sometimes tinted reddish-violet on the surfaces and margin, paler green on lower surface; basal few to numerous, the lamina of outer 10–70 × 6–26 mm, mainly broadly elliptical or ovate, broadly rounded-obtuse to subacute at apex, sometimes apiculate, entire or denticulate and cuneate or shortly narrowed at base, the lamina of inner 30–90 × 5– 22 mm, rigid and coriaceous, elliptical, narrowly elliptical or oblanceolate, mainly acute, often acuminate or occasionally subacute at the frequently twisted apex, usually entire, sometimes undulate, or serrate-dentate, with a few, shallow

to narrowly mammiform teeth and long attenuate or occasionally shortly narrowed at base, the majority usually completely glabrous on the entire upper surface, but with numerous to dense, long to very long, wavy, white, dark-based simple eglandular hairs on the margins and petioles, with scattered to numerous, pale or dark-based simple eglandular hairs on the lower surface, few very short, yellowish glandular hairs on the lower surface and margin and sometimes a few stellate hairs on the margins, the petioles more or less (often broadly) winged and frequently reddish-violet tinted at the base; cauline 1–2(–3), the lowest semipatent, the lamina 10–60(–90) × 1.5–10 mm, linear-oblanceolate to narrowly elliptical, acute or acuminate, rarely obtuse at apex, entire and attenuate at the sessile or petiolate base, glabrous on the upper surface but with numerous, longish simple eglandular hairs and some stellate hairs on the margin and lower surface, the remainder usually linear, entire, acute at apex or bract-like, occasionally with a rudimentary bud in the axil. Capitula usually solitary, sometimes 2, or with 2 or more stems from the basal rosette each bearing a capitulum, 40–55 mm in diameter, the involucre dark greenish-grey or greenish in appearance, narrowly campanulate, shortly narrowed or rounded at base; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green, the inner with the margins paler green; inner 10–16 × 1–2 mm, appressed or loosely appressed, linearlanceolate, sharply acute or abruptly acute and acuminate and sometimes violet-tinted at apex; outer shorter and subacute or acute at apex; all with long, very wavy, interwoven, white or somewhat greyish, black-based simple eglandular hairs, being sparse in zones but invariably numerous overall, and often very long (–5 mm) at the base, many very short to short, pale or discoloured glandular hairs which are only partially obscured by the simple eglandular hairs and without stellate hairs, but the inner with some very short hairs at the apex. Ligules medium yellow, the outer up to 4–6 mm wide, slightly discrete, slightly upturned and often deeply lobed at apex, with very few to several, short to medium, pale or slightly discoloured simple eglandular hairs on the lower surface and a few very short or short (to medium) ones, or none, projecting beyond the apex. Styles pure yellow to dirty yellow, often darkening when dried. Receptacle pits with margins shortly dentate. Achenes 3.5–4.1 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Native. Extremely local and very scarce in crevices and on ledges of shelving coarse-grained granite rocks and boulders between 800 and 920 m. Confined to the Cairngorms on Carn Crom and Derry Cairngorm on the west side of Glen Derry in Aberdeenshire and on the east side of Cairn Gorm in Inverness-shire. Endemic. 101. H. leptodon P. D. Sell & D. J. Tennant Narrow-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 12–30 cm, medium to slender, sometimes flexuous, usually reddish-violet-tinted at base, with scattered to numerous, slender, wavy, medium to very long (–5 mm), pale, black-based simple eglandular hairs, few pale and darker extremely short to very short glandular hairs and few stellate hairs in the lower part and with numerous long simple

39. Hieracium eglandular hairs with long, black bases, very short to short, blackish glandular hairs and several to numerous but not very dense stellate hairs in the upper part. Leaves rather pale to medium green, subglaucous matt or only faintly glossy above often with reddish-violet tints, frequently so on the teeth and margin, paler green beneath; basal numerous, the lamina of outer 8–60 × 5–25 mm, broadly elliptical to ovate, rounded-obtuse, sometimes apiculate, occasionally retuse at apex, denticulate to subentire, sometimes with several mammiform teeth and cuneate or shortly narrowed, sometimes attenuate at base, the lamina of inner 25–90 × 9–22 mm, narrowly to broadly elliptical, occasionally oblanceolate, obtuse to acute, often apiculate at the sometimes twisted apex, denticulate, dentate or serratedentate, sometimes undulate, with regular or irregular, often ascending, narrowly or occasionally broadly mammiform teeth, with long, rounded-apiculate apices but often entire in the upper third of the margins and shortly narrowed or attenuate at base, all with numerous, short, medium and long (–3 mm), white simple eglandular hairs with pale or dark bases uniformly distributed on both surfaces of most leaves and dense long, ones on the margins, and with scattered, very short, yellowish glandular hairs, the petioles usually short, winged and with rather dense, very long, wavy, simple eglandular hairs and often reddish-tinted on the underside; cauline 2–4(–5), the lowest semipatent, the lamina 10– 50(–70 cult.) × 2–20 mm, narrowly elliptical or oblongelliptical, acute or subacute at apex, often serrate-dentate or serrate, with irregular, very long (–10 mm), narrow, often forwardly curved, apiculate teeth which are frequently spinulose or at least with a few, sharp, narrow teeth, occasionally entire, narrowed at base, sessile and semiamplexicaul or subpetiolate, the second lowest often with a few, sharp, narrow teeth or entire, the remainder erect, linear, bract-like or filamentous, entire and acute at apex. Capitula 1–3(–4), 35–50(–60) mm in diameter, the involucre greenish-grey to dark grey in appearance, cylindrical or campanulate and rounded or truncate at base; peduncles long and slender, clothed like the upper stem. Involucral bracts incumbent in bud, blackish-green; inner 12–16 × 0.8–1.0(–1.5) mm, appressed, linear-lanceolate, mostly gradually and sharply acute, sometimes abruptly acute at apex and the innermost sometimes filamentous; outer much shorter, semilax, erect, subacute at apex; all with numerous to dense, medium to long (–3 mm), white or greyish, black-based simple eglandular hairs usually completely obscuring the numerous, very short and short, dark glandular hairs, without stellate hairs but the inner with a few very short hairs at the apex. Ligules pale, bright or occasionally medium yellow, the outer up to 4 mm wide, numerous but slightly discrete, upturned and usually deeply and irregularly lobed at the apex, with numerous to very numerous, very short, to medium pale simple eglandular hairs on the lower surface and numerous very short and some short ones projecting beyond the apex. Styles deep greenish-grey. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Shows little variation but the Argyllshire plants rarely develop the long characteristic spinulose teeth on the lower cauline leaves and have wider involucral bracts and ligules.

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Native. Cliff ledges and crevices in large boulders on muscovite-albite pelitic schist and other hard mica-schist rocks at 710–890 m in the north-east corrie of Ben More in Perthshire; and scarce locally on granitic rocks and scree at 500–600 m in Glen Coe, Argyllshire. Endemic. 102. H. macrocarpum Pugsley Large-fruited Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–24(–30) cm, pale green, often reddish-violet-tinted towards the base, slender to medium, rarely somewhat robust, sometimes flexuous, often striate, with numerous long or very long, wavy, dark-based simple eglandular hairs, few, very short glandular hairs and scattered stellate hairs in the lower part, the upper part moderately to densely clothed with mainly medium, sometimes very long (over 4 mm), slender, unequal, wavy, white to somewhat greyish simple eglandular hairs with long thickened black bases, numerous very short to short, dark glandular hairs and moderately to densely clothed with stellate hairs. Leaves usually pale green, matt or barely glossy, sometimes tinted with reddishviolet, especially on the midrib, margin and teeth, occasionally blotched, paler and whitish-green on lower surface and minutely granulate in appearance; basal usually numerous, often in a more or less neat rosette, the lamina of outer 10– 50 × 5–20 mm, mainly broadly elliptical, sometimes obovate, often undulate, broadly rounded-obtuse to obtuse and sometimes mucronulate at apex, entire to sinuate-dentate with shallowly mammiform teeth and cuneate or shortly narrowed at base, the lamina of inner 15–90 × 3–20 mm, not coriaceous but often very rigid, often slightly involute and undulate at the margins and slightly hooded at apex, mainly narrowly elliptical or oblanceolate, sometimes oblong-elliptical or linear-elliptical, sometimes asymmetrical, broadly obtuse to acute, often mucronate or apiculate and sometimes twisted at apex, subentire or more or less uniformly denticulate to serrate-dentate, with often irregular, narrowly mammiform, mammiform or sometimes large, acute, cusped teeth, the larger often confined to the lower half of the margin and attenuate or shortly narrowed at the base, the majority moderately to rather densely and uniformly clothed on the upper surface with medium or long, wavy or apically curved, subrigid, white simple eglandular hairs with mainly pale, bulbous bases, sometimes very long on the innermost leaves, with numerous, mainly medium ones on the margin and moderately clothed with shorter, softer, very slender, very wavy ones on the lower surface, with scattered, very short, yellowish glandular hairs on both surfaces and the margin and occasionally a few stellate hairs on the midrib of the lower surface, the petioles shortish, seldom very long, slender or winged, usually reddishviolet-tinted at least at the base and with numerous medium or long simple eglandular hairs, which become dense and very long and wavy towards the base and scattered, very short, yellowish glandular hairs; cauline (0–)1–2(–3), the lowest suberect, the lamina 5–60 × 2–8(–12) mm, linear or linear-oblanceolate, acute at apex, usually entire, very occasionally dentate, very attenuate at base, shortly petiolate or sessile and often semiamplexicaul, the remainder small, appressed, bract-like and very sharply acute at apex and the

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uppermost filamentous. Capitula usually solitary, occasionally 2, very rarely 3, often with a rudimentary bud in the axil of a cauline leaf, 25–45 mm in diameter, occasionally with an additional stem from the base bearing a capitulum, the involucre lead grey to greyish-black in appearance, sometimes cylindrical, frequently narrowly campanulate, very rounded at base and constricted in the middle; peduncles when present clothed like the upper stem. Involucral bracts incumbent in bud, the outer blackish-green, the inner paler and greenish; inner (8–)10–15(–17) × 0.9–1.5(–1.7) mm, often more or less uniform, rather closely appressed, narrowly linear-lanceolate or lanceolate-subulate, the innermost often very narrow, attenuate to a sharply acute or very occasionally subacute or acuminate apex; outer shorter, often more or less appressed and indistinct, acute at apex; all with dense, medium to long, very wavy, slender, white to somewhat greyish, sometimes semiappressed, often interwoven, black-based simple eglandular hairs, numerous very short to short, fine yellowish or darker glandular hairs, which are nearly always fully obscured by the simple eglandular hairs, or occasionally with a few obvious ones, without stellate hairs but the inner sometimes with a distinct tuft of very short hairs at the apex; Ligules medium yellow, sometimes with a slight greenish tinge, the outer usually not very broad (3.2–3.8(–4.1) mm), few and very discrete and not contiguous for the greater part of their length, deeply lobed at apex or occasionally the lobe partially developed and the 2 outermost very large and the remainder very small, with numerous short to medium, pale, sometimes rather appressed simple eglandular hairs on the lower surface and numerous, very short to short and often medium ones projecting beyond the apex. Styles pure yellow to faintly discoloured (olive yellow). Receptacle pits with margins longdentate. Achenes usually long (3.0–)3.7–5.3 mm, purplishblack. Flowers 7–8. Apomictic. 2n = 36, 37. Native. Rare or local on cliff ledges, in rock crevices and amongst turf on rocky slopes and stream gullies between 610 and 1,000 m, mainly on schist formations and quartz-biotite-feldspar gneiss but also spreading to adjacent, coarse-grained granite. Cairngorm Mountains with an outlying locality to the south-west; Coire Etchachan, several places on Cairn Toul, above the Lairig Ghru Pass and on Craig an Dail Bheag (Little Craigandal) in Aberdeenshire and several localities towards the head of Glen Einich, the south side of Coire Garbhlach in Glen Feshie and Coire Chuirn near Dalwhinnie in Inverness-shire. Plants with dark styles occur in Glen Carron Forest, Ross-shire which otherwise cannot be satisfactorily distinguished from H. macrocarpum, but which also have some characters with H. eximium. Endemic. 103. H. eximium Backh. fil. Hirsute Hawkweed H. nigrescens subsp. eximium (Backh. fil.) Zahn; H. halleri auct. Phyllopodous perennial herb with a branched stock. Stem 9–28(–35) cm, slender to somewhat robust, sometimes striate and flexuous, occasionally pale reddish-violet-tinted at base, with numerous to dense, unequal, long to very long, wavy, white or slightly grey, blackish-based simple eglandular hairs throughout or occasionally with only shortish to

medium ones in upper part, nearly always with numerous to very dense, long to very long, wavy ones at base, with few to numerous, slender, very short to short, and very occasionally a few medium, pale and dark glandular hairs, often very numerous above and with a moderate to dense clothing of stellate hairs in the upper part with rather few towards the base. Leaves rather pale to medium bright green, occasionally drab, rather deep green, only faintly glossy or matt on upper surface, seldom tinted with reddish-violet, the outer often somewhat coriaceous, but the inner not or only slightly so, sometimes rigid; basal few to numerous, the lamina of outer 10–50 × 5–20 mm, elliptical or broadly oblanceolate, occasionally oblong, usually obtuse-apiculate, very occasionally retuse, sometimes twisted at the apex, entire, denticulate or with a few larger teeth, occasionally with several, regular, evenly spaced, narrow aquiline teeth, and usually more or less attenuate at the base, the lamina of inner 20–100 × 5–25 mm, rarely coriaceous, usually narrow, either oblanceolate, linear-oblanceolate or narrowly elliptical, sometimes undulate or twisted, and frequently and markedly twisted sideways at the acute, often apiculate, sometimes subacute, only very occasionally obtuse at apex, denticulate or frequently serrate-dentate, with irregular, often widely spaced, long, narrow, aquiline-mammiform or occasionally spinulose teeth, occasionally all leaves entire, very attenuate at the base often into a more or less winged petiole, all with numerous to very dense, coarse, long to very long, white simple eglandular hairs with blackish bulbous bases, occasionally with more slender, very wavy, subrigid, shorter ones, invariably uniformly distributed on both surfaces and on the margin, except the outermost which are sometimes without hairs on the upper surface except near the margins, the hairs usually softer and rather shorter on the lower surface, often with many, very short, yellowish glandular hairs on the surfaces and margins, the petioles short to very long (up to 9 cm), with numerous, long to very long simple eglandular hairs which are frequently very dense, coarse and wavy towards the base, only occasionally with fewer, medium, subrigid ones throughout; cauline 1–3(–5), rarely none, the lamina of the lowest 10–110 × 1–15 mm, linear-lanceolate or linear-elliptical to oblanceolate, invariably markedly acute at apex, usually entire, occasionally denticulate or infrequently with several, often narrow, acute teeth, very attenuate at base, often decurrent and occasionally petiolate, the remainder much smaller, often suberect, more or less bract-like, linear, entire, or filamentous and acute at apex, all clothed with numerous long to very long, coarse, white, black-based simple eglandular hairs, uniformly distributed on the surfaces and the margins, occasionally with only scattered, medium, subrigid ones, frequently with dense, long, wavy ones around the base. Capitula usually solitary, very rarely 2, 30–50 mm in diameter, occasionally with more than 1 stem from the basal rosette each bearing a capitulum, the involucre variable, narrow to somewhat robust, cylindricalcampanulate or campanulate, rounded, somewhat narrowed or occasionally subtruncate at base, medium greenish-grey, rarely blackish-grey in appearance; peduncles when present clothed like upper stem. Involucral bracts incumbent in bud, greenish-black; inner 9–20 × 0.7–1.6 mm, numerous

39. Hieracium in well-developed plants, usually somewhat lax, occasionally rather appressed, more or less equal or unequal, narrowly linear-lanceolate or linear, the innermost somewhat filamentous, attenuate-acute, acuminate or seldom subacute at apex; outer much shorter, subacute or acute at apex, lax and erect; densely clothed with wavy, long to very white or infrequently greyish, black-based simple eglandular hairs, often very dense at the base, usually fully obscuring the very short and short, pale and dark, slender glandular hairs, or occasionally with some obvious pale glandular hairs, without stellate hairs, but the inner with a tuft of short hairs at the apex. Ligules bright, pale or occasionally medium yellow, the outer up to 4.1 mm wide, discrete to slightly discrete, more or less upturned and more or less deeply lobed at the apex, with several to numerous, very short to longish, pale simple eglandular hairs on the lower surface and few to numerous mainly very short and short ones (–0.9 mm) projecting beyond the apex. Styles pure yellow, or slightly discoloured, or greenish-grey to very dark grey; stigmas often somewhat paler. Receptacle pits with margins dentate. Achenes 3.0–4.6 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 36. Variable in degree of dentation of leaves, colour and length of petioles and colour and diversity of simple eglandular hairs. A variant with coriaceous, sometimes subglabrous leaves and often some obvious involucral glandular hairs occurs locally in the eastern part of the Cairngorms. All appear to be tetraploid. (1) Forma eximium Styles greenish-grey or very dark grey. 2n = 36. (2) Forma tenellum (Backh. fil.) P. D. Sell & C. West H. eximium var. tenellum Backh. fil.; H. alpinum subsp. alpinum var. pumilum subvar. tenellum (Backh. fil.) Zahn Styles pure yellow to slightly discoloured. 2n = 36. Native. On cliff ledges, in rock crevices and amongst turf between 620 and 1,050 m. Widespread in the Scottish Highlands although not detected from the extreme northwestern Highlands where it appears to be replaced by H. marginatum. Frequent throughout the Cairngorms and the glens of Forfarshire; in Glen Callater and on Lochnagar in south Aberdeenshire; on Cairnwell in east Perthshire and a few hills in mid Perthshire, including Ben Lawers; in Glen Etive and Glen Coe in Argyllshire; in Glen Strathfarrar and in the Loch Laggan area in Inverness-shire. Endemic. Both forms are widespread. 104. H. pentaploideum P. D. Sell & D. J. Tennant Pentaploid Hawkweed Phyllopodous perennial herb with a branched stock. Stem 7–22 cm, rather pale green, medium to rather slender, often flexuous and markedly thickened just below the involucre, with numerous, often dense, medium and long to very long (to 5 mm), very wavy, white, black-based, simple eglandular hairs throughout, few to many, very short, fine glandular hairs below, becoming more or less numerous, very short and short (to 0.5 mm) and dark in upper part, and scattered, very short stellate hairs below, which become numerous to rather dense in upper part. Leaves medium to deepish, often

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somewhat drab (greyish-olive) green, rather glossy on upper surface, paler and often slightly glossy on lower surface, the outermost and the apices occasionally tinted with dull, deep reddish-violet; basal few to numerous, the outer with lamina 5–30 × 3–16 mm, mainly obovate or broadly elliptical, broadly rounded-obtuse, often mucronate, sometimes slightly retuse at apex, entire, denticulate or sometimes with a few small, seldom large teeth and cuneate, rounded or shortly narrowed at the base into often shortish, sometimes winged petioles, the inner with lamina 10–70 × 2–15 mm, often slightly coriaceous and mainly narrowly elliptical or sometimes oblanceolate or lanceolate, the innermost often linear-elliptical to linear, mainly acute or subacute, the middle ones of the inner group occasionally obtuse-mucronate at apex, with the margin often very undulate, sometimes twisted, especially towards the apex, subentire, denticulate or with a few shallow teeth, sometimes serrate-dentate with several acute, shallowly to very narrowly mammiform teeth, and cuneate to more or less attenuate at base into a long and slender, sometimes short and winged, occasionally reddish-violet-tinted petioles, all usually with numerous, erect, long, wavy, pale-based, white simple eglandular hairs on the upper surface, sometimes uniformly distributed but more frequently restricted towards the margins, the outermost sometimes with few, often with rather dense and very wavy ones on the margins, and with more or less uniformly scattered to very numerous, softer, usually palebased, occasionally black-based ones on the lower surface, all with scattered, very short, yellowish glandular hairs on the surfaces and on the margin and without stellate hairs, the petioles with numerous, often dense, very wavy, long to very long simple eglandular hairs on the margins; cauline 1–2(–4), the lowest semipatent, or suberect, with lamina 8– 30 × 1–7 mm, linear-oblanceolate, linear-elliptical or linear and bract-like, usually sharply acute at apex, occasionally more or less obtuse, entire or with a few, acute teeth and attenuate at base into a usually short petiole, with scattered to numerous simple eglandular hairs on the upper surface, and numerous, often black-based ones on the margin and lower surface, becoming denser and longer at the base (to 5 mm), the remainder linear, bract-like and usually more or less appressed. Capitula solitary (to 3 in cult.), 35–50 mm in diameter, occasionally with a second stem arising from the basal rosette bearing a capitulum, the involucre very dark silvery olive-grey in appearance, narrowly campanulate tapering to a narrowly rounded or narrowly subtruncate base; peduncle when present clothed as the upper stem. Involucral bracts incumbent in bud and blackishgreen; the inner unequal, the longest (11–)13–15 × 1.0–1.6 mm, linear-lanceolate, the majority sharply acute or acuminate at apex, the innermost long and finely acuminate, the outer much shorter, lax, erect and acute at apex; all very densely clothed with long to very long (to 6 mm), slender, very wavy, interwoven (woolly), white or pale silvery grey simple eglandular hairs with long, thickened blackish bases, sometimes becoming longer (to 9 mm) at the base, and very short and short (a few to 0.6 mm), fine rather pale and dark glandular hairs which are more or less completely obscured by the simple eglandular hairs, without stellate hairs but with a tuft of short, branched hairs at the apex.

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Ligules rather deep (to medium) yellow, sometimes with a faint greenish tint, the inner numerous, the outer up to 4.0(–4.3) mm wide, shallowly to very upturned, few and nearly always very discrete and often with narrow lobes at apex, with scattered to many, short to longish, pale or slightly discoloured simple eglandular hairs on the lower surface and more or less numerous, very short, short and a few medium ones projecting beyond the apex. Styles discoloured, medium to deepish olive-grey. Receptacle pits not seen. Achenes (3.7–)4.0–4.5 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 45. The only known pentaploid in the genus Hieracium (exclud. Pilosella). H. pentaploideum shares several characters with H. holosericeum, which accompanies it, and may have arisen as a cross involving it. Native. Local on Torridonian Sandstone, among turf and fine scree on rocky slopes between 600 and 700 m. Only known on the An Teallach Range in Ross-shire. Endemic. 105. H. pseudopetiolatum (Zahn) Roffey Narrow-leaved Hawkweed H. nigrescens subsp. pseudopetiolatum Zahn; H. petiolatum auct. Phyllopodous perennial herb with a branched stock. Stem 7–25(–30) cm, pale green, pale reddish-violet at base, slender, often flexuous, with few to many, delicate, slender, wavy, patent, unequal, medium to long, white to somewhat greyish simple eglandular hairs throughout, often numerous and with long, thickened, blackish bases in the upper part, scattered, very short, pale glandular hairs becoming numerous, very short and short and dark above and scattered stellate hairs often becoming dense in the upper part. Leaves drab pale green to deepish brighter green, matt or occasionally faintly glossy on the upper surface, paler on the lower surface with the midrib often tinted reddish-violet; basal few, typically 2–4, very rarely over 6, the outermost withering before flowering, the lamina of outer 8–30 × 4– 15 mm, obovate-spathulate or narrowly obovate, occasionally asymmetrical, broadly rounded-obtuse at apex, subentire and attenuate, narrowly cuneate or shortly narrowed at base, the lamina of inner 30–75 × 5–12 mm, somewhat rigid, slightly coriaceous, sometimes with conspicuously impressed veins, spathulate, narrowly elliptical or oblanceolate, obtuse or subacute, usually apiculate and markedly twisted sideways at apex, entire, sometimes slightly undulate or denticulate, rarely with larger teeth and long attenuate at base, all usually with numerous, slender, wavy, medium and long, pale-based, white simple eglandular hairs, which are confined towards and at the margin on the upper surface, sometimes with none except for a few on the margin or with uniformly distributed short to medium ones becoming long on the innermost leaves, the lower surface with scattered short to medium ones, or none, and several, very short, yellowish glandular hairs on both surfaces and the margins, occasionally with very few, stellate hairs on the midrib of either surface, the petioles up to 60 mm, very slender to winged and either lacking simple eglandular hairs or with several to numerous, medium to long ones on the margin; cauline 1–3(–6), the lowest frequently near the base of the stem, semipatent, the lamina

10–65(–90) × 2–18 mm, often similar to inner basal, oblanceolate, broadly linear or sometimes spathulate, subacute or acute, apiculate and markedly twisted at apex, entire, sometimes twisted throughout, long attenuate and often petiolate but more frequently broadly decurrent at base and sometimes semiamplexicaul, the remainder variable but usually much smaller, linear-oblanceolate to linear or bractlike, erect to semipatent, acute at apex, entire and occasionally with a few stellate hairs. Capitulum always solitary, 30–40(–45) mm in diameter, occasionally with a second stem arising from the basal rosette bearing a capitulum, the involucre usually narrowly cylindrical-campanulate, sometimes narrowly campanulate, cuneate or narrowed at base, greyish-green to very dark grey in appearance. Involucral bracts incumbent in bud, blackish-green, the inner with paler margins; inner 9–14 × 0.9–1.7 mm, linear-lanceolate, unequal, appressed or semiappressed and sharply acute, acuminate or subacute-mucronate at apex; outer much shorter, semilax, erect and acute or subacute at apex; all sparsely to rather densely clothed with patent or semiappressed, slender, delicate, wavy, pale to somewhat greyish, black-based, medium, long or occasionally very long (– 6.0 mm) simple eglandular hairs, very short or short rather pale to darkish glandular hairs, which are completely or almost completely obscured by the simple eglandular hairs, without stellate hairs and often without obvious short hairs at the apex. Ligules bright, pale yellow, often rather narrow (up to 4.2 mm), the outermost often rather few and fairly discrete, often straight, sometimes slightly upturned or slightly recurved and often deeply lobed at apex, moderately clothed with very short or short simple eglandular hairs on the lower surface, or occasionally few, which are often semiappressed or slightly discoloured and few to several extremely short ones projecting beyond the apex, or none. Styles medium greenish-grey, the stigmas frequently discoloured yellow. Receptacle pits with margins incise-dentate. Achenes 3.4– 4.2 mm, purplish-black. Flowers 7–8. Apomictic. 2n = 27. This species is uniform in the Cairngorms, but in the single locality in west Inverness-shire it has more hairy leaves. Native. Stabilised scree below cliffs or near lake margins, rocky slopes amongst turf or short Calluna, stream banks or on large boulders, infrequently on cliff ledges, between 750 and 1,060 m and always on granite in the Cairngorms. A rare and local species in the Cairngorms: in Coire Etchachan, at Loch Etchachan, on Derry Cairngorm, Cairn Toul and Beinn a’ Bhuird in Aberdeenshire, in Glen Einich in Inverness-shire; also on mica-schist in Coire Ardair on Creag Meagaidh on the north side of Loch Laggan in Inverness-shire. Endemic. 106. H. alpinum L. Alpine Hawkweed H. alpinum var. melanocephalum auct.; H. melanocephalum auct.; H. alpinum var. insigne auct. Phyllopodous perennial herb with a branched stock. Stem 4–20(–25) cm, slender to robust, often flexuous, sometimes reddish-violet-tinted at base, with many to very numerous, unequal, medium to very long (up to 5.5 mm), slender to somewhat coarse, wavy, white or greyish simple eglandular hairs with very long, dark or black bases throughout, scattered, very short, yellowish and dark glandular

39. Hieracium hairs which become mainly very short to short (up to 0.9 mm) and darker in the upper part, and moderately clothed with stellate hairs throughout which become dense in the upper part. Leaves bright deep green, very rarely paler, slightly to very glossy on the upper surface, somewhat paler beneath, sometimes with slight reddish-violet tinges mainly on the margin and beneath; basal numerous to occasionally very numerous, sometimes in a rather neat rosette, the lamina of outer 5–80 × 4–27 mm, subrotund, obovatespathulate, obovate-oblong or broadly elliptical, broadly rounded-obtuse to more or less truncate or slightly retuse or sometimes mucronate at apex, entire, sometimes denticulate or occasionally dentate and shortly narrowed to narrowly cuneate at base, the lamina of inner 10–120 × 2–20 mm, often slightly channelled, rather rigid and somewhat coriaceous, occasionally slightly undulate, obovatespathulate, spathulate, oblong-spathulate, oblanceolate or narrowly elliptical, occasionally elliptical, rounded-obtuse to subacute and sometimes apiculate at the usually downwardly twisted apex, entire, denticulate or sometimes dentate or serrate-dentate with shallowly mammiform teeth, rather rarely with larger, very irregular, narrowly mammiform teeth which are occasionally interspersed with denticulations, or very rarely sublaciniate and often gradually, sometimes shortly narrowed at base, all with numerous medium and long, slender to rather coarse, wavy or straight, subrigid, white simple eglandular hairs often with pale, sometimes discoloured or black bulbous bases on the upper surface which are usually confined towards and at the margins, or at least with fewer or often none about the midrib, the innermost often more densely and uniformly clothed with long to very long ones, at least on the margins, the outer often with few or none except sometimes on the margins, with more or less uniformly scattered, shorter, softer, wavy, slender ones, or with none on the lower surface, and with numerous, very short, yellowish glandular hairs on both surfaces and the margins, the petioles short to long, slender to somewhat winged and sparsely to densely clothed with long to very long, coarse or slender, wavy, dark-based or blackish-based simple eglandular hairs and numerous, very short, yellowish glandular hairs and faintly to strongly tinted with reddish-violet; cauline 1–3, rarely more, the lowest usually more or less patent, the lamina 5–70(–90) × 2–13 mm, often wavy, sometimes recurved in apical part, linear-oblong, narrowly oblanceolate or occasionally narrowly spathulate, obtuse or subacute at the usually twisted apex, usually entire, very occasionally denticulate and gradually narrowed to a sessile or occasionally shortly petiolate base, the second lowest also sometimes large and similar, the remainder small, linear or linear-oblong, often semipatent and obtuse or subacute at apex, very rarely filamentous. Capitulum always solitary, 20–45(–53) mm in diameter, the involucre greyishblack to greenish-grey in appearance, almost globular in bud, campanulate to broadly campanulate, rounded, sometimes broadly rounded, rarely more or less truncate at base. Involucral bracts incumbent in bud, blackish-green, usually not very numerous; inner 9–15 × 1.5–2.0(–2.5) mm, somewhat unequal, rather lax, often curved, usually lanceolatesubulate, sometimes broadly linear-lanceolate, attenuate to

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a sharply acute and sometimes incurved apex, the majority with a long, finely acuminate point; outer very lax, patent, often recurved, broadly linear-lanceolate, abruptly acute to occasionally obtuse, or often acuminate at apex, rarely subfoliaceous; all with numerous to dense, long (up to 4.5 mm), slender to somewhat coarse, very wavy, white to greyish simple eglandular hairs with long, black bases, numerous, very short (to short), dark and yellowish glandular hairs which are almost fully obscured by the simple eglandular hairs, or with several obvious ones and without stellate hairs, the inner with or without an indistinct tuft of short hairs at the apex. Ligules bright rather pale to medium yellow, the outer up to 4.1 mm wide, slightly discrete or very occasionally densely arranged, straight or slightly recurved, rarely slightly upturned at apex, usually with long, slender lobes, with few to numerous, very short or short to medium, pale simple eglandular hairs which are sometimes greyishblack towards their bases, and with few to several, short and very short ones projecting beyond the apex, or none. Styles pure pale yellow or seldom with a greenish tinge; stigmas always yellow. Margins of the receptacle pits shortly dentate. Achenes 2.8–4.0 mm, reddish-brown or reddish-black. Flowers 7–8. Apomictic. 2n = 27. This species is remarkably variable in size and very large plants with leaf-teeth have been mistaken for H. insigne. Native. Cliff ledges, crevices in sloping rocks and sometimes on rocky slopes from 600 to 1,100 m, most frequently on coarse-grained granite, more local on quartz-biotitefeldspar gneiss, quartz-biotite, quartz-porphyry or micaschist. It is widely distributed in the Cairngorms and also occurs in Stirlingshire, Perthshire, Inverness-shire, Argyllshire, Dunbartonshire, Ross-shire and Sutherland. Greenland and Iceland, Scandinavia and central Europe east to the Tatra Mountains. In Continental Europe it has mostly been recorded as a triploid, but there are diploid counts. This may be the only taxon in Great Britain of the Section Alpina which is not endemic. 107. H. holosericeum Backh. fil. Shaggy Hawkweed H. alpinum subsp. alpinum var. genuinum subvar. villosissimum forma holosericeum (Backh. fil.) Zahn Phyllopodous perennial herb with a branched stock. Stem 4–23(–37) cm, pale green often tinted reddish-violet at base, slender to robust, sometimes flexuous, usually with very numerous to dense, very long (up to 12 mm), delicate, slender, very wavy, unequal, white or occasionally greyish black-based simple eglandular hairs throughout, scattered, very short to short yellowish and dark glandular hairs, becoming numerous in the upper part and scattered stellate hairs becoming numerous but rarely dense in upper part. Leaves rather pale to medium or sometimes deepish green, more or less glossy above, sometimes tinted with reddishviolet, paler beneath; basal numerous, sometimes in a neat rosette, the lamina of outer 5–40 × 4–15 mm, obovatespathulate, subrotund or elliptical, broadly rounded-obtuse, often slightly retuse at the sometimes twisted apex, usually entire and shortly narrowed at base into frequently winged petioles, the lamina of inner 8–120(–210) × 2–15 mm, often rather rigid and somewhat coriaceous, sometimes slightly channelled (shallowly cymbiform), narrowly

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oblong-spathulate, spathulate, linear-oblong or narrowly oblanceolate, broadly rounded-obtuse or occasionally subacute at the usually twisted, sometimes mucronulate apex, usually entire, sometimes remotely denticulate, seldom with a few, shallow, more or less sinuate or undulate teeth, very rarely with larger teeth and very attenuate at base, all with the upper surface frequently without simple eglandular hairs about the midrib, but with numerous, medium to very long, coarse or slender, wavy, subrigid, white ones with pale or occasionally dark bulbous bases, towards and on the margins, the innermost often uniformly clothed, the outer sometimes with few except on the margin, with few to numerous medium to long ones on the lower surface and with numerous, very short, yellowish glandular hairs on both surfaces and on the margins, the petioles shortly winged to rather long and slender, with numerous long or very long, coarse, wavy simple eglandular hairs and very short, yellowish glandular hairs; cauline 0–2(–3), very rarely up to 11, the lowest patent or semipatent, the lamina 5–70 × 2–6 mm, often wavy, linear-oblong, linear-oblanceolate or linear, obtuse or subacute at the often twisted apex, entire and sessile, the upper often semipatent, small, linear, sometimes linear-oblong, obtuse or subacute at apex and never filamentous. Capitulum always solitary, 20–40(–47) mm in diameter, the involucre silvery greyish-green or occasionally dark grey in appearance, the hairs giving it a shaggy, bearded appearance at the base, narrowly campanulate, cuneate or cuneate-subtruncate at base. Involucral bracts incumbent in bud, blackish-green, the outer frequently pure green; inner 9–17 × 0.5–1.5 mm, semiappressed, more or less linear, linear-lanceolate, sometimes lanceolate-subulate, often much narrower than the outer, the innermost often very narrow, acute, acuminate or subacute and occasionally reddish-tinted at apex; outer broader, unequal, sometimes long (about 10 mm) and subfoliaceous, very lax, often patent, sometimes very slightly recurved, linear-oblong, usually rounded-obtuse at apex; all with dense, very long, very wavy, white or sometimes greyish, black-based simple eglandular hairs, becoming markedly longer (up to 10 mm) about the base, with numerous, very short and short fine, dark and yellowish glandular hairs which are completely obscured by the simple eglandular hairs and without stellate hairs, but the inner with or without a distinct tuft of short hairs at the apex. Ligules medium or sometimes rather pale yellow, often densely arranged, the outer not broad (less than 4 mm), barely discrete, usually upturned and often turned up or ascending towards the usually narrowly and deeply lobed, very occasionally reddish-tinged apex, the lobes sometimes partially developed and rounded, usually with numerous to dense, unequal, very short to longish, erect, wavy, pale simple eglandular hairs on the lower surface and numerous short to medium ones protruding beyond the apex. Styles pure pale yellow, very rarely with a faint greenish tinge. Receptacle pits with margins shortly dentate. Achenes 3.2–4.3 mm, purplish-black or occasionally reddish-black. Flowers 6–8. Apomictic. 2n = 27, 36. Varies mainly in size, small plants in exposed situations, large robust plants on rock ledges. Native. Exposed rocky knolls, stable scree, crevices in boulders and shelving rocks, cliff ledges and amongst turf

on rocky slopes, from 550 to 1,100 m, on granite, schist, hornblende and slate. Frequent in the western and northwestern Scottish Highlands, sporadic throughout the Cairngorms, rather local and less common elsewhere in Scotland, the Lake District and Caernarvonshire. Recorded from Scandinavia and the mountains of central Europe, but almost certainly not this species. The tetraploid count is from Strath Nethy in the Cairngorms, but this specimen requires confirmation. Endemic. Section 9. Subalpina Pugsley Subsection Hyparctica F. N. Williams Phyllopodous perennial herbs. Stems up to 60 cm, usually with numerous simple eglandular hairs, especially below and often mixed with glandular ones above, rarely with glandular hairs only. Leaves often more or less bluish-green and usually with simple eglandular hairs, often with some very short glandular hairs, the cauline 1–4, sometimes petiolate and sometimes more or less amplexicaul. Inflorescence usually with few capitula, often on long peduncles which are usually straight and variably clothed. Capitula small to large. Involucral bracts usually incumbent in bud, clothing variable from hairs being entirely eglandular to entirely glandular with every kind of mixture in between. Ligules often with hairs at tip. Styles yellow to discoloured. Receptacle pits with margins dentate, subulate-dentate or fimbriate-dentate. Achenes dark. Flowers (6–)7–8. Upland and mountainous areas on cliffs, rocky slopes and particularly by streams which sometimes carry the seed down to lower altitudes. Widespread in Scotland, rare in northern England. Frequent in mountainous areas of Continental Europe. This section is artificial: all its species almost certainly arose from species of the Section Alpina having hybridised with species of other sections while they were sexual, segregated and then became apomictic. They are usually recognisable by their few, dark capitula on straight peduncles, hairy ligule tips and few leaves on which are often found a few, very short glandular hairs. Some species like H. callistophyllum, cremnanthes, lingulatum and sinuans are near to the species of the Section Alpina itself. Others like H. aristidens, arranense, cumbriense, eustales, fissuricola, glandulidens, oenophyllum, ovaliforme, petrocharis, pseudanglicoides, pseudanglicum, sannoxense and thalassinum all seem to be small, dark-headed relatives of the Section Cerinthoidea and have been keyed out with that section as well as with the group with few cauline leaves. H. senescens may possibly have arisen from a cross between a species of the Section Alpina and a species of the Section Oreadea. All the remaining species have probably some connection with the species of the Section Hieracium and are keyed out with them, except for a few with several cauline leaves which are keyed out with the Section Vulgata. 108. H. lingulatum Backh. fil. ex Hook. & Arn. Oblong-leaved Hawkweed H. nigrescens subsp. lingulatum (Backh. ex Hook. & Arn.) Zahn; H. nigrescens var. lingulatum (Backh. ex Hook. & Arn.) Hook. fil.; H. divaricatum G. Don nom. nud.; H. saxifragum auct.

39. Hieracium Phyllopodous perennial herb with a branched stock. Stem (15–)25–40(–50 cult.), pale yellowish-green, sometimes flushed brownish-purple below, slender and flexuous to more or less robust, with numerous, medium to long, pale, sometimes dark-based simple eglandular hairs throughout and numerous stellate and occasional very short, dark glandular hairs above. Leaves yellowish to slightly bluish medium green on upper surface, paler beneath; basal few, the lamina 2–20 × 1–3 cm, oblong, narrowly ellipticaloblong, lanceolate or elliptical, obtuse-mucronate to longacute at apex, entire, distally denticulate or with a few, larger, often hooked teeth, cuneate to attenuate at base to a short to rather long, long-hairy, often winged petiole; cauline (1–)2–3(–4), the lower and sometimes median-like basal and petiolate or sessile, the upper linear, acute, entire and sessile; all with numerous short to long, pale, subrigid simple eglandular hairs on the upper surface and numerous, medium to long, pale simple eglandular hairs beneath and on the margin, and a few, minute glandular hairs on the margins. Inflorescence with 1–4 capitula, furcate-corymbose; peduncles usually rather long, with few to numerous stellate hairs, numerous medium to long, pale, dark-based simple eglandular hairs and few to numerous, very short, dark glandular hairs. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–13 × 1.0–1.5 mm, blackish-green, the inner with slightly paler margins, all linear-lanceolate, acute at apex, with numerous, medium to long, pale, dark-based simple eglandular hairs, numerous, minute, dark glandular hairs and a few stellate hairs near the base. Ligules yellow, with a few, very short simple eglandular hairs at apex. Styles yellow to discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. One of the easiest species of Hieracium to recognise with its oblong basal and cauline leaves and dark capitula. There is a tendency for the northern plants to have yellow styles and the more southerly ones discoloured, but neither the style colour nor the distribution is clear-cut. Native. Rocky beds and banks of mountain streams and less commonly on grassy banks and exposed cliff ledges from 300 to 900 m. Common in the mountains of central and northern Scotland south to Dunbartonshire. Endemic. 109. H. cremnanthes (F. Hanb.) Pugsley Cliff Hawkweed H. marshallii var. cremnanthes F. Hanb.; H. callistophyllum var. cremnanthes (F. Hanb.) F. Hanb.; H. conspurcans subsp. callistophyllum var. cremnanthes (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–30(–45) cm, pale yellowish-green, sometimes purplish at base, slender and flexuous to fairly robust, striate, with numerous, medium to long, pale, waxy simple eglandular hairs throughout, sometimes dense below, and numerous stellate and few, short, dark glandular hairs above. Leaves pale green on upper surface, even paler beneath and occasionally tinted purplish; basal usually few, the outer when present, with lamina 2–4 × 1.5–3.0 cm, elliptical, rounded-obtuse at apex and with large mammiform teeth, the remainder with lamina 5–8 × 2–4 cm, elliptical, oblong

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and oblong-elliptical to lanceolate, gradually narrowed to an obtuse or more or less acute apex, jaggedly and unequally dentate, the teeth large and mammiform or elongate-mammiform or aquiline mammiform, sometimes interspersed with smaller teeth, sometimes with long, very narrow teeth extending onto the petioles from the cuneate base, the petioles short to medium, winged and with long, pale, wavy simple eglandular hairs; cauline 1–2, the lower usually similar to the inner basal with a winged petiole, the upper bract-like; all with numerous, medium to long, pale simple eglandular hairs on both surfaces, particularly on the midrib beneath and on the margins, or glabrous above or hairy only round the edges, with a few, minute, yellowish glandular hairs. Inflorescence with 1–4 capitula, furcate; peduncles usually rather long, with numerous stellate hairs, numerous short to medium, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs. Capitula 30–40 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 6–12 × 1.0– 1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an acute apex, with dense, medium to long, pale, dark-based simple eglandular hairs and few, short glandular hairs, stellate hairs almost absent. Ligules deep yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers. 7–8. Apomictic. Native. Rocky ledges of cliffs between 800 and 900 m. Mountains of Perthshire, Forfarshire, Aberdeenshire, Inverness-shire and Argyllshire. Endemic. 110. H. sinuans F. Hanb. Wavy-leaved Hawkweed H. atratum subsp. sinuans (F. Hanb.) Zahn; H. olivaceum subsp. pseudolangwellense Zahn; H. pulmonarium auct. Phyllopodous perennial herb with a branched stock. Stem 10–40(–50) cm, pale yellowish-green, often purplish below, slender to more or less robust, striate, with few to numerous, medium to long, pale simple eglandular hairs below, few or none above, and few stellate and few to numerous, short, dark glandular hairs above. Leaves bright metallic green on upper surface, paler and often tinted purplish beneath; basal few to numerous, the lamina 2–9 × 1–3 cm, elliptical, lanceolate, oblong-lanceolate, elliptic-oblong or oblong, usually more or less rounded-obtuse at apex, rarely more or less acute, denticulate to characteristically deeply mammiform-dentate, the teeth blunt and cuneate at base to a fairly long petiole with numerous long hairs; cauline 1–2, similar to basal with a short, winged petiole or bractlike; all sometimes glabrous or nearly so on both surfaces or with numerous medium to long, pale simple eglandular hairs beneath and on the margins, and with scattered, very short, yellowish glandular hairs. Inflorescence with (1–)2–6 capitula, furcate-corymbose; peduncles long, suberect, with numerous stellate hairs, numerous, very unequal, slender, very short to short, dark glandular hairs and few, medium, dark or dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 6–12 × 1.0–1.5 mm, almost black, the inner with paler margins and rather broadly linear-lanceolate, the outer obtuse, the inner long acute, all with numerous, very

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short and short, slender, dark glandular hairs and numerous short and medium dark simple eglandular hairs, without stellate hairs. Ligules golden yellow, with very short simple eglandular hairs at apex. Styles darkly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Ravines of mountain streams and moist cliff ledges from 400 to 700 m. Perthshire, Inverness-shire, Argyllshire, Ross-shire and Barra in the Outer Hebrides. Endemic. 111. H. callistophyllum F. Hanb. Beautiful-leaved Hawkweed H. conspurcans subsp. callistophyllum (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–40(–60) cm, pale yellowish-green, often suffused brownish-purple especially below, with scattered, short to long, pale simple eglandular hairs, those in the upper part of the stem dark-based, and numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves pale yellowish-green on upper surface, paler beneath, often flushed purplish; basal often numerous, the outer with lamina 2–6 × 2–4 cm, subrotund, ovate or elliptical, one of them often balloon-like, rounded-apiculate at apex, dentate or serrate-dentate especially in the lower half, the teeth mammiform, and subtruncate at base with rather short petioles, the inner with lamina 4–7 × 0.8–3.0 cm narrowly elliptical or lanceolate, acute at apex, dentate or serrate-dentate, often deeply, the teeth mammiform or aquiline-mammiform and often narrow and upcurved, cuneate at base to fairly long, long-hairy petioles; cauline 0–1, like inner basal but smaller and shortly petiolate, or bract-like; all glabrous on upper surface or with short to medium, rather stiff, pale simple eglandular hairs on the upper surface and medium to long, pale simple eglandular hairs on the under surface and margin, apparently without minute glandular hairs. Inflorescence with 1–6 capitula, furcate-corymbose, sometimes with a long, lower branch; peduncles rather short, suberect, with dense stellate hairs, numerous, short to medium, darkbased simple eglandular hairs and few, short, dark glandular hairs. Capitula 30–50 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 6–13 × 1.0–1.5 mm, blackish-green, linear-lanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs and stellate hairs at the base, along the margins and at the apex. Ligules deep yellow, with a few, very short hairs at apex when young, becoming glabrous. Styles yellow to slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Rock-ledges and streamsides from 600 to 900 m. Inverness-shire, Argyllshire, Ross-shire and Sutherland. Endemic. 112. H. hastiforme P. D. Sell & C. West Hastate-leaved Hawkweed H. callistophyllum var. cremnanthes auct. Phyllopodous perennial herb with a branched stock. Stem 15–45 cm, pale yellowish-green, sometimes suffused reddish below, slender, flexuous, striate, with more or less

numerous, medium to long, pale, sometimes dark-based simple eglandular hairs throughout, few to fairly numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves pale green on upper surface, even paler beneath; basal few, the outer with lamina 2.5–5.0 × 1.0–2.5 cm, elliptical, obtuse-mucronulate at apex, subentire to denticulate and abruptly contracted at base to medium petioles, the inner with lamina 4–7 × 2–3 cm, lanceolate or oblonglanceolate, the basal teeth often making them appear hastate, more or less acute at apex, dentate, mostly in the lower half, the teeth mammiform or aquiline-mammiform, sometimes extending on the petiole, abruptly contracted or subtruncate at base with short to medium, long-hairy petioles; cauline 1–2, lower similar to inner basal and petiolate, the upper bract-like; all with upper surface glabrous or nearly so, with few to numerous, medium to long, pale simple eglandular hairs beneath and on the margin, and sometimes a few stellate hairs beneath and an occasional, minute glandular hair. Inflorescence with 1–3 capitula, furcate-corymbose; peduncles rather long, suberect, with numerous stellate hairs, numerous short, dark glandular hairs and few, short and medium, dark or dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, narrowed or rounded at base. Involucral bracts incumbent in bud, 5–13 × 1.0–1.5 mm, blackish-green, the inner with a paler margin, all linearlanceolate, obtuse or subacute at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs and few stellate hairs. Ligules deep yellow, glabrous or with few, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with shortly dentate margins. Achenes 3.0–3.5 mm, reddishblack. Flowers 7–8. Apomictic. This is the Ross-shire plant which was called H. callistophyllum var. cremnanthes, which differs from H. cremnanthes by its much sharper leaf-teeth. Native. Rock-ledges and streamsides up to 650 m. Argyllshire and Ross-shire. 113. H. isabellae E. S. Marshall Isabel’s Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, often reddish below, slender to robust, flexuous, striate, with numerous, long, pale simple eglandular hairs at the base and few, shorter ones for the rest of the stem, and numerous stellate hairs and some short, dark glandular hairs above. Leaves medium green on upper surface, paler and sometimes tinted purplish beneath; basal few, the outer with lamina 3–7 × 2.0–3.5 cm, lanceolate, ovate or ovate-lanceolate, obtuse-mucronate at apex, often broadly rounded, dentate, sometimes deeply so, the teeth often mammiform, and cuneate at base to short petioles, the inner with lamina 6–10 × 2–3 cm, lanceolate or oblong-lanceolate, acute or acuminate at apex, sharply denticulate to serrate, the teeth narrow and sharp, cuneate or attenuate at base to a long, long-hairy petiole; cauline 1–2, the lower similar to inner basal but often narrower, with a long acuminate apex and with a short, winged petiole, or bract-like; all with numerous, short to long hairs on both surfaces and the margins and a few, scattered, minute, yellowish glandular hairs. Inflorescence with 2–6 capitula, furcatecorymbose; peduncles fairly long, suberect, with numerous to dense stellate hairs, numerous, short, dark glandular hairs

39. Hieracium and few to numerous, short or medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, narrowed at base. Involucral bracts incumbent in bud, 5–14 × 1.0–1.3 mm, dark olive-green, the inner with paler margins, all narrowly linear-lanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous short, dark glandular hairs, and a few stellate hairs towards the base. Ligules deep yellow, glabrous tipped. Styles discoloured. Receptacle pits with margins more or less fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. Frequent by alpine and subalpine streams between 600 and 800 m on gneiss and granite, often in shady situations. Perthshire and Inverness-shire in the mountains of central Scotland. Endemic. Named after Fanny Isabel Marshall, wife of the author of the species. 114. H. senescens Backh. fil. Tufted-hair Hawkweed H. senescens var. brevifurcatum Pugsley; H. nigrescens subsp. chrysanthum var. senescens (Backh. fil.) Nyman; H. nigrescens var. senescens (Backh. fil.) Hook. fil. Phyllopodous perennial herb with a branched stock. Stem 15–30(–45) cm, pale yellowish-green, often brownishpurple towards the base, slender to fairly robust, striate, with few to numerous, long, pale simple eglandular hairs near the base, upwards becoming fewer and shorter or almost absent, and numerous stellate hairs and short, slender, dark glandular hairs in the upper part. Leaves medium bright green on upper surface, paler and sometimes tinged brownish-purple beneath; basal often numerous, the lamina 3–10 × 1–3 cm, elliptical, elliptic-oblong, lanceolate or oblong-lanceolate, obtuse to acute at apex, entire to denticulate with a few larger teeth, or in larger plants mammiformdentate, cuneate at base to fairly long, long-hairy petioles; cauline 1–3, the lower like the basal and petiolate, the upper bract-like; all with numerous, short to medium, rather stiff, pale simple eglandular hairs on the upper surface, with numerous, medium to long, pale simple eglandular hairs beneath and on the margin, with a few minute glandular hairs, sometimes with some stellate hairs on the underside of the cauline leaves. Inflorescence with 1–6 capitula, furcate-corymbose; peduncles usually rather long, with numerous stellate hairs, numerous, very unequal, very short and short, dark glandular hairs and few to numerous, short to medium, dark or dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.0–1.5 mm, blackish, the inner with paler margins, all linear-lanceolate, gradually narrowed to a more or less acute apex, with a dense tuft of hairs at apex especially in bud, numerous, very unequal, very short and short, dark glandular hairs and few to fairly numerous, short to medium, dark or dark-based simple eglandular hairs. Ligules golden yellow, usually with very short simple eglandular hairs at apex. Styles usually yellow, sometimes slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. On grassy mountain ledges, in rock crevices and by streamsides, usually on mica-schist, rarely on granite, mainly between 800 and 900 m. Widespread in the Scottish mountains south to Dunbartonshire. Recorded from the

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Mourne Mountains in Co. Down, but the specimens have not been seen. Endemic. 115. H. molybdochroum Omang Mountain Hawkweed H. nigrescens var. commutatum Lindeb.; H. nigrescens subsp. commutatum (Lindeb.) Lindeb.; H. commutatum (Lindeb.) Elfstr., non J. Gay ex Monnier; H. tumescens subsp. molybdochroum Dahlst. nom. illegit.; H. atratum subsp. molybdochroum Zahn nom. illegit. Phyllopodous perennial herb with a branched stock. Stem 15–45 cm, pale yellowish-green, sometimes tinted purplish below, slender to more or less robust, striate, with few, long, pale simple eglandular hairs especially below and dense stellate and few, short, dark glandular hairs above. Leaves bright green on upper surface, paler beneath; basal few to fairly numerous, the outer with lamina 2–7 × 1.5– 3.0 cm, elliptical, elliptic-lanceolate or ovate-lanceolate, rounded-mucronulate at apex, denticulate to undulatedentate especially in the lower half, the teeth more or less mammiform and attenuate at base to rather long, slender petioles, the inner with lamina 5–10 × 1–4 cm, narrowly to broadly elliptical, lanceolate or ovate-lanceolate, gradually narrowed to an acute apex, deeply and irregularly dentate to incise-dentate, the teeth mammiform, narrowly mammiform or aquiline-mammiform, sometimes unequal, sometimes interspersed with denticulations, attenuate at base, the teeth sometimes descending the long, slender, long-hairy petioles; cauline 1–3, the lower similar to the inner basal with a short to long, sometimes winged petiole; all with few to numerous, short to medium, pale, rather stiff simple eglandular hairs on the upper surface, numerous, short to long simple eglandular hairs beneath and on the margin, stellate hairs sometimes present on the under surface of the cauline, and scattered, minute, yellowish glandular hairs. Inflorescence with 1–5 capitula, furcate-corymbose; peduncles often long, suberect, with dense stellate hairs, few to numerous, short, dark glandular hairs and few to numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 30–35 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.0–1.5 mm, blackish-green, linear-lanceolate, gradually narrowed to a narrow, acute apex, with numerous, short to long, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs, some stellate hairs towards the base and a tuft of hairs at the apex. Ligules yellow, with very short simple eglandular hairs at the apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish black. Flowers 7–9. Apomictic. Native. Cliff ledges up to over 1,000 m at least sometimes on schist. Dubh Lochan on Beinn a’ Bhuird, near the Black Spout on Lochnagar, both in Aberdeenshire; Coire Kander, Coire Garbhlach in Glen Feshie and Sgoran Dubh in Inverness-shire, all localities in the Cairngorms. Sweden. 116. H. laetificum P. D. Sell & C. West Black-headed Hawkweed H. nigrisquamum P. D. Sell & C. West (1967), non Hyl. (1943) Phyllopodous perennial herb with a branched stock. Stem 35–55 cm, pale yellowish-green, sometimes slightly

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purplish towards the base, more or less robust, striate, with rather sparse, medium to long, pale simple eglandular hairs, those above dark-based, and few stellate and short, dark glandular hairs above. Leaves yellowish-green on upper surface, paler beneath; basal few, the outer with lamina 3– 10 × 2.5–4.5 cm, elliptical to broadly elliptical, roundedmucronulate at apex, denticulate or shortly dentate and cuneate at base to rather long petioles, the inner with lamina 5–10 × 2–4 cm, elliptical, lanceolate or ovate, acute at apex, denticulate or dentate, the teeth shallowly mammiform or sometimes larger and narrower and cuneate at base to more or less long, long-hairy petioles; cauline 0–2, lower lanceolate, long-acute or acuminate at apex, dentate, the teeth in the lower half often long, narrow and curved upwards, with a short, winged petiole; all with few, short or medium, pale simple eglandular hairs on upper surface or glabrous, numerous, short to long, pale simple eglandular hairs on lower surface and the margins, few, minute, yellowish glandular hairs scattered over the surfaces and sometimes a few stellate hairs beneath. Inflorescence with 2–8 capitula, furcate-corymbose; peduncles long, suberect, with numerous to dense stellate hairs, numerous short and medium, dark glandular hairs and few, short to medium, pale, darkbased simple eglandular hairs. Capitula 45–52 mm in diameter, with broad, rounded bases. Involucral bracts porrect or incumbent in bud, 5–14 × 1.0–1.2 mm, blackish, the inner with pale margins, all linear-lanceolate, drawn out to a very narrow, acute apex, with dense, very short to medium, very unequal dark glandular hairs, numerous, short to long, pale, dark-based simple eglandular hairs, and very few stellate hairs near the base. Ligules pale yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 4.0– 4.2 mm, black. Flowers 7–8. Apomictic. Native. Cliff ledges, scree slopes and riverside rocks. Mountainous areas of central and northern Scotland and Stirlingshire. Endemic. 117. H. marshallii E. F. Linton Marshall’s Hawkweed H. chrysanthum var. marshallii (E. F. Linton) F. N. Williams; H. senescens subsp. marshallii (E. F. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–40(–45) cm, pale yellowish-green, often flushed brownish-purple towards the base, slender to more or less robust, striate, with medium to long, pale, often darkbased simple eglandular hairs, dense at the base and few to numerous above and numerous stellate hairs above. Leaves yellowish-green on upper surface, paler beneath, rarely slightly flushed purplish; basal few to fairly numerous, the outer with lamina 3–8 × 2–3 cm, subrotund or elliptical, obtuse-mucronate at apex, denticulate to shortly and broadly dentate, and cuneate or rounded at base to fairly long, densely long-hairy petioles, the inner with lamina 6–10 × 2–3 cm, elliptical, lanceolate, oblong-lanceolate, ovate-lanceolate or ovate, gradually narrowed to a more or less acute apex, denticulate or dentate in the lower half, the teeth narrowly mammiform, cuneate at base to long, long-hairy petioles; cauline 1–2, the lower like the inner basal with a short, winged petiole, the upper linear or

bract-like, entire and sessile; all with few to numerous, short to medium, pale, rather stiff simple eglandular hairs on the upper surface, and numerous, medium to long, pale, sometimes bulbous-based hairs beneath and on the margins, and a few, minute, yellowish glandular hairs. Inflorescence with 1–3(–5 cult.) capitula, furcate-corymbose; peduncles usually long, suberect, with numerous to dense stellate hairs, few, minute, dark glandular hairs and few to numerous, short to long, pale, dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, broad and subtruncate at base. Involucral bracts incumbent to porrect in bud, 6–15 × 1.2– 1.8 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with dense, medium to long, pale, dark-based simple eglandular hairs intermixed with a few, very short, dark glandular hairs and sometimes a tuft of hairs at apex. Ligules bright golden yellow, with short simple eglandular hairs at apex. Styles yellowish to discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Moist rocky ledges of cliffs and shallow ravines of alpine streams. Perthshire, Forfarshire, Aberdeenshire, Banffshire, Inverness-shire, Argyllshire and Mull. Endemic. Named after Edward Shearburn Marshall (1858– 1919). 118. H. anfractiforme E. S. Marshall Jagged-toothed Hawkweed H. subanfractum E. S. Marshall nom. illegit.; H. caesium subsp. neomarshallianum Zahn; H. neomarshallianum (Zahn) Roffey nom. illegit. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, often reddish towards the base, slender and flexuous to more or less robust, striate, more or less glabrous or with a few, long, pale simple eglandular hairs at the base and a few stellate hairs and an occasional, short, dark glandular hair above. Leaves bright green on upper surface, paler and sometimes tinted reddish beneath; basal few to numerous, the outer with lamina 4–9 × 1.5–3.0 cm, oblanceolate, oblong-oblanceolate or elliptical, rounded-mucronulate at apex, dentate or serratedentate and attenuate at base to fairly long petioles, the inner with lamina 6–10 × 1.5–3.0 cm, narrowly elliptical or lanceolate, acute at apex, dentate to incise-dentate, the teeth unequal and narrowly mammiform, sometimes cusped, often interspersed with denticulations, narrowed at base to rather long, slender, long-hairy petioles on which the teeth sometimes descend and are long-linear; cauline 0–1, when present often similar to inner basal but smaller and petiolate, sometimes bract-like; all glabrous on upper surface, with scattered, pale, medium to long simple eglandular hairs beneath, especially on the midrib and margins, and scattered, minute, yellowish glandular hairs. Inflorescence with 1–8 capitula, furcate-corymbose; peduncles rather long, suberect, with numerous to dense stellate hairs, few to numerous, short, dark glandular hairs and few to numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0– 1.3 mm, blackish-green, all linear-lanceolate, obtuse at

39. Hieracium apex, with numerous short to medium, pale, dark-based, simple eglandular hairs, fewer, short, dark glandular hairs and numerous stellate hairs on the margins and at the base. Ligules yellow, usually with some very short simple eglandular hairs at the apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 7–8. Apomictic. Native. Cliff ledges, rocky slopes and particularly streamside rocks, perhaps mostly on granite and mica-slate, between 500 and 900 m. Cruach Ardrain (Corrie Ardran), Killin and Ben Laoigh in Perthshire; Fersit Forest, Roy Bridge and Beinn a’ Chlianaig (Ben Chlinaig) in Invernessshire; frequent in Argyllshire; and Coire an t-Slugain and Meall a’ Chrasgaidh in Ross-shire. Endemic. 119. H. vorlichense P. D. Sell Vorlich Hawkweed Phyllopodous perennial herb with a branched stock. Stem 40–50 cm, pale yellowish-green, sometimes slightly tinted brownish-red below, slender, striate, with few to numerous, medium to long, pale simple eglandular hairs in the lower half and occasional ones above and numerous stellate hairs and few to numerous, short, dark glandular hairs above. Leaves bright medium green on upper surface, paler beneath and sometimes reddish tinted; basal few to numerous, the outer with lamina 2.0–5.5 × 1.5–2.5 cm, elliptical or ovate, rounded-mucronulate at apex, dentate in the lower half with mammiform teeth and abruptly contracted at base, the inner with lamina narrowly elliptical, oblong, lanceolate or oblong-lanceolate, acute to long-acute at apex, entire or denticulate in the upper half, dentate in the lower half, the teeth mammiform or narrowly mammiform, and attenuate at base to a fairly long, slender, long-hairy petiole, the teeth sometimes extending on the petiole; cauline 0–2, the lower when present like the inner basal and shortly petiolate, the upper bract-like; all with numerous, short to long, pale simple eglandular hairs on both surfaces and the margin and occasional minute glandular hairs. Inflorescence with 2–3 capitula, furcate-corymbose; peduncles suberect, with numerous stellate hairs, numerous unequal, very short to medium, slender, dark glandular hairs and occasional, medium, pale, dark-based simple eglandular hairs. Capitula 35–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–13 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually drawn out to a narrow, acute apex, the inner almost filamentous, all with numerous very short and short, unequal, slender, dark glandular hairs, numerous, medium to rather long, pale, dark-based simple eglandular hairs and fairly numerous stellate hairs on the margins and towards the base. Ligules yellow, with numerous, very short simple eglandular hairs on the tips. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 7–8. Apomictic. Formerly included in H. anfractiforme which it resembles in general appearance, but differs in its more hairy, slightly differently toothed leaves and longer, narrower, acute involucral bracts. Native. Known only from rocks by a stream at about 660 m on the east side of Ben Vorlich in Dunbartonshire. Endemic.

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120. H. centripetale F. Hanb. Velvet-headed Hawkweed H. atratum subsp. centripetale (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, rarely slightly reddish below, slender to robust, striate, with scattered, medium to long, pale simple eglandular hairs below and shorter, darkbased ones above, and numerous stellate and few, short glandular hairs above. Leaves clear apple green to dark green on upper surface, paler beneath; basal numerous, the outer with lamina 2–7 × 1.5–3.0 cm, elliptical, obtuse-mucronate at apex, entire, denticulate or sinuate-dentate and subtruncate or abruptly contracted to short, densely long-hairy petioles, the inner with lamina elliptical, lanceolate, oblonglanceolate, oblong-elliptical or oblong, acute to acuminate at apex, undulate-denticulate to sinuate-dentate the teeth mammiliform and cuneate at base; cauline 0–1, lanceolate, acuminate at apex, serrate, the teeth small; all glabrous or with few to numerous, short and medium simple eglandular hairs on the upper surface, numerous medium to long, pale simple eglandular hairs on the lower surface and with scattered, minute glandular hairs. Inflorescence with 3–15 capitula, furcate-corymbose, usually with a long lower branch; peduncles rather short, with dense stellate hairs, with numerous, short, dark glandular hairs and few, short to medium, pale, dark-based simple eglandular hairs. Capitula 25–35 mm in diameter, narrowed at base. Involucral bracts porrect in bud, 4–13 × 0.8–1.0 mm, blackish-green, the inner with pale margins, all narrowly linear-lanceolate, gradually narrowed to an acute apex, with numerous, very short to short, unequal, dark glandular hairs, numerous, short and medium, pale, dark-based or dark simple eglandular hairs and occasional stellate hairs along the margin. Ligules deep yellow, with numerous short hairs at apex. Styles discoloured. Receptacle pits with margins incisedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–8. Apomictic. This species seems to be morphologically close to H. duriceps F. Hanb. in the Section Hieracium, but the furcate inflorescence and glandular hairs on the leaves suggest it is better placed in the Section Subalpina. Native. Cliff ledges and streamsides often between 350 and 460 m. Dumfries-shire, Peebles-shire, Perthshire, Inverness-shire, Isle of Arran and Kintyre. Endemic. 121. H. breacense P. D. Sell Breac Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, slightly reddish at base, slender to robust, striate, with few to fairly numerous, medium to long, pale simple eglandular hairs throughout and numerous stellate hairs and few to numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal few to numerous, the lamina 3–10 × 1.0–3.5 cm, elliptical or elliptical-oblong, rarely subrotund, obtuse-mucronate to acute at apex, regularly dentate, the teeth mammiform or aquiline-mammiform, cuneate at base, the petioles up to 7 cm, with dense, long, pale simple eglandular hairs; cauline 1(–3), linear-lanceolate, long-acute at apex, sharply dentate and shortly petiolate, when more than 1 bract-like;

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all glabrous or nearly so on upper surface, with numerous, short or medium, pale simple eglandular hairs beneath and on the margin. Inflorescence with 1–11 capitula, furcatecorymbose; peduncles medium to short, suberect, with dense stellate hairs, numerous, very short and short, dark glandular hairs and a very occasional pale simple eglandular hair. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–14 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to a long, slender, acute apex, with numerous, very short, short and medium, slender, dark glandular hairs, numerous short and medium pale dark-based simple eglandular hairs, and stellate hairs along the margin and at the apex. Ligules deep yellow, with fairly numerous, very short hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 7. Apomictic. Very close to H. centripetale but distinguished by its greyish, not black capitula, more stellate hairs on the involucral bracts and larger leaves with mostly larger teeth. Native. Known only from Meall Breac in Ross-shire. Endemic. 122. H. kingshousense P. D. Sell Kingshouse Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often purplish towards the base, slender and flexuous, striate, with more or less numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and few to numerous stellate hairs and sometimes an occasional short, dark glandular hair above. Leaves medium yellowish-green on upper surface, paler beneath; basal with lamina 4–12 × 2.5–4.0 cm, elliptical, broadly elliptical or ovate-lanceolate, acute to longacuminate at apex, rarely obtuse, denticulate to shallowly dentate, 1 leaf often with longer, aquiline-mammiform teeth, abruptly contracted or cuneate at base, the petioles always long, up to 8 cm and long-hairy; cauline 0–1, lanceolate, long-acuminate at apex, sharply dentate, cuneate at base and petiolate; all with more or less numerous, short and medium simple eglandular hairs on both surfaces and the margins. Inflorescence with 4–7 capitula, furcate-corymbose; peduncles short to medium, with dense stellate hairs, numerous, short and medium, dark-based simple eglandular hairs and short, dark glandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.2–1.5 mm, blackishgreen, inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous to dense, short to medium, dark or dark-based simple eglandular hairs, few to numerous, short or very short, dark glandular hairs and numerous stellate hairs along the margins. Ligules deep yellow, glabrous or with a few very short hairs at tip. Styles yellowish, becoming discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7. Apomictic. Native. Known only from three more or less identical gatherings from and by streams in the vicinity of Kingshouse in Argyllshire. Endemic.

123. H. nigrifactum P. D. Sell Dusky-headed Hawkweed H. nigrescens auct.; H. subnigrescens auct.; H. atratum subsp. subnigrescens auct.; H. gracilifolium auct. Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, often tinted pink or purple at base, slender and rather flexuous, striate, with few to fairly numerous, medium to long, pale simple eglandular hairs in the lower part, becoming fewer and shorter upwards, with numerous stellate hairs and few, very short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal few to numerous, the outer with lamina 2–7 × 1.5–3.5 cm, subrotund, elliptical or broadly elliptical, obtuse-mucronate at apex, subentire, denticulate or shallowly dentate, the teeth mamillate and cuneate or subtruncate at base, the inner with lamina 3–8(–10) × 1–3 cm, lanceolate or narrowly elliptical, more or less pointed at apex, dentate especially in the lower half, the teeth mammiform or aquiline-mammiform, cuneate or abruptly contracted at the often asymmetrical base, the petioles medium to long, with dense, long, pale simple eglandular hairs; cauline 0–1, when present lanceolate or rarely linear, long-acute at apex, usually sharply toothed at base, the teeth aquiline-mammiform with a long-cuspidate apex, sessile or petiolate; all with short to medium, pale simple eglandular hairs on both surfaces and the margin and with a few, very short, yellowish glandular hairs. Inflorescence with 1–6 capitula, furcate-corymbose; peduncles short to long, with more or less dense stellate hairs, numerous very short and short, dark glandular hairs and very occasionally a medium simple eglandular hair. Capitula 25–30 mm in diameter, rounded-based. Involucral bracts incumbent in bud, 11–13 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous to dense, very short to medium, dark glandular hairs, a few stellate hairs along the margin, and very rarely an occasional simple eglandular hair. Ligules deep yellow, with a few, very short hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 6–8. Apomictic. Native. Rocky streamsides and slopes from 600 to 1,000 m. Perthshire, Argyllshire and Inverness-shire and Ben Vorlich in Dunbartonshire. Endemic. 124. H. gracilifolium (F. Hanb.) Pugsley Slender-leaved Hawkweed H. nigrescens var. gracilifolium F. Hanb.; H. atratum subsp. subnigrescens var. gracilifolium (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–50 cm, pale yellowish-green, often tinted purplish-red towards the base, slender to robust, striate, with numerous, long, pale simple eglandular hairs throughout or becoming fewer and shorter and dark-based above, with numerous stellate hairs and few to numerous, short, dark glandular hairs in the upper part. Leaves pale green, sometimes slightly bluish on upper surface, paler beneath; basal few, the outer with lamina 2–6 × 1.5–3.0 cm, elliptical, obtusemucronate at apex, subentire, denticulate or shallowly dentate and cuneate at base to a fairly long petiole, the inner

39. Hieracium with lamina 6–10(–12) × 1.0–3.5 cm, narrowly elliptical to elliptical or lanceolate, more or less acute at apex, denticulate to dentate especially in the lower half, cuneate to attenuate at base and with fairly long petioles; cauline 1–2, the lower with lamina 3–10 × 1–2 cm, lanceolate or linear, acute at apex, entire or with teeth at the cuneate base and sessile or shortly petiolate, the upper always bractlike, sometimes both bract-like, or absent; all with short to medium, pale simple eglandular hairs on both surfaces and the margins and scattered, very short, yellowish glandular hairs. Inflorescence with 1–5 capitula, furcate-corymbose; peduncles long, suberect, with dense stellate hairs, numerous very short and short, unequal, dark glandular hairs and a few, medium, dark-based simple eglandular hairs. Capitula 20–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, very short and short, dark glandular hairs and several, medium, dark-based simple eglandular hairs. Ligules yellow, with few, very short hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. Streamside rocks from 500 to 900 m. Frequent in Perthshire, Feith am Laoigh in Banffshire, Aonach Beg in Inverness-shire and Allt nan Giubhas in Argyllshire. When originally describing the plant Hanbury said it was found throughout the Breadalbane Range, but it has not recently been much collected. Endemic. H. W. Pugsley (1948) put together H. gracilifolium, plants called H. atratum, now named H. atriglandulosum, and plants called H. nigrescens, now called H. nigrifactum. 125. H. atriglandulosum P. D. Sell Black-glandular Hawkweed H. atratum auct.; H. gracilifolium auct. Phyllopodous perennial herb with a branched stock. Stem 25–40 cm, pale yellowish-green, often reddish or purplish below, rather slender and flexuous, striate, with numerous to dense, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and more or less dense stellate hairs and few to numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal with lamina 4–9 × 2–4 cm, narrowly to broadly elliptical, sometimes ovate, the outer obtuse, the inner acute at apex, subentire, denticulate or very shallowly dentate, the teeth mamillate, and more or less cuneate at base, the petioles usually long and with dense, long, pale simple eglandular hairs; cauline 0– 1, when present lanceolate, acute at apex, shortly toothed, cuneate at base and petiolate; all with numerous, short to medium, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 2–6(–9 cult.) capitula, furcate-corymbose; peduncles long, suberect, with numerous to dense stellate hairs, numerous, very short, short and medium, dark glandular hairs and rarely a solitary eglandular hair. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.0–1.4 mm, blackish-green, the inner with paler margins, all linearlanceolate, more or less acute at apex, with numerous to

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dense, very short, short and medium, slender, dark glandular hairs, without or with very occasional simple eglandular hairs and a few stellate hairs on the margin and at the apex. Ligules deep yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Rock-ledges and streamsides between 600 and 950 m. A few records in Perthshire, Aberdeenshire, Inverness-shire and Argyllshire. 126. H. dissimile (Lindeb.) T. Durand & B. D. Jacks. Scandinavian Hawkweed H. murorum var. dissimile Lindeb.; H. furculatum subsp. dissimile (Lindeb.) Elfstr. nom. illegit.; H. caesium subsp. dissimile (Lindeb.) Zahn nom. illegit.; H. porrigens subsp. floccifrons Elfstr. Phyllopodous perennial herb with a branched stock. Stem 20–45 cm, pale yellowish-green, sometimes tinted reddish at base, slender, flexuous, striate, with a few, medium to long, pale simple eglandular hairs mainly in the lower half and few stellate and very short, dark glandular hairs in the upper part. Leaves yellowish-green, paler beneath; basal few, with lamina 5–8 × 2.5–3.0 cm, lanceolate or ovate-lanceolate, acute at apex, sharply serrate, the teeth long, narrow, acute and curved upwards, cuneate at base to long, long-hairy petioles; cauline 0–1, similar to the basal and long petiolate; all glabrous or nearly so on upper surface, with more or less numerous, pale simple eglandular hairs beneath and on the margins and some stellate hairs beneath the cauline. Inflorescence with 3–5 capitula; peduncles rather long and spreading, with numerous to dense stellate hairs and few to fairly numerous, very short, dark glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous short, dark glandular hairs, and a few stellate hairs on the margins. Ligules yellow, with numerous, very short simple eglandular hairs on the tips. Styles discoloured. Receptacle pits with subulate-dentate margins. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–8. Apomictic. Native. Recorded only from Loch an Dughaill in Kintyre and Beinn Liath Mor Fannaich in Ross-shire. Sweden. 127. H. cuspidens P. D. Sell & C. West Cusped-toothed Hawkweed H. dissimile var. majus Pugsley Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, yellowish-green, often tinted reddish in lower half, slender to robust, flexuous, striate, with more or less numerous, medium to long, pale simple eglandular hairs, and fairly numerous, short, dark glandular hairs and stellate hairs in the upper part. Leaves yellowish-green on upper surface, paler beneath and sometimes tinted purplish; basal few, with lamina 4–14 × 1.5–6.0 cm, lanceolate, elliptical, ovate-lanceolate, or oblong-lanceolate, more or less acute at apex, deeply and irregularly dentate or

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incise-dentate, the teeth mammiform or long and narrow, aquiline-mammiform and very acute, sometimes interspersed with denticulations, cuneate at base to long, longhairy petioles on which the teeth sometimes occur; cauline 0–1, up to 10 × 5 cm, very like basal, apex sometimes acuminate, petiolate; all glabrous or with scattered, short to medium simple eglandular hairs on the upper surface, more or less numerous, short to medium, pale simple eglandular hairs beneath and on the margin and a few minute glandular hairs. Inflorescence with 1–5(–9) capitula, furcatecorymbose; peduncles long, suberect, with numerous to dense stellate hairs, numerous, short and medium, pale, dark-based simple eglandular hairs and more or less numerous, short, dark glandular hairs. Capitula 30–40 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 5–13 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous stellate hairs on the margins and apex, more or less numerous, short and medium, pale, dark-based simple eglandular hairs and more or less numerous, very short, dark glandular hairs. Ligules deep yellow, with minute hairs at apex. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Rock-ledges and streamsides. Frequent in Perthshire and also in Argyllshire. Endemic. 128. H. oxyodus (W. R. Linton) W. R. Linton Moffat Hawkweed H. longilobum (Dahlst. ex Zahn) Roffey; H. stenophyes var. oxyodus W. R. Linton; H. murorum subsp. longilobum Dahlst. ex Zahn Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, often suffused purplish towards the base, slender, striate, without or with a few, scattered simple eglandular hairs, and few stellate and short, dark glandular hairs in the upper part. Leaves bright medium green on upper surface, paler and sometimes tinted purplish beneath; basal few to fairly numerous, with lamina 2–10 × 1–3 cm, lanceolate or elliptical, obtuse to acute or acuminate at apex, denticulate to irregularly dentate or incise-dentate, the teeth often long, narrow, acute and curved upwards, even more deeply incised in cultivated plants, cuneate at base to rather short, sparsely hairy petioles; cauline 1–2, similar to basal but with long-acuminate apex and with a short petiole or sessile; all with a few, medium simple eglandular hairs on the lower surface and margin and quite a number of minute, yellowish glandular hairs. Inflorescence with 2–10(–15 cult.) capitula, furcatecorymbose; peduncles rather long, suberect with dense stellate hairs and few to numerous, fine, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–11 × 1.0–1.3 mm, blackish green, inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, unequal, very short and short, dark glandular hairs, a few, medium, dark simple eglandular hairs and a few, scattered stellate hairs. Ligules yellow, with very short, simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–8. Apomictic.

Native. Known only from south-facing cliffs by the Midlaw Burn near Moffat in Dumfries-shire. Originally found in Herjedalen and Jemtland in Sweden. 129. H. vennicontium Pugsley Vennicontes’s Hawkweed H. euprepes var. glabratum E. F. Linton, non H. glabratum Hoppe ex Willd.; H. duplicatum auct.; H. sagittatum subsp. euprepes var. glabratum (E. F. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 25–60 cm, pale yellowish-green, often suffused reddish at the base, slender to more or less robust, often flexuous, striate, with numerous, long, pale simple eglandular hairs towards the base and glabrous or with occasional ones upwards, and few to numerous stellate hairs and occasional, short, dark glandular hairs in the upper part. Leaves yellowish or bright green on upper surface, paler and slightly bluish beneath; sometimes tinted reddish, basal few, the outer with lamina 3–8 × 1.5–3.0 cm, narrowly elliptical to elliptical, obtuse-mucronate at apex, denticulate or shallowly dentate and asymmetrically cuneate at base to a long petiole, the inner with lamina 4–14 cm, lanceolate, ovate-lanceolate or oblong-lanceolate, acute to long-acuminate at apex, dentate, the teeth shallowly mammiform, cuneate, asymmetrically dentate at base to very long, long-hairy petioles; cauline 1–2(–3), the lower like the inner basal and petiolate; all glabrous or with a few, short to medium, pale, scattered simple eglandular hairs on both surfaces, and few to numerous, short to medium ones on the margins, sometimes an occasional minute, yellowish glandular hair. Inflorescence with 2–12 capitula, laxly furcate-corymbose, sometimes with a long lower branch; peduncles medium to long, suberect, with numerous to dense stellate hairs, numerous, dark or dark-based, short to medium simple eglandular hairs and few to numerous, very short or short, dark glandular hairs. Capitula 30–40 mm in diameter, narrowed towards the base. Involucral bracts incumbent in bud, 6–13 × 1.5– 2.0 mm, blackish-green, the inner with paler margins, all broadly linear-lanceolate, obtuse at apex, with numerous, short and medium, dark simple eglandular hairs, numerous, short, dark glandular hairs, and stellate hairs at the base and apex. Ligules golden yellow, glabrous or with a few, very short simple eglandular hairs at apex. Styles yellow to slightly discoloured. Receptacle pits with margins subulatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. Mainly on grassy and rocky streamsides in the mountains up to 850 m, but recorded from river shingle and in Kintyre on rocks by the shore. Yorkshire, Cumberland and the mountains of central Scotland. Endemic. Named after the Vennicontes tribe. 130. H. westii P. D. Sell West’s Hawkweed H. submurorum auct. Phyllopodous perennial herb with a branched stock. Stem 28–60 cm, pale yellowish-green, slender, striate, with scattered stellate hairs mainly in the upper part, few short to medium, pale, sometimes dark-based simple eglandular hairs throughout, few yellowish, minute glandular hairs throughout and few short, dark glandular hairs in the upper part. Leaves bluish-green, sometimes slightly purple

39. Hieracium tinted on the upper surface, paler beneath; basal few, with lamina 2–10 × 1.5–5.0 cm, ovate, lanceolate or elliptical, rounded-obtuse to acute at apex, regularly dentate, the teeth often more or less mammiform, usually asymmetrical and cuneate, but sometimes truncate at base, with short or medium, long-hairy petioles; cauline 1–2(–3), the lower lanceolate, narrowly acute or acuminate at apex, dentate, the teeth usually rather narrow and often cusped, with short or medium petioles or sessile and semiamplexicaul, the upper usually bract-like; all with numerous, short and medium, pale simple eglandular hairs on both surfaces and the margins, a few, longer ones on the midrib beneath, and scattered, very short, yellowish glandular hairs throughout. Inflorescence with 2–10 capitula, furcatecorymbose; peduncles straight and suberect, the acladium sometimes very short, with numerous to dense stellate hairs, numerous short and very short, dark glandular hairs and few to numerous, short and medium, dark simple eglandular hairs. Capitula 10–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–13 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, long tapering to a narrow, more or less acute apex, the inner almost filamentous, with numerous, unequal, very short, short and medium, dark glandular hairs, few or numerous, short or medium, dark simple eglandular hairs, and sometimes a few stellate hairs towards the base. Ligules golden to nearly orange-yellow, rather shallowly dentate, with numerous, very short, pale simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, dark. Flowers 7–8. Apomictic. Native. Apparently recorded from at least two localities in the mountains around Kingshouse in Argyllshire, but now known only from the side of the rocky stream of Allt nan Giubhas. Endemic. Named after Cyril West (1887–1986). 131. H. ochthophilum P. D. Sell Streamside Hawkweed Phyllopodous perennial herb with a branched stock. Stem 24–37 cm, pale yellowish-green, sometimes slightly purplish at base, slender, flexuous, striate, with few, scattered, short or medium, pale, dark-based simple eglandular hairs and few to numerous stellate hairs above. Leaves pale greyish-green on upper surface, paler beneath; basal normally few, with lamina 2–6 × 1.5–3.0 cm, elliptical, lanceolate, ovate or subrotund, obtuse-mucronate to more or less acute at apex, subentire or more usually with shallow, spreading teeth in the lower half, cuneate at base to a medium, slender, glabrous to fairly hairy petioles; cauline 0–1, when present with lamina lanceolate, acute at apex, dentate in lower half, petiolate; all glabrous or nearly so on upper surface, with few to numerous, medium to long, pale simple eglandular hairs beneath and on the margin, and scattered, minute, yellowish glandular hairs. Inflorescence with 2–4 capitula, furcate-corymbose; peduncles rather long, suberect, with numerous to dense stellate hairs, few to numerous, short, dark glandular hairs and few, short to medium, dark-based simple eglandular hairs. Capitula 30– 35 mm in diameter, broadly rounded or subtruncate at base. Involucral bracts incumbent in bud, 6–14 × 1.5–2.0 mm, blackish-green, broadly linear-lanceolate, gradually nar-

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rowed to an obtuse apex, with numerous to rather dense, short to medium, pale, dark-based simple eglandular hairs, few to fairly numerous, short, dark glandular hairs, and some stellate hairs at the base and apex. Ligules golden yellow, glabrous or nearly so at apex. Styles discoloured. Receptacle pits shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. H. ochthophilum has been confused with H. westii (i.e. H. submurorum auct.) with which it grows, but can be distinguished by the fewer, more obtuse, shallower teeth of the leaves and broader, less glandular-hairy capitula. Native. Streamside rocks at about 650 m. Known only from Allt nan Giubhas, near Kingshouse in Argyllshire. Endemic. 132. H. melanochloricephalum Pugsley Sombre-headed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–45 cm, pale yellowish-green, often tinted brownishpurple towards the base, with scattered, short or medium, pale simple eglandular hairs and few stellate hairs and occasional, short, dark glandular hairs in the upper part, or nearly glabrous. Leaves bluish-green on upper surface, paler beneath, and sometimes suffused metallic purple; basal leaves fairly numerous, the outer with lamina 2–6 × 1.5–2.5 cm, ovate or elliptical, obtuse-mucronate at apex, irregularly dentate, the teeth mammiform or aquilinemammiform and subtruncate at base with a short petiole, the inner with lamina 5–10 × 1.5–2.5 cm, lanceolate or ovatelanceolate, acute at apex, dentate especially in the lower half, the teeth mammiform to aquiline-mammiform and cuneate at base to short or long petioles; cauline 1–2, the lower similar to inner basal with a winged petiole, or bractlike; all nearly glabrous or with few to numerous, short to medium, pale simple eglandular hairs on both surfaces and the margins, and a few, minute, yellowish glandular hairs. Inflorescence with 2–6 capitula, furcate-corymbose; peduncles short, suberect, with dense stellate hairs, few to fairly numerous, short to medium, pale simple eglandular hairs and a few, short glandular hairs. Capitula 35–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 6–13 × 1.0–1.3 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous, unequal, short to medium, dark glandular hairs, and some stellate hairs towards the base. Ligules deep yellow, with numerous, very short simple eglandular hairs at the tips and sometimes also on the backs. Styles discoloured. Receptacle pits with margins more or less dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Rock-ledges on schistose and granite cliffs up to 1,100 m. Dubh Lochan on Beinn a’ Bhuird and in Coire Etchachan in Aberdeenshire, and Sgoran Dubh Mor and A’ Phocaid in Glen Einich in Inverness-shire, all in the Cairngorm Mountains. Endemic. 133. H. raveniorum P. D. Sell Ravens’ Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–35 cm, pale yellowish-green, often flushed brownish-purple, slender but rigid, flexuous, striate, with

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numerous, long, pale simple eglandular hairs near the base and fewer, shorter and dark-based ones upwards and few to numerous stellate hairs and sometimes an occasional, short glandular hair in the upper part. Leaves dark green on upper surface, paler beneath and sometimes tinted reddish; basal numerous, with lamina 2–7 × 1.5–2.5 cm, elliptical, oblanceolate, obovate or lanceolate, usually all obtuse at apex, rarely more or less acute, regularly sinuate-serrulate, the teeth small and sharp, cuneate or attenuate at base to a short or medium, densely long-hairy petiole; cauline 0– 2, when present like the basal but smaller and petiolate, or bract-like; all with numerous, short to long, pale simple eglandular hairs on both surfaces and the margins, those on the upper surface rather stiff, and with scattered, minute, yellowish glandular hairs. Inflorescence with 1–3 capitula, furcate-corymbose; peduncles rather short, suberect, with dense stellate hairs, numerous short to medium, pale, darkbased simple eglandular hairs and fewer, short, dark glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.8–1.2 mm, dark greyish-green, the inner with paler margins, all linearlanceolate, subobtuse to more or less acute at apex, with numerous, short to long, pale, dark-based simple eglandular hairs, few to fairly numerous, short, dark glandular hairs and a small tuft of hairs at apex. Ligules yellow, glabrous-tipped. Styles yellowish. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–8. Apomictic. Native. Known only from Argyllshire on north-facing cliffs of Beinn na Cille and a shallow ravine on Allt a’ Choire Dhuibh, both in Morvern, and Meall a’ Bhuiridh and Meal Garbh, Loch Cretan. Endemic. Named after Charles Earle Raven (1885–1964) and his son John Earle Raven (1914–80). 134. H. diversidens P. D. Sell & C. West Variable-toothed Hawkweed H. caesium var. alpestre auct.; H. caesium var. insulare auct.; H. submurorum auct. Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, rarely tinted reddish towards the base, slender, markedly striate, with few, medium to long, pale simple eglandular hairs towards the base and occasional ones upwards and a few stellate hairs and occasional, minute, dark glandular hairs in the upper part. Leaves pale green on upper surface, even paler beneath; basal few, the outer with lamina 2.5–7.0 × 1.0–3.0 cm, broadly elliptical or subrotund, rounded-mucronulate at apex, denticulate to shallowly dentate, and abruptly contracted at base to medium, long-hairy petioles, the inner with lamina 4–8 × 2–4 cm, elliptical, lanceolate or ovatelanceolate, acute at apex, with unequal mammiform or aquiline-mammiform teeth and denticulations, cuneate at base to a fairly long, long-hairy petiole; cauline 0–1, like the inner basal or bract-like; all glabrous on upper surface or with short, rather stiff, pale simple eglandular hairs, with few to numerous, medium to long, pale simple eglandular hairs beneath and on the margin and a few, minute, yellowish glandular hairs. Inflorescence with 1–4 capitula, furcatecorymbose; peduncles fairly long, suberect, with numerous stellate hairs, few, short or medium, dark simple eglandu-

lar hairs and few, short glandular hairs. Capitula 30–35 mm in diameter, narrowed to a rounded base. Involucral bracts porrect in bud, 4–10 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with few stellate hairs, numerous, short or medium, dark simple eglandular hairs and fewer, very short or short, dark glandular hairs. Ligules deep yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, dark. Flowers 7–8. Apomictic. Native. Known only from shaded rocks at 525 m. Allt Odhar, near Fortingal and on micaceous cliffs of Cruach Ardrain (Corrie Ardran), Glen Dochart, both in Perthshire and Coire Coille, Glen Spean in Inverness-shire. The Corrie Ardran plant was originally included in H. insulare. Endemic. 135. H. clovense E. F. Linton Clova Hawkweed H. praecox subsp. clovense (E. F. Linton) Zahn; H. leucograptum auct. Phyllopodous perennial herb with a branched stock. Stem (15–)20–35(–40) cm, pale yellowish-green, often brownish-red towards the base, slender to more or less robust, flexuous, striate, with a few stellate hairs above, sometimes with scattered, medium to long, pale simple eglandular hairs throughout and sometimes with an occasional, short, dark glandular hair above. Leaves bluishgreen and usually spotted and blotched brownish-purple on upper surface, paler beneath and sometimes flushed purple, the outer with lamina 1.5–4.0 × 1.5–2.5 cm, ovate, obtuse-mucronate at apex, denticulate or shortly dentate and truncate at base to a short petiole, the inner with lamina 3–9 × 2–5 cm, lanceolate, ovate-lanceolate or ovate, acute to acuminate at apex, dentate or incise-dentate, with mammiform, flattened mammiform or long and narrow mammiform, subtruncate at base to a rather short, longhairy petiole; cauline 0 or 1, with lamina narrowly lanceolate, acuminate at apex; all with medium to long, pale, simple eglandular hairs below and on the margins and glabrous or nearly so on upper surfaces, and with very occasional small, yellowish glandular hairs. Inflorescence with 1–9 capitula, furcate-corymbose; peduncles rather short, suberect, with numerous to dense stellate hairs, numerous, unequal, short and very short, dark glandular hairs and occasional, short to medium, dark simple eglandular hairs. Capitula 40–50 mm in diameter, broadly rounded below. Involucral bracts porrect in bud, 5–14 × 1.0–1.3 mm, dark green, the inner with paler margins, all linear-lanceolate, the outer obtuse, the inner acute at apex, with dense, unequal, short and very short, dark glandular hairs, few, short to medium, dark-based simple eglandular hairs, and a few stellate hairs towards the base. Ligules orange-yellow, glabrous or with a few minute hairs at tip. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. Cliff ledges at elevations between 360 and 750 m. Fairly common in the Clova and Perthshire mountains with solitary records from Glen Callater in Aberdeenshire and Glen Einich in Inverness-shire. Endemic.

39. Hieracium 136. H. lakelandicum P. D. Sell Lakeland Hawkweed H. clovense auct.; H. silvaticum var. prolixum auct. Phyllopodous perennial herb with a branched stock. Stem 15–35 cm, pale yellowish-green, often brownish-purple towards the base, slender, flexuous, striate, with numerous, long, pale simple eglandular hairs near the base and sometimes scattered ones upwards, with a few stellate hairs and an occasional, short, dark glandular hair in the upper part. Leaves bluish-green and often spotted and blotched brownish-purple on the upper surface, paler and sometimes tinted reddish beneath; basal often numerous, with lamina 1–6 × 1–3 cm, subrotund, broadly elliptical, lanceolate or ovate, often all obtuse at apex, sometimes 1–2 more or less acute, denticulate to dentate, the teeth small and rounded, subtruncate or abruptly contracted at base to short, long-hairy petioles; cauline 0–1, when present the lamina lanceolate, acute or acuminate at apex, dentate, petiolate; all glabrous or nearly so on upper surface, with numerous, short to long, pale simple eglandular hairs beneath, especially on the midrib and on the margin, and a few, scattered, minute, yellowish glandular hairs. Inflorescence with 1–4 capitula, furcate-corymbose; peduncles rather short, suberect, with dense stellate hairs and numerous, unequal, short and very short to medium, dark glandular hairs. Capitula 30–40 mm in diameter. Involucral bracts incumbent in bud, 4–12 × 1.0–1.3 mm, dark green, the inner with paler margins, all linear-lanceolate, gradually drawn out to a more or less acute apex, with dense, very short to medium, dark glandular hairs and numerous stellate hairs on the margin and base, without simple eglandular hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Gullies and rock-ledges up to 790 m. Central and eastern mountains of the Lake District with an outlying station in the Shap Fells. Endemic. 137. H. chrysolorum P. D. Sell & C. West Golden-flowered Hawkweed H. clovense auct. Phyllopodous perennial herb with a branched stock. Stem 18–13 cm, pale yellowish-green, often suffused brownishpurple, especially near the base, slender, flexuous, striate, with fairly numerous, long, pale simple eglandular hairs near the base and fewer, shorter ones upwards, and few to fairly numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves bluish-green on upper surface, paler beneath and sometimes tinted purplish; basal few to numerous; the outer with lamina 2–7 × 1.5–4.0 cm, subrotund or broadly elliptical, obtuse-mucronate at apex, denticulate or sinuate-dentate, with shallow teeth and shortly cuneate or rounded at base to a short petiole, the inner with lamina 3–11 × 1.5–4.5 cm, lanceolate, ovate or elliptical, acute or acuminate at apex, dentate, the teeth mammiform or narrowly mammiform, abruptly contracted or subtruncate at base with short, densely long-hairy petioles; cauline 0–1, with lamina small, lanceolate, acute or acuminate at apex, with few teeth, petiolate; all glabrous or nearly so on upper surface, with few to numerous, medium to long, pale simple eglandular hairs beneath, particularly on the midrib and

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on the margin, numerous stellate hairs on the under surface of the cauline, and scattered, minute, yellowish glandular hairs. Inflorescence with 2–6 capitula, furcate-corymbose; peduncles long, suberect, with numerous to dense stellate hairs and numerous, short, unequal, dark glandular hairs. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–13 × 1.0–1.5 mm, blackishgreen, the inner with paler margins, all linear-lanceolate, the inner very narrow, more or less acute at apex, with numerous to dense, very short, short and medium, dark glandular hairs, a few, short to medium, dark-based simple eglandular hairs and stellate hairs frequent on the margins and at the base. Ligules deep yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Known only from rocks by the Midlaw Burn near Moffat in Dumfries-shire and at Hen Hole in Northumberland. Endemic. 138. H. aristidens P. D. Sell Aristate-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale green, often purplish below, flexuous, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards, and few to numerous stellate hairs and few, very short, dark glandular hairs in the upper part. Leaves medium green on upper surface, paler beneath; basal rather few, the outer with lamina 1.5–4.5 × 1.0–2.5 cm, elliptical, rounded-mucronulate at apex, sparsely denticulate, and narrowed or rounded at base, the inner with lamina 6– 8 × 2–3 cm, narrowly elliptical to elliptical or lanceolate, acute at apex, serrate-dentate, with small, sharply spinulose teeth, and cuneate at base, the petioles up to 5 cm, with numerous, long, pale simple eglandular hairs; cauline 0–1, when present with lamina lanceolate, acute at apex, sharply toothed, narrowed at base and semiamplexicaul, or bract-like; all with few to numerous, short to medium simple eglandular hairs on both surfaces and the margin and some minute glandular hairs. Inflorescence with 1– 5 capitula, sometimes with a long lower branch; peduncles long and slender, with dense stellate hairs, numerous, very short, dark glandular hairs and a few, longer, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–15 × 1.0–1.3 mm, blackish-green, the inner with narrow pale margins, all linear-lanceolate, narrowed to a more or less acute apex, with numerous to dense, short or medium dark-based simple eglandular hairs, few to fairly numerous but not obvious, short, dark glandular hairs and few stellate hairs towards the base. Ligules yellow, with numerous, short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges. An Coileachan in Ross-shire, Carn Dearg in Sutherland and the Isle of Skye. Endemic. 139. H. dasythrix (E. F. Linton) Pugsley Dense-haired Hawkweed H. pictorum var. dasythrix E. F. Linton; H. rivale var. dasythrix (E. F. Linton) W. R. Linton; H. sagittatum subsp. rivale var. dasythrix (E. F. Linton) Zahn

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Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, sometimes brownish-red in lower half, usually slender, flexuous, striate, with numerous, medium to long, pale simple eglandular hairs below and fewer, shorter, dark-based ones above, and few to numerous stellate hairs above. Leaves medium green on upper surface, paler and often suffused purplish beneath; basal few, with lamina 3–10 × 2.0–3.5 cm, elliptical, lanceolate, ovate-lanceolate, sometimes all obtuse, sometimes all more or less acute, sometimes the outer obtuse and the inner acute at apex, subentire or denticulate, sometimes with small, sharp teeth and cuneate or subtruncate at base to rather long, long-hairy petioles; cauline 0–1, lanceolate, long-acute or acuminate at apex and shortly petiolate, or bract-like; all with short to long, pale simple eglandular hairs on both surfaces and the margins or the upper surface glabrous or nearly so, lower surface of cauline with numerous stellate hairs, without minute glandular hairs. Inflorescence with 2–7 capitula, furcate-corymbose; peduncles fairly long, suberect, with dense stellate, short to medium, pale, darkbased simple eglandular hairs and fewer, short, dark glandular hairs. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–10 × 1.0–1.3 mm, dark green, the inner with paler margins, all short and broad, linear-lanceolate, obtuse at apex, with dense, short to long, pale, dark-based simple eglandular hairs, few to fairly numerous, dark glandular hairs, and a few scattered stellate hairs which are sometimes more numerous on the margins. Ligules deep yellow, glabrous-tipped or nearly so. Styles yellowish to slightly discoloured. Flowers 7–8. Apomictic. Closely allied to H. petrocharis from which it is easily distinguished by its short, obtuse involucral bracts which are less stellately hairy. Native. Cliff ledges, scree and streamsides in the mountains. Widespread in central Scotland and on Ben Vorlich in Dunbartonshire. Endemic. 140. H. petrocharis (E. F. Linton) W. R. Linton Crag Hawkweed H. caesium var. petrocharis E. F. Linton; H. insulare var. petrocharis (E. F. Linton) F. Hanb.; H. caesium subsp. petrocharis (E. F. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–45 cm, pale yellowish-green, often purplish towards the base, slender to more or less robust, striate, with numerous, long, pale simple eglandular hairs towards the base and fewer upwards, and numerous stellate hairs and a few, very short, dark glandular hairs in the upper part. Leaves rather pale green, sometimes purple-spotted on upper surface, paler and often tinted purplish beneath; basal few to fairly numerous, with lamina 2.5–9.0 × 1.5–3.5 cm, mainly elliptical, sometimes lanceolate or ovate, usually obtusemucronate, sometimes the inner acute at apex, entire to sinuate-denticulate, rounded or cuneate at base to rather long petioles; cauline 0–1, when present with lamina lanceolate, acute at apex, entire or denticulate and petiolate, or bract-like; all glabrous or nearly so on upper surface, with few to numerous, short to long, pale simple eglandular hairs beneath, especially on the midrib and on the margin and petioles and a few, minute glandular hairs on the margin. Inflorescence with 1–5(–7) capitula, furcate; peduncles with

numerous to dense stellate hairs, few to numerous, short or medium, pale simple eglandular hairs and a few, short, dark glandular hairs. Capitula 30–40 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.5 mm, dark greyish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous stellate hairs all over, but particularly on the margins, numerous, short to long, pale, dark-based simple eglandular hairs and few, short dark glandular hairs. Ligules deep yellow, usually with a few, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Cliff ledges and streamsides up to 800 m. Several localities in Perthshire, Dalmally and Ben Laoigh in Argyllshire and Cnoc Reamhar in Kintyre. Endemic. 141. H. pseudanglicum Pugsley Upland Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–45 cm, pale yellowish-green, often tinted reddish towards the base, striate, slender, with few to numerous, short to long, pale, dark-based simple eglandular hairs and few to numerous stellate and short, dark glandular hairs in the upper part. Leaves rather pale green on upper surface, even paler beneath; basal few, with lamina 2–8(–11) × 1.5–3.0(–4) cm, lanceolate, narrowly elliptical or oblong-elliptical, obtuse to acute at apex, entire or with a few denticulations, or with few, slightly larger teeth, cuneate or attenuate at base to a long-hairy, fairly long petiole; cauline 0–2, the lower like basal but sessile and often semiamplexicaul, the upper bract-like; all with short to medium, rather stiff, pale simple eglandular hairs on the upper surface or nearly glabrous, few to numerous, medium to long, pale simple eglandular hairs on the lower surface, particularly the midrib and numerous short to medium, pale simple eglandular hairs and a few minute glandular hairs on the margin. Inflorescence with 1–3 capitula, furcate; peduncles with numerous stellate hairs, numerous, short to medium, very unequal glandular hairs and a few, medium, dark simple eglandular hairs. Capitula 30–45 mm in diameter, narrowed or rounded at base. Involucral bracts porrect in bud, 5–13 × 1.0–1.4 mm, blackish, the innermost with pale margins, all narrowly linear-lanceolate, gradually narrowed from base to a very fine, acute apex, with numerous, short and medium, very unequal, black glandular hairs and few to numerous dark or dark-based simple eglandular hairs, sometimes with very few stellate hairs. Ligules yellow with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Rock-ledges, slopes and streamsides on micaschist, limestone and granite up to 950 m. Common in central Scotland, on Ben Vorlich in Dunbartonshire and in a few localities in northern Scotland. Endemic. 142. H. oenophyllum P. D. Sell Wine-leaved Hawkweed H. subplanifolium auct. Phyllopodous perennial herb with a branched stock. Stem 20–35 cm, pale yellowish-green, usually strongly

39. Hieracium wine-coloured below, slender to fairly robust, striate, with few to numerous, medium to long, pale simple eglandular hairs below and occasional ones or none upwards, and few to numerous stellate hairs above. Leaves medium yellowish-green or slightly bluish-green, sometimes suffused brownish-purple or with a few blotches on the upper surface, usually deeply suffused purplish beneath; basal few to fairly numerous, with lamina 1.5–6.0 × 1–3 cm, subrotund, elliptical, elliptic-oblong or lanceolate, mostly rounded-obtuse-mucronate, sometimes some of the inner more or less acute at apex, subentire, denticulate or with a few shallow teeth at the base, rounded, abruptly contracted or shortly cuneate at the base with short, purplish, longhairy petioles; cauline 0–1, when present with lamina lanceolate, acute at apex, entire or denticulate, cuneate at base and shortly petiolate, or sometimes bract-like; all glabrous or with few to numerous, short to medium, pale simple eglandular hairs on the upper surface and with numerous, medium to long, pale simple eglandular hairs beneath and on the margin and especially on the midrib. Inflorescence with 2–7 capitula, furcate-corymbose, sometimes with a lower branch; peduncles with more or less dense stellate hairs, numerous short to medium, pale simple eglandular hairs and a few, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect to slightly incumbent in bud, 5–12 × 1.0–1.3 mm, dark greyish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to an acute apex, the inner very narrow, all with numerous stellate hairs, often dense on the margins, numerous, short to medium, pale, dark-based simple eglandular hairs and few, short, dark glandular hairs. Ligules medium to deep yellow, with numerous, very short simple eglandular hairs at apex. Styles yellow to slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–7. Apomictic. Native. Common on rock outcrops and ledges, possibly all limestone. Pass of Melfort, near Kilmelfort in Argyllshire and An Stuchd, near Loch Fuar-Bheinne and Cnoc na Seamraig all in Knapdale, Kintyre. Endemic. 143. H. cumbriense F. Hanb. Cumbrian Hawkweed H. atratum subsp. cumbriense (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–40 cm, pale yellowish-green, sometimes flushed brownish-purple towards the base, more or less slender, striate, glabrous, scaberulous or with few simple eglandular hairs and scattered stellate hairs. Leaves pale to medium green on upper surface, paler beneath and sometimes tinted pinkish; basal few to fairly numerous, the outer with lamina 3–7 × 2–4 cm, subrotund, elliptical, ovate or oblongelliptical, rounded-obtuse at apex, subentire or denticulate and rounded or subtruncate at base, the inner with lamina 6–9 × 2.5–3.5 cm, lanceolate, ovate-lanceolate, oblonglanceolate or oblong-elliptical, more or less acute at apex, subentire to shortly and sharply dentate in the lower half and abruptly contracted or shortly cuneate at base, all with petioles up to 5 cm with dense, long, pale simple eglandular hairs; cauline 0–1, with lamina lanceolate, acute at apex, entire or nearly so, sessile or rarely shortly petiolate; all glabrous or with an occasional simple eglandular hair on the

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upper surface, few to fairly numerous, short or medium simple eglandular hairs beneath, particularly on the midrib, and numerous, short and medium, pale simple eglandular hairs and occasional, minute glandular hairs on the margin. Inflorescence with 2–7 capitula, cymose-corymbose; peduncles short, with dense stellate hairs, few to numerous, short, dark glandular hairs and few to numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 25–35 mm in diameter, narrow, rounded at base. Involucral bracts porrect in bud, 6–13 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, short and medium, dark or dark-based simple eglandular hairs, numerous, short, dark glandular hairs and more or less numerous stellate hairs on the margins and apex. Ligules yellow, glabrous or with very short simple eglandular hairs at apex. Styles discoloured, possibly yellow when fresh. Receptacle pits with margins fimbriatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. It is possible that this species originated from a cross between H. ampliatum and H. subgracilentipes. It differs from H. ampliatum, with which it has been confused, by its different inflorescence and smaller narrower, more glandular capitula. Native. A rare species occurring on cliff ledges in a few localities in Cumberland and Westmorland. Endemic. 144. H. glandulidens P. D. Sell & C. West Glandular-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often suffused purplish towards the base, more or less slender, markedly striate, with numerous, medium and long, pale simple eglandular hairs near the base, becoming fewer upwards, and with few to numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves deep green on upper surface, paler beneath, sometimes tinted purplish; basal usually few, the outer with lamina 2–7 × 1.5–4.0 cm, broadly elliptical or subrotund, obtuse-mucronate at apex, entire, denticulate or shallowly toothed and cuneate at base to fairly long, long-hairy petioles, the inner with lamina 4–8 × 2.0–3.5 cm, lanceolate or ovate, subobtuse to acute at apex, denticulate and often with sharp, apiculate-tipped teeth and cuneate at base to long, long-hairy petioles; cauline 0–2, lower sometimes like inner basal and petiolate, upper usually bract-like; all glabrous or with a few simple eglandular hairs on the upper surface, with few to numerous, medium to long, pale simple eglandular hairs beneath, particularly on the midrib, and on the margin, and very occasional, minute glandular hairs on the margin. Inflorescence with 1– 3 capitula, furcate; peduncles rather long, with numerous to dense stellate hairs, numerous, short, dark glandular hairs and occasional, pale, dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, narrowed to a rounded base. Involucral bracts incumbent in bud, 5–12 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linearlanceolate, more or less acute at apex, with few, short, dark glandular hairs and numerous, short to medium, pale, darkbased simple eglandular hairs, and stellate hairs on the margins and at the apex. Ligules yellow, with numerous, short, pale simple eglandular hairs at apex. Styles discoloured.

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Receptacle pits with margins long-dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. Cliff ledges, slopes and by streams. Common in the west of Scotland, including the islands, from sea level to at least 800 m. Endemic. It is sometimes difficult to separate from H. pseudanglicum but usually has broader, more ovate leaves and fewer glandular hairs on the involucral bracts. 145. H. sannoxense P. D. Sell Sannox Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–55 cm, pale green, slightly purplish towards the base, slender, striate, with numerous, long, wavy, pale simple eglandular hairs below, becoming fewer and shorter upwards, and with numerous minute glandular hairs and stellate hairs in the upper part. Leaves medium green on upper surface, paler beneath; basal numerous, all similar, the lamina 3.0–6.5 × 1.5–3.5 cm, ovate to elliptical, roundedmucronulate to subacute at apex, denticulate or with a few, narrow and sharp teeth and abruptly contracted or subtruncate at base, the petioles up to 4 cm, with dense, long, pale simple eglandular hairs; cauline 0–1, when present with lamina small, lanceolate, acute at apex, semiamplexicaul; all glabrous or nearly so on upper surface, with numerous medium to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 1–15 capitula, sometimes with a long lower branch; peduncles long and slender, with numerous to dense stellate hairs, numerous unequal, very short and short, dark glandular hairs and a few short and medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded to subtruncate at base. Involucral bracts incumbent in bud, 5–13 × 1.0–1.2 mm, blackishgreen, the inner with paler margins, all narrowly linearlanceolate, acute at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, fairly numerous, short, dark glandular hairs and a few stellate hairs along the margin and at the base. Ligules with numerous, short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–6. Apomictic. Native. Known only from Sannox Bridge in Arran. Endemic. 146. H. insulare (F. Hanb.) F. Hanb. Ben More Hawkweed H. caesium var. insulare F. Hanb.; H. flocculosum var. insulare (F. Hanb.) F. N. Williams; H. subramosum subsp. insulare (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–30(−40) cm, pale yellowish-green, often flushed brownish-purple especially towards the base, slender, striate, with few to numerous, medium to long, pale simple eglandular hairs especially below, and numerous stellate hairs and an occasional, very short, dark glandular hair in the upper part. Leaves rather deep green on upper surface, paler beneath; basal few, the outer with lamina 2– 5 × 2.0–2.5 cm, elliptical, oblong-elliptical or subrotund, obtuse-mucronate at apex, subentire to denticulate and cuneate or subtruncate at base, the inner with lamina 5–6 × 1.5–2.0 cm, lanceolate, acute at apex, dentate, with short teeth and cuneate at base, the petioles rather long and

slender with dense, long simple eglandular hairs; cauline 0–1, similar to inner basal, petiolate; all with short to medium, rather stiff, pale simple eglandular hairs on the upper surface to almost glabrous, and numerous short to long, pale simple eglandular hairs beneath, especially on the midrib, and on the margins. Inflorescence with 1– 2(−6) capitula, furcate-corymbose; peduncles rather long, with numerous to dense stellate hairs, with few to numerous, short to medium, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 10–12 × 1.0–1.5 mm, greyish- to blackish-green, linear-lanceolate, obtuse at apex, with numerous to dense stellate hairs especially at the base, margins and apex, numerous, short, unequal, dark glandular hairs and numerous short to medium, pale, dark-based simple eglandular hairs. Ligules deep yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins dentate. Achenes 3.0–3.5 mm, blackish. Flowers 7–8. H. insulare is closely allied to H. eustales with which it grows on Ben More and can be distinguished by its more dentate inner leaves, obtuse involucral bracts and hairy tips to the ligules. Native. The only specimens seen of this rare species were obtained from mica-schist cliffs of Am Binnein on Ben More in Perthshire at 950 m. Endemic. 147. H. fissuricola P. D. Sell Fisher Ghyll Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, often purplish below, slender, striate, with few to numerous, long, pale simple eglandular hairs below, becoming shorter and fewer or absent upwards, and dense stellate hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath, sometimes slightly tinted purplish; basal more or less numerous, the lamina 2–10 × 1–4 cm, elliptical, lanceolate, oblong-lanceolate, oblong-elliptical or subrotund, most of them obtuse, but some broadly rounded and others narrowed at apex and occasionally acute, entire or remotely denticulate and rounded or cuneate at base, the petioles up to 6 cm and with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, when present with lamina narrowly linear, entire and sessile; all glabrous or with scattered, short, stiff simple eglandular hairs on the upper surface, and with numerous, short to medium, stiff simple eglandular hairs beneath and on the margins and scattered stellate hairs beneath. Inflorescence with 1–11 capitula, furcatecorymbose, sometimes with a long lower branch; peduncles rather short and straight, with dense stellate hairs, numerous, short or medium, pale simple eglandular hairs and few to numerous, short, dark glandular hairs. Capitula 25– 30 mm, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.0–1.2 mm, dark olive-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to an obtuse apex, with numerous, short and medium, dark-based simple eglandular hairs, few to fairly numerous, short, dark glandular hairs and some stellate hairs towards the base. Ligules yellow, with very short simple eglandular hairs at apex. Styles yellowish to discoloured. Receptacle

39. Hieracium pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only from Fisher’s Ghyll gully, Helvellyn, Cumberland at about 430 m. Endemic. 148. H. eustales E. F. Linton Floccose-bracted Hawkweed H. caesium subsp. eustales (E. F. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 25–45 cm, yellowish-green, rarely slightly reddish below, more or less slender, striate, with numerous, long, pale simple eglandular hairs near the base and a few upwards, and numerous to dense stellate hairs and few, short, dark glandular hairs in the upper part. Leaves dull green on upper surface, paler beneath; basal few to numerous, with lamina 3–6 × 1.5–2.5 cm, elliptical, lanceolate or ovate-lanceolate, obtuse-mucronate to rarely acute at apex, entire, denticulate or very shallowly dentate, rounded or subtruncate at base to medium petioles; cauline leaves 0–1, with lamina lanceolate, acute at apex, entire and petiolate, or bract-like; all with short simple hairs to nearly glabrous on upper surface, with numerous short to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 1–6 capitula, furcate-corymbose; peduncles with dense stellate hairs, few to numerous, short, dark glandular hairs and an occasional short or medium, pale simple eglandular hair. Capitula 30–40 mm in diameter, narrowed or rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linearlanceolate, more or less acute at apex, with dense stellate hairs especially towards the base and on the margins, numerous, short to medium, dark glandular hairs and few to numerous, medium to long, dark-based simple eglandular hairs. Ligules yellow, with numerous, very short simple eglandular hairs at the tips. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. Streamsides and rock-ledges up to 800 m. A rare species found only in a few scattered localities in Perthshire. Endemic. 149. H. ovaliforme P. D. Sell Oval-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 15–25 cm, pale green, sometimes purplish towards the base, slender, with long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and a few stellate and minute glandular hairs in the upper part. Leaves rather pale green on upper surface, sometimes with faint blotches, paler beneath; basal numerous, the lamina 1.5–5.0 × 1.0– 3.0 cm, subrotund or elliptical, rounded at apex, rarely subacute, entire or remotely denticulate, rounded or abruptly contracted at base, the petioles up to 2 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, when present with lamina small, linear-lanceolate or lanceolate, entire, acute and semiamplexicaul, or bract-like; all with few simple eglandular hairs on the upper surface or glabrous, with numerous medium to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 1–2 capitula; peduncles slender, with dense stellate hairs, few to numerous, short, dark glandular hairs and an

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occasional simple eglandular hair. Capitula 35–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.2–1.5 mm, dark greyish-green, the inner with paler margins, all linear-lanceolate, narrowed but obtuse at apex, with numerous, medium, dark-based simple eglandular and few glandular hairs and numerous stellate hairs along the margin. Ligules yellow, with short, simple eglandular hairs at the apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 6–7. Apomictic. Native. Cliff ledges. Ben Hope and Meallan Liath in Sutherland and An Teallach and Couldoran in Ross-shire. Endemic. 150. H. arranense P. D. Sell Arran Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–40 cm, pale yellowish-green, sometimes slightly reddish at base, slender to slightly robust, striate, with numerous, long, pale simple eglandular hairs below, with few or none above, and with few to numerous stellate hairs above. Leaves yellowish-green and blotched and spotted brownishpurple on upper surface, paler beneath and sometimes suffused violet-purple; basal with lamina 1.5–5.5 × 1–3 cm, mainly ovate, sometimes elliptical or lanceolate, most of them obtuse-mucronate but the inner sometimes more or less acute at apex, entire or minutely denticulate, more or less rounded at base to rather short, often reddish, densely long-hairy petiole; cauline absent or linear and bract-like; all glabrous or nearly so on upper surface, with numerous, medium to long, pale simple eglandular hairs beneath and on the margin, and a few, minute glandular hairs on the margin. Inflorescence with 1–5 capitula, furcate-corymbose; peduncles with numerous to dense stellate hairs, sometimes a few, short to medium, pale simple eglandular hairs and occasionally a few, short, dark glandular hairs. Capitula 30–40 mm in diameter, broadly rounded at base. Involucral bracts more or less porrect in bud, 4–12 × 1.2–1.5 mm, dark greyishgreen, the inner with paler margins, all linear-lanceolate, the inner very narrow, more or less acute at apex, with numerous stellate hairs all over, but particularly on the margin, numerous short to medium, pale simple eglandular hairs and few, short, dark glandular hairs. Ligules deep yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes about 3.0 mm, reddishblack. Flowers 5–6. Apomictic. H. arranense is closely allied to H. petrocharis, from which it differs in its broader, generally more ovate leaves on rather short petioles and shorter indumentum of the involucral bracts. Native. Sandstone rocks in several places between Brown Head and Dougarie on the west coast of the island of Arran. Endemic. 151. H. thalassinum P. D. Sell Hairy-bracted Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, often flushed purplish towards the base, with numerous, long, pale simple eglandular hairs below, becoming shorter upwards, and with numerous stellate hairs in the upper part. Leaves medium

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green on upper surface, paler beneath and sometimes tinted reddish or purplish; basal few, the lamina 1.5– 8.0(−10) × 1.0–3.5(−5) cm, ovate, obovate or elliptical, obtuse-mucronate to acute at apex, subentire, sparsely denticulate or with an occasional tooth and rounded, cuneate or attenuate at base, the petioles up to 7 cm and with dense, long, pale, wavy simple eglandular hairs; cauline 1– 3, the lower like basal and with a winged petiole and semiamplexicaul, the upper narrower, and more or less sessile; all with numerous, short to long, pale simple eglandular hairs on both surfaces and on the margin. Inflorescence with 2–6 capitula, furcate-corymbose, sometimes with a lower branch; peduncles rather long and slender, with dense stellate hairs and numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4– 12 × 1.0–1.2 mm, dark green, the inner with paler margins, all oblong-lanceolate, acute at apex, with dense, short to long, pale dark-based simple eglandular hairs, some stellate hairs on the margins and a very occasional short, dark glandular hair. Ligules yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 2.5–3.0 mm, reddish-brown. Flowers 6–7. Apomictic. Native. Sea-cliffs at Ballygroggan in Kintyre and rocks by the River Oude in the pass of Melfort, near Kilmelfort in Argyll. Endemic. 152. H. hyparcticoides Pugsley ex P. D. Sell Traligill Hawkweed H. hyparcticum auct. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes reddish below, slender to robust, striate, with scattered medium to long, pale, sometimes dark-based simple eglandular hairs and with few stellate and short, dark glandular hairs above. Leaves yellowish-green on upper surface, paler beneath; basal few, the outer with lamina 2–7 × 1.5–3.0 cm, obovate, oblong-obovate or elliptical, obtuse-mucronate at apex, entire, denticulate or rarely shallowly dentate and abruptly contracted or cuneate to rather long, densely long-hairy petioles, the inner with lamina narrow elliptical or lanceolate, more or less acute at apex, mostly denticulate, cuneate at base to fairly long petioles; cauline 0–2, the lower like inner basal and petiolate, the rest linear and bract-like; all glabrous or nearly so on upper surface, with few to numerous, short to medium, pale simple eglandular hairs beneath and on the margin, numerous to dense stellate beneath and sometimes above, and a few, minute, yellowish glandular hairs also present. Inflorescence with 2–9 capitula, furcatecorymbose; peduncles long, suberect, with numerous to dense stellate hairs, numerous, short, dark glandular hairs and occasionally few, short to medium, pale, dark-based simple eglandular hairs. Capitula 30–35 mm in diameter, narrowed at the base. Involucral bracts porrect in bud, 5– 13 × 1.0–1.5 mm, blackish-green, linear-lanceolate, gradually narrowed to an obtuse apex, with numerous, unequal, very short and short, dark glandular hairs, few, medium, blackish simple eglandular hairs and numerous stellate hairs especially on the margins and at the apex. Ligules yellow, with numerous, very short simple eglandular hairs

at apex. Styles yellowish to discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 6–7. Apomictic. Native. Known only from limestone banks of the Traligill Burn and other streams near Inchnadamph, and from Ben Hope in Sutherland and Maoile Lunndaidh, Ross-shire. Endemic. 153. H. pseudanglicoides J. E. Raven, P. D. Sell & C. West Entire-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–30 cm, pale yellowish-green, often tinted reddish below, very striate, slender, often flexuous, with few to numerous stellate hairs throughout, few to fairly numerous, medium to long, dark or dark-based simple eglandular hairs and few to fairly numerous, short, dark glandular hairs in the upper part. Leaves pale green on upper surface, even paler beneath and rarely purple tinted; basal rather few, with lamina 2–7 × 1.0–2.5 cm, narrowly elliptical or oblongelliptical, obtuse to more or less acute at apex, usually entire or nearly so, sometimes with a few, small teeth, cuneate to attenuate at base to a fairly long, slender, long-hairy petiole; cauline 0–2, rarely like inner basal, usually linearlanceolate or absent; all with more or less numerous stellate hairs on both surfaces, with few to numerous, short to long, pale simple eglandular hairs beneath and on the margin and a few, minute glandular hairs on the margin and midrib beneath. Inflorescence with 1–2 capitula, furcate; peduncles with numerous to dense stellate hairs, numerous short to medium, dark or dark-based simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Capitula 20–35 mm in diameter, narrowed or rounded at base. Involucral bracts porrect in bud, 6–13 × 1.0–1.4 mm, blackish, the inner with pale margins, all very narrow linear-lanceolate, gradually narrowed from base to apex, with numerous, short to medium, dark or dark-based simple eglandular hairs, few to numerous, short, dark glandular hairs and a few stellate hairs. Ligules yellow, with very short simple eglandular hairs at apex. Styles dark. Receptacle pits with margins subulate-dentate. Achenes 4.5–5.0 mm, reddish-black. Flowers 7–9. Apomictic. Native. Rock-ledges and slopes, perhaps always on granite up to 290 m. In a few localities in Perthshire, Forfarshire, Aberdeenshire, Inverness-shire and Kintyre. Endemic. Section 10. Lanatella (Arv.-Touv.) Zahn Taxon Lanatella Arv.-Touv.; Taxon Aurellinia Arv.-Touv.; Subsection Aurellinia (Arv.-Touv.) Rouy Plant with long hairs on underside of leaves, stem and involucre, some of which are more or less plumose. Leaves mostly basal, spotted and marbled, cauline usually one. Inflorescence cymose-corymbose. Styles yellow. Achenes dark. Receptacle pits with slightly dentate margins. Native of the western Alps, Italy and Balkans. 154. H. rionii Gremli Marbled Hawkweed H. caesioides subsp. rionii (Gremli) Zahn; H. subincisum var. rionii (Gremli) Arv.-Touv. Phyllopodous perennial herb with a branched stock. Stems 30–50 cm, pale green, often tinted purplish, striate, flexuous, slender, with stellate hairs throughout, densely so

39. Hieracium above, with numerous, long, pale simple or subplumose eglandular hairs below and scattered ones above, without glandular hairs. Leaves glaucous green and densely spotted, blotched and marbled on the upper surface with brownishpurple, paler beneath, glabrous or nearly so on the upper surface, nearly glabrous on the lower surface but with long, pale, wavy simple or subplumose eglandular hairs on the midrib beneath and on the margin; basal numerous, with lamina 4–8 × 2–5 cm, the outer broadly elliptic-oblong, rounded-mucronulate at apex, deeply divided into large mammiform teeth with smaller teeth in between, and subtruncate to shortly cuneate at base, the inner with lamina lanceolate to ovate, acute at apex, deeply laciniate-dentate, the teeth large and mammiform, or aquiline-mammiform, or very narrow and acute and usually with some long, narrow teeth free at the subtruncate or cuneate base, the petioles long with very long, wavy, pale simple eglandular hairs; cauline usually one, with lamina more or less lanceolate or lanceolate in outline, long acute or acuminate at apex, deeply laciniate, often nearly to the midrib, the teeth unequal, long and narrow, more or less petiolate. Inflorescence cymose-corymbose with a rather long acladium and sometimes a long lower branch; peduncles long and slender, with dense stellate hairs, numerous, long, pale, wavy simple eglandular hair and a very occasional minute glandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts 6–13 × 0.5–1.0 mm, greyish-green, the inner with paler margins, linear-lanceolate, gradually narrowed to a fine, acute apex, the innermost almost filamentous, with numerous stellate hairs, numerous to dense, long, wavy, pale simple or subplumose eglandular hairs, without or with an occasional minute glandular hair. Ligules medium yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins fibrillous-dentate. Achenes 2.5–3.5 mm, reddish-black. Flowers 5–6, and again 9–10 when it still keeps its characters. Introduced. Planted in the Cambridge Botanic Garden from which it has spread and become a weed on beds and rockeries and has been recorded in other gardens. Its handsome marbled leaves may mean it is grown widely. It should not be mistaken for H. scotostictum which is also grown in gardens and which can be distinguished by its sparsely dentate leaves and numerous glandular as well as simple eglandular hairs on the involucral bracts. Native of the Alpes Maritimes, Basses-Alpes, HautesAlpes and Corsica. Section 11. Andryaloidea Monnier Taxon Lanata Arv.-Touv.; Section Lanata (Arv.-Touv.) Zahn; Taxon Tomentosa N¨ageli & Peter; Section Tomentosa (N¨ageli & Peter) Zahn 155. H. lanatum Vill. Woolly Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–50 cm, yellowish-green, fairly robust, striate with long, pale simple eglandular to more or less plumose hairs, very dense at the base and dense to numerous upwards, and numerous stellate hairs. Leaves medium green on upper surface, paler beneath, the basal in a rosette, 2–11 × 1.0–3.5 cm, ovate, elliptical or elliptical-oblong, obtuse-mucronate

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or acute at apex, entire or denticulate, and narrowed at base to a short or medium winged petiole, the cauline 2–5, ovate to oblong, more or less acute at apex, rounded and more or less amplexicaul at base, all with numerous medium to long, pale, more or less plumose hairs on both surfaces and the margins. Inflorescence with 2–7 capitula, solitary at the ends of stems and branches; peduncles long, with numerous stellate hairs and numerous to dense, medium to long, pale, wavy, simple eglandular or subplumose hairs and sometimes with a few, slender glandular hairs. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts 11– 15 × 1.5–1.8 mm, blackish-green, linear-lanceolate, with a gradually narrowed, acute apex, with dense, medium to long, pale simple eglandular or subplumose hairs, with a few, minute glandular hairs. Ligules pale yellow, with minute glandular hairs at apex. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, blackish-red. Flowers 6–7. Introduced. Grown in gardens and established at Canterbury in Kent, the North Denes at Great Yarmouth in Norfolk and Littleferry in Sutherland. Native of the mountains of the Jura to the Alpi Apuane. Section 12. Villosa (Griseb.) Gremli Taxon Villosa Gremli; Subsection Villosa (Griseb.) Rouy; Taxon Heterodonta Arv.-Touv.; Section Heterodonta (Arv.-Touv.) Arv.-Touv. Phyllopodous perennial herbs densely hairy throughout. Leaves medium green, the basal numerous, the cauline (2–)3–6, oblong or oblong-lanceolate, entire or denticulate. Inflorescence 2–4 capitula on long, straight peduncles. Capitula large. Involucral bracts acute, with dense, long simple eglandular hairs, few stellate hairs and no glandular hairs. Ligules with very short hairs at tip. Styles yellow. Receptacle pits with margins fimbriate-dentate. Achenes reddish-black. Common in the mountains of central and south Europe. 156. H. pilosum Schleich. ex Froel. Fimbricte-pitted Hawkweed Phyllopodous perennial herb with a branched stock. Stem 15–30(−40) cm, pale yellowish-green, sometimes flushed brownish-purple below, slender, flexuous, with numerous to dense, pale, medium to long simple eglandular hairs and dense stellate hairs in the upper part, without glandular hairs. Leaves medium green on upper surface, paler beneath; basal numerous, 4–8 × 1.5–2.0 cm, oblong or oblong-lanceolate, obtuse to subacute at apex, entire or denticulate, narrowed to base to a short, winged petiole; cauline (2–)3–6, often rather small, lanceolate or ovate, acute at apex, entire or denticulate, sessile and sometimes semiamplexicaul, with numerous to dense, medium to long, pale simple eglandular hairs on both surfaces. Inflorescence with 2–4 capitula, furcate; peduncles long, with dense stellate hairs and numerous medium to long, pale simple eglandular hairs, without glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts 5–15 × 2– 3 mm, blackish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous to dense, medium to long simple eglandular hairs, sometimes with a few stellate hairs below, without glandular hairs. Ligules

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pale yellow, with very short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins fimbriatedentate. Achenes 3.5–4.5 mm, reddish-black. Flowers 7–8. Introduced. Naturalised on the beach at Dungeness in Kent. Native in mountains from the Jura and the Carpathians to the south-west Alps, central Apennines and Macedonia. Section 13. Cerinthoidea Monnier Section Oxylepida Froel. nom. illegit.; Subsection Cerinthella Scheele; Subsection Olivacea Scheele; Subsection Eumurorum F. N. Williams; Subsection Cerinthea Rouy Perennial herbs. Stems up to 60 cm, usually with numerous simple eglandular hairs, especially below and few glandular ones above. Leaves often more or less glaucous and with long simple eglandular hairs, the basal in a rosette, the cauline 1–6(−9) and more or less amplexicaul. Inflorescence often with long peduncles. Capitula large. Involucral bracts usually incumbent in bud, simple eglandular hairs dominant, glandular hairs often few. Ligules usually with hairs at tip, often pale. Styles discoloured. Receptacle pits with margins dentate or fimbriate-dentate. Achenes dark. Flowers 6–7. Upland and mountainous areas on cliffs, rocky slopes and by streams, and sea-cliffs in the north. Widespread in the north of England, Scotland and Ireland. Faeroes, Iceland, western Europe and the south-west Alps but absent from Scandinavia. Usually recognised by their hairy appearance with long peduncles, large heads and amplexicaul cauline leaves. Similar but smaller plants with smaller, darker heads are considered to originate from crosses of species of this section with those of the Section Alpina, and have been put in the Section Subalpina. 157. H. iricum Fr. Erin Hawkweed H. mougeotii subsp. iricum (Fr.) Zahn; H. peramplum Dahlst.; H. lapeyrousii auct. Phyllopodous perennial herb with a branched stock. Stem (10–)30–60 cm, pale yellowish-green, often reddish towards the base, usually more or less robust, striate, with numerous, very long(−6 mm), pale, wavy simple eglandular hairs throughout and usually with tufts of these hairs at the nodes, and stellate hairs mainly in the upper part. Leaves bluish-green, sometimes tinted red; basal few to numerous, the lamina 3–17 × 1.5–5.0 cm narrowly to broadly elliptical, lanceolate or oblong-lanceolate, mostly more or less acute at apex, remotely denticulate to sharply serratedentate, the teeth narrow and upturned, cuneate to attenuate at base to a long, long-hairy petiole; cauline 3–6(−9 cult.), decreasing in size upwards, the lamina lanceolate or panduriform, long-acute or acuminate at apex, denticulate to sharply serrate-dentate, narrowed or rounded to a more or less amplexicaul base; all with numerous medium to long, pale, wavy simple eglandular hairs on both surfaces and the margins. Inflorescence with 1–20 capitula, furcatecorymbose; peduncles long, suberect, with dense stellate hairs, numerous short to medium, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs. Capitula 30–40 mm in diameter, broad and

subtruncate at base. Involucral bracts incumbent in bud, 4–13 × 1.0–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, broad below, suddenly narrowing above to a more or less acute apex, with dense, medium to long, pale, dark-based simple eglandular hairs, few to numerous short, dark glandular hairs and a few stellate hairs towards the base. Ligules yellow, with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with the margins strongly fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Cliff ledges, rocky outcrops, and streamsides. Northern England, Scotland and much of Ireland in upland areas, but sometimes coming down to the coast. Faeroes. 158. H. scarpicum Pugsley Scarp Hawkweed Phyllopodous perennial herb with a branched stock. Stem 10–35 cm, pale yellowish-green, often suffused purplish, more or less slender, striate, with numerous, long, pale, wavy simple eglandular hairs, dense at the very base and numerous stellate hairs in the upper part. Leaves pale green on upper surface, even paler beneath and sometimes suffused purplish; basal with lamina 2–9 × 2–4 cm, the outer elliptical to subrotund, obtuse-mucronate at apex, denticulate or dentate and narrowed at base, the inner mostly elliptical, more acute at apex and with more numerous, sharper teeth and narrowed at base, the petioles up to 8 cm and longhairy; cauline 2–4, the lower with lamina similar to the inner basal, often with coarse teeth and with a winged, semiamplexicaul petiole, the upper lanceolate, acute at apex, sharply dentate, narrowed at base and semiamplexicaul; all glabrous or nearly so on upper surface, with medium to long, pale simple eglandular hairs beneath particularly on the midrib and margin, and a few stellate hairs beneath. Inflorescence with 2–6 capitula, furcate-corymbose, peduncles with dense stellate hairs and numerous, medium to long, pale, dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, broadly rounded at base. Involucral bracts porrect in bud, 6–15 × 1.2–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, gradually attenuate to a subacute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs intermixed with few to fairly numerous, short, dark glandular hairs, with stellate hairs along the margins. Ligules yellow, with a few, short hairs at apex. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 4.0–4.5 mm, blackishpurple. Flowers 6–7. Apomictic. Native. Cliffs and streamsides. Scarp, Harris, Lewis and Little Bernera in the Outer Hebrides. Endemic. 159. H. cerinthiforme F. Hanb. Clasping-leaved Hawkweed H. anglicum var. cerinthiforme (F. Hanb.) F. Hanb.; H. mougeotii subsp. anglicum var. cerinthiforme (F. Hanb.) Zahn; H. anglicum var. amplexicaule Backh.; H. anglicum var. decipiens Syme nom. illegit.; H. perampliforme Dahlst.; H. patens Dahlst.; H. cerinthoides auct.; H. lawsonii auct. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes slightly reddish towards the base, slender to robust, striate, with dense,

39. Hieracium long, pale simple eglandular hairs at the very base and few to numerous upwards, and few to numerous stellate hairs in the upper part. Leaves bluish-green on upper surface, paler beneath; basal few, the lamina 3–10 × 2.0–3.5 cm, elliptical, lanceolate, oblong-lanceolate or oblong, obtuse to acute at apex, entire to shallowly dentate, narrowed below to fairly long petioles which have dense, long, pale simple eglandular hairs; cauline 1–4, the lower similar to basal with a winged amplexicaul petiole, the upper lanceolate or oblonglanceolate, acute at apex, entire to sparsely and shallowly dentate, rounded-amplexicaul at base; all with numerous, short to long, pale simple eglandular hairs on the upper surface and more numerous similar hairs beneath and on the margins, sometimes with some stellate hairs on the underside of the cauline. Inflorescence with 1–7 capitula, furcatecorymbose; peduncles with numerous stellate hairs, numerous, pale, dark-based simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Capitula 30–50 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 4–13 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually attenuate to an acute apex, with numerous, medium to long, pale dark-based simple eglandular hairs intermixed with few, short, dark glandular hairs and with few stellate hairs. Ligules pale yellow, with numerous, short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 6–8. Apomictic. Differs from H. anglicum mainly in its narrower leaves the cauline of which are more numerous and amplexicaul. In the north of Scotland and the Orkney Islands this species is common and uniform. Just occasionally some plants are difficult to distinguish from H. iricum. In central Scotland it is much scarcer and most plants, which have its same general appearance have darker, narrower, more glandular capitula and are referable to H. pseudanglicum. Other plants which have a similar appearance, but do not have semiamplexicaul leaves are referable to H. glandulidens. Irish plants are like north Scottish plants. Native. Cliff ledges and slopes and by streams from sea level to at least 700 m. Common in central and north Scotland and the Orkney Islands, apparently scarce in the western islands; a few records south to Westmorland; scattered records over much of Ireland. Endemic. 160. H. hartii (F. Hanb.) P. D. Sell & C. West Hart’s Hawkweed H. cerinthiforme var. hartii F. Hanb.; H. anglicum var. cerinthiforme forma hartii (F. Hanb.) W. R. Linton; H. mougeotii subsp. anglicum var. cerinthiforme forma hartii (F. Hanb.) Zahn; H. anglicum var. hartii (F. Hanb.) F. N. Williams Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, yellowish-green, sometimes slightly reddish at base, more or less robust, striate, with dense, long, pale simple eglandular hairs at the base and few to fairly numerous upwards, with few to fairly numerous stellate hairs and minute glandular hairs in upper part. Leaves medium green on upper surface, paler beneath; basal 3–5, often disappearing before anthesis, the lamina more or less obovate,

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obtuse-mucronate, subentire, fairly long petiolate; cauline 3–5, the lamina ovate to obovate, slowly decreasing in size, obtuse-mucronate to acute at apex, more or less entire, the lower petiolate, the upper sessile and amplexicaul; all with upper surface glabrous or nearly so, with few to numerous, short to medium, pale simple eglandular hairs beneath and on the margins. Inflorescence with 2–5 capitula, furcatecorymbose; peduncles with numerous stellate hairs, numerous, short and medium, pale simple eglandular hairs, occasional short glandular hairs and some minute glandular hairs. Capitula 30–40 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 6–15 × 1.5–2.0 mm, green, the inner with paler margins, all broadly linearlanceolate, obtuse at apex, with numerous, short to medium, pale simple eglandular hairs, stellate hairs along the margin, and a few, short, dark glandular hairs. Ligules pale yellow, with numerous short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins dentatefimbriate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only from Ben Bulben in Co. Sligo and Slieve League in Co. Donegal in Ireland. In 1891 it was said to be the Hieracium of the mountain as regards Slieve League. Although it has been searched for in both localities it has not been found since. Endemic. Named after Henry Chichester Hart (1847–1908). 161. H. magniceps P. D. Sell & C. West Large-headed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 40–60 cm, pale yellowish-green, sometimes flushed purplish below, more or less robust, striate, with dense, long, pale, wavy simple eglandular hairs at the very base and few to fairly numerous upwards, with numerous stellate hairs in the upper part. Leaves medium to dark green on the upper surface, paler beneath; basal few, the lamina 3– 15 × 2–7 cm, broadly elliptical to oblong-elliptical, obtuse to acute at apex, subentire to shallowly undulate-dentate, attenuate at base to petioles which are up to 8 cm, winged and long-hairy; cauline 2–4, similar to basal or the lamina ovate or lanceolate and usually long-acute, with a short winged petiole, or sessile and semiamplexicaul; all with few to numerous, short, pale simple eglandular hairs or glabrous on the upper surface, with few to numerous, short to long, pale simple eglandular hairs beneath, particularly on the veins, and on the margins. Inflorescence with 2–6 capitula, furcate-corymbose; peduncles with dense stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and occasional short, dark glandular hairs. Capitula 35–50 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 6–15 × 1.5–2.0 mm, blackish-green, inner with paler margins, all linearlanceolate, gradually narrowed from the base to the obtuse or subacute apex, with numerous stellate hairs, particularly on the margins, numerous to dense, short to medium, pale, dark-based simple eglandular hairs and few to fairly numerous, short, dark glandular hairs intermixed. Ligules pale yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins

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fimbriate-dentate. Achenes 3.5–4.0 mm, blackish-purple. Flowers 6–7. Apomictic. Native. Found only on grassy banks and pebbly margins of the Shee Water near the Spittal of Glen Shee, Perthshire. Endemic. 162. H. langwellense F. Hanb. Langwell Hawkweed H. olivaceum subsp. langwellense (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale yellowish-green, rarely slightly reddish at base, striate, slender to robust, with dense long hairs at the very base, fewer upwards, and numerous, stellate hairs in the upper part. Leaves yellowish-green on upper surface, paler beneath; basal few to numerous, the outer with lamina 3–8 × 2.0–3.5 cm, elliptical, oblong-elliptical or subrotund, rounded-mucronulate at apex, subentire, denticulate or very shallowly undulate-dentate and narrowed below to short petioles, the inner with lamina 6–12 × 2.0– 4.5 cm, elliptical, lanceolate, ovate-lanceolate or ovateoblong, acute at apex, denticulate or shallowly undulatedentate, and narrowed at base to fairly long petioles; cauline 1–3, the lower like inner basal and petiolate, the upper lanceolate, acute at apex, entire or very shallowly dentate and sessile; all glabrous or nearly so on upper surface, with fairly numerous, medium to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 3–13 capitula, furcate-corymbose, often with a long, lower branch; peduncles usually long, with numerous to dense stellate hairs, numerous short to medium, pale, darkbased simple eglandular hairs and few, short, dark glandular hairs. Capitula 30–45 mm in diameter, with broad rounded bases. Involucral bracts incumbent in bud, 6–13 × 1.3–1.5 mm, olive-green, the inner with paler margins, all linearlanceolate, obtuse or subacute at apex, with numerous stellate hairs especially on the margins, numerous, short to medium, pale, dark-based simple eglandular hairs and a few, small, dark glandular hairs. Ligules yellow, with a few, very short, simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins dentate and long-ciliate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Cliff ledges and slopes and streamside rocks from sea level up to 800 m. Scattered localities in central and north Scotland and one record from Galway. Endemic. 163. H. oxybeles P. D. Sell Pointed-leaved Hawkweed H. cerinthoides var. acutifolium Backh. fil., non H. acutifolium Vill.; H. anglicum var. acutifolium (Backh. fil.) Backh. fil.; H. mougeotii subsp. anglicum forma acutifolium (Backh. fil.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale yellowish-green, often tinted reddish towards the base, usually more or less robust, striate, with dense, long, pale simple eglandular hairs at the very base and fewer upwards, and numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves pale green to bluish-green on upper surface, paler beneath; basal rather few, the outer with lamina 4–8 × 2–4 cm, broadly elliptical, obtuse-mucronate, dentate at least in the lower half, the teeth tending to be cusped, narrowed to base, with the teeth sometimes extending on to the petiole, the inner with

lamina 5–14 × 2–7 cm, broadly elliptical, ovate or oblongovate, gradually acute at apex, dentate, particularly in the lower half, the teeth aquiline-mammiform, cusped, sometimes long and narrow and often extending on to the fairly long, densely long-hairy petiole; cauline (0–)1–2, sometimes like inner basal, sometimes lanceolate, acute at apex, entire or with a few teeth; all with few to numerous, short to medium, pale simple eglandular hairs on upper surface, more numerous and longer on the lower surface, especially the midrib and the margin. Inflorescence with 1–6 capitula, furcate-corymbose, sometimes with a long lower branch; peduncles with dense stellate hairs, numerous short, dark glandular hairs and few to numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 30–50 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 6–15 × 1.0–1.5 mm, olive-green, the inner with paler margins, all linear-lanceolate, gradually drawn out to a narrow, acute apex, with numerous, short to long, pale, dark-based simple eglandular hairs intermixed with a few to fairly numerous, short dark glandular hairs and a few stellate hairs. Ligules pale yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. This species has been confused with H. langwellense by all previous botanists, chiefly because it can have pale green or bluish-green leaves. It is distinguished from H. langwellense by its long, cusped teeth in the lower half of the leaf and more glandular-hairy inflorescences. Native. Cliff ledges and slopes and streamsides from sea level to at least 800 m. Common in north and central Scotland and scattered records in Ireland. Endemic. 164. H. anglicum Fr. English Hawkweed H. cerinthoides var. anglicum (Fr.) Backh. fil.; H. mougeotii subsp. anglicum (Fr.) Zahn; H. subimpressum Dahlst.; H. perampliforme auct. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes slightly reddish towards the base, striate, robust, with very dense, long, wavy, pale simple eglandular hairs at the very base, and more or less numerous similar hairs above, and few to numerous stellate hairs and few, very short, dark glandular hairs in the upper part. Leaves bluish-green on upper surface, paler beneath; basal numerous with lamina 2.5– 10.0 × 2–5 cm, broadly elliptical or ovate, the earliest sometimes subrotund, rounded-mucronulate to more or less acute, subentire to denticulate or shallowly undulatedentate, cuneate or abruptly contracted at base to short or medium, densely long-hairy petioles; cauline (0–)1(−2), with lamina lanceolate or ovate, acute at apex, entire or denticulate, narrowed to the sessile base, the upper if present bract-like; all with short to medium, subsetiform, pale simple eglandular hairs on the upper surface, and medium to long, pale simple eglandular hairs beneath, particularly on the midrib and on the margins, sometimes the cauline with a few stellate hairs beneath. Inflorescence with 1–7 capitula, furcate-corymbose, sometimes with a long, lower branch; peduncles with numerous stellate hairs, numerous, medium to long, pale, dark-based simple eglandular hairs and few

39. Hieracium to fairly numerous, short, dark glandular hairs. Capitula 40–60 mm in diameter, broadly rounded-based. Involucral bracts incumbent in bud, 6–13 × 1.2–1.5 mm, blackishgreen, the inner with paler margins, all narrowly linearlanceolate, gradually narrowed to a long, acute apex, with dense, medium to long, pale, dark-based simple eglandular hairs with few to fairly numerous, short, dark glandular hairs intermixed, and a few stellate hairs. Ligules pale yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Cliff ledges, rocky slopes and streamsides up to 800 m, often on limestone or basalt. Common in northern England and central Scotland, and occasionally further north in Sutherland and Orkney Islands; widespread in Ireland where it is often a short plant. Endemic. 165. H. jaculifolium (F. Hanb.) Roffey Javelin-leaved Hawkweed H. anglicum var. jaculifolium F. Hanb.; H. olivaceum subsp. jaculifolium (F. Hanb.) Zahn; H. ampliatum var. jaculifolium (F. Hanb.) Pugsley Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, yellowish-green, sometimes reddish towards the base, robust, striate, with dense, long, pale, wavy simple eglandular hairs at the very base and few upwards, and numerous stellate hairs in the upper part. Leaves slightly bluish-green on upper surface, paler beneath; basal numerous, the outer with lamina 3.5–10.0 × 2–6 cm, broadly elliptical, ovate or subrotund, obtuse-mucronate to subacute, subentire to mammiform-dentate in the lower half and subtruncate or abruptly contracted, the inner with lamina 8–12 × 3–6 cm, lanceolate to broadly ovate, more or less acute at apex, with mammiform or aquiline-mammiform teeth in the lower half and abruptly contracted at base, all with very long, slender petioles up to 14 cm with dense, long, wavy simple eglandular hairs; cauline 1–2, with lamina lanceolate, acuminate at apex, subentire to dentate, with a short petiole; all glabrous or with a few simple eglandular hairs on the upper surface, with numerous, short to long, pale simple eglandular hairs on the lower surface. Inflorescence with 2–10 capitula, furcate-corymbose, sometimes with a long lower branch; peduncles with dense stellate hairs, more or less numerous, short to medium, pale simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Capitula 15–20 mm in diameter, with broad rounded bases. Involucral bracts porrect in bud, 6– 12 × 1.0–1.5 mm, greyish-green, the inner with paler margins, all linear-lanceolate, subacute at apex, with dense stellate hairs to the very tip of the bracts, with numerous to dense, short to medium, pale, dark-based simple eglandular hairs and few to numerous, short glandular hairs intermixed. Ligules poorly developed (stylose), with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Anna Well and Tilberthwaite Gill in Westmorland and Walton Crag, Keswick in Cumberland. Endemic.

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166. H. ampliatum (W. R. Linton) Ley Shaggy-stalked Hawkweed H. sarcophyllum var. ampliatum W. R. Linton; H. sagittatum subsp. sarcophyllum var. ampliatum (W. R. Linton) Zahn; H. anglicum var. calcaratum E. F. Linton; H. olivaceum subsp. jaculifolium subvar. calcaratum (E. F. Linton) Zahn; H. jaculifolium var. calcaratum (E. F. Linton) Roffey Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, yellowish-green, sometimes suffused brownishpurple below, slender to robust, striate, with dense, long, pale, wavy simple eglandular hairs at the very base and with very few up the remainder of the stem and numerous stellate hairs in the upper part. Leaves rather bluish-green on upper surface, paler beneath; basal few to numerous, the outer with lamina 2–5 × 2–5 cm, broadly elliptical to subrotund, rounded-mucronulate at apex, entire to denticulate or rarely dentate and rounded or subtruncate at base, the inner 5–9 × 2–5 cm, elliptical, oblong or lanceolate, obtuse to acute at apex, obscurely denticulate to dentate and subtruncate or abruptly contracted at base, the petioles up to 7 cm, with dense, long, wavy simple eglandular hairs; cauline 0–1(−2), with lamina lanceolate, acuminate at apex, entire to sharply dentate, abruptly contracted at base, petiolate; all glabrous or with scattered, short to medium simple eglandular hairs on upper surface, with numerous short to long, pale simple eglandular beneath, especially on the veins, and on the margins. Inflorescence with 2–10 capitula, furcate-corymbose, sometimes with a long lower branch; peduncles with dense stellate hairs, numerous short to medium, pale, dark-based simple eglandular hairs and sometimes an occasional, short glandular hair. Capitula 35–50 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 7–15 × 1.5–1.8 mm, dark greyishgreen, the inner with paler margins, all linear-lanceolate, obtuse to subacute at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs and fairly numerous stellate hairs particularly on the margins. Ligules yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and slopes and by streams. Common on the limestone of northern England. Endemic. 167. H. cyclicum P. D. Sell Round-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and few to numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves rather pale bluish-green on upper surface, paler beneath; basal numerous, the outer with lamina 2–8 × 2–5 cm, subrotund or broadly ovate, rounded-mucronate at apex, subentire or denticulate in the upper half, dentate below, often with mammiform or aquiline-mammiform teeth, and subtruncate or broadly cuneate at base, the inner with lamina 5–10 × 3.0–5.5 cm, lanceolate, ovate or elliptical, usually acute at apex, entire or denticulate in the upper half, usually

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dentate with narrow teeth below, broadly cuneate or subtruncate at the base, the petioles up to 5 cm, with dense, long, pale simple eglandular hairs; cauline 1–2, the lower like the inner basal with the petiole semiamplexicaul, the upper bract-like; all with few, medium, pale simple eglandular hairs on both surfaces. Inflorescence with 1–9 capitula, furcate-corymbose; peduncles long, with dense stellate hairs, numerous, short, dark glandular hairs and numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 30–35 mm in diameter, with rounded bases. Involucral bracts porrect in bud, 5–13 × 1.0–1.2 mm, blackish-green, the inner with narrow, paler margins, all linear-lanceolate, gradually narrowed to a more or less acute apex, with numerous medium, pale, dark-based simple eglandular hairs, few to fairly numerous, short, dark glandular hairs, and few stellate hairs at the base. Ligules yellow, with numerous, short simple eglandular hairs at apex. Styles yellow or slightly discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 6–7. Apomictic. Native. Mountain and sea cliffs. Sutherland and Rossshire probably extending into Argyllshire. Endemic. 168. H. ampliatiforme P. D. Sell Small-flowered Hawkweed H. ampliatum var. gracilius Pugsley Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes purplish below, slender to robust, striate, with dense, long, pale simple eglandular hairs at the very base and few, medium but similar hairs upwards, few to numerous, minute glandular hairs and occasional stellate hairs upwards. Leaves slightly bluish-green on upper surface, paler beneath; basal few to numerous, the outer with lamina 4–7 × 2.5–4.0 cm, narrow to broad elliptical or ovate, rounded-obtuse at apex, subentire, denticulate or with a few shallow teeth and abruptly contracted at base, the inner with lamina 5–10 × 2.5–4.0 cm, lanceolate, lanceolate-oblong or elliptical, obtuse to more or less acute, subentire to shallowly mammiform-dentate and abruptly contracted at base, all with medium to long petioles up to 7 cm with few to numerous, long, pale simple eglandular hairs; cauline usually 1, the lamina lanceolate, acute at apex, entire or with a small tooth, shortly petiolate; all glabrous or with a few simple eglandular hairs on the upper surface, with numerous short to medium, pale simple eglandular hairs on the lower surface and margin. Inflorescence with 1–5 capitula, furcate-corymbose; peduncles with dense stellate hairs, numerous, short to medium, dark or dark-based simple eglandular hairs and few to fairly numerous, short glandular hairs. Capitula 15–20 mm in diameter, with broad, rounded bases. Involucral bracts porrect in bud, 6–15 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all rather narrow linear-lanceolate, gradually narrowed to a more or less acute apex, with numerous, short to medium, dark or dark-based simple eglandular hairs and numerous, short, dark glandular hairs intermixed, with stellate hairs on the margins. Ligules yellow, with numerous very short simple eglandular hairs at apex, poorly developed (stylose). Styles discoloured. Receptacle pits with mar-

gins fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Upper Teesdale in Co. Durham, Dove Crag, Skelwith Force, Striding Edge and Dollywaggon Pike in Westmorland. Endemic. 169. H. hebridense Pugsley Hebridean Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, sometimes tinted purplish at the base, more or less slender, with dense, long, pale simple eglandular hairs at the very base, becoming shorter and much fewer upwards, with numerous stellate hairs in the upper part. Leaves pale green on upper surface, paler and slightly bluish beneath and sometimes flushed purplish; basal few to fairly numerous, the outer with lamina 5–8 × 3– 4 cm, broadly elliptical, obtuse-mucronate at apex, subentire or denticulate and rounded or shortly cuneate at base, the inner with lamina 4–10 × 3.0–4.5 cm, lanceolate, ovatelanceolate or elliptical, more or less acute at apex, denticulate or shallowly sinuate-dentate and sometimes with 2 larger teeth towards the cuneate or abruptly contracted base, the petioles short to fairly long and densely longhairy; cauline 0–2, the lower often like the inner basal and petiolate, the upper with lamina linear-lanceolate and sessile; all with simple eglandular hairs. Inflorescence with 2–9 capitula, furcate-corymbose; peduncles with numerous to dense stellate hairs, few to numerous, short to medium, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 6–13 × 1.5–1.8 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, medium to long, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs. Ligules medium yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and streamsides; in the few localities where the rock is recorded it is limestone. Probably common in the Hebrides and in scattered localities in central Scotland, Sutherland and in Co. Leitrim and Co. Down in Ireland. Endemic. 170. H. flocculosiforme P. D. Sell Small-toothed Hawkweed H. flocculosum var. denticulatum Pugsley, non H. denticulatum Sm. Phyllopodous perennial herb with a branched stock. Stem 20–55 cm, pale yellowish-green, sometimes reddish at base, slender to slightly robust, flexuous, striate, with rather scattered, medium to long, pale simple eglandular hairs and stellate hairs mainly in the upper part. Leaves bluish-green, sometimes spotted and blotched on upper surface, paler beneath; basal few, the lamina 1.5–10 × 1.0–4.5 cm, elliptical to broadly elliptical to subrotund, rounded-obtuse to subacute at apex, denticulate to sparsely dentate, cuneate at base to fairly short, long hairy petioles; cauline 1–2, the lamina lanceolate or ovate, acute at apex, subentire to denticulate, sometimes with a few teeth, with a short,

39. Hieracium winged, semiamplexicaul petiole; all with numerous stellate hairs on both surfaces or only beneath, few to numerous, short to long, pale simple eglandular hairs beneath, mainly on the midrib and on the margin. Inflorescence with 2–5 capitula, furcate-corymbose; peduncles long, suberect, with dense stellate hairs, fairly numerous, short to medium, pale, dark-based simple eglandular hairs and occasional to few, short, dark glandular hairs. Capitula 30–35 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 4–13 × 1.0–1.5 mm, blackish-green, linear-lanceolate, drawn out to a more or less acute apex, with numerous to dense, short to medium, pale, dark-based simple eglandular hairs, and numerous stellate hairs at the base and on the margins, and sometimes a few, short, dark glandular hairs. Ligules pale yellow, glabrous at tip. Styles yellow. Receptacle pits with margins fibrillous-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Streamsides and one locality on limestone cliffs, up to 850 m. A few scattered localities in Aberdeenshire, Banffshire, Inverness-shire and the Outer Hebrides. Endemic. 171. H. flocculosum Backh. ex Bab. Stellate-haired Hawkweed H. olivaceum subsp. flocculosum (Backh. ex Bab.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often brownish-purple towards the base, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, numerous to dense stellate hairs throughout and a few, short, dark glandular hairs above. Leaves dull glaucous green, sometimes spotted and blotched brownish-purple on upper surface, paler and sometimes suffused purplish beneath; basal few, with lamina 3–9 × 1.0–4.0 cm, subrotund, broadly elliptical to narrowly elliptical, ovate or lanceolate, rounded-obtuse to acute at apex, entire to denticulate, sometimes with a few, small teeth at the base, rounded or cuneate at base to medium, densely long-hairy petioles; cauline 0–2, the lamina lanceolate to ovate, acute at apex, entire or denticulate, with a short, winged semiamplexicaul petiole; all with dense stellate hairs on both surfaces and with fairly numerous, short to long, pale simple eglandular hairs on the lower surface and some with a few on the upper. Inflorescence with 1–8 capitula, furcate-corymbose; peduncles long, suberect, with dense stellate hairs, numerous, short and medium, pale, dark-based simple eglandular hairs and numerous, short glandular hairs. Capitula 40–60 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 5–16 × 1.0–1.5 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to a long-acute apex, with dense, medium to long, pale dark-based simple eglandular hairs, few to fairly numerous, short, dark glandular hairs and stellate hairs near the base and apex. Ligules pale yellow, with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. Cliff ledges and slopes and by streams. A few localities on the Yorkshire limestone and scattered over central and north Scotland. Endemic.

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172. H. flocculipubens P. D. Sell Floccose Hawkweed H. flocculosum var. angustifolium Pugsley, non H. angustifolium Hoppe; H. eustales auct. Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, sometimes brownishpurple at base, slender to more or less robust, with few to fairly numerous, long, pale simple eglandular hairs, small stellate hairs throughout, and rarely a solitary small glandular hair above. Leaves bluish-green on upper surface, paler beneath; basal few, the outer with lamina 2–5 × 1.5–2.5 cm, elliptical, obtuse at apex, entire or denticulate and rounded at base to a short petiole, the inner with lamina 7–11 × 2.5– 3.5 cm, ovate-lanceolate, lanceolate or elliptical, subobtuse to more or less acute at apex, subentire or denticulate, sometimes with small teeth and cuneate or attenuate at base to long, long-hairy petioles; cauline 1–2, the lower with lamina lanceolate, acute at apex, entire, denticulate or with an occasional tooth and with a long, winged petiole, the upper small or bract-like; all with more or less numerous, small stellate hairs on both surfaces, few, medium to long, pale simple eglandular hairs on the lower surface and more numerous on the margins. Inflorescence with 2–5 capitula, furcatecorymbose; peduncles long, suberect, with numerous to dense stellate hairs, numerous, short to medium, pale, sometimes dark-based simple eglandular hairs and sometimes an occasional, short, dark glandular hair. Capitula 30–40 mm in diameter, with a broad rounded base. Involucral bracts incumbent in bud, 6–12 × 1.0–1.5 mm, dark greyish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with dense, short to medium, pale, dark-based simple eglandular hairs, numerous stellate hairs especially at the base and on the margins and sometimes an occasional, short, dark glandular hair. Ligules pale yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins fimbriatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7–8. Apomictic. Native. Known only from rocks by the Spinkwee River in the Mourne Mountains, and a stream in Tollymore Park, both in Co. Down, and a few scattered localities by streams in Perthshire, Aberdeenshire, Argyllshire, Ross-shire and Sutherland. Endemic. 173. H. shoolbredii E. S. Marshall Shoolbred’s Hawkweed H. anglicum var. longibracteatum F. Hanb.; H. olivaceum subsp. flocculosum subvar. longibracteatum (F. Hanb.) Zahn; H. mougeotii subsp. skyense Zahn; H. skyense (Zahn) Roffey Phyllopodous perennial herb with a branched stock. Stem 15–50 cm, pale yellowish-green, occasionally slightly purplish towards the base, slender to robust, striate, with few to numerous stellate hairs particularly above, few to numerous, medium to long, wavy, pale simple eglandular hairs, dense near the very base, few to numerous in the lower half and few or absent above, sometimes an occasional, small glandular hair above. Leaves bluish-green to medium green on upper surface, paler and greyish or bluish beneath and sometimes flushed purplish; basal few to numerous, with lamina

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4–11 × 2–7 cm, the outer broadly elliptical to subrotund, rounded or obtuse-mucronate at apex, entire or denticulate and rounded or subtruncate at base, the inner broadly elliptical, lanceolate or ovate-lanceolate, subobtuse to acute at apex and denticulate or shallowly dentate towards the base which is rounded, subtruncate or cuneate, the petioles up to 5 cm and with long, wavy simple eglandular hairs; cauline 0–1, the lamina ovate, lanceolate or linear, finely acute at apex, entire or denticulate, sessile and semiamplexicaul; all glabrous or nearly so on upper surface, with numerous, short to medium, pale simple eglandular hairs beneath, especially on the midrib, and on the margin, usually with numerous stellate hairs beneath. Inflorescence with 1–6(−15 cult.) capitula, furcate-corymbose, sometimes with a long lower branch; peduncles with numerous stellate hairs and numerous, short, dark glandular hairs. Capitula 30–50 mm in diameter, broadly rounded at base. Involucral bracts porrect in bud, 8–15 × 1.2–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, gradually narrowed to a fine, acute apex, with numerous stellate hairs especially in the lower part, numerous, unequal, short, dark glandular hairs and fewer, short or medium, pale, dark-based simple eglandular hairs. Ligules pale yellow, with shortly hairy tips. Styles discoloured. Receptacle pits with margins dentatefimbriate. Achenes 3.0–3.5 mm, blackish-purple. Flowers 6–7. Apomictic. Native. Rock-ledges and boulders, stream banks, river gorges, grassy banks and old quarries on granite, schist and limestone up to 750 m. A widespread and common species of central and north Scotland and the Hebrides. Endemic. Named after William Andrew Shoolbred (1852–1928). Section 14. Oreadea (Fr.) Dahlst. Taxon Oreadea Fr.; Subsection Oreadea (Fr.) Rouy Phyllopodous perennial herbs. Leaves often more or less bluish-green, the basal usually few, the cauline 2–8, all with margins with rigid simple eglandular hairs which often extend on to the upper surface. Inflorescence with 1–9 capitula on long, straight, erect peduncles. Capitula usually large. Involucral bracts often porrect in bud, up to 14 mm, mostly obtuse at apex and rather broad. Ligules usually glabrous-tipped. Styles mostly yellow. Receptacle pits with margins usually subulate-dentate. Achenes reddish-black. Flowers 5–7. Cliff ledges, scree, riverside rocks, grassy slopes and sandhills, usually on basic soils. Mountains and coastal areas in Scotland and Wales and Cos. Kildare, Wicklow, Dublin, Donegal, Down and Antrim in Ireland. Occurs widely in Europe and extends into south-west Asia. The section is characterised by its few basal leaves and 2–8 cauline leaves, all with rigid simple eglandular hairs, few large heads on long, erect peduncles and usually yellow styles. The species are similar in general appearance to the species of the Section Cerinthoidea which differ in their usually more numerous basal leaves, softer hairs and darker styles. 174. H. scoticum F. Hanb. Scottish Hawkweed H. scoticiforme Dahlst. Phyllopodous perennial herb with a branched stock. Stem 25–60 cm, pale yellowish-green, often suffused with

reddish-purple, slender and flexuous, striate, with numerous, medium to long, pale, deflexed simple eglandular hairs below, becoming few or absent upwards, and numerous stellate hairs in the upper part. Leaves medium to bluishgreen on upper surface, paler and bluish beneath, sometimes tinted reddish-purple; basal few and often shrivelled at anthesis, the lamina 5–9 × 2.5–4.0 cm, ovate or elliptical, rounded or mucronulate at apex, subentire or denticulate, cuneate or attenuate at base, the petioles up to 5 cm, with numerous, long, pale, deflexed simple eglandular hairs; cauline 3–8(−12 cult.), the lamina 2–14 × 1–6 cm, gradually decreasing in size upwards, elliptical, ovate or lanceolate, acute at apex, characteristically sharply denticulate or shallowly dentate, rarely with a few larger teeth, cuneate at base, the lower petiolate, the upper sometimes bract-like; all glabrous or nearly so on upper surface, with scattered, medium or long, pale simple eglandular beneath, especially on the midrib and medium to long, pale, stiff simple eglandular hairs on the margin. Inflorescence with 1–8(−12 cult.) capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, numerous, short and medium, pale simple eglandular hairs and a few, minute, dark glandular hairs. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4– 12 × 1.0–1.5 mm, dark green, the inner with paler margins, all broadly linear-lanceolate, obtuse at apex, with numerous short and medium, pale, dark-based simple eglandular hairs, few, minute, dark glandular hairs and fairly numerous stellate hairs on the margins. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulatedentate. Achenes 4.0–4.5 mm, reddish-black. Flowers 6–8. Apomictic. Native. Ledges of low cliffs and riverside rocks. Coastal areas of Sutherland and Caithness in north Scotland and Co. Donegal in Ireland. Faeroes. 175. H. subscoticum P. D. Sell Tapered-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–60 cm, pale yellowish-green, often tinted reddish below, slender to more or less robust, with numerous, medium to long, pale, wavy simple eglandular hairs below, sometimes throughout and sometimes fewer and shorter upwards, and more or less numerous stellate hairs in the upper part. Leaves dark green to slightly glaucous on upper surface, paler beneath; basal few, often shrivelled at anthesis, the lamina 5–10 × 1–3 cm, elliptical, oblong-elliptical or lanceolateoblong, narrowed to an obtuse or acute apex, subentire or denticulate, attenuate at base, the petioles up to 5 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 2–6, gradually decreasing in size upwards, the lower with lamina lanceolate, elliptical, oblong-elliptical or broadly elliptical, usually more or less acute at apex, dentate, with small, sharp teeth, attenuate at base and with winged petioles up to 6 cm long, the upper with lamina lanceolate, ovate or elliptical, long-acute at apex, with small, sharp teeth, and rounded or cuneate to a sessile base; all glabrous or nearly so on upper surface, with numerous, short to long, pale, subrigid simple eglandular hairs below and on the margin. Inflorescence with 1–5(−7) capitula, furcatecorymbose; peduncles long and straight, with dense stellate

39. Hieracium hairs, numerous short or medium, pale, dark-based simple eglandular hairs and few, very short, dark glandular hairs. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 5–13 × 1.2–1.7 mm, dark green, the inner paler at the margins, all linear-lanceolate, obtuse at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, few, very short, dark glandular hairs and a few stellate hairs towards the base. Ligules lemon yellow, glabrous-tipped. Styles dull yellow. Receptacle pits with margin shortly dentate. Achenes 3.5– 4.0 mm, reddish-black. Flowers 6–7. Apomictic. Plants of this species have previously been called H. scoticum, but they differ from that species by their narrower, attenuate-based leaves. The Shetland plant has hairy upper surfaces to the leaves and may be slightly different. Native. Rocks, grassy slopes and sandy places. Coastal areas of Sutherland and Caithness, the Orkney Islands and a grassy field at Heylor in the Shetland Islands. Endemic. 176. H. apheles P. D. Sell Hepste Hawkweed H. septentrionale var. simplex Ley, non H. simplex Viv. Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, sometimes purplish towards the base, with few to numerous, medium to long, pale simple eglandular hairs below, becoming fewer and shorter or absent upwards, and numerous stellate hairs in the upper part. Leaves medium yellowish-green on the upper surface, paler beneath; basal few, the lamina 6–12 × 2.0– 4.5 cm, lanceolate to broadly or narrowly elliptical, gradually narrowed to a more or less acute apex, denticulate to remotely dentate, the teeth more or less mammiform, sometimes aquiline-mammiform, attenuate at base, the petioles up to 6 cm, with numerous, long, pale simple eglandular hairs; cauline 2–5, the lower similar to basal but narrower, usually shortly petiolate, rarely longer, the upper with lamina lanceolate or linear, acute at apex, entire or dentate, shortly petiolate or sessile; all glabrous or nearly so on upper surface, with numerous, medium to long, subrigid simple eglandular hairs beneath and on the margin. Inflorescence with 2–6 capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs and a few, short to medium, pale simple eglandular hairs. Capitula 20–30 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 5–12 × 1.5–2.0 mm, dark green, the inner with paler margins, all broadly linear-lanceolate, narrowed to an obtuse apex, with numerous stellate hairs especially on the margins, numerous, short or medium, pale, dark-based simple eglandular hairs and a few, dark glandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, blackishred. Flowers 6–7. Apomictic. Native. Known only from rocks in the Hepste Glen, Breconshire. Endemic. 177. H. sowadeense P. D. Sell Sowa Dee Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, usually much suffused with brownish-purple, slender and flexuous to robust, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and dense stel-

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late hairs in the upper part. Leaves medium green on upper surface, paler and often purplish beneath; basal few and sometimes shrivelled at anthesis, the lamina 2–8 × 1.5–4.0 cm, elliptical, lanceolate or ovate, obtuse to acute at apex, entire or remotely denticulate and cuneate or attenuate at base, the petiole up to 3 cm, with dense, long, wavy simple eglandular hairs; cauline 3–6, the lower like the basal, sessile or shortly petiolate, the remainder gradually becoming smaller upwards, the uppermost with lamina linear or bract-like; all glabrous or nearly so on upper surface, with numerous, short to long, pale, rather stiff simple eglandular hairs beneath and on the margins. Inflorescence with 1–6 capitula, furcate-corymbose; peduncles long and straight, with numerous stellate hairs and fairly numerous, short to long, pale simple eglandular hairs. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 5–13 × 1.5–2.0 mm, dark olive-green, the inner paler at the margin, all lanceolate, obtuse at apex, with numerous, short to medium, pale simple eglandular hairs and a few, scattered stellate hairs along the margin. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margin shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only from grassy crags by the Sowa Dee near Sandwick in a remote part of Mainland in the Orkney Islands. Endemic. 178. H. pusillifolium P. D. Sell Penwyllt Hawkweed H. scoticum var. microphyllum Pugsley, non H. microphyllum Hyl. Phyllopodous perennial herb with a branched stock. Stem 20–50(−60) cm, pale yellowish-green, often suffused purplish, slender, flexuous, striate, with medium to long, pale, rather stiff simple eglandular hairs below, becoming fewer and shorter upwards. Leaves medium green on upper surface, pale beneath, sometimes tinted red; basal few to numerous, the lamina 4–8 × 2–5 cm, elliptical, broadly elliptical, ovate, broadly ovate or subrotund, roundedmucronulate or abruptly acute at apex, subentire, denticulate or sinuate-dentate, the teeth apiculate, abruptly contracted or cuneate at base, the petioles short, with dense, long, wavy, pale simple eglandular hairs; cauline 4–7, gradually decreasing in size upwards, lower like basal and sessile, the upper with lamina ovate, acute at apex, sharply denticulate, sessile; all glabrous or few to numerous, short to medium, stiff simple eglandular hairs on the upper surface and margins, and few to numerous, short to medium simple eglandular hairs beneath particularly on the midrib. Inflorescence with 2–9(numerous in cult.) capitula, furcatecorymbose; peduncles long and straight, with dense stellate hairs and numerous, short to medium, pale simple eglandular hairs. Capitula 25–30 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 5– 10 × 1.0–1.2 mm, dark green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, occasional minute glandular hairs and numerous stellate hairs on the margins towards the base. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 6–7. Apomictic.

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Native. Known only from limestone cliffs at Penwyllt in Breconshire. Endemic. 179. H. aggregatifolium P. D. Sell Rosette Hawkweed H. scoticum var. rosulare Pugsley, non H. rosulare Brenner; H. scoticum var. submaculatum Dahlst.; H. carpathicum subsp. euchrysostylum auct. Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, often tinted purple, slender and flexuous, striate, with numerous to dense, long, pale simple eglandular hairs below, becoming fewer or absent upwards, and with few to numerous stellate hairs upwards. Leaves medium green on upper surface sometimes slightly spotted, paler beneath; basal numerous, the lamina 4–8 × 1.5–4.0 cm, ovate, narrowly elliptical, lanceolate or oblong-lanceolate, mainly acute, but sometimes obtuse at apex, shallowly dentate, the teeth short and apiculate, rarely with a few larger teeth, cuneate or abruptly contracted at base, with a short, winged petiole with dense, long, pale simple eglandular hairs; cauline 2–4, rapidly decreasing in size, the lower very near the basal and similar to them, the upper with lamina lanceolate, acute at apex, denticulate and petiolate or bract-like; all more or less glabrous on upper surface or with a few hairs, with few to medium, pale simple eglandular hairs beneath and longer ones on the midrib, and short to medium, rather stiff simple eglandular hairs on the margin. Inflorescence with 2–6 capitula, furcatecorymbose; peduncles long and straight, with numerous stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and an occasional, minute glandular hair. Capitula 30–40 mm in diameter, sometimes flowers poorly developed, subtruncate at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.5 mm, dark green, the inner with paler margins, all broad linear-lanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs and few, minute, dark glandular hairs. Ligules yellow, often poorly developed, glabroustipped. Styles yellow or slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rocky outcrops, grassy slopes and sandhills by the sea. Coasts of Sutherland, Caithness and Orkney Islands. Endemic. 180. H. breconicola P. D. Sell Beacons Hawkweed Phyllopodous perennial herb with a branched stock. Stem 35–55 cm, pale yellowish-green, sometimes reddish below, slender, striate, with numerous, medium or long, pale simple eglandular hairs below, which become fewer upwards or sometimes none at all, and numerous stellate hairs in the upper part, without glandular hairs. Leaves medium green on upper surface, paler beneath, often suffused reddishpurple, subcoriaceous; basal few to fairly numerous, the lamina 3–10 × 2–4 cm, elliptical, lanceolate, ovate or oblong-elliptical, narrowed at apex, but obtuse-mucronate, entire to remotely denticulate, rarely with an occasional tooth, cuneate or attenuate at base, the petioles up to 4 cm and with numerous, long, pale, wavy simple eglandular hairs; cauline 2–5, the lower similar to inner basal and long petiolate, the upper with lamina ovate, acute at apex,

denticulate and sessile; all with short to medium, stiff, pale simple eglandular hairs on the upper surface and margin or with the upper surface glabrous or nearly so, and more numerous, longer hairs beneath, especially on the midrib. Inflorescence with 1–7 capitula, furcate-corymbose, sometimes with a long, lower branch; peduncles long and straight, with dense stellate hairs, numerous, short and medium, pale simple eglandular hairs and a few, minute, dark glandular hairs. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts porrect in bud, 3–12 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all broad linear-lanceolate, obtuse at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, very few, short, dark glandular hairs and a few stellate hairs on the margin. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 4.0–4.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliffs of Fan Nedd and above Llyn-y-fan-fawr in Breconshire and Llyn-y-fan-fechan in Carmarthenshire. Endemic. 181. H. pseudozetlandicum Roffey Sand Hawkweed H. caledonicum var. pseudozetlandicum (Roffey) Pugsley; H. shoolbredii Zahn nom. nud., non E. S. Marshall Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, sometimes purplish at the very base, slender and flexuous, striate, with numerous, long, pale simple eglandular hairs below, becoming shorter and fewer upwards, and numerous stellate and occasional short, dark glandular hairs in the upper part. Leaves dull dark green on upper surface, paler beneath; basal few, the lamina 4–10 × 2–4 cm, broadly elliptical or ovate, acute at apex, shallowly dentate, the teeth with long apiculate tips, cuneate at base, petioles up to 4 cm, with numerous, long, pale simple eglandular hairs; cauline 2–3, the lower like the basal and shortly petiolate, the upper with lamina lanceolate or linear, acute at apex, denticulate and sessile; all glabrous or nearly so on upper surface and with few to numerous, medium to long, pale, subrigid simple eglandular hairs below and on the margins. Inflorescence with 2–3 capitula, furcate; peduncles straight, with dense stellate hairs, numerous short to medium, pale, dark-based simple eglandular hairs and an occasional, dark, minute glandular hair. Capitula 30–45 mm in diameter, truncate at base. Involucral bracts incumbent in bud, 5–14 × 1.2– 1.5 mm, blackish-green, the inner with paler margins, all broadly linear-lanceolate, obtuse at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, few, very short, dark glandular hairs and a few stellate hairs along the margins. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Sandy, grassy slopes and sea-cliffs. The Bettyhill to Farr area of Sutherland. Endemic. 182. H. scottii P. D. Sell Scott’s Hawkweed Phyllopodous perennial herb with a branched stock. Stem 15–30 cm, pale yellowish-green, often purplish at base, slender and flexuous, striate, with numerous, medium to

39. Hieracium long, pale, wavy simple eglandular hairs which become fewer and shorter upwards, and dense stellate and a few, minute, dark glandular hairs in the upper part. Leaves medium yellowish-green on the upper surface, paler beneath, sometimes tinted reddish; basal few, the lamina 4–9 × 1.5–3.5 cm, lanceolate or elliptical, roundedmucronulate to acute at apex, remotely denticulate, cuneate to attenuate at base, the petioles up to 4 cm, with numerous, long, wavy, pale simple eglandular hairs; cauline 2– 4, quickly decreasing in size upwards, the lower like the basal and petiolate, the upper with lamina lanceolate or linear, acute at apex, denticulate or entire and sessile; all with medium to long, pale, subrigid simple eglandular hairs on both surfaces and the margins and a few stellate hairs on the under surface. Inflorescence with 1–8 capitula, furcate; peduncles erect and straight, with dense stellate hairs, numerous short or medium, pale simple eglandular hairs and numerous, minute, dark glandular hairs. Capitula 30–40(−50 cult.) mm in diameter, broadly rounded to subtruncate at base. Involucral bracts incumbent in bud, 5–14 × 1.5–2.0 mm, dark green, the inner with paler margins, all broadly lanceolate, obtuse at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, numerous very short and short, dark glandular hairs and numerous stellate hairs especially on the margins. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 7–8. Apomictic. Native. Rocky geos below Windy Scord, Voe of Snarraness and rocks and pastures west of Bousta, Sandness, both in the Shetland Islands. Endemic. Named after Walter Scott for his large contribution to the flora of Shetland. 183. H. farrense F. Hanb. Farr Hawkweed H. caesium subsp. farrense (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–55 cm, pale yellowish-green, sometimes reddish below, with numerous, medium to long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and with dense stellate hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal few to several, the lamina 5–11 × 2–4 cm, lanceolate, elliptical, oblong-ovate, more or less acute at apex, subentire or denticulate, sometimes with an occasional larger tooth, attenuate at base, the petioles up to 6 cm, with long, pale simple eglandular hairs; cauline 2–3, the lower like the basal, the upper with lamina lanceolate, acute at apex and petiolate, the uppermost with the lamina lanceolate, acute at apex and entire or bract-like; all with short and medium, pale, subrigid simple eglandular hairs on the upper surface, and longer ones beneath and on the margin. Inflorescence with 1–4 capitula, furcate-corymbose; peduncles straight, with dense stellate hairs, numerous short and medium, pale simple eglandular hairs and a few, very short, dark glandular hairs. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.3 mm, dark green, the inner with paler margins, all linearlanceolate, obtuse at apex, with dense stellate hairs particularly on the margins, numerous short to medium, pale, dark-based simple eglandular hairs and a few, short, dark

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glandular hairs. Ligules deep yellow, glabrous-tipped. Styles yellow to slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Banks above the River Naver at Bettyhill, near Farr, Sutherland. Endemic. 184. H. angustisquamum (Pugsley) Pugsley Red-tinted Hawkweed H. holophyllum var. angustisquamum Pugsley Phyllopodous perennial herb with a branched stock. Stem 30–65 cm, pale yellowish-green, often suffused with reddish-purple, more or less robust, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter above, scattered stellate hairs throughout and an occasional minute glandular hair in the upper part. Leaves bright green on upper surface, paler and slightly bluish beneath, often suffused reddish-purple and sometimes also above; basal few, the outer with lamina 3.5–8.0 × 1.5–3.0 cm, elliptical, rounded-obtuse to acute at apex, entire or remotely denticulate and cuneate at base, the inner with lamina 5–14 × 2–6 cm, lanceolate, ovate, elliptical or oblongelliptical, narrowed to an obtuse or subacute apex, subentire, denticulate or with a few larger teeth and cuneate at base, the petiole up to 6 cm, with numerous, long simple eglandular hairs; cauline 2–4, the lower often large and similar to inner basal and usually more acute at apex and shortly petiolate, the upper with lamina lanceolate or ovate, acute at apex, denticulate or dentate, sessile or with a very short petiole; all glabrous or with a few scattered hairs on upper surface, with scattered, medium, pale simple eglandular hairs beneath, numerous short to medium, pale, rather stiff hairs on the margin and sometimes with stellate hairs on the cauline beneath. Inflorescence with 2–10 capitula, furcate-corymbose; peduncles usually long and straight, with dense stellate hairs and scattered short, dark glandular hairs and dark-based, medium simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all narrow linear-lanceolate, narrowed above, but obtuse, with numerous to dense stellate hairs especially on the margins and tufted at the apex, numerous, short to medium, darkbased simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Native. Cliff ledges and scree, perhaps always on limestone or at least basic rock. Cheddar Gorge, Somerset; Duncton Hill near Petworth in Sussex; near Talla Reservoir in Peebles-shire; Creag nam Fitheach in Argyllshire; near Ballochantuy in Kintyre; near Ledmore in Sutherland; Poulaphuca in Co. Wicklow and near Lucan in Co. Dublin. Endemic. 185. H. proximum F. Hanb. Caithness Hawkweed H. hanburyanum Zahn nom. illegit.; H. orarium var. erythraeum E. F. Linton nom. invalid.; H. erythraeum E. F. Linton ex Pugsley; H. orarium var. fulvum auct.

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Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often suffused brownishpurple towards the base, usually rather slender and flexuous but sometimes more or les robust, striate, with numerous, medium to long, pale, wavy simple eglandular hairs towards the base, becoming fewer and shorter upwards, and with dense stellate hairs in the upper part. Leaves pale or medium yellowish-green on the upper surface, paler beneath, some reddish-purple tinged or rimmed round the margin; basal few, the lamina 4.5–9.0 × 1.5–4.0 cm, ovate, lanceolate, elliptical or oblong-lanceolate, obtuse-mucronate to more or less acute at apex, subentire or denticulate to shallowly mammiform-dentate, cuneate at base, the petioles up to 6 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 2–4, lower like basal, acute at apex and petiolate, the upper with lamina lanceolate, long acute at apex, entire to dentate, cuneate at base and shortly petiolate or sessile, or bract-like; all with numerous, short to long, pale, rigid simple eglandular hairs on the upper surface and margins and similar longer, less rigid ones beneath. Inflorescence with 1–7(many cult.) capitula, furcate-corymbose; peduncles erect and straight, with dense stellate hairs and numerous, short and medium, pale simple eglandular hairs. Capitula 35–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 5–13 × 1.5–1.8 mm, dark green, the inner with paler margins, all broad linear-lanceolate, more or less acute at apex, with dense, short to long, pale simple eglandular hairs, numerous stellate hairs, often dense on the margins and a few hidden, very short glandular hairs. Ligules deep yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. The Melness plants have been called H. erythraeum, but no difference can be seen in plants from that locality. They have not, however, been seen in the field. Native. Rocky gorge near the sea at Reay and cliffs near the sea at Thurso in Caithness. The Melness plants of Sutherland occur on sandhills. The Irish material from Co. Kildare, Co. Donegal and Co. Antrim placed here by H. W. Pugsley has not been seen. Endemic. 186. H. subrude Arv.-Touv. Tain Hawkweed H. onosmoides subsp. subrude (Arv.-Touv.) Zahn; H. buglossoides var. subrude (Arv.-Touv.) Arv.-Touv.; H. buglossoides auct.; H. onosmoides var. buglossoides auct.; H. onosmoides auct. Phyllopodous perennial herb with a branched stock. Stem 30–70 cm, pale yellowish-green, rarely reddish at base, rather slender and flexuous, striate, with numerous, long, rather stiff simple eglandular hairs below becoming fewer upwards, and a few stellate hairs and minute glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal few to fairly numerous, the outer with lamina 4.0–7.5 × 1.0–2.5 cm, narrowly elliptical, rounded-obtuse at apex, subentire to remotely dentate and attenuate at base, the inner with lamina 7–15 × 2–5 cm, lanceolate, oblong-lanceolate, oblong-elliptical or oblong, narrowed to an obtuse-mucronate to acute apex, denticulate to dentate, with mammiform teeth, attenuate below; petioles usually long, up to 9 cm, with numerous, long, pale, wavy

simple eglandular hairs; cauline 1–5, the lower like the inner basal and long petiolate, the upper with lamina lanceolate or linear, long-acute at apex, dentate and to a short petiole or sessile; all with medium to long, pale, rigid simple eglandular hairs on both surfaces and the margin. Inflorescence with 2–12 capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, numerous very short and short, dark glandular hairs and occasional, short to medium simple eglandular hairs. Capitula 30–35 mm in diameter, truncate at base. Involucral bracts porrect in bud, 5–12 × 1.0–1.2 mm, dark green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, very short or short, dark glandular hairs, fairly numerous, short to medium, pale, dark-based simple eglandular hairs and some stellate hairs. Ligules yellow, glabroustipped. Styles yellow or slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Probably introduced as mainly on railways and roadsides. Railway banks at Tain and sandhills south of Balintore in Ross-shire, a railway embankment near Tyndrum in Perthshire and a dry bank of roadside above Uig in Skye. An E. S. Marshall herbarium sheet from Balintore as well as containing a typical plant of this species has one of H. beebyanum. Native of France (Basses-Alpes), Piedmont, Valais and west Germany, which also suggests the north Scotland plants are not native. 187. H. deganwyense Pugsley Deganwy Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale yellowish-green, often brownish-purple towards the base, slender and flexuous to more or less robust, striate, with numerous, long, rather stiff, pale simple eglandular hairs below, becoming much fewer above or almost absent, with a few stellate hairs in the upper part. Leaves dull medium bluish-green on upper surface, paler beneath; basal numerous, with lamina 2.5–9.0 × 1–4 cm, elliptical, ovate or lanceolate, narrowed to an obtuse-mucronate apex, subentire, denticulate or dentate, the teeth narrowly mammiform, cuneate or attenuate at base, the petioles up to 5 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 2–3, the lower with lamina lanceolate, ovate or elliptical, more or less acute at apex, dentate, narrowed at base and shortly petiolate or sessile, the upper with lamina linear, linear-lanceolate or bract-like and sessile; all with medium to long, rigid, bulbous-based, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 1–8 capitula, furcate-corymbose; peduncles long and straight, with numerous stellate hairs, numerous, minute, dark glandular hairs and few to numerous, short to medium, pale simple eglandular hairs. Capitula 25–35 mm in diameter, truncate at base. Involucral bracts 4.0–11.5 × 1.0–1.2 mm, olive to dark green, the inner with paler margins, linear-lanceolate, obtuse at apex, with numerous, short to medium, pale simple eglandular hairs, a few, minute, dark glandular hairs and a few stellate hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic.

39. Hieracium Native. Rock ledges. Cardiganshire and Caernarvonshire in Wales, Isle of Man, Northumberland in northern England and scattered localities in Scotland, from Kirkcudbright north to Ross-shire, probably always on basic rocks. Endemic. 188. H. ascendentidens P. D. Sell Harris Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, rarely slightly purplish at base, slender and flexuous, with few to numerous, medium to long, pale simple eglandular hairs and numerous stellate hairs and numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal numerous, with lamina 1.5–9.0 × 1.0– 3.5 cm, lanceolate to ovate, acute to acuminate at apex often with a long acumen, sharply dentate, the teeth apiculate and ascending, mostly very narrow, but some of them mammiform, cuneate to attenuate at base, the petioles up to 6 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–2, lower when present like basal and petiolate, upper with lamina narrowly lanceolate, acute and entire or bract-like; all with numerous, medium to long, pale, rigid simple eglandular hairs on both surfaces and the margins. Inflorescence with 2–6 capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, numerous very short and short (0.2–0.7 mm), dark glandular hairs and occasional, medium, pale simple eglandular hairs. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts 5–12 × 1.0–1.5 mm, olive-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to a slender, more or less acute apex, with numerous stellate hairs especially on the margins, numerous unequal, short to medium (0.3–1.0 mm), dark glandular hairs and more or less numerous, short to medium, pale, dark-based simple eglandular hairs. Ligules yellow, with very short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 6–7. Apomictic. The specimens from the two localities are very slightly different in that the Maarig River plant has rather more simple hairs on the involucral bracts than the holotype from Lingadale. Native. Riverside rocks. Lingadale and Maarig River on North Harris in the Outer Hebrides. Endemic. 189. H. beebyanum Pugsley Beeby’s Hawkweed H. schmidtii var. crinigerum auct. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, often purplish towards the base, slender and flexuous, striate, with dense, medium to long, pale, wavy simple eglandular hairs below, becoming fewer and shorter above, and numerous stellate hairs in the upper part. Leaves bluish-green on upper surface, paler beneath; basal rather numerous, with lamina 3.5–8.0 × 1.5– 3.0 cm, ovate or elliptical, rounded-mucronulate to acute at apex, subentire, denticulate or remotely dentate, cuneate at base, the petioles up to 3 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 2–4, lower similar to basal and shortly petiolate, upper with lamina lanceolate, long acute at apex, subentire or denticulate and shortly petiolate,

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or bractlike; all with numerous, medium to long, pale, rigid simple eglandular hairs on the upper surface and margins and numerous similar, but softer ones beneath. Inflorescence with 2–6 capitula, forming a corymb; peduncles straight and erect, with dense stellate hairs, numerous short to medium, pale simple eglandular hairs and an occasional, minute glandular hair. Capitula 20–30 mm in diameter, subtruncate at base. Involucral bracts 3–12 × 0.8–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short and medium, pale simple eglandular hairs, few stellate hairs and few or no, minute glandular hairs. Ligules yellow, glabrous or with few, very short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Craggy outcrops, cliff ledges and rocky streamsides. Shetland Islands, the Hebrides, Kintyre and Co. Wicklow, Co. Donegal and Co. Down in Ireland. Endemic. Named after William Hadden Beeby (1849–1910). 190. H. griffithii (F. Hanb.) F. Hanb. Griffith’s Hawkweed H. clovense var. griffithii F. Hanb.; H. sommerfeltii subsp. sommerfeltii var. griffithii (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, often purplish towards the base, slender and flexuous, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards, and a few stellate hairs and minute, black glandular hairs in the upper part. Leaves rather pale green and more or less spotted and blotched with dark purple on upper surface, paler beneath; basal numerous, with lamina 2–5(−8) × 1.0–2.5 cm, ovate, lanceolate or rarely elliptical, obtuse-mucronate to sharply acute at apex, denticulate or with a few, irregular, mammiform teeth, cuneate to attenuate at base, the petioles up to 3 cm, with numerous, long, pale, stiff simple eglandular hairs; cauline 2–4, the lower like the basal, the upper with lamina lanceolate, long-acute at apex, dentate and sessile; all with medium to long, pale, rigid simple eglandular hairs on both surfaces and the margins. Inflorescence with 2–8 capitula, furcate-corymbose; peduncles straight, with numerous stellate hairs, numerous short to medium, pale simple eglandular hairs and few to fairly numerous, very short, dark glandular hairs. Capitula 35–40 mm in diameter, subtruncate at base. Involucral bracts porrect in bud, 5–11 × 1.0–1.2 mm, dark green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, a few, short, dark glandular hairs and a few stellate hairs. Ligules yellow, with a very few, short hairs at apex. Styles yellow. Receptacle pits incise-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Native. Rocks and mountain streamsides. Known only from the Nant Francon Pass, Caernarvonshire. Endemic. Named after John Edwards Griffith (1843–1933). 191. H. orimeles F. Hanb. ex W. R. Linton Pale Hawkweed H. orimeles F. Hanb. nom. invalid.; H. saxifragum var. orimeles F. Hanb. nom. nud.; H. saxifragum var. orimeles

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(F. Hanb. ex W. R. Linton) F. Hanb.; H. saxifragum subsp. orimeles (F. Hanb. ex W. R. Linton) Zahn; H. orimeles var. obscurum Pugsley Phyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale yellowish-green, sometimes purplish below, usually slender but sometimes more or less robust, striate, with numerous, medium to long, pale, wavy, simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and sometimes a few, minute glandular hairs in the upper part. Leaves rather pale green, sometimes slightly bluish on upper surface, even paler beneath and sometimes flushed purplish; basal few, with lamina 1.5–12.0 × 1.0–3.5 cm, elliptical, oblong-elliptical or lanceolate-elliptical, mostly acute at apex but sometimes obtuse-mucronate, subentire or with a few, small teeth, cuneate to attenuate at base, the petioles up to 5 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 2–5, the lower large and similar to basal, the upper gradually getting smaller, with lamina oblong-lanceolate, drawn out to an acute apex, denticulate, narrowed at base and sessile, or bract-like; all with few to numerous, short to medium, pale, stiff simple eglandular hairs on the upper surface, more numerous and softer beneath and numerous, short to long, pale, stiff simple eglandular hairs on the margins, the cauline sometimes with stellate hairs beneath. Inflorescence with 1–6 capitula, furcate-corymbose; peduncles long and erect, with dense stellate hairs, fairly numerous, short to medium, pale simple eglandular hairs and a few, very short, dark glandular hairs especially just below the capitulum. Capitula 18–30 mm in diameter, subtruncate at base. Involucral bracts porrect in bud, 4–11 × 1.0– 1.2 mm, olive-green, at least the inner with paler margins, all linear-lanceolate, obtuse to acute at apex, with numerous, very short to medium, pale simple eglandular hairs, few stellate hairs towards the base and a few, minute glandular hairs. Ligules yellow, glabrous or with a few, very short simple eglandular hairs at the apex. Styles yellow to slightly discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–8. Apomictic. Small plants on scree look very different from those elsewhere in size. Large plants in Carmarthenshire look very like H. substrigosum, with which they grow. Native. Cliff ledges, streamside rocks and scree. Llyny-fan-fechan, Carmarthenshire, common in north Wales, scattered over northern England and central and western Scotland north to the Shetland Islands; recorded from Londonderry by H. W. Pugsley. Endemic. 192. H. filisquamum P. D. Sell Narrow-bracted Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–40 cm, pale yellowish-green, often purplish towards the base, slender and flexuous, striate, with dense, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards and with dense stellate hairs in the upper part, without glandular hairs. Leaves slightly bluish-green on upper surface, paler beneath and sometimes with a slight purplish tint; basal few, with lamina

6–10 × 2–3 cm, narrowly elliptical or lanceolate, acute or obtuse-mucronulate at apex, subentire or sinuatedenticulate, cuneate to attenuate at base, the petioles up to 3 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 2–3, lower like basal and often large, with a winged petiole, the upper with lamina narrower, long acute at apex, denticulate, narrowed at base and semiamplexicaul, or bract-like; all with numerous, short to long, pale, rigid simple eglandular hairs on the upper surface and the margin, longer, more numerous and less rigid beneath. Inflorescence with 2–3 capitula forming a corymb; peduncles slender and erect, with dense stellate hairs, numerous short and medium, pale simple eglandular hairs and sometimes a few, minute glandular hairs just below the capitulum. Capitula 15–20 mm in diameter, rounded at base. Involucral bracts porrect in bud, 3–10 × 0.6–1.0 mm, blackish-green, the inner with paler margins, linear or linear-lanceolate, drawn out to a filamentous apex, with numerous, short to medium, darkbased simple eglandular hairs, a few stellate hairs towards the base and a few, very short, dark glandular hairs. Ligules yellow, glabrous-tipped, very short and poorly developed. Styles yellow. Achenes not known. Flowers 7. Apomictic. Native. Known only from Rigg Head, near Seatoller in Cumberland. Endemic. 193. H. chloranthum Pugsley ex P. D. Sell Green-flowered Hawkweed H. onosmoides auct.; H. buglossoides auct.; H. saxifragum var. pseudonosmoides auct.; H. saxifragum subsp. pseudonosmoides auct.; H. pseudonosmoides auct.; H. chloranthum Pugsley sine diagn. lat. Phyllopodous perennial herb with a branched stock. Stem 20–55 cm, pale yellowish-green, often purplish towards the base, usually slender and flexuous, striate, with numerous long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards and with numerous stellate hairs throughout. Leaves medium bluish-green on upper surface, paler beneath; basal few, the outer with lamina 2–6 × 1.0–2.5 cm, usually elliptical, rarely ovate, obtusemucronulate at apex, subentire, denticulate or shallowly dentate and rounded or cuneate at base, the inner with lamina 7–11 × 1.5–2.5 cm, narrowly elliptical or oblongelliptical, gradually narrowed from near the middle to an acute apex, denticulate to shallowly dentate, sometimes with a few, larger teeth, gradually narrowed at the base, the petioles up to 4 cm, winged, with numerous, long, pale, wavy simple eglandular hairs; cauline 2–4, the lower like inner basal and petiolate, the upper with lamina lanceolate or linear, gradually narrowed to an acute apex, dentate and sessile; all glabrous or nearly so on upper surface or rarely with scattered stiff hairs; with numerous, but scattered, stiff, pale, short to medium simple eglandular hairs beneath and numerous, short to long, more or less setiform simple eglandular hairs on the margin, stellate hairs also often present beneath especially on the cauline. Inflorescence with (1–)3–5(−8) capitula, furcate-subcorymbose, sometimes with a long lower branch; peduncles rather long and slender, with dense stellate hairs, numerous, short and medium simple eglandular hairs and sometimes an occasional, short, dark glandular hair. Capitula 25–35 mm in diameter,

39. Hieracium rounded at base. Involucral bracts more or less porrect in bud, 4–12 × 0.8–1.2 mm, dark green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous stellate hairs, numerous short to medium, pale simple eglandular hairs and a few, inconspicuous, minute, dark glandular hairs. Ligules dull greenish-yellow, glabrous or with very short simple eglandular hairs at apex, usually illdeveloped or more or less tubular. Styles dull yellow. Receptacle pits with margins fimbriate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. This is the only species of hawkweed in Great Britain and Ireland with greenish-yellow, poorly developed ligules. Native. Cliff ledges, rocky streamsides and grassy slopes. Common in central and west Scotland and absent from most of the north and south. Endemic. 194. H. cacuminum (Ley) Ley Summit Hawkweed H. diaphanum var. cacuminum Ley; H. vulgatum var. cacuminum (Ley) F. Hanb.; H. demissum var. cacuminum (Ley) W. R. Linton; H. angustatum subsp. cacuminum (Ley) Zahn Phyllopodous or pseudophyllopodous perennial herb with a branched stock, the basal leaves dying early. Stem 15–35(−50) cm, pale yellowish-green, usually suffused brownish-purple below, with dense, long, wavy, pale simple eglandular hairs below, becoming fewer and shorter upwards and numerous stellate hairs in the upper part. Leaves bluish-green on upper surface, paler beneath, sometimes tinted reddish; basal often absent at flowering, or in a false rosette, with lamina 3–7 × 2.0–2.5 cm, lanceolate or elliptical, more or less acute at apex, subentire or denticulate, cuneate at base, the petioles up to 2 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 4–6, with lamina narrowly elliptical or oblong, acute at apex, subentire or with a few denticulations, cuneate or attenuate at base, often with long, tinted petioles which are semiamplexicaul, the upper linear and sessile; all sometimes nearly glabrous, but usually with a few, scattered simple eglandular hairs. Inflorescence with 1–3 capitula, furcate-corymbose; peduncles long and slender, with numerous stellate hairs and few to fairly numerous, short to medium, pale simple eglandular hairs. Capitula 20–30 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–10 × 1.5–1.9 mm, blackish-green, the inner with paler margins, all lanceolate, obtuse at apex, with short to medium, numerous, pale, darkbased simple eglandular hairs, few, short, dark glandular hairs and a few stellate hairs near the base. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 7. Apomictic. Native. When first described as H. diaphanum var. cacuminum the specimens referred to are a mixture of taxa. The original description, however, fits only the plants from the central cliff of the Brecon Beacons and is typified by the original material collected in 1888, not 1886 (see Jour. Bot. (London) 47: 51 (1909)). H. siluriense with which it has been confused also grows there as some mixed sheets of the Lintons’ Set no. 116 show, and the Y Fan Big and Craig Cerrig-gleisiad (Craig-y-Gledsiau) records

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originally included under H. cacuminum are also H. siluriense. Endemic. Section 15. Stelligera Zahn Taxon Pallida Backh.; Series Pallida (Backh.) P. D. Sell & C. West comb. inval.; Section Suboreadea Pugsley Phyllopodous perennial herbs. Leaves often more or less bluish-green, the basal usually fairly numerous, the cauline 0–1, sometimes 2, the margins usually with rigid simple eglandular hairs which often extend on to the upper surface, several species with numerous stellate hairs beneath. Inflorescence with 1–6(−12) capitula on long, straight and erect peduncles. Capitula medium to large. Involucral bracts often porrect in bud, usually over 11 mm, but up to 16 mm, mostly narrow and acute at apex. Ligules usually glabroustipped. Styles mostly yellow. Receptacle pits with margins sharply dentate. Achenes reddish-black. Flowers 5–7. Cliff ledges, scree, riverside rocks and grassy places usually on basic soils. Wales, western and northern England and Scotland and scattered over Ireland, mostly in mountainous areas. Occurs widely in Europe and extends into south-west Asia. The section is characterised by its numerous basal leaves, 0–1(−2) cauline ones with rigid simple eglandular hairs and medium to large capitula on long, erect peduncles. 195. H. subrubicundum Dahlst. Large-leaved Hawkweed H. olivaceum subsp. subrubicundum (Dahlst.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often suffused brownishpurple, slender to more or less robust, with numerous, medium to long, pale, wavy simple eglandular hairs towards the base, becoming fewer and shorter upwards and with numerous stellate and some short glandular hairs in the upper part. Leaves medium bluish-green on upper surface, paler beneath, often suffused with reddish-purple; basal numerous, with lamina 6–13 × 3–6 cm, broadly ovate, broadly elliptical or oblong-elliptical, narrowed to an obtuse-mucronate apex, subentire, remotely denticulate or with a few, broad, shallow teeth, cuneate at base, the petioles up to 8 cm, with dense, long, pale, wavy simple eglandular hairs; cauline usually 2, the lower large, as big as and similar to the basal and often long-petiolate, the upper with lamina ovate or lanceolate, gradually narrowed to an acute apex, denticulate or with an occasional tooth, sessile; all glabrous on upper surface or with a few, medium, subrigid, pale simple eglandular hairs near the margin, with few to numerous, short to medium, subrigid simple eglandular hairs beneath and on the margin. Inflorescence with 2–12 capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, numerous, short and medium, pale simple eglandular hairs and few, short, dark glandular hairs. Capitula 30–40 mm in diameter, truncate at base. Involucral bracts porrect in bud, 5–13 × 1.2– 1.5 mm, dark greyish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an obtuse or subacute apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs, and numerous stellate hairs along the margins.

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Ligules yellow, glabrous-tipped. Styles yellow or slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 4.0–4.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rocky streamsides, grassy places and pebbly shores. Perthshire, Argyllshire, Ross-shire, Sutherland, Caithness and Orkney. Faeroes. 196. H. hesperium P. D. Sell Donegal Hawkweed H. scoticum var. occidentale F. Hanb. Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, often flushed purplish below, slender to slightly robust, striate, with few to fairly numerous, long, pale simple eglandular hairs below, absent or nearly so above, with a few stellate and sometimes an occasional, short, dark glandular hair in the upper part. Leaves bright green on upper surface, paler and bluish-green beneath; basal several, with lamina 4–12 × 3–6 cm, subrotund, very broadly elliptical or broadly ovate, roundedobtuse at apex, subentire or denticulate, or with an occasional larger tooth, rounded or abruptly contracted at base, the petioles up to 9 cm, with few to fairly numerous, long, pale simple eglandular hairs; cauline usually 2, the lower like the basal but acute at apex and petiolate, the upper with lamina ovate, more or less acute, entire or with few teeth and sessile; all glabrous on upper surface, with few to fairly numerous, medium to long, pale simple eglandular hairs below, especially on the midrib and on the margins. Inflorescence with 2–4 capitula, furcate-corymbose; peduncles rather short and straight, with numerous stellate hairs, few, short, pale simple eglandular hairs and few, minute, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.0–1.5 mm, dark green, most with paler margins, all broad linearlanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous, short, dark glandular hairs and few stellate hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 4.0–4.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only from the rocky banks of the River Carrick and River Calabber in Co. Donegal. Endemic. 197. H. dentulum (E. F. Linton) P. D. Sell Thick-leaved Hawkweed H. holophyllum var. dentulum E. F. Linton Phyllopodous perennial herb with a branched stock. Stem 30–55(−70) cm, pale yellowish-green, sometimes purplish towards the base, slender to robust, striate, with numerous, long, pale simple eglandular hairs towards the base, with fewer upwards and sometimes none above, with few to numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves dull medium green on upper surface, paler beneath and often tinted reddish or purplish, thick; basal few to fairly numerous, with lamina 4.5– 11.0 × 2.5–5.5 cm, ovate, elliptical or subrotund, roundedmucronulate to more or less acute at apex, sharply denticulate or with a few, small teeth, subtruncate or rounded at base, the petioles up to 4 cm, with dense, long, pale simple eglandular hairs; cauline 1–2, the lower like the basal, acute

at apex and petiolate, the upper with lamina ovate or lanceolate, acute at apex, denticulate and sessile, or bract-like; all with short to medium, subrigid, pale simple eglandular hairs on the upper surface and margin, similar ones beneath but long ones on the midrib. Inflorescence with 3–7 capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, numerous, short or medium, pale simple eglandular hairs and fairly numerous, short, dark glandular hairs. Capitula 35–45 mm in diameter, rounded to subtruncate at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.5 mm, greyish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, numerous, short, dark glandular hairs and numerous stellate hairs on the margins and along the base. Ligules yellow, with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Grassy banks and cliff ledges on limestone. Yorkshire and Derbyshire, just extending into Staffordshire. Endemic. 198. H. holophyllum W. R. Linton Undivided-leaved Hawkweed H. vulgatum var. cinereum Backh.; H. subramosum subsp. holophyllum (W. R. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 25–70 cm, pale yellowish-green, sometimes purplish towards the base, slender to robust, striate, with more or less numerous, medium to long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards, and few to numerous stellate hairs and an occasional, short glandular hair in the upper part. Leaves deep green on upper surface, paler and bluish-green beneath, sometimes flushed with purple, rather thick; basal few to numerous, with lamina 3–11(−21 cult.) × 2–4(−9 cult.), lanceolate, ovate or narrowly elliptical, gradually narrowed to an acute or obtuse apex, entire to sparsely denticulate, abruptly contracted or cuneate at base, the petioles up to 3 cm, with numerous, long, wavy, pale simple eglandular hairs; cauline 0–2, the lower like the basal, usually acute at apex and petiolate, the upper usually bract-like; all with short to medium, rather stiff, pale simple eglandular hairs on both surfaces and the margin. Inflorescence with 2–7 capitula, furcate-corymbose; peduncles long, slender and straight, with dense stellate hairs, few to numerous, short to medium, pale simple eglandular hairs and a few, minute, dark glandular hairs. Capitula 20–30 mm in diameter, truncate-based. Involucral bracts incumbent in bud, 5–12 × 1.5–1.7 mm, greyish-green, the inner with paler margins, all broad linearlanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous stellate hairs along the margins and few, short, dark glandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins laciniate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rock-ledges and slopes on limestone. Great Ormes Head, Caernarvon, Cefn Fedw, Denbigh and Derbyshire Dales, extending into Staffordshire. Endemic.

39. Hieracium 199. H. caledonicum F. Hanb. Caledonian Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green with a purplish base, usually more or less slender and flexuous, striate, with numerous, medium to long, pale, wavy simple eglandular hairs below, becoming shorter and fewer upwards, and numerous stellate and occasional, short, dark glandular hairs in the upper part. Leaves dull medium green on upper surface, paler and bluish beneath and sometimes becoming purple; basal usually numerous, with lamina 3.5–9.0 × 2.5– 4.0 cm, mostly ovate, but sometimes elliptical or subrotund, rounded-mucronulate to acute at apex, with few to fairly numerous, small apiculate teeth, abruptly contacted or truncate at base, the petioles up to 4 cm, but often short, with numerous to dense, long, pale, wavy simple eglandular hairs; cauline 0–1, occasionally 2, with lamina lanceolate or ovate, acute at apex, entire or denticulate, sessile or shortly petiolate, the upper if present usually bract-like; all with short to medium, pale, rather rigid simple eglandular hairs on both surfaces and the margins. Inflorescence with 1–4 capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, few to fairly numerous, short, dark glandular hairs and a few, short or medium, pale simple eglandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–11 × 1.2–1.5 mm, dark green, the inner with paler margins, all broad linear-lanceolate, narrowed but obtuse at apex, with dense stellate hairs on the margin and apex, numerous short to medium, pale, dark-based simple eglandular hairs and few to fairly numerous, minute, dark glandular hairs. Ligules orange-yellow, glabroustipped. Styles yellow. Receptacle pits with margins incisedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges, streamsides, grassy slopes and dunes. Common in the coastal areas of Sutherland and Caithness, and recorded for Inverness-shire, Ross-shire and the Orkney Islands. Endemic. 200. H. veterascens Dahlst. Coldbackie Hawkweed H. caledonicum subsp. veterascens (Dahlst.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, sometimes slightly purplish at base, slender and flexuous, striate, with numerous, long, wavy, pale simple eglandular hairs at the base, becoming fewer and shorter upwards, sometimes sparse throughout, and numerous stellate hairs in the upper part. Leaves medium yellowish-green, often spotted and blotched on upper surface, paler and bluish-green beneath; basal numerous, with lamina 5–11 × 2–6 cm, broadly ovate, lanceolate or elliptical, more or less acute at apex, sometimes caudate, irregularly toothed, the teeth sharp and apiculate, sometimes narrowly mammiform and apiculate, abruptly contracted or cuneate at base, the petioles up to 5 cm, with dense, wavy, pale simple eglandular hairs; cauline 1–2, the lower near the base and like the basal, the upper usually bract-like; all glabrous or with scattered, short to medium, pale, subrigid hairs on the upper surface and numerous medium to long, pale simple eglandular hairs beneath, particularly on the

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midrib, and on the margin. Inflorescence with 2–4 capitula, furcate-corymbose; peduncles long and straight, with numerous, short, pale simple eglandular hairs and few to numerous, very short, dark glandular hairs. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–12 × 1.2–1.5 mm, dark green, the inner with paler margins, all broad linear-lanceolate, long attenuate at apex, but with obtuse tips, with dense stellate hairs on the margin and apex, numerous short and medium, pale, dark-based simple eglandular hairs and a few, short, dark glandular hairs. Ligules orange-yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, reddish-black. Apomictic. Native. Coastal rocks and streamsides around Coldbackie, near Tongue, Sutherlandshire. Faeroes.

201. H. rubicundiforme (Zahn) Roffey Rubicund Hawkweed H. caledonicum subsp. rubicundiforme Zahn; H. rubicundum F. Hanb., non Peter Phyllopodous perennial herb with a branched stock. Stem 20–40(−60 cult.) cm, pale yellowish-green, often purplish towards the base, usually slender, striate, with few to numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and minute glandular hairs in the upper part. Leaves characteristically deep bluish-green clouded with purple and more or less coriaceous; basal few to numerous, with lamina 3–11 × 2–5 cm, mostly ovate, sometimes elliptical or subrotund, rounded-obtuse or narrowed to an obtuse or acute apex, subentire, remotely denticulate or with an occasional tooth, rounded or cuneate at base, the petioles up to 5 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–2, the lower sometimes like basal and petiolate, sometimes with lamina lanceolate and acute at apex, the upper and sometimes lower linear and bract-like; all glabrous or nearly so on upper surface, with medium to long, pale simple eglandular hairs below, particularly on the midrib, and on the margins. Inflorescence with 1– 6 very broad capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and a few, short, dark glandular hairs. Capitula 35–45 mm in diameter, broadly truncate at base. Involucral bracts incumbent in bud, 5–13 × 1.5–1.7 mm, dark green, the inner with paler margins, often red-tipped, all broad linearlanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few short, dark glandular hairs and numerous stellate hairs on the margins. Ligules rather deep yellow, glabrous or with a few, very short simple eglandular hairs at the tip. Styles yellow or slightly livid. Receptacle pits with margins subulatedentate. Achenes 4.0–4.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges, rocky streamsides and grassy slopes. Mountainous areas of north Wales south to Cardiganshire, northern England and Scotland. Endemic. Most of the Scottish plants called H. rubicundum are probably H. boswellii.

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202. H. boswellii E. F. Linton Boswell’s Hawkweed H. rubicundum var. boswellii (E. F. Linton) F. N. Williams; H. caledonicum subsp. rubicundiforme var. boswellii (E. F. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 25–40 cm, pale yellowish-green, often brownish-purple towards the base, slender, striate, flexuous, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and a few, minute, dark glandular hairs in the upper part. Leaves medium bluish-green on upper surface, paler beneath and often tinted purplish; basal usually few, with lamina 3–9 × 2–4 cm, lanceolate, ovate or elliptical, obtusemucronate to acute at apex, dentate, the teeth mammiform and sometimes larger and irregular, cuneate at base, the petioles up to 5 cm, with numerous, long, pale simple eglandular hairs; cauline 1–2(−3), the lower like the basal and petiolate, the upper with lamina lanceolate, acute at apex, dentate and sessile, or bract-like; all glabrous or nearly so on upper surface, rarely with scattered hairs, with numerous, medium to long, pale simple eglandular hairs below, especially on the midrib, and on the margin. Inflorescence with 1–6 capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, numerous, short to medium, pale simple eglandular hairs and numerous, very short, dark glandular hairs. Capitula 35–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5– 12 × 1.2–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, narrowed at apex to an obtuse point, with numerous, short and medium, pale, dark-based simple eglandular hairs, numerous very short, dark glandular hairs and very few stellate hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 4.0–4.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges, scree and streamside rocks. Northern and central Scotland and the western islands. Endemic. Named after John Thomas Irvine Boswell-Syme (1822–88). 203. H. leyanum (Zahn) Roffey Fat-leaved Hawkweed H. sagittatum subsp. leyanum Zahn; H. pollinarium var. platyphyllum Ley; H. caledonicum var. platyphyllum (Ley) Ley; H. platyphyllum (Ley) W. R. Linton, non (Arv.-Touv.) Arv.-Touv. Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often purplish below, usually robust, rarely slender, striate, with numerous, medium to long, pale simple eglandular hairs throughout and numerous stellate hairs in the upper part. Leaves dull dark green on upper surface, paler beneath and often suffused purplish, coriaceous; basal rather few, with lamina 5–10 × 3–7 cm, ovate to broadly triangular-ovate or subrotund, roundedmucronulate to acute at apex, denticulate with a few larger teeth in the lower half, the teeth more or less mammiform or aquiline-mammiform and the lowest pair sometimes retrorse, rounded, cuneate or truncate at base, the petioles up to 6 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 1–2, the lower often large and

similar to the basal, often with large teeth and petiolate or sessile, the upper usually bract-like; all glabrous or nearly so on the upper surface, with numerous, medium to long, pale simple eglandular hairs beneath, especially on the midrib, and on the margin. Inflorescence with 2–9 capitula, furcate-corymbose; peduncles long and straight, with dense stellate hairs, numerous short to medium, pale, darkbased simple eglandular hairs and an occasional, short, dark glandular hair. Capitula 25–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.5– 1.7 mm, dark green, the inner with paler margins, all broadly linear-lanceolate, obtuse at apex, with dense stellate hairs especially on the margins and at the apex, numerous short and medium, pale, dark-based simple eglandular hairs and few, short, dark glandular hairs. Ligules usually poorly developed, but see note, glabrous-tipped. Styles discoloured. Receptacle pits with margins dentate-fimbriate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. The plants from the Hatteral Hills and the Brecon Beacons which have been called H. rubicundum (i.e. H. rubicundiforme) with normally developed ligules have the broader, more toothed leaves and the dense stellate hairs on the involucral bracts of H. leyanum and are both included in that species. Native. Cliff ledges, scree and grassy slopes. The mountains of Breconshire, Monmouthshire and Herefordshire. The plant from quarry debris on Portland, Dorset referred to this species seems to be a good match for the plant formerly called H. rubicundum and discussed above. Endemic. Named after Augustine Ley (1842–1911). 204. H. angustatiforme P. D. Sell & C. West Small-leaved Hawkweed H. angustatum auct. Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, usually suffused brownish-purple towards the base, slender, striate, often with some, medium to long, pale simple eglandular hairs near the base, occasionally with a few of them higher up the stem, but usually with them absent, with a few stellate hairs above. Leaves dull medium green on upper surface, paler beneath; basal numerous, mostly rather small, with lamina 2–7 × 1–2 cm, ovate-elliptical, elliptical, lanceolate or oblong-elliptical, obtuse to acute at apex, finely and regularly serrate, the teeth sharp and apiculate, cuneate at base, the petioles up to 5 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 0–2, the lower with lamina lanceolate to linear-lanceolate, acute at apex, finely serrate or denticulate, sessile or petiolate, the upper usually bractlike; all more or less glabrous on upper surface, with few to fairly numerous, short to medium, pale simple eglandular hairs beneath, and stiff, medium to long, pale simple eglandular hairs on the margin. Inflorescence with 1–4 capitula, furcate-corymbose; peduncles long and slender, with numerous stellate hairs and few, short to medium, pale simple eglandular hairs and a very occasional glandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, dark green, the inner paler, all linear-lanceolate, subacute at apex,

39. Hieracium with numerous, short to medium, pale, dark-based simple eglandular hairs, few, very short, dark glandular hairs and a few stellate hairs on the margins. Ligules yellow, glabroustipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Native. Cliff ledges, grassy banks and streamsides. Breconshire. Endemic. 205. H. angustatum (Lindeb.) Lindeb. Lesser-leaved Hawkweed H. caesium var. angustatum Lindeb.; H. orarium var. angustatum (Lindeb.) F. N. Williams Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often purplish towards the base, slender and flexuous, with a few, long, pale simple eglandular hairs below, becoming scarce or absent upwards, and with stellate hairs and an occasional very short, dark glandular hair in the upper part. Leaves bluishor greyish-green on upper surface, paler beneath; basal few to fairly numerous, with lamina 3–12 × 1.5–3.0 cm, lanceolate, oblong and narrowly elliptical, obtuse to acute at apex, with a few, small, mammiform teeth mainly towards the base and cuneate or attenuate at base, the petioles up to 9 cm, and with numerous, long, pale, wavy simple eglandular hairs; cauline 1–2, sometimes like basal but sessile, or small and linear; all with scattered, short to medium, stiff, pale simple eglandular hairs on upper surface, more numerous softer simple eglandular hairs beneath and on the margin. Inflorescence with 1–5 capitula, furcate-corymbose; peduncles long, slender and erect, with dense stellate, numerous, short to medium simple eglandular hairs and an occasional, short, dark glandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 0.8–1.2 mm, dark green, the inner with paler margins, all linear-lanceolate, gradually narrowed to a long, narrow, subacute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs and a few, short, dark glandular hairs, without or with very few stellate hairs. Ligules yellow, glabrous-tipped. Styles yellowish or slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Very like H. argenteum in appearance, but with a different clothing of the involucral bracts and peduncles. Native. Cliff ledges and scree. Known only from a few high-level gullies in Cumbria. Scandinavia. 206. H. siluriense (F. Hanb.) P. D. Sell Silurian Hawkweed H. nitidum var. siluriense F. Hanb.; H. saxifragum subsp. nitidum var. siluriense (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, often brownish-purple below, slender to slightly robust, striate, with numerous, long, pale, wavy simple eglandular hairs below becoming fewer and shorter upwards, and numerous stellate hairs in the upper part. Leaves bright green on upper surface, pale beneath, sometimes tinted purple; basal few to numerous, with lamina 2.5–9.0 × 2.0–4.0 cm, narrow to broadly

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elliptical, lanceolate to ovate or oblong, obtuse-mucronate to acute at apex, subentire to sharply dentate, the teeth small and well spaced, abruptly contracted or cuneate at base, the petioles up to 7 cm and with numerous, long, pale, wavy simple eglandular hairs; cauline 0–2, the lower like the basal, often large and petiolate, the upper linear and bract-like, acute at apex and shortly petiolate or sessile; all usually glabrous on upper surface with few to numerous, short to medium, pale simple eglandular hairs beneath and on the margin. Inflorescence with 2–5 capitula, furcatecorymbose; peduncles long and slender, with numerous stellate hairs and numerous, short to medium, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0– 1.5 mm, blackish-green, the inner with a paler margin, all linear-lanceolate, narrowed to an obtuse or subacute apex, with numerous, short to medium, pale simple eglandular hairs, a few, short, dark glandular hairs and stellate hairs frequent on the margins towards the base. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and scree. Abundant on all the higher ridges of the Brecon Beacons from Glyn Collwng westward to Craig Du; at Llanwychllyn in Merionethshire and Craigy-llyn in Glamorganshire. Endemic. 207. H. hypophalacrum P. D. Sell Berriedale Hawkweed H. oreades var. subglabratum F. Hanb.; H. argenteum var. subglabratum (F. Hanb.) Pugsley; H. saxifragum subsp. argenteum var. subglabratum (F. Hanb.) Zahn; H. oreades auct.; H. saxifragum var. oreades auct. Phyllopodous perennial herb with a branched stock. Stem 30–50(−60 cult.) cm, pale yellowish-green, sometimes a little purplish at base, striate, rather robust, flexuous, with a few, rarely numerous, long, pale simple eglandular hairs towards the base and rarely higher up, few stellate hairs in the upper part and rare or occasional, very short glandular hairs above. Leaves glaucous, paler beneath, sometimes flushed purplish; basal few to numerous, with lamina 4– 13 × 1–4 cm, narrowly to broadly elliptical, lanceolate or oblong, obtuse-mucronate to acute at apex, entire to sharply denticulate or serrate, rounded to cuneate at base, the petioles short to medium, with numerous to dense, long, pale simple eglandular hairs; cauline 0–2(−3), the lower like the basal but usually acute at apex, petiolate and standing out at right angles, the upper usually bract-like but sometimes long-linear; all glabrous or with an occasional, medium or long, rigid, pale, simple eglandular hair on the upper surface, with few to numerous, medium to long, pale, more or less rigid simple eglandular hairs beneath (especially on the midrib) and on the margins. Inflorescence laxly furcatecorymbose, with 2–6(−10 cult.) capitula; peduncles straight and erect, with dense stellate hairs, with few to numerous, medium to long, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs. Capitula 35–45 mm in diameter, rounded to subtruncate at base. Involucral bracts porrect in bud, 7–15 × 1.0–1.5 mm, dark to blackish-green, the inner with paler margins,

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all linear-lanceolate, gradually narrowed above to an acute apex, the inner almost filamentous, all with numerous to dense, medium to long, pale, dark-based simple eglandular hairs, few to numerous, short, dark glandular hairs and few, scattered stellate hairs. Ligules pale yellow, with more or less numerous, short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.8–4.0 mm, blackish-red. Flowers 6–7. Apomictic. Although this species has the glaucous leaves and spreading cauline leaf of H. argenteum and H. argentiforme it differs from both in its larger capitula and more sharply toothed leaves. Native. Originally described from rocky places near the sea at Berriedale in Caithness, it also occurs in Uig in Lewis and Lochmaddy in North Uist in the Hebrides. Plants from the Kyle of Tongue and the Vagastie Burn, Altnaharra in Sutherland match exactly except that they have glabroustipped ligules. Endemic. 208. H. argentatum (Pugsley) P. D. Sell Killarney Hawkweed H. orimeles var. argentatum Pugsley Phyllopodous perennial herb with a branched stock. Stem 20–35 cm, pale yellowish-green, sometimes suffused reddish-purple towards the base, slender to more or less robust, striate, flexuous, glabrous or with a few, long, pale simple eglandular hairs towards the base and a few stellate hairs above. Leaves silvery-glaucous, paler beneath and often tinted reddish, fleshy; basal few to numerous, with lamina 3–7 × 1.5–3.0(−4.0) cm, elliptical, lanceolate or ovate, obtuse to acute at apex, entire to denticulate, cuneate or rounded at base, the petioles rather short, with numerous, long, pale, rather stiff eglandular hairs; cauline 0–2(−3), the lower like basal with a long-acute to acuminate apex, standing out at right angles, the upper sometimes like lower, but usually bract-like and sessile; all glabrous or nearly so on upper surface, with few to numerous, medium and long, stiff, pale simple eglandular hairs below and on or near the margin. Inflorescence laxly furcate-corymbose, with 2–8 capitula; peduncles straight and erect, with few to numerous stellate hairs and sometimes a few, very short, dark glandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–11 × 1.0–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, narrowed to an obtuse or rarely for the inner an acute apex, with numerous, unequal, very short and short, dark glandular hairs, few, short to medium, pale, dark-based simple eglandular hairs and few stellate hairs near the base. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins dentate. Achenes 3.5–4.0 mm, nearly black. Flowers 5–7. Apomictic. This species is clearly more closely allied to H. argenteum than to H. orimeles in its glaucous leaves and spreading cauline leaves. It differs from H. argenteum in its broader leaves and from H. argentiforme in its almost entire leaves and more glandular capitula. Native. Sandstone rocks by the upper lake, Killarney and rocks below Coolyard Bridge on the Slakeney River, Kilgarvan in Co. Kerry. It is likely that the records of H. argenteum, from the River Sheen south of Kenmare from near Ashgrove to above Dromagorteen Bridge and along

its tributary to Coomeolan stream for a mile or more, and of H. orimeles on the Inchee Bridge of the upper Roughty valley, both sides of the Caragh River, Glencar and about Killarney lakes up to 450 m in R. W. Scully, Flora of County Kerry (1916), all belong to this species. Endemic. 209. H. argentiforme P. D. Sell Sutherland Hawkweed H. argenteum var. septentrionale F. Hanb., non (Norrl.) Norrl.; H. saxifragum subsp. argenteum var. septentrionale (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale green, sometimes tinted purplish, striate, sometimes flexuous, often glabrous or nearly so, or with medium to long, pale simple eglandular hairs below or scattered above, with some minute glandular hairs and occasional stellate hairs above. Leaves silvery glaucous, paler glaucous and sometimes tinted purplish beneath; basal usually numerous, with lamina 3–12 × 1.2–4.0 cm, mostly broadly elliptical, sometimes lanceolate or ovate, the outer sometimes rounded-obtuse-mucronate, the inner acute, or sometimes all acute at apex, regularly denticulate, the teeth mucronulate, sometimes with a few, larger teeth, the petioles usually short, with numerous to dense, long, pale simple eglandular hairs; cauline often absent, when present with lamina lanceolate, acute at apex, denticulate, shortly petiolate and spreading, or bract-like; all glabrous or nearly so on the upper surface, with numerous, medium to long, pale simple eglandular hairs beneath and numerous, mostly long, subrigid, pale simple eglandular hairs on the margin. Inflorescence furcate-corymbose, with 2–5(−11) capitula; peduncles usually rather long, straight and erect-spreading, with numerous to dense stellate hairs, few to numerous, short, dark glandular hairs and a few, short to long simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–11 × 1.0–1.5 mm, dark green, the inner with pale margins, all linear-lanceolate, gradually narrowed to an obtuse apex, with numerous to dense, short to long, pale, darkbased simple eglandular hairs, few, short glandular hairs and occasional stellate hairs. Ligules pale yellow, glabroustipped. Styles yellow. Receptacle pits with margins dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. In the field the broad, mostly elliptical, glaucous rosette of leaves is distinctive, but in the herbarium it is difficult to separate it from some variants of H. nitidum. It is easily distinguished from H. argenteum by its much broader leaves and more dense simple eglandular hairs on the involucral bracts. Native. Cliff ledges, rocks and sandhills. North coastal region of Sutherland. Endemic. 210. H. argenteum Fr. Silvery Hawkweed H. pallidum var. persicifolium auct.; H. saxifragum subsp. argenteum (Fr.) Zahn; H. microdon Dahlst.; H. saxifragum subsp. microdon (Dahlst.) Dahlst.; H. pseudomicrodon Dahlst.; H. stenotum auct. Phyllopodous perennial herb with a branched stock. Stem 15–45(–60) cm, pale green, often tinted reddish, especially below, slender to more or less robust, striate, nearly

39. Hieracium glabrous or with scattered, medium to long, pale simple eglandular hairs especially below and a few stellate and very short, dark glandular hairs in the upper part. Leaves silvery glaucous, paler beneath, sometimes spotted and blotched, sometimes flushed purplish beneath; basal usually few, rarely numerous, with lamina 1.5–11.0 × 1.0–2.5 cm, linear, linear-lanceolate or narrowly elliptical, obtusemucronate to acute at apex, entire to denticulate, occasionally with larger, cusped teeth, cuneate to attenuate at base, the petioles short to quite long, pale green, sometimes tinted purplish, with few to numerous, medium to long, pale simple eglandular hairs; cauline 0–2 (rarely more), similar to basal or bract-like, usually spreading; all glabrous on upper surface, often glabrous or with short to long, scattered, pale simple eglandular hairs beneath and on the margin. Inflorescence laxly furcate-corymbose, with 1–4(−14) capitula; peduncles straight and erect, with few to numerous, very short, dark glandular hairs and a few, rarely numerous stellate hairs and few, medium, pale simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–12 × 1.0–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, gradually narrowed to the apex, the outer obtuse, the inner acute, with numerous, unequal, short and very short, dark glandular hairs, few to numerous, short to medium, pale, dark-based simple eglandular hairs, and few stellate hairs mostly on the margin. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins dentate. Achenes 3.5–4.0 mm, nearly black. Flowers 6–7. Apomictic. As understood here, H. argenteum is easily recognised in the field by its narrow, glaucous leaves the cauline of which tend to stand out at right angles from the stem and its involucral bracts have numerous glandular hairs. In some colonies there are broader-leaved plants, as well as the narrow-leaved ones, which approach H. argentiforme, but they have fewer simple eglandular hairs on the involucral bracts. In cultivation H. argenteum sometimes turns much greener and the spreading cauline leaves are more erect, but it still does not look like any other species. Native. Cliff ledges, rocks, streamsides, heathy hillocks and river shingle, on basalt, limestone and mica-schist, also on granite where it is often the only hawkweed, from sea level to 800 m. In mountainous areas of Wales, northern England and Scotland including the islands, and coming down to the coast where there are cliffs; in Ireland it occurs in Co. Galway, Co. Donegal and Co. Down, but the Co. Kerry plants are described as a different species, H. argentatum. Norway and Iceland. 211. H. vagense (F. Hanb.) Ley Wye Hawkweed H. britannicum var. vagense F. Hanb.; H. wiesbaurianum subsp. vagense (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, striate, slender, with more or less numerous stellate hairs in the upper part and few below, with scattered, short to medium, pale simple eglandular hairs throughout and few to numerous, short, dark glandular hairs above. Leaves greyish-green, paler and sometimes tinted purplish beneath; basal numerous, with lamina 2–14 × 1.5–4.5 cm, the primordial small, elliptical, rounded-mucronulate at apex, entire or denticulate,

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and rounded at base, the later extremely variable, elliptical, lanceolate, ovate, oblong, oblong-lanceolate or oblongelliptical, obtuse-mucronate to sharply acute at apex, denticulate or dentate to deeply laciniate-dentate, the teeth spreading or slightly ascending, mammiform, aquilinemammiform, long and narrow or cusped and sometimes descending the petiole, cuneate or subtruncate at base, the petioles usually long, narrow and densely clothed with long, pale simple eglandular hairs; cauline leaves usually absent, when present usually narrowly lanceolate or bractlike, rarely like basal and petiolate; all with the upper surface glabrous or nearly so, with a few, scattered, short to medium, pale simple eglandular hairs on the lower surface but with numerous long ones on the midrib and numerous, long, stiff, pale simple eglandular hairs on and near the margin. Inflorescence furcate-corymbose with 2–6 capitula; peduncles short to long, with numerous to dense stellate hairs and more or less numerous, short glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–13 × 1.0–1.5 mm, grey-green, the inner with paler margins, all narrowly linear-lanceolate, the inner almost filiform, long drawn out to a narrow, acute apex, with scattered stellate hairs, numerous, unequal, short and medium, dark glandular hairs and occasional to fairly numerous, medium, pale, dark-based simple eglandular hairs. Ligules glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, blackish. Flowers 6–8. Apomictic. Native. Crevices in limestone cliffs and on banks. Banks of the River Wye in Breconshire and Radnorshire between Builth Wells and Glasbury; Gloddaeth, Llandudno Junction and a roadside near Llanberis in Caernarvonshire; Bryn Euryn, Colwyn Bay in Denbighshire; and banks of the River Dee at Tongland in Kirkudbrightshire. A specimen whose origin is said to be Craig-y-cilau (Craig Cille) in Breconshire may be mislabelled as nobody else has seen it there. Endemic. 212. H. cambricum (Baker) F. Hanb. Welsh Hawkweed H. caesium var. cambricum Baker; H. flocculosum var. cambricum (Baker) F. N. Williams; H. wiesbaurianum subsp. cambricum (Baker) Zahn Phyllopodous perennial herb with a branched stock. Stem 10–40 cm, pale green, often tinted purplish at base, slender, striate, glabrous. Leaves glaucous green, sometimes blotched deeper green often with brownish-purple spots and blotches; basal numerous, with lamina 2–10 × 0.7–3.0 mm, elliptical, lanceolate, oblong, or oblong-lanceolate, narrowly obtuse, acute or acuminate at apex or sometimes caudate, sharply denticulate to deeply laciniate-dentate especially in the lower half, the teeth irregular, mammiform, narrowly mammiform or aquiline-mammiform and often cusped and sometimes long and narrow, attenuate at base and the teeth often descending the long, slender, narrow, often purplish, glabrous or slightly hairy petioles; cauline 0–1, long linear or rarely linear-lanceolate, acute at apex, entire or with a few, sharp teeth, sessile; all glabrous or with a few, short simple eglandular hairs on the margins. Inflorescence furcate-corymbose, with 2–7 capitula; peduncles short to long, erect-spreading, with few stellate hairs,

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without simple eglandular or glandular hairs. Capitula 25– 35 mm in diameter, rounded at base. Involucral bracts more or less porrect in bud, 5–12 × 1.0–1.5 mm, greyish-green to dark green, the inner with paler margins, all linearlanceolate, the inner very narrow, gradually narrowed to an obtuse apex, with numerous stellate hairs especially on the margins, with numerous, very short and occasional, short and medium, dark glandular hairs and occasional to few, short or medium, rather dark simple eglandular hairs. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins irregularly and sharply dentate. Achenes 2.5–3.0 mm, blackish. Flowers 5–7. Apomictic. Native. Limestone cliffs and rubble. Various places on the Great Ormes Head, near Llandudno in Caernarvonshire. At its other two localities at Craig Fawr near Treorchy in Glamorganshire and Cefn Fawr near Llangollen in Denbighshire it has not been seen for many years. Endemic. 213. H. amydrostictum P. D. Sell Pale-spotted Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, often purplish at the base, slender and flexuous, striate, with few to fairly numerous, medium to long, pale simple eglandular hairs below, becoming even fewer and shorter upwards, and rather few stellate and an occasional short, dark glandular hair in the upper part. Leaves slightly glaucous green and blotched and spotted brownish-purple on upper surface, paler beneath and sometimes tinted purple; basal few to numerous; with lamina 2–8 × 1.0–2.5 cm, mostly lanceolate or ovate, sometimes oblong-elliptical, obtuse-mucronate to more or less acute at apex, denticulate or sometimes with a few, larger teeth towards the base, cuneate at base, the petioles up to 4 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1, often like basal and petiolate, or linear-lanceolate, acute at apex and entire to dentate; all with numerous, short to medium, subrigid simple eglandular hairs on the upper surface and more numerous, short to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 1–4 capitula, furcate-corymbose; peduncles long, slender and erect, with more or less dense stellate hairs, fairly numerous, short, dark glandular hairs and few to fairly numerous, short to medium, dark-based simple eglandular hairs. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts porrect in bud, 5– 13 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an acute apex, with numerous, short to medium, dark-based simple eglandular hairs, numerous, short, dark glandular hairs, and some stellate hairs along the margins. Ligules yellow, with a few, very short hairs at tips. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 6–7. Apomictic. Native. Cliff ledges and scree. Mountains of central Scotland. Endemic. 214. H. fucatifolium P. D. Sell Painted-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often brownish-purple at base, more or less robust, striate, with few to numerous,

long, pale, wavy simple eglandular hairs near the base, glabrous or nearly so or with a few stellate hairs above. Leaves bluish-green with dark brownish-purple blotches and spots on the upper surface, paler beneath; basal few, large, with lamina 4–12 × 2–4 cm, elliptical, lanceolateelliptical or oblong-elliptical, rounded-mucronate to acute at apex, denticulate to dentate, sometimes with large, hooked teeth, cuneate to attenuate at base, the petioles long, up to 10 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 0–2, one usually large like the basal ones and long-petiolate, but sometimes bract-like; all glabrous or nearly so on the upper surface, with few to numerous, medium to long, pale simple eglandular hairs beneath, especially on the midrib, and on the margin. Inflorescence with 2–6 capitula, furcate-corymbose; peduncles long, slender and erect, with numerous stellate hairs, fairly numerous, very short or short, dark glandular hairs and an occasional, medium, pale simple eglandular hair. Capitula 30–40 mm in diameter, rounded to subtruncate at base. Involucral bracts porrect in bud, 5–14 × 1.0–1.5 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, gradually attenuate to a narrow, acute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, more or less numerous, short, dark glandular hairs, and fairly numerous stellate hairs along the margins. Ligules yellow, glabrous-tipped. Styles yellow or slightly discoloured. Receptacle pits with margins incise-dentate. Achenes 3.5– 4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges, scree and riverside rocks. Perthshire, Aberdeenshire and Argyllshire, local. Endemic. 215. H. sommerfeltii Lindeb. Sommerfelt’s Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, often suffused brownishpurple below, slender and flexuous, striate, with a few, long, pale simple eglandular hairs below, becoming absent above, or absent or nearly so throughout, with some stellate hairs in upper part. Leaves medium green, blotched and speckled with brownish-purple on upper surface, paler and sometimes tinted purplish beneath; basal few to fairly numerous with lamina, 2–8 × 1–3 cm, elliptical, oblong-elliptical, lanceolate or lanceolate-oblong, narrowed to an obtusemucronulate or acute apex, denticulate or dentate, with small, sharp teeth, cuneate to attenuate at base, the petioles up to 3 cm, often reddish and with few to numerous, long, pale simple eglandular hairs; cauline 1–2, the lower like the basal and sessile, the upper usually bract-like; all glabrous or nearly so on upper surface, with few to numerous, short to medium, pale simple eglandular hairs beneath, especially on the midrib, and on the margin. Inflorescence with 1–3 capitula, furcate-corymbose; peduncles long and straight, with fairly few stellate hairs, and a few, small glandular hairs just beneath the capitulum. Capitula 30–35 mm in diameter, rounded to subtruncate at base. Involucral bracts porrect in bud, 5–13 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, short to long, pale, dark-based simple eglandular hairs, few, short glandular hairs, and fairly numerous stellate hairs along the margins and at the apex. Ligules yellow, glabrous-tipped. Styles discoloured.

39. Hieracium Receptacle pits with margins incise-dentate. Achenes 3.5– 4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rocky ledges and slopes and streamsides, rather scarce. Aberdeenshire, Inverness-shire and Argyllshire. Norway. Named after Søren Christian Sommerfelt (1794– 1838). 216. H. carneddorum Pugsley Carnedd Hawkweed H. sommerfeltii var. splendens F. Hanb., non H. splendens Elfstr.; H. sommerfeltii subsp. sommerfeltii var. griffithii subvar. splendens (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, slender, striate, with few stellate hairs below becoming numerous upwards, with numerous, medium to long, pale, wavy simple eglandular hairs and an occasional, short, dark glandular hair above. Leaves glaucous green, usually heavily blotched with dark brownish-purple on upper surface, paler and sometimes suffused purplish beneath; basal numerous, with lamina 2–12 × 1.0–3.2 cm, ovate, elliptical, oblong or lanceolate, mostly acute at apex but the outer sometimes obtusemucronate, almost entire to denticulate, sometimes with a few, larger teeth towards the base, cuneate or attenuate at the base, the petioles short to rather long, with numerous to dense, medium to long simple eglandular hairs; cauline 0–1 (−3 cult.), when present lanceolate, acute at apex, entire or denticulate, sessile or shortly petiolate when cultivated; all usually glabrous above, but sometimes with a few, medium, pale, stiff hairs, with numerous, medium to long, pale simple eglandular hairs beneath especially on the midrib, and often (especially the cauline) with few to numerous stellate hairs beneath, with numerous long, pale, rigid simple eglandular hairs on the margin. Inflorescence furcate-corymbose, with 1–6(−8 cult.) capitula; peduncles slender, with dense stellate, numerous short to medium, pale simple eglandular and numerous, unequal, dark glandular hairs. Capitula 25–35 mm in diameter, rounded or subtruncate at base. Involucral bracts porrect in bud, 7–12 × 1.0–1.2 mm, dark green, the inner with paler margins, all narrow linear-lanceolate, gradually narrowed to an acute apex, with stellate hairs mostly on the margins, numerous to dense, medium to long, pale simple eglandular hairs and few to numerous, obscure, short, dark glandular hairs. Ligules yellow, strongly pilose-tipped. Styles yellow, becoming discoloured when dry. Receptacle pits with margins incise-dentate. Achenes 3.0–3.5 mm, blackish. Flowers 6–8. Apomictic. Native. Basic rock-ledges, crevices and scree. Burfa Bank Camp in Radnorshire, upper reaches of the River Clywedog and Craig Breidden in Montgomeryshire, Llyn Styllan (Tewstyllon) and Cwm Croesor in Merionethshire, the area of Carnedd Dafydd, Carnedd Llewellyn and the Nant Francon Pass and Cwm Glas Fach on Snowdon in Carnarvonshire, the Berwyn Mountains in Denbighshire and Fleetwith Pike in Cumberland. A plant seen from the eastern Pyrenees labelled H. vestitum Gren. & Godr. seems to be this species and their relationship needs further investigation. 217. H. basicrinum (Zahn) Roffey Fionn Glenn Hawkweed H. praecox subsp. basicrinum Zahn; H. sommerfeltii var. tactum F. Hanb.

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Phyllopodous perennial herb with a branched stock. Stem 10–30 cm, pale yellowish-green, usually suffused with brownish-purple, slender and flexuous, with numerous, long, pale simple eglandular hairs below, becoming fewer upwards, and numerous stellate hairs in the upper part. Leaves bluish-green, blotched and spotted with dark purple on upper surface, paler beneath; basal few and small, with lamina 2–6 × 1–2 cm, lanceolate or elliptical, roundedmucronulate to obtuse at apex, sharply dentate or serratedentate, the teeth narrow and apiculate, cuneate at base, the petioles up to 5 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 0–2, similar to basal and petiolate or linear and bract-like; all glabrous on upper surface, with few to numerous, pale, short to medium simple hairs beneath and on the margin. Inflorescence with 1–4 capitula, furcate-corymbose; peduncles long, slender and erect, with dense stellate hairs, few to numerous, short to medium, pale simple eglandular hairs and some short, dark glandular hairs. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 5–12 × 1.0–1.5 mm, dark greyish-green, the inner with paler margins, all broad linear-lanceolate, obtuse at apex, with numerous stellate hairs especially towards the base, numerous, short to medium, pale, dark-based simple eglandular hairs and few, short, dark glandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7. Apomictic. Native. Known only from granite cliffs at about 800 m in Fionn Ghleann, Glen Etive, Argyllshire. Endemic. 218. H. pseudoleyi (Zahn) Roffey Purple-flushed Hawkweed H. saxifragum subsp. pseudoleyi Zahn; H. caesium var. decolor auct.; H. decolor auct. Phyllopodous perennial herb with a branched stock. Stem 15–40(−55) cm, pale yellowish-green, sometimes suffused brownish-purple, slender, striate, with stellate hairs throughout, dense in the upper part, with few to numerous, short to medium, pale simple eglandular hairs below and few similar simple eglandular and short, dark glandular hairs above. Leaves dull bluish- or greyish-green with purplish tints on upper surface, paler and often suffused purplish beneath; basal few to numerous, with lamina 4–13 × 1.1– 5.0 cm, narrowly to broadly elliptical or ovate, usually rounded-mucronulate at apex but sometimes the inner acute, subentire, denticulate or dentate in the lower half, widely cuneate to attenuate at base, teeth in larger plants sometimes descending the short to long petioles which are often purplish; cauline 0–1(−2), sometimes linear-lanceolate to lanceolate, acute at apex, slightly dentate and cuneate at base to the petiole, sometimes linear, acute at apex and sessile; glabrous or with an occasional simple hair on the upper surface, few to numerous, short to medium, pale simple eglandular hairs on the lower surface and longer more numerous hairs on the midrib, with few to numerous, short to long, often subrigid simple eglandular hairs on the margin or with only the base of the hairs left. Inflorescence furcatecorymbose, with 1–7(−18 cult.) capitula, the acladium usually long, sometimes with long lower branches; peduncles suberect, with dense stellate hairs, more or less numerous, short, dark glandular hairs and an occasional, medium, pale

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simple eglandular hair. Capitula 25–30 mm in diameter, with a rounded or subtruncate base. Involucral bracts 10– 12 × 1.0–1.5 mm, greyish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to a subobtuse apex, with dense stellate hairs, numerous, short, dark glandular hairs and few to numerous, pale, medium simple eglandular hairs. Ligules clear, bright yellow, glabroustipped. Styles pure yellow. Receptacle pits with margins subulate-dentate, with long, narrow teeth. Achenes 3.5–4.0 mm, blackish. Flowers 5–7. Apomictic. The original material of H. pseudoleyi was that cultivated by E. F. Linton (whose origin was Ysgolion Duon in the Carnedd Mountains of Caernarvonshire) and sent out in his Set of British Hieracia as no. 128, H. leyi F. Hanb. It is not H. leyi but is identical with plants growing on Great Ormes Head, Caernarvonshire which were long known as H. decolor. The type of H. decolor is a Yorkshire plant here recognised as a different species. H. pseudoleyi has never been refound on Ysgolion Duon and it may be that there is a mistake in the origin of the garden plant. H. pseudoleyi is however, the only available name for the Great Ormes Head plant. Native. Ledges on cliffs and cracks in pavement on limestone. Great Ormes Head, Gloddaeth and Pydew all near Llandudno in Caernarvonshire. Endemic. Named after Augustin Ley (1842–1911). 219. H. tricolorans (Zahn) Pugsley Three-coloured Hawkweed H. silvaticum var. tricolor W. R. Linton; H. bifidum subsp. tricolorans Zahn; H. cordigerum var. tricolorans (Zahn) Roffey Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, usually suffused reddishor brownish-purple below, slender to robust, striate, with a few, long, pale simple eglandular hairs below and an occasional one upwards, and more or less numerous stellate hairs in the upper part. Leaves bright green sometimes slightly blotched brown and purplish-brown on upper surface, paler and often suffused purple beneath; basal usually numerous, the outer with lamina 2–9 × 1–5 cm, broadly elliptical or oblong-elliptical, rounded-obtuse at apex, subentire, denticulate or shortly sinuate-dentate and rounded or subtruncate at base, the inner with lamina 5–10 × 2.5–6.0 cm, ovate or elliptic-ovate, gradually narrowed to an acute or obtuse apex, denticulate to sinuate-dentate and often with 2 spreading or deflexed teeth at the subtruncate or cordate base, the petioles medium or long with numerous to dense, very long, pale simple eglandular hairs; cauline 0–1, usually small and linear, rarely like inner basal and petiolate; all with short to medium, pale simple eglandular hairs on both surfaces and the margins or glabrous or nearly so on the upper surface. Inflorescence with 1–7 capitula, furcate-corymbose, sometimes with a long, lower branch; peduncles long, suberect, with dense stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Capitula 30–45 mm in diameter, broadly rounded at base. Involucral bracts porrect in bud, 6–14 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, the inner almost filamentous, gradually narrowed to an obtuse or acute apex,

with dense stellate hairs especially on the margins and at the apex, numerous, short to medium, pale, dark-based simple eglandular hairs and few, short, dark glandular hairs. Ligules yellow, sometimes with few, very short simple eglandular hairs at apex. Styles yellowish. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, reddish-brown. Flowers 6–7. Apomictic. Native. Rock-ledges, scree, and grassy slopes. Common on the Yorkshire limestone. Endemic. 220. H. subcyaneum (W. R. Linton) Pugsley Porrect-bracted Hawkweed H. silvaticum var. subcyaneum W. R. Linton; H. bifidum subsp. subcyaneum (W. R. Linton) Zahn; H. cordigerum var. subcyaneum (W. R. Linton) Roffey Phyllopodous perennial herb with a branched stock. Stem 20–45 cm, pale yellowish-green, often suffused purple below, more or less robust, striate, without or with an occasional, pale simple eglandular hair, with numerous stellate hairs in the upper part. Leaves dull green sometimes with suffused blotches of purple on upper surface, paler and often suffused purple beneath; basal usually numerous, the outer with lamina 2–5 × 1.0–2.5 cm, elliptical, obtuse at apex, subentire or with a few, small teeth towards the base and abruptly contracted or subtruncate at base, the inner with lamina 5–9 × 1.0–2.5 cm, lanceolate, acute or acuminate to a narrow point at apex, subentire towards the apex, dentate below with narrow, aquiline-mammiform teeth, the basal pair often deflexed and sometimes on the petiole, the petioles medium to long, with numerous, very long simple eglandular hairs; cauline usually absent, but sometimes 1 and linear, entire and acute at apex; all glabrous or nearly so on upper surface, with scattered to fairly numerous, short to medium, pale simple eglandular hairs below and numerous ones on the margin, and scattered stellate hairs sometimes present below. Inflorescence with 1–6 capitula, furcate-corymbose; peduncles rather short, with numerous stellate, numerous, short, dark glandular hairs and a few, medium simple eglandular hairs. Capitula 30–35 mm in diameter rounded at base. Involucral bracts porrect in bud, 4–13 × 1.0–1.5 mm, dark greyish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with dense stellate hairs particularly on the margins, numerous short to medium, dark-based simple eglandular hairs and occasional dark glandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 5–7. Apomictic. Native. Rock-ledges, scree and grassy slopes. Common on the Yorkshire limestone. Endemic. 221. H. decolor (W. R. Linton) Ley Shade Hawkweed H. caesium var. decolor W. R. Linton; H. caesium var. smithii auct. Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often suffused purplish towards the base, slender to more or less robust, striate, with few to fairly numerous, medium to long, pale simple eglandular hairs towards the base and few upwards, with numerous stellate hairs and an occasional glandular hair in the upper part. Leaves medium bluish-green on upper

39. Hieracium surface, paler beneath; basal few to numerous, the outer with lamina 3–8 × 1.5–3.0 cm, elliptical, ovate or lanceolate, obtuse at apex, denticulate or with small teeth at the base and abruptly contracted at the base, the inner with lamina lanceolate, oblong-lanceolate or oblong, more or less acute at apex, regularly denticulate or dentate, usually with some larger, narrower aquiline-mammiform teeth at base and abruptly contracted or subtruncate at base, the petioles mostly rather long, with numerous to dense, long, pale simple eglandular hairs; cauline 0–1, when present usually linear, acute at apex and entire, rarely like inner basal and petiolate; all glabrous or with a few, scattered, short or medium, pale simple eglandular hairs, numerous, medium to long, pale simple eglandular hairs beneath, especially on the midrib and numerous stiff ones on the margin. Inflorescence with 1–5 capitula, furcate-corymbose; peduncles long and erect, with numerous to dense stellate hairs, few, short to medium simple eglandular hairs and few, short, dark glandular hairs. Capitula 40–50 mm in diameter, rounded or subtruncate at base. Involucral bracts porrect in bud, 4–14 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, the inner filamentous, acute at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs and numerous, stellate hairs. Ligules bright yellow, glabrous or with few, very short hairs at apex. Styles yellow. Receptacle pits with margins subulatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rock-ledges, scree and grassy slopes. Frequent on the Yorkshire limestone. Endemic. 222. H. inaequilaterum P. D. Sell Uneven-toothed Hawkweed H. silvaticum var. asymmetricum Ley; H. cordigerum var. asymmetricum (Ley) Roffey; H. silvaticum subsp. cordigerum var. asymmetricum (Ley) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, often tinted reddish-purple below, fairly robust, striate, glabrous or nearly so below, with fairly numerous stellate and a few, dark-based simple eglandular and short, dark glandular hairs above. Leaves dull green, usually purplish-brown blotched, paler beneath and often suffused purple; basal numerous and large, the outer with lamina 2–9 × 2–5 cm, broadly elliptical to oblong-elliptical, obtuse to broadly rounded at apex, subentire to shallowly dentate and abruptly contracted or cuneate at base, the inner with lamina 6–11 × 2–5 cm, elliptical, lanceolate, oblong-lanceolate or ovate-oblong, more or less acute to acuminate, asymmetrically dentate, the teeth large and aquiline-mammiform or mammiform especially in the lower half, abruptly contracted or subtruncate at base, the petioles medium with numerous, very long, pale simple eglandular hairs; cauline usually absent, rarely linear, entire and long acute at apex; all glabrous on upper surface or rarely with a few, scattered simple eglandular hairs, and a few similar hairs beneath especially on the midrib and more numerous on the margin. Inflorescence with 2–5 capitula, furcate-corymbose; peduncles rather short, suberect, with dense stellate hairs, few to fairly numerous, short, dark glandular hairs and few, medium, dark-based simple

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eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–12 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to an acute apex, with numerous, short to medium, dark simple eglandular hairs, few, short, dark glandular hairs, and numerous stellate hairs on the margin. Ligules yellow, with short hairs at apex. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rock-ledges, scree and grassy slopes. Frequent on the Yorkshire limestone. Endemic. 223. H. promontoriale P. D. Sell Humphrey Head Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, sometimes slightly purplish at base, slender to more or less robust, striate, with few, medium to long, pale simple eglandular hairs below, becoming absent or nearly so upwards, and fairly numerous stellate hairs in the upper part. Leaves pale green, occasionally faintly blotched on upper surface, paler beneath; basal numerous, with lamina 4–9 × 1.0–3.0 cm, elliptical, oblong-elliptical or lanceolate, obtuse-mucronate to acute, acuminate or cuspidate at apex, very sharply dentate or serrate, the teeth narrow and very apiculate, mostly cuneate at base, the petioles up to 2 cm and with numerous, long, pale simple eglandular hairs; cauline 0–2, when present more or less linear or linear-lanceolate, with a long, narrow, acute apex and usually long, narrow teeth and shortly petiolate, sometimes bract-like; all more or less glabrous on upper surface or with a few hairs near the margin, with numerous, short to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 3–5 (numerous in cultivation) capitula, furcate-corymbose; peduncles straight and erect, with dense stellate hairs, few to numerous, short, dark glandular hairs and an occasional short to medium, pale simple eglandular hair. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 5.0–13.5 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to a narrow, subacute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs, and numerous stellate hairs on the margins and towards the base. Ligules yellow, glabroustipped. Styles yellowish or slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 4.0–4.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only from limestone sea-cliffs at Humphrey Head, near Grange-over-Sands, Lancashire. Endemic. 224. H. cyathis (Ley) W. R. Linton Cup-shaped Hawkweed H. hypochaeroides var. cyathis Ley; H. wiesbaurianum subsp. cyathis (Ley) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes tinted purplish, slender to robust, striate, with usually rather few, sometimes rather numerous, short to long, pale simple eglandular hairs throughout, numerous stellate and sometimes a few,

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minute, dark glandular hairs above. Leaves dark glaucousgreen, paler and even more bluish beneath and sometimes tinted purplish, often spotted or blotched with brownishpurple; basal few to numerous, with lamina 2.0–12.0 × 1.5– 4.0 cm, the outer ovate, obovate, oblong-elliptical, elliptical or lanceolate, rounded-mucronulate or obtuse at apex, often entire or nearly so in upper part sometimes becoming denticulate to sinuate-dentate below and abruptly contracted or rounded at base, the inner often broader and more pointed at apex; the petioles short to long, with dense, long, wavy, pale simple eglandular hairs; cauline 0–1, when present narrowly lanceolate, acute at apex, and dentate, or linear and more or less bractlike, sessile or shortly petiolate; all glabrous or nearly so on upper surface, with numerous, short to long, pale simple eglandular hairs beneath particularly on the midrib, the margins with numerous to dense, medium to long, pale, usually rather stiff simple eglandular hairs. Inflorescence cymose-subcorymbose, with short to medium acladium and in large plants a long lower branch; peduncles erect-spreading, with numerous stellate hairs, numerous, minute, dark glandular hairs and few to numerous, pale, short to medium simple eglandular hairs. Capitula 30– 45 mm in diameter, subtruncate at base. Involucral bracts porrect with spreading tips in bud, 6–12 × 1.0–1.5 mm, dark green, the inner with broad, pale margins, all linearlanceolate, gradually narrowed to an acute apex, with fairly numerous stellate hairs, but mainly on the margins, numerous, short to medium, pale simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, blackish. Flowers 5–7. Apomictic. There is slight variation. The Cheddar Gorge plants are sometimes smaller and neater and the Carmarthenshire ones rather more hairy than those from Breconshire. The species can be recognised by its tendency to produce obovate outer leaves and the porrect, out-turned involucral bracts in bud. Native. Ledges and crevices on limestone cliffs. Cheddar Gorge in Somerset, Craig-y-cilau (Craig Cille) where it is common in the central and eastern sectors, Daren Fach and Craig-y-nos in Breconshire and Llyn-y-fan-fechan in Carmarthenshire. Endemic. 225. H. arvonense P. D. Sell Caernarvon Hawkweed Phyllopodous perennial herb with a branched stock. Stem 15–35 cm, pale yellowish-green, often suffused brownishpurple below, slender and flexuous, striate, with few to fairly numerous, long, pale simple eglandular hairs below, becoming fewer and shorter or absent upwards, and with numerous stellate hairs and an occasional, very short, dark glandular hair in the upper part. Leaves pale to medium green with purplish-brown speckles on the upper surface, paler and sometimes tinted purplish beneath; basal usually numerous, with lamina 1.5–7.0 × 1–3 cm, lanceolate or elliptical, rounded-obtuse to sharply acute at apex, sinuate-denticulate usually with a few larger teeth at the abruptly contracted or subtruncate, often asymmetrical base, the petioles up to 4 cm and with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, when present ovate, with a long, narrow,

acute apex, entire or dentate in the lower half, cuneate at base and usually petiolate; all glabrous or nearly so on upper surface, with numerous, short to long, pale simple eglandular hairs beneath and on the margin, the marginal hairs subrigid. Inflorescence with 1–5 capitula, furcate-corymbose; peduncles slender and erect, with numerous stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and numerous, short, fine, dark glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–13 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually tapering to a narrow, more or less acute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous, short, dark glandular hairs, and numerous stellate hairs along the margin. Ligules yellow, glabrous or with an occasional very short hair at the apex. Styles yellow. Receptacle pits with margin subulatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. This species has been confused with H. leyi (with which it often grows), saxorum and petrocharis. It is nearest to H. leyi but can immediately be recognised in the field by its pale to medium green, spotted leaves which are smaller and more contracted at the base. Native. Rock-ledges and rocky slopes. Frequent in the mountains of Caernarvonshire. Endemic. 226. H. sublasiophyllum P. D. Sell Slender-bracted Hawkweed H. leyi var. vestitum Ley ex E. F. Linton, non H. vestitum Gren. & Godron; H. lasiophyllum var. semiglabrum auct. Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, usually purplish towards the base, slender, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and with numerous stellate hairs and short, dark glandular hairs in the upper part. Leaves bluish-green on upper surface, paler beneath; basal usually numerous, the lamina 2–11 × 1.5–4.0 cm, oblong, lanceolate-oblong, elliptical, ovate or lanceolate, obtuse-mucronate to acute at apex, entire to shallowly sinuate-dentate, and abruptly contracted or rounded at base, the petioles up to 6 cm, with numerous, long, pale simple eglandular hairs; cauline usually 1, with lamina linear and bract-like, sometimes lanceolate, acute at apex, subentire and petiolate; all glabrous or nearly so on upper surface, with numerous, short to medium, pale simple eglandular hairs beneath and stiff ones on the margin. Inflorescence with 2–12 capitula, furcatecorymbose, often with a long lower branch; peduncles fairly long and curved, with dense stellate hairs, numerous short, dark glandular hairs and few, short to medium, dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.3 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an obtuse or acute, long, narrow apex, with numerous, short and medium, dark or dark-based simple eglandular hairs, numerous, short, fine, dark glandular hairs, and very few stellate hairs near the base. Ligules yellow, with very short simple eglandular hairs at the apex. Styles yellow.

39. Hieracium Receptacle pits with margins shortly dentate. Achenes 3.5– 4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rock-ledges. Falcon Clints in Durham, Dollywaggon Pike and Snaka Rocks near Ambleside in Westmorland, near Blairgowrie in Perthshire, and Glen Canness, Unich Water and Craig Maskeldie in Forfarshire. A specimen from Norway collected and labelled H. schmidtii* crinelloides Omang by Omang is probably identical with this species but we cannot find a publication of that name, which would be very appropriate as it is closely allied to H. crinellum Omang with which it sometimes grows. Another specimen from Dresden in Germany appears to be the same thing. 227. H. crinellum Omang Blunt-leaved Hawkweed H. pallidum subsp. crinellum (Omang) Zahn; H. lasiophyllum var. semiglabrum auct. Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, sometimes purplish at the base, slender and flexuous, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer above and sometimes almost absent, and with numerous stellate hairs and some, short, dark glandular hairs in the upper part. Leaves bluish-green on upper surface, paler beneath, thick; basal few to numerous, with lamina 2– 8 × 1.5–3.0 cm, lanceolate, ovate, elliptical and oblongelliptical, mostly rounded-mucronulate at apex, but an inner leaf often acute, subentire, denticulate or with a few, small teeth near the base, rounded or truncate, rarely cuneate at base, the petioles up to 7 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, lanceolate or linear-lanceolate, acute or acuminate at apex, subentire or remotely dentate, petiolate or sessile; all glabrous or nearly so on upper surface, with numerous, short to long, pale simple eglandular hairs beneath, and numerous, medium to long, rigid, pale ones on the margin. Inflorescence with 1–5 capitula, furcate-corymbose; peduncles long, slender and erect, with dense stellate hairs, numerous, very short and short, dark glandular hairs and fairly numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 20–35 mm in diameter, subtruncate at base. Involucral bracts porrect in bud, 4–12 × 0.8–1.4 mm, dark green, the inner with paler margins, all narrow linear-lanceolate, gradually narrowed to a long, thin, acute apex, with numerous, very short and short, dark glandular hairs, numerous short to medium, pale, dark-based simple eglandular hairs and stellate hairs especially on the margins. Ligules yellow, with very short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cronkley Fell in Yorkshire; Falcon Clints in Durham, Dollywaggon Pike in Westmorland and Inveraray, Aoineadh Beag and Creag an Fitheach in Argyllshire. Scandinavia. 228. H. peroblongum P. D. Sell Westmorland Hawkweed Phyllopodous perennial herb with branched stock. Stem 12– 30 cm, pale yellowish-green, purplish towards the base, slender, with few to fairly numerous, pale, short to long

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simple eglandular hairs to almost none, and rather few stellate hairs and an occasional, short, dark glandular hair in upper part. Leaves bluish-green on upper surface, paler beneath and often suffused purple; basal numerous, the lamina 2–8 × 1–2 cm, oblong, narrowly elliptical or narrowly lanceolate, obtuse-mucronate to more or less acute at apex, entire or remotely denticulate and rounded or cuneate at base, the petioles up to 4 cm and with long, pale simple eglandular hairs; cauline 0–1, bract-like; all glabrous on the upper surface; with numerous, short to medium, rather stiff simple eglandular hairs beneath and on the margin. Inflorescence with 1–4 capitula, furcate-corymbose; peduncles straight and slender, with rather few stellate hairs, numerous, fine, short and very short, dark glandular hairs and numerous, short or medium, dark or dark-based simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all narrow linear-lanceolate, very narrow and acute at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous, short, dark glandular hairs and few stellate hairs along the margin. Ligules yellow, with very short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins fimbriate-dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. This species is very near to H. argenteum, with which it grows, but can be distinguished by its oblong, more or less entire leaves, narrow involucral bracts, which have more simple and less obvious glandular hairs, and ciliate ligules. Native. Known only from the Snaka Rocks near Ambleside, Dollywaggon Pike and Coniston Old Man, all in Westmorland. Endemic. 229. H. leyi F. Hanb. Ley’s Hawkweed H. pallidum subsp. leyi (F. Hanb.) Zahn; H. anglicum var. brevifurcatum F. N. Williams Phyllopodous perennial herb with a branched stock. Stem 15–40(−50) cm, pale yellowish-green, sometimes flushed brownish-purple towards the base, usually slender and flexuous, striate, with few to fairly numerous, medium to long simple eglandular hairs towards the base, becoming shorter and sparser upwards or with them absent, and with few to numerous stellate hairs and some, short, dark glandular hairs above. Leaves dull bluish or dark green on upper surface, paler and sometimes flushed purplish beneath; basal numerous, with lamina 2–9 × 1.0–3.5 cm, mostly lanceolate or ovate, sometimes elliptical or oblong, obtuse-mucronulate to acute at apex, often entire in the upper part, denticulate to dentate in lower half and sometimes throughout, the teeth short and sharp and sometimes narrowly mammiform near the base, occasionally on the petiole, cuneate or attenuate at base, the petioles up to 5(−9) cm, and with numerous, long, pale, wavy simple eglandular hairs; cauline 0–2, often with the lowest like the basal and petiolate, or the only 1 or the second bract-like; all glabrous or nearly so on upper surface, with more or less numerous, pale simple eglandular hairs beneath, especially on the midrib, and numerous, short to medium, pale, subrigid hairs on the margin, the cauline sometimes with stellate hairs beneath. Inflorescence with 1–4(−7) capitula,

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furcate-corymbose; peduncles rather long and slender, with dense stellate hairs, numerous, short, dark glandular hairs and few to fairly numerous, pale, medium simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–12 × 0.8–1.0 mm, dark green, the inner with a paler margin, all narrowly linear-lanceolate, gradually narrowed to an acute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, many short, dark glandular hairs, and numerous stellate hairs along the margins. Ligules yellow, glabroustipped. Styles yellow or slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rock-ledges and rocky slopes. Common in the mountains of northern Wales and in Cumbria and Hareheugh Crags, Berwickshire. Endemic. Named after Augustin Ley (1842–1911). 230. H. grampianum P. D. Sell Grampian Hawkweed H. jovimontis auct.; H. pallidum subsp. jovimontis auct.; H. leyi auct.; H. schmidtii var. superbum auct. Phyllopodous perennial herb with a branched stock. Stem 15–45(−50) cm, yellowish-green, sometimes purplish towards the base, rather slender, striate, sometimes flexuous, with few, medium or long, pale simple eglandular hairs scattered throughout or more numerous towards the base and a few stellate and minute, dark glandular hairs above. Leaves more or less bluish-green, especially beneath, rarely slightly spotted; basal with lamina 4–11 × 1.2–3.5 cm, the outer elliptical, the inner more or less lanceolate or slightly oblong-lanceolate, the outer obtuse, the inner acute at apex, all denticulate to dentate, the teeth triangularmucronate and gradually larger towards the base and sometimes with very narrow ones continued down one side of the petiole, the base abruptly narrowed or asymmetrically subtruncate, the petioles usually medium, with numerous, long, wavy, pale simple eglandular hairs; cauline 0–1, when present generally small and bract-like but sometimes like inner basal; all glabrous or with an occasional, pale simple eglandular hair on the upper surface, numerous, short to long, pale, slightly stiff simple eglandular hairs beneath and also on and near the margin and often numerous stellate hairs on the lower surface especially the cauline. Inflorescence with 1–6 capitula, furcate-corymbose, sometimes with a long, lower branch, the acladium often quite long; peduncles long, straight and erect, with numerous to dense stellate hairs, numerous very short, dark glandular hairs and occasional, pale simple eglandular hairs. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 6–14 × 1.0–1.5 mm, greyish-green, the inner with paler margins and sometimes reddish at tips, all narrow linear-lanceolate, the inner almost filamentous, all gradually narrowed to an acute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, without or with few, short, dark glandular hairs, and with dense stellate hairs on the margins and a few on the surface. Ligules pale yellow, glabrous or slightly hairy at tip. Styles more or less yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, blackish. Flowers 7–8. Apomictic.

This species has been known as H. jovimontis of which there are no type specimens. The original description does not fit our plant and, as H. W. Pugsley remarked, it is doubtful if our plant is the same as that from Germany. It has therefore been described as a new species. Native. Cliff ledges and crevices, rocks and scree on micaceous limestone and other basic rocks up to 1,160 m. Mountains of Perthshire, Forfarshire, Aberdeenshire, Banffshire, Inverness-shire and Argyllshire. Endemic. 231. H. carenorum F. Hanb. Pale-leaved Hawkweed H. saxifragum subsp. carenorum (F. Hanb.) Zahn; H. nitidum var. carenorum (F. Hanb.) Pugsley Phyllopodous perennial herb with a branched stock. Stem 25–40 cm, pale yellowish-green, more or less slender, striate, flexuous, with scattered, short to long, pale simple eglandular hairs especially below and a few stellate and occasional, very short glandular hairs in the upper part. Leaves pale green, paler beneath; basal few, with lamina 2–20 × 1.0–4.0 cm, usually narrowly elliptical to elliptical, rarely lanceolate, obtuse-mucronate to sharply acute, entire to denticulate, sometimes with a few, sharp teeth, cuneate at base, the petioles long, clothed with medium or long, wavy, pale simple eglandular hairs; cauline usually 1, sometimes absent, lanceolate, long-acute or acuminate at apex, entire or denticulate, rarely with a few, small teeth, cuneate at base and petiolate; all nearly glabrous, or with short to long, pale simple eglandular hairs beneath, occasionally with a few hairs above, and with rather stiff ones on the margin. Inflorescence laxly furcate-corymbose, with 2–7(−11 cult.) capitula; peduncles long, straight, erect and slender, with more or less numerous stellate hairs, numerous, very short, dark glandular hairs and few, rarely numerous, pale, short to medium simple eglandular hairs. Capitula 25–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–12 × 1.0–1.2 mm, dark green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to a more or less acute apex, with numerous, short to long, pale, dark-based simple eglandular hairs, much fewer, short, dark glandular hairs, and numerous stellate hairs on the margins, particularly of the outer. Ligules pale yellow, glabroustipped. Styles yellow. Receptacle pits with margin subulatedentate. Achenes 3.8–4.0 mm, reddish-black. Flowers 6–8. Apomictic. Closely allied to H. nitidum, from which it differs in its pale green, entire or denticulate, long-petioled leaves and stellately hairy margins to at least the outer involucral bracts. Both species grow together on low cliffs between Bettyhill and Farr Church in Sutherland. Native. Cliff ledges and sides of river gorges on basic rock. Allt Giubhas Beag, Beallach Corrie in Wyvis Forest and Allt na Leacachain in Ross-shire and between Bettyhill and Farr Church, Ben Hope, near Tongue and Cashel Dhu in Sutherland. Endemic. 232. H. nitidum Backh. fil. Shining-leaved Hawkweed H. saxifragum subsp. nitidum (Backh. fil.) Zahn; H. saxifragum var. vimineum auct. Phyllopodous perennial herb with a branched stock. Stem 20–40(−55) cm, pale yellowish-green, sometimes suffused

39. Hieracium purplish towards the base, slender to fairly robust, striate, sometimes flexuous, with scattered, short to long, pale simple eglandular hairs particularly below, scattered to few stellate hairs above and occasional to fairly numerous, short, dark glandular hairs near the inflorescence. Leaves bright medium green, paler beneath, sometimes tinted purplish beneath; basal rather few, with lamina 2.0–10.0 × 1.2–3.5 cm, narrowly to broadly elliptical, ovate or lanceolate, rarely subrotund, obtuse-mucronate to acute at apex, denticulate to serrate-dentate the teeth usually ascending, in cultivation becoming long and cusped, cuneate at base, the petioles short, with dense, long, pale simple eglandular hairs; cauline 0–1, lanceolate, acute at apex, denticulate or serrate-dentate, cuneate at base, ascending, or small and bract-like; all nearly glabrous or with scattered, short to medium, sometimes stiff, pale simple eglandular hairs on the upper surface and more numerous ones below and on the margins. Inflorescence laxly furcate-corymbose, with 2–7 capitula; peduncles straight and erect, with numerous, but not dense stellate hairs, numerous, short, dark glandular hairs and a few, pale, dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–12 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to the obtuse apex, the inner sometimes acute, all with numerous, unequal, short and very short, dark glandular hairs, numerous, short, medium and long, pale, dark-based simple eglandular hairs, and few, stellate hairs near the base. Ligules pale yellow, glabrous or slightly hairy at tip. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–3.8 mm, reddish-black. Flowers 6–8. Apomictic. In the field the bright green leaves easily distinguish H. nitidum from H. argenteum and its allies, but in the herbarium they can be confused, especially Sutherland plants with H. argentiforme. Native. Basic cliff ledges, rocky outcrops and riverside rocks and gorges. Widespread in central, northern and western Scotland, including Raasay. Endemic. 233. H. cillense Pugsley Cille Hawkweed H. hypochaeroides var. griseum Ley, non H. griseum Forman; H. wiesbaurianum subsp. hypochaeroides var. griseum (Ley) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, sometimes slightly tinted purplish below, striate, usually slender, with minute stellate hairs throughout, but not densely, with few to numerous, medium, pale simple eglandular hairs below, becoming sparse or absent upwards. Leaves greyish-glaucous, often with purplish spots and blotches, paler beneath and sometimes flushed purplish; basal with lamina 1.5–19.0 × 1.0– 4.5 cm, oblong, ovate or elliptical, the outer obtusemucronate, the inner acute or acuminate at apex which is often curved or twisted, sometimes nearly entire except for a tooth at the base, but usually dentate in the lower part with irregular mammiform teeth, which sometimes descend on to the petiole, abruptly contracted or subtruncate at the asymmetrical base, the petioles often purplish, with numerous stellate and long simple eglandular hairs; cauline 0–1, linear to linear-lanceolate, acute to acuminate at apex, sometimes

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with an occasional tooth; all with numerous to dense stellate hairs on both surfaces, with few to numerous, pale, stiff simple eglandular hairs on the margins. Inflorescence furcate-corymbose, with 2–4 capitula, usually with a long acladium, in large plants with a long, lower branch; peduncles long and erect, with numerous to dense, very small stellate hairs and scattered, minute, dark glandular hairs. Capitula 25–45 mm in diameter, rounded at base. Involucral bracts porrect, 7–13 × 1.0–1.5 mm, greyish-green, the inner with wide pale margins, all linear-lanceolate, acute at apex, the inner very narrow at apex, with numerous to dense stellate hairs and scattered, short, pale simple eglandular hairs and short, dark glandular hairs. Ligules clear yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.5–4.0 mm, reddish-brown. Flowers 6–7. Apomictic. This rare but very distinct species is easily recognised by the abundant stellate hairs on both surfaces of its leaves. Native. Limestone rock crevices. Known only from Craig-y-cilau (Craig Cille) especially below Waun Rudd and the eastern end of Daren Cilau quarry in Breconshire. Endemic. 234. H. lasiophyllum W. D. J. Koch Stiff-haired Hawkweed H. schmidtii forma lasiophyllum (W. D. J. Koch) F. N. Williams; H. pallidum subsp. lasiophyllum (W. D. J. Koch) Zahn; H. cinerascens auct. Phyllopodous perennial herb with a branched stock. Stem 15–30(−40) cm, pale yellowish-green, rarely purplish towards base, striate, flexuous, with numerous, medium to long, pale simple eglandular hairs throughout, and few to numerous stellate hairs and few, very short, dark glandular hairs in the upper part. Leaves thick, glaucous, paler beneath, sometimes spotted; basal few to numerous, with lamina 2–11 × 1.5–4.0 cm, mostly oblong or narrowly elliptical to elliptical, sometimes lanceolate, obtuse-mucronate to acute at apex, entire or undulate-denticulate, often with 1–2 larger, broad teeth near the base, rounded or asymmetrically truncate at base, the petioles usually rather short, with dense, medium to long, pale simple eglandular hairs; cauline usually absent, but when present linear or lanceolate, acute or acuminate at apex, entire or with broad teeth at base, shortly petiolate; all with numerous to dense stellate hairs on both surfaces, with numerous, medium to long, pale simple eglandular hairs beneath, especially on the midrib and numerous to dense, medium to long, very rigid, pale simple eglandular hairs on and near the margin, a few also sometimes on the upper surface. Inflorescence furcate-corymbose, with 1–7 capitula; peduncles medium to long, straight and erect, with numerous to dense stellate hairs, numerous short and very short, dark glandular hairs and few to numerous, medium to long, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–11 × 1.0–1.3 mm, dark greyish-green, the inner with paler margins, all narrow linear-lanceolate, the inner almost filamentous, gradually narrowed to an acute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few to numerous, short, dark glandular hairs and numerous to dense stellate

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hairs especially on the margins. Ligules pale yellow, usually glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Native. Cliff ledges, crevices and scree on dolerite, basic gneiss, basalt, Silurian grit, slate and other basic rocks but apparently not on limestone. Breconshire, Radnorshire, Cardiganshire, Montgomeryshire, Merionethshire and Caernarvonshire in Wales, Shropshire, Herefordshire, Monmouthshire, Cos. Durham and Cumberland in England, Perthshire, Forfarshire, Aberdeenshire and Argyllshire in Scotland and Co. Down, Antrim and Londonderry in Ireland. Central Europe from France to Roumania, but some records are probably H. schmidtii. 235. H. brigantum (F. Hanb.) Roffey Yorkshire Hawkweed H. lima var. brigantum F. Hanb.; H. anglicum var. brigantum (F. Hanb.) W. R. Linton; H. anglicum × lancifolium auct. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, yellowish-green, sometimes purplish towards the base, more or less robust, flexuous, striate, with few to numerous, medium to very long, pale, wavy simple eglandular hairs especially towards the base, and few stellate and occasional, short, dark glandular hairs above. Leaves thick, dull, dark green, paler beneath, sometimes with a few spots or blotches above and tinted purplish beneath; basal usually numerous, with lamina 3–10(−13) × 1.5–6.0 cm, ovate, elliptical to broadly elliptical, oblong or subrotund, rounded, obtuse or the inner acute at apex, entire or denticulate in the lower half and often with a few cusped teeth near the base which sometimes extend on the petiole, abruptly contracted, rounded or shortly cuneate at base, the petioles short to fairly long, usually with numerous to dense, very long, wavy, pale simple eglandular hairs but sometimes with only a few; cauline 0–1(−2), when present lanceolate, acute at apex, entire or denticulate, sessile or shortly petiolate, or bract-like; all with numerous short to medium, pale, rigid, bulbous-based simple eglandular hairs on the upper surface and margin and even longer ones beneath especially on the midrib. Inflorescence furcate-corymbose, with 1–6(−8 cult.) capitula; peduncles long, erect, straight, with numerous to dense stellate hairs, numerous, short to medium, rather dark or pale, dark-based simple eglandular hairs and numerous, short, unequal, dark glandular hairs. Capitula 35–50 mm in diameter, very large and handsome, ventricose at base. Involucral bracts porrect in bud, 5– 15 × 0.5–1.2 mm, dark greyish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to an acute apex, with numerous stellate hairs especially on the margin and at the apex, numerous to dense, medium to long, wavy, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs obscured by the simple hairs. Ligules pale yellow, sometimes with a few hairs at tip. Styles yellow. Receptacle pits with margins dentate-fimbriate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. This species has been placed with H. anglicum in the Section Cerinthoidea, but its thick leaves clothed with rigid

hairs and yellow styles clearly place it in the Section Stelligera. It is easily recognised by its thick, hairy nearly entire leaves with large capitula on long stems making it look top-heavy. Native. Cliff ledges, scree, rubble and pavement. Confined to the Yorkshire limestone where it is sometimes abundant. Endemic. 236. H. schmidtii Tausch Schmidt’s Hawkweed H. pallidum subsp. schmidtii (Tausch) Zahn; H. lima F. Hanb.; H. pallidum subsp. lima (F. Hanb.) Zahn; H. dentifex E. F. Linton; H. lasiophyllum auct.; H. cinerascens auct.; H. lasiophyllum var. euryodon auct. Phyllopodous perennial herb with a branched stock. Stem 10–40(−50) cm, yellowish-green, sometimes purplish towards the base, striate, slender to robust, with numerous, medium to long, pale, wavy simple eglandular hairs, and numerous stellate hairs and sometimes a few, short, dark glandular hairs in the upper part. Leaves more or less glaucous, thick, often tinted purplish beneath; basal numerous, with lamina 2–9(−10) × 1–4 cm, subrotund, elliptical, oblong-elliptical, ovate, obovate or oblong, roundedmucronulate to more or less acute at apex, entire, denticulate or with a few mammiform teeth near the base (more numerous in cultivation), sometimes only on 1 side, rounded or subtruncate at base, the petioles up to 40 mm, with dense, long, pale simple eglandular hairs; cauline 0 or 1, when present oblong-lanceolate or lanceolate, acute at apex, entire or denticulate, petiolate or sessile; all rough with medium to long, rigid, curved, pale simple eglandular hairs on the upper surface and margins and similar, rather softer hairs beneath. Inflorescence furcate-corymbose, with 1–10(−15) capitula, sometimes with a long lower branch; peduncles usually rather long, straight and erect, with numerous, short, dark glandular hairs, numerous to dense stellate hairs and few to numerous, medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect to incumbent in bud, 5– 12 × 0.8–1.2 mm, greyish to dark green, the inner with paler margins, all narrow linear-lanceolate, gradually narrowed to a long-attenuate, acute apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, few to numerous, short, dark glandular hairs and numerous stellate hairs especially on the margins. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins dentate-fimbriate. Achenes 3.5–4.0 mm, dark reddish-black. Flowers 5–7. Apomictic. All authors have found H. schmidtii difficult to define. Some of the plants from Cheddar Gorge, Somerset, described as H. lima, match perfectly the type material of H. schmidtii. The Cheddar population is, however, very variable especially in leaf shape. This variation has been used as a guide as to the limits of H. schmidtii. As here defined H. schmidtii is a species with glaucous or glaucous-green leaves, entire in the upper two-thirds, often with a few large teeth near the base and covered on the upper surface with curved, rigid simple eglandular hairs, the involucres having more simple eglandular than glandular hairs and the styles are yellow. No specimens actually labelled H. dentifex have

39. Hieracium been found, but what appears to be the original material is typical H. schmidtii. Native. Cliff ledges and scree on limestone and other basic rock. Scattered localities in Wales, western and northern England and Scotland, and perhaps Ireland. It is the most widespread and variable species of the Section Stelligera found in Great Britain. It is probably also widespread in Continental Europe. Named after Franz Wilibald Schmidt (1764–96). 237. H. bettyhillense P. D. Sell Bettyhill Hawkweed H. schmidtii auct. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale green, often purplish towards the base, slender and flexuous to fairly robust, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter or absent upwards, and numerous stellate and sometimes an occasional, short, dark glandular hair in the upper part. Leaves bluish-green on upper surface, paler beneath, thick; basal numerous, the lamina 4– 10 × 1.5–3.0 cm, mostly ovate, sometimes oblong-ovate or elliptical, mostly acute at apex, sharply dentate or serratedentate, abruptly contracted or cuneate at base, shortly petiolate; cauline 0–1(−2), when present usually bract-like, rarely lanceolate, sharply acute at apex, serrate and cuneate at base to a short, winged petiole; all glabrous or with a few, stiff hairs on the upper surface, numerous, medium and long, pale, stiff simple eglandular hairs on the margin and numerous softer ones beneath. Inflorescence with 2– 4(−8) capitula, furcate-corymbose; peduncles rather short, straight, with dense stellate hairs, numerous, very short, fine glandular hairs and few, short or medium, pale simple eglandular hairs. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts porrect in bud 4–12 × 1.0– 1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, medium, pale, dark-based simple eglandular hairs, more or less numerous, very short and short, dark glandular hairs and a few stellate hairs. Ligules yellow, with a few very short hairs at tip. Styles yellow. Receptacle pits with margins dentate-fimbriate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only from low cliffs and rocks by the River Naver, at Bettyhill in Sutherland. Endemic. 238. H. stictum P. D. Sell Lance-leaved Hawkweed H. hypochaeroides var. lancifolium W. R. Linton, non H. lancifolium Vuk. Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often purplish towards the base, with numerous, long, pale simple eglandular hairs at the base, becoming fewer and shorter upwards, and numerous stellate and occasional, short, dark glandular hairs in the upper part. Leaves pale or medium yellowish-green with scattered spots and blotches on upper surface, paler beneath and often suffused purple; basal few to fairly numerous, the lamina 4–14 × 2.5–6.0 cm, ovate, elliptical or lanceolate, rounded-mucronulate to acute at apex, subentire, denticulate or with a few longer teeth near the base and abruptly contracted or cuneate at base, the petioles up to 8 cm and with numerous, long, pale, wavy simple eglandular

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hairs; cauline 1–2, often large, the lower similar to basal and petiolate, the upper linear or linear-lanceolate, acute at apex, toothed and sessile; all glabrous or nearly so or with more or less numerous, stiff hairs on upper surface, with numerous, short to medium, pale simple eglandular hairs beneath and more numerous, longer ones on the midrib, and long, stiff ones on the margin, although they are sometimes rather sparse. Inflorescence with 2–12 capitula, paniculatecorymbose, often with a long, lower branch; peduncles fairly long and robust, with more or less dense stellate hairs, numerous, short to medium, pale simple eglandular hairs and occasional, short, dark glandular hairs. Capitula 30– 40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 5–12 × 1.5–2.0 mm, dark green, the inner with paler margins, all very broad linear-lanceolate, obtuse at apex, with numerous, short and medium, pale simple eglandular hairs, few, short, dark glandular hairs and numerous stellate hairs along the margins and towards the base. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliffs, ledges and rocks by streams. Scattered localities on the Yorkshire limestone. Endemic. 239. H. hypochaeroides S. Gibson Cat’s-ear Hawkweed H. gibsonii Backh. fil. nom. illegit.; H. wiesbaurianum subsp. hypochaeroides (S. Gibson) Zahn; H. caesium var. hypochaeroides (S. Gibson) Bab. Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale green, often purple-tinted especially below, striate, sometimes flexuous, with few, sometimes rather numerous, medium to long, pale simple eglandular hairs below becoming occasional or absent upwards, few, minute, dark glandular hairs in the upper part and rarely an occasional stellate hair. Leaves pale yellowish-green, beautifully spotted and marbled with brownish-purple on the upper surface, paler and often tinted or suffused purplish on the lower surface; basal usually rather numerous, with lamina 1.5–9.0 × 1–5 cm, subrotund, ovate, lanceolate or oblong, rounded-obtuse-mucronate to acute, entire, sinuate, denticulate or sometimes with sharp or mammiform teeth towards the base, subcordate, rounded or abruptly contracted at base, the petioles 10–50 mm with dense, long, pale simple eglandular hairs; cauline 0 or 1, when present narrowly linear-lanceolate, acute at apex, entire or denticulate and shortly petiolate, or bract-like; all glabrous or nearly so on the upper surface, with short to long, few to numerous, pale simple eglandular hairs on the lower surface and numerous, subrigid ones on the margin. Inflorescence furcate-corymbose, with (1–)2–4(−6) capitula; peduncles short to fairly long, straight and suberect, with more or less dense stellate hairs, more or less numerous, very short or minute, dark glandular hairs and occasional to few, pale or dark simple eglandular hairs. Capitula 30–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4– 11 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, the outer gradually narrowed to an acute apex, the inner almost filiform above with an acute apex, with dense stellate hairs on the margins but sparse elsewhere, numerous, very short, dark glandular hairs and

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few to numerous, short to medium, pale, dark-based simple eglandular hairs. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, dark reddish-brown. 3.5–4.0 mm. Flowers 5–8. Apomictic. Native. Ledges, rubble and crevices on limestone crags, quarries and pavement. Near Tutshill in Gloucestershire, Dyffryn Crawnon in Breconshire, Craigiau Eglwyseg near Llangollen in Denbighshire, frequent in mid and north-west Yorkshire and the English Lake District, the Burren in Co. Clare and Murlough Bay in Co. Antrim. Endemic. 240. H. repandulare Druce Repand-leaved Hawkweed H. ciliatum var. repandum Ley; H. repandum (Ley) Ley, non Schrank; H. pseudorepandum Pugsley Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes tinted purplish near the base, slender, striate, with few to numerous stellate hairs particularly in the upper part, few to numerous, medium to long, pale simple eglandular hairs below and occasional shorter ones higher up the stem. Leaves deep bluish-green on upper surface, paler beneath, sometimes slightly tinted purplish; basal numerous, with lamina 3–12 × 2.0–6.5 cm, the outer elliptical or oblong-elliptical, rounded-mucronulate at apex, entire or denticulate or with an occasional larger tooth and rounded at base, the inner sometimes similar to outer but often ovate, acute at apex, and repand-denticulate to subdentate, subtruncate at base, often with 2 spreading basal teeth; cauline often absent, sometimes resembling inner basal and petiolate; all usually glabrous on the upper surface, with few to numerous, short and medium, pale simple eglandular hairs on the lower surface, particularly the midrib and more numerous, stiffer ones on and near the margin; petioles up to 60(−110) mm, with numerous to dense, medium to long, pale, curly simple eglandular hairs. Inflorescence furcate-corymbose, with 1–6 capitula, with rather long acladium; peduncles with dense stellate hairs, short glandular hairs and few, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded below. Involucral bracts porrect in bud, 10– 13 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, drawn out to an acute apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs and numerous stellate hairs especially on the margins. Ligules pale yellow, with numerous, short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins fibrillousdentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–7. Apomictic. Native. Cliff ledges and rubble on limestone. Common on a number of Breconshire cliffs and on a rocky streamside in Cwm Tarell. There is an isolated record from Carnedd Dafydd in Carnarvonshire, the specimens exactly matching the Breconshire plants. Endemic. 241. H. riddelsdellii Pugsley Riddelsdell’s Hawkweed H. ciliatum var. venosum Ley, non H. venosum L.; H. repandum var. venosum (Ley) Ley Phyllopodous perennial herb with a branched stock. Stem (10–)25–40 cm, pale yellowish-green, often purplish towards the base, slender and flexuous, striate, with numer-

ous, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards, and with numerous stellate hairs and an occasional, short, dark glandular hair in the upper part. Leaves deep green on the upper surface, paler and often suffused reddish-purple beneath, rather thick; basal usually numerous, the lamina 2.5–8.0 × 1.5–3.5 cm, elliptical, ovate, ovate-lanceolate, or elliptic-oblong, rounded-obtuse to acute or long acuminate, subentire to denticulate or with a few, sharp teeth near the base and rounded or subtruncate at base, the petioles up to 5 cm and with numerous, long, wavy, pale simple eglandular hairs; cauline 0–1, when present linear-lanceolate, acuminate at apex, subentire and petiolate or bract-like; all with few to fairly numerous, short to medium simple eglandular hairs to nearly glabrous on the upper surface and numerous, short to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 2–9 capitula, compactly paniculatecorymbose, sometimes with a long lower branch; peduncles short and straight, with dense stellate hairs, numerous short to medium, pale simple eglandular hairs and numerous, short, dark glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5– 12 × 1.0–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, with an obtuse to more or less acute apex, with dense, short to long, dark or dark-based simple eglandular hairs, fairly numerous, short, dark glandular hairs, and dense stellate hairs on the margins and at the apex. Ligules yellow, with dense, short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins fimbriatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Native. Limestone cliff ledges and scree along the Craigy-nos ridge in Breconshire and at Burnmouth in Berwickshire. Endemic. Named after Harry Joseph Riddelsdell (1866–1941). 242. H. ebudicum Pugsley Ebudes Hawkweed Phyllopodous perennial herb with a branched stock. Stem 15–30 cm, pale yellowish-green, sometimes purplish towards the base, usually slender, with more or less numerous, long, pale simple eglandular hairs at the base becoming shorter upwards, and numerous stellate hairs and a few, very short, fine, dark glandular hairs in the upper part. Leaves medium glaucous green on upper surface, paler beneath; basal few to fairly numerous, the lamina 2–6 × 1.5–3.0 cm, ovate to elliptical or subrotund, rounded-mucronate to more or less acute at apex, denticulate or with few, small teeth towards the rounded or truncate base, the petioles up to 3 cm and with dense, long, pale, wavy simple eglandular hairs; cauline 0 or 1, when present usually linear and bract-like, sometimes ovate-lanceolate, acute at apex, dentate and narrowed to a broadly winged petiole; all with short to long, pale, rather stiff simple eglandular hairs on both surfaces and the margins. Inflorescence with 1–3(−4) capitula, furcate-corymbose; peduncles long, straight to curved, with dense stellate hairs, numerous unequal, short and very short, dark glandular hairs and a few, short to medium, pale simple eglandular hairs. Capitula 30–45 mm in diameter, rounded below. Involucral bracts porrect in bud, 5–14 × 1.2–1.5 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, long and narrow to

39. Hieracium an acute apex, the inner filamentous, with numerous, very short, short and medium, slender glandular hairs, numerous, short to medium, pale, dark-based simple eglandular hairs, and stellate hairs on the margins towards the base. Ligules clear yellow, with numerous, very short simple eglandular hairs at the apex. Styles yellow. Receptacle pits with margins fimbriate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rocky outcrops and stream banks. Outer Hebrides. Endemic. 243. H. eucallum P. D. Sell & C. West Spreading-toothed Hawkweed H. praetermissum P. D. Sell & C. West, non Juxip Phyllopodous perennial herb with a branched stock. Stems 10–50 cm, pale yellowish-green, often suffused purplish, especially below, slender but rigid to more or less robust, striate, with few, rarely more numerous, medium to long simple eglandular hairs and few to numerous stellate hairs and an occasional, short, dark glandular hair in the upper part. Leaves medium bluish-green and often faintly blotched and spotted on upper surface, paler beneath and sometimes suffused brownish-purple; basal numerous, 2–9 × 1–5 cm, ovate, elliptical or lanceolate, obtuse-mucronate to acute at apex, usually dentate or dentate-serrate, rarely denticulate, the teeth narrow and apiculate, cuneate or subtruncate at base, the teeth sometimes descending the short, densely long-hairy petioles; cauline 0–1(−2 cult.), lanceolate, longacute at apex, sharply and often deeply toothed, shortly petiolate or sessile, sometimes linear and bract-like; all glabrous or with scattered, short to medium, pale, subrigid simple eglandular hairs on upper surface and numerous, medium to long, pale simple eglandular hairs beneath and on the margins and especially the midrib. Inflorescence with 2– 10 capitula, narrowly cymose, sometimes with a long lower branch; peduncles medium to long, suberect, with numerous stellate hairs, few, pale, dark-based simple eglandular hairs and few, very short, dark glandular hairs. Capitula 30–50 mm in diameter, broadly rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse to subacute at apex, with numerous stellate hairs especially on the margins and at the apex, more or less dense, pale simple eglandular hairs and few, very short and short dark glandular hairs. Styles dirty yellow to discoloured. Ligules deep yellow, with very short simple eglandular hairs at apex. Receptacle pits with margins long-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and rocky streamsides. Western Scotland from Kirkcudbrightshire to Sutherland. Endemic. 244. H. saxorum (F. Hanb.) P. D. Sell & C. West Rock Hawkweed H. hypochaeroides var. saxorum F. Hanb.; H. wiesbaurianum subsp. hypochaeroides var. saxorum (F. Hanb.) Zahn; H. olivaceum subsp. llynense Zahn; H. sommerfeltii var. subtruncatum Pugsley Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes flushed purplish, slender to robust, with few, medium, pale simple eglandular hairs throughout or almost absent and few stel-

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late and sometimes a few, short, dark glandular hairs above. Leaves dark or bluish-green with faint to intense brownishpurple spots and blotches on the upper surface, paler and rather glaucous beneath; basal numerous, with lamina 2–10 × 1–4 cm, elliptical, lanceolate or ovate, mostly rounded-mucronate, but the inner sometimes acute, subentire, denticulate or with a few larger teeth towards the rounded or subtruncate base, the petioles short to long with dense, pale, curly simple eglandular hairs; cauline 0–1, when present lanceolate, acute at apex, often dentate, and sessile or petiolate or bract-like; all with the upper surface glabrous or nearly so, the lower surface with sparse, medium to long, pale simple eglandular hairs which are much more numerous on the midrib, the margins with numerous, short to long, pale, stiff simple eglandular hairs. Inflorescence furcate-corymbose, with 1–5 capitula; peduncles with few to numerous stellate hairs, more or less numerous, short, dark glandular hairs and occasional to few, short to medium, pale simple eglandular hairs. Capitula 30– 40 mm in diameter, rounded or subtruncate at base. Involucral bracts porrect in bud, 5–13 × 1.0–1.2 mm, blackishgreen, the inner with paler margins, linear-lanceolate, gradually narrowed to an acute apex, with stellate hairs mainly on the margins towards the base, numerous to dense, short and medium, pale to dusky simple eglandular hairs and few to numerous, inconspicuous, short glandular hairs. Ligules yellow, glabrous-tipped or with sparse hairs at tip. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, blackish. Flowers 6–8. Apomictic. Native. Rock-ledges, crevices and scree on basic cliffs. Scattered localities in Wales, Derbyshire, northern England and Scotland. Endemic. 245. H. dalense P. D. Sell Dales Hawkweed H. subplanifolium auct.; H. lasiophyllum var. planifolium auct. Phyllopodous perennial herb with a branched stock. Stem 20–45 cm, pale yellowish-green, often tinted purplish below, slender but rigid, striate, with a few, long, pale simple eglandular hairs below and numerous stellate and numerous, short, dark glandular hairs above. Leaves glaucousgreen on upper surface, paler and often tinted reddish beneath; basal usually numerous, the outer with lamina 3–9 × 1.5–4.0 cm, broadly elliptical or elliptical, usually obtuse-mucronate at apex, subentire or denticulate, and rounded or shortly cuneate at base, the inner with lamina 4–11 × 1.5–4.5 cm, elliptical, ovate or lanceolate, more or less acute at apex, subentire, denticulate or with a few, small teeth near the base and abruptly contracted or shortly cuneate at base, the petioles short to medium, with dense, long, pale simple eglandular hairs; cauline 0–1, when present similar to inner basal and petiolate or linear and acute at apex; all glabrous on upper surface, with numerous, short to medium, pale simple eglandular hairs beneath and those of the margin stiff. Inflorescence with 2–6 capitula, furcate-corymbose; peduncles rather short, more or less erect, with dense stellate hairs, numerous, short, dark glandular hairs and rarely a simple eglandular hair. Capitula 30– 35 mm in diameter, rounded at base. Involucral bracts more or less porrect in bud, 4–13 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate,

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the inner filamentous, all more or less acute at apex, with numerous, very unequal, short to medium dark glandular hairs, few, medium, dark-based simple eglandular hairs, and few stellate hairs mostly towards the base. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Native. Ledges on cliffs and rocky outcrops on the limestone. Fairly common in the Derbyshire Dales and extending into Staffordshire. Endemic. 246. H. subplanifolium Pugsley Flat-leaved Hawkweed H. lasiophyllum var. planifolium F. Hanb., non H. planifolium Brenner; H. pallidum subsp. lasiophyllum var. planifolium (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–40(−60 cult.) cm, pale yellowish-green, often pinkishred tinted towards the base, slender, striate, with few, scattered, short to medium, pale simple eglandular hairs and numerous stellate and few to numerous, short, dark glandular hairs in the upper part. Leaves thick, bluish-green on upper surface, paler beneath; basal usually numerous, the outer with lamina 2–5 × 1.5–3.0 cm, elliptical, ovate, or subrotund, rounded-obtuse at apex and entire, denticulate or with a few, small teeth at the rounded, subtruncate or abruptly contracted base; the inner with lamina 3–8 × 1.5– 4.0 cm, ovate or lanceolate, rarely oblong, acute or acuminate at apex, subentire, denticulate or with a few teeth at the subtruncate base; petioles usually rather short but can be up to 6 cm, with dense, medium to long, pale simple eglandular hairs; cauline usually 0, occasionally 1, like the inner basal and petiolate; all glabrous or with scattered, medium to long, rigid, pale simple eglandular hairs on upper surface, numerous softer hairs beneath especially on the midrib and numerous, rigid hairs on the margin. Inflorescence with 2– 5 capitula, furcate-corymbose; peduncles long and slender, erect, with dense stellate hairs, numerous, very short, fine, dark glandular hairs and an occasional short or medium, pale simple eglandular hair. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 6– 14 × 1.0–1.3 mm, greyish-green, the inner with paler margins, all narrow linear-lanceolate, narrowly drawn out to an acute apex, the inner very long and filamentous, with numerous, short to medium, dark-based simple eglandular hairs and fewer, short, dark glandular hairs, and numerous stellate hairs mainly along the margin and at the apex. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 4–6. Apomictic. Normally this species is quite distinct from H. lasiophyllum by its broad, acute inner leaves, but some plants generally resemble that species in having narrower obtuse inner leaves, but they never have the dense stellate hairs on the under surface as in H. lasiophyllum. Native. Limestone rocks. Symonds Yat and Coldwell Rocks in Gloucestershire, Piercefield Park in Monmouthshire and Great Doward in Herefordshire. Plants growing on a limestone wall at Tutshill in Gloucestershire were long known as H. schmidtii var. devoniense, but they are typical H. subplanifolium. Plants from Torrs Cliffs, Colvend in Kirkcudbrightshire and Cartland Crags,

Lanarkshire seem indistinguishable from H. subplanifolium. Endemic. 247. H. basalticola Pugsley Basalt Hawkweed H. stenolepis auct.; H. murorum var. pachyphyllum auct.; H. britannicum auct. Phyllopodous perennial herb with a branched stock. Stem (15–)30–50 cm, yellowish-green, sometimes purplish at the base, more or less robust, striate, with a few, long, pale simple eglandular hairs especially near the base and numerous stellate hairs in the upper part. Leaves bright or bluishgreen, paler beneath, often tinged purplish beneath; basal numerous, with lamina 2–11 × 1.2–6.5 cm, the outer and sometimes all elliptical, ovate or subrotund, rounded-obtuse at apex, entire or with a few teeth in the basal half and rounded at base, the inner sometimes more acute, with more prominent, narrowly mammiform teeth and their bases subtruncate and sometimes with retrorse teeth; petioles up to 5 cm, with numerous to dense, medium and long, pale, curly simple eglandular hairs; all glabrous or with an occasional simple hair on the upper surface, few to numerous, medium, pale simple eglandular hairs on the lower surface, numerous, sometimes subrigid simple eglandular hairs on the margin and numerous stellate hairs beneath. Inflorescence furcate-corymbose, often with fairly a long acladium, sometimes with a long lower branch, peduncles 1.5– 7.0 cm, erect, with dense stellate hairs, few to numerous, short, dark glandular hairs and sometimes an occasional, pale simple eglandular hair. Capitula 30–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 7– 12 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, abruptly narrowed to a drawn-out but obtuse apex, with numerous to dense stellate hairs especially on the margins, numerous short and medium, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs intermixed. Ligules pale yellow, with very short simple eglandular hairs at apex. Styles yellow or slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7(−8). Apomictic. Some plants with more pointed and more dentate leaves were included by H. W. Pugsley (1948) in H. britannicum, but they grow mixed with H. basalticola and all the Irish plants have more or less stellately hairy leaves, obtuse, stellately hairy involucral bracts and hairy ligules. Native. Ledges on limestone and basalt cliffs, Co. Clare, Co. Galway, Co. Sligo, Co. Leitrim, Co. Fermanagh, Co. Donegal, Co. Down, Co. Antrim and Co. Londonderry. Endemic. 248. H. apiculatidens P. D. Sell Apiculate-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–55 cm, pale yellowish-green, often brownish-purple towards the base, slender to fairly robust, striate, with a few, long, pale simple eglandular hairs below becoming fewer upwards, and fairly numerous stellate hairs in the upper part. Leaves dark bluish-green on upper surface, paler beneath, thick and rigid; basal usually numerous, and large and cabbagy, with lamina 2–10 × 1.5–6.0 cm, broadly ovate or broadly elliptical, sometimes nearly subrotund, mostly

39. Hieracium shortly acute, rarely rounded-mucronulate at apex, regularly dentate or serrate-dentate, the teeth sharp, apiculate, and sometimes curved, truncate, abruptly contracted or shortly cuneate at base, the petioles usually long, especially in Julyflowering plants, up to 11 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1, when present similar to the basal but much smaller, petiolate, or narrower and more deeply toothed; all glabrous on upper surface or with few, short or medium, pale simple eglandular hairs usually near the margin, and with more or less numerous, medium to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 4–7 capitula, furcatecorymbose; peduncles long, slender and erect, with dense stellate hairs, without simple eglandular or glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–11 × 1.0–1.5 mm, greyish-green, the inner paler at the margin, all linear-lanceolate, gradually narrowed but obtuse at apex, with dense stellate hairs at least on the margins but often all over, numerous short to medium, pale, dark-based simple eglandular hairs and few, short, dark glandular hairs. Ligules yellow, with few, very short hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 5–7. Apomictic. Native. Grassy banks and cliff ledges on limestone. In a few localities in Yorkshire. Endemic. 249. H. breconense P. D. Sell Brecon Hawkweed H. britannicum var. ovale Ley; H. dicella auct. Phyllopodous perennial herb with a branched stock. Stem 20–35 cm, pale yellowish-green, sometimes reddish-purple below, more or less slender, striate, with rather numerous, medium to long, pale simple eglandular hairs below and occasional ones upwards, and numerous to dense stellate hairs and few to numerous, very short, dark glandular hairs above. Leaves medium green and sometimes spotted and blotched brownish-purple on upper surface, paler and often suffused purplish beneath; basal numerous, the outer with lamina 3.5–8.5 × 2.5–4.5 cm, elliptical or broadly elliptical, rounded-mucronulate at apex, denticulate or sinuatedentate, rounded or abruptly contracted and often asymmetrical at base, the inner with lamina 4–12 × 2–5 cm, ovate, elliptical or slightly oblong-ovate, more or less acute at apex, denticulate to shortly sinuate-dentate and sometimes with a few, larger teeth at base, the petioles mainly short, but up to 5 cm and with dense, long, pale simple eglandular hairs; cauline sometimes large, lanceolate or ovate, acute or acuminate at apex, sinuate-dentate, sometimes with large teeth, usually petiolate; all glabrous or nearly so on upper surface, with numerous, long, pale simple eglandular hairs beneath, particularly on the midrib and numerous to rather dense, medium to long, rather stiff simple eglandular hairs on and near the margin. Inflorescence with 2–5(−8) capitula, in a rather tight corymb, sometimes with a lower branch; peduncles rather short, straight and more or less erect, with dense stellate hairs, numerous, short, unequal, dark glandular hairs and occasional or few, medium simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–15 × 1.0–1.3 mm, dark greyish-green, the inner with paler margins, all narrowly linear-lanceolate and grad-

347

ually drawn out from base to an acute apex, with numerous, medium, long, dark-based simple eglandular hairs, numerous, unequal, short to medium, dark glandular hairs and numerous stellate hairs particularly on the margins. Ligules bright yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6(−7). Apomictic. Native. Known only from limestone cliffs at Craig-ycilau and Dan-y-darren in Breconshire. Endemic. 250. H. dicella P. D. Sell & C. West Forked Hawkweed H. furcilliferum Dahlst., non H. furcelliferum Omang; H. britannicum auct.; H. britannicum var. glaucinum Pugsley Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, sometimes flushed reddish below, slender to robust, striate, glabrous or with a few, scattered, medium, pale simple eglandular hairs and a few stellate hairs above. Leaves thick, bluish-green on upper surface, pale beneath, sometimes tinted reddish-purple; basal often numerous, the outer with lamina 2–7 × 2.0–3.5 cm, elliptical or oblong-elliptical, rounded-mucronulate at apex, entire or denticulate in the upper half, denticulate to sinuatedentate below, the lower teeth slightly hooked, the base abruptly contracted or subtruncate, usually a few inner with lamina ovate, lanceolate, or ovate-oblong, acute at apex, dentate and truncate at base; petioles usually rather short, with numerous, long, pale simple eglandular hairs; cauline usually 0, rarely 1, linear, acute at apex, entire; all glabrous on upper surface, with numerous, medium, pale simple eglandular hairs beneath and very numerous, medium to long, rather stiff, pale simple eglandular hairs on the margin. Inflorescence with 2–6 capitula, furcate-corymbose; peduncles straight, rather long and suberect, with dense stellate hairs, few, short to medium, dark-based simple eglandular hairs and few, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 6–13 × 1.0–1.2 mm, dark greyish-green, the inner with paler margins, narrow linear-lanceolate, the inner filamentous, acute at apex, with numerous stellate hairs especially on the margins, numerous short to medium, pale simple eglandular hairs and few, short glandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 5–7. Apomictic. Native. Cliff ledges and scree on limestone, basalt and other calcareous rocks. Derbyshire Dales, Yorkshire limestone and scattered records throughout Scotland. Endemic. 251. H. sarcophylloides Dahlst. Fleshy-leaved Hawkweed H. dasypodum Dahlst.; H. pycnodon var. acutidens Dahlst.; H. sinclairii Dahlst.; H. pallidum auct.; H. britannicum auct.; H. sarcophyllum var. expallidiforme auct. Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, often tinted purplish towards the base, rather slender, nearly glabrous or with few to fairly numerous, medium to long, pale simple eglandular hairs and numerous stellate hairs in the upper part. Leaves deep green on upper surface, paler and often suffused

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238. H. stictum P. D. Sell (Yorkshire)

240. H. repandulare Druce (Breconshire)

242. H. ebudicum Pugsley (North Harris)

239. H. hypochaeroides S. Gibson (Yorkshire)

241. H. riddelsdellii Pugsley (Breconshire)

243. H. eucallum P. D. Sell & C. West (Sutherland)

Outer and inner leaves of species of the Section Stelligera Zahn

39. Hieracium

349

245. H. dalense P. D. Sell (Derbyshire) 244. H. saxorum (F. Hanb.) P. D. Sell & C. West (Carmarthenshire)

246. H. subplanifolium Pugsley (Gloucestershire) 247. H. basalticola Pugsley (Co. Leitrim)

248. H. apiculatidens P. D. Sell (Yorkshire)

249. H. breconense P. D. Sell (Breconshire)

Outer and inner leaves of species of the Section Stelligera Zahn

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purplish beneath; basal usually numerous, the outer with lamina 1.5–5.0 × 1–3 cm, subrotund or broadly elliptical, rounded at apex, subentire or denticulate and rounded at base, the inner with lamina 4–8 × 1.5–4.0 cm, ovate, more or less acute at apex, denticulate to more or less dentate, with rather sharp, narrowly mammiform teeth, especially at the more or less truncate base, the petioles up to 4 cm with dense, long, pale simple eglandular hairs; cauline 0–1, when present a smaller version of the inner basal or linear, shortly petiolate; all with fairly numerous, medium to long, pale simple eglandular hairs on both surfaces and especially the midrib beneath, the margins with dense, rather stiff hairs. Inflorescence with (1–)2–4(−6) capitula, furcate-corymbose; peduncles straight, erect-spreading, with dense stellate hairs, numerous, medium to long, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded below. Involucral bracts porrect in bud, 4– 12 × 0.6–1.0 mm, dark olive-green, the inner with paler margins, all narrow linear-lanceolate, obtuse to subacute at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, very short glandular hairs and stellate hairs along the margin and at the apex. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishbrown. Flowers 6–8. Apomictic. Native. Rocky crags, cliff ledges and rocky slopes. Creag nam Fitheach in Argyllshire; near Campbel town in Kintyre; Cul Beg in Ross-shire; near Cape Wrath, Foinaven, Durness, near Tongue and Ben Hope in Sutherland and several localities in the Orkney Islands. Faeroes. 252. H. fratrum Pugsley Shale Hawkweed H. sordidum W. R. Linton ex E. F. Linton, non Gillies Phyllopodous perennial herb with a branched stock. Stem 15–30(−35 cult.) cm, pale yellowish-green, sometimes purplish below, striate, rather slender, flexuous, with scattered stellate hairs particularly above, and occasional, medium, pale simple eglandular hairs mainly above. Leaves dull bluish-green and often spotted brownish-purple on upper surface, paler and more bluish beneath and sometimes tinted purplish; basal few to numerous, with lamina 3–6 × 2–3 cm, the outer ovate or subrotund, rounded-mucronulate at apex, subentire, denticulate or shortly dentate and rounded or subtruncate at base, the inner similar but often more or less acute at apex and more often toothed, the teeth sharply pointed and the basal pair sometimes reflexed and usually subtruncate at base; cauline often absent, when present small and bract-like; petioles up to 40 mm, often flushed purplish, with dense, long, curly simple eglandular hairs; all glabrous or with an occasional simple eglandular hair above, with medium, pale simple eglandular hairs scattered on the surface beneath and numerous on the midrib, numerous, short and medium, pale, subrigid (very rigid in cultivation) simple eglandular hairs on and near the margin and often numerous stellate hairs below. Inflorescence furcate-corymbose, with fairly long acladium; peduncles 5– 50 mm, with few to numerous stellate hairs, few to numerous, dark glandular hairs and usually a few, medium, pale, dark-based simple eglandular hairs. Capitula 30–47 mm in diameter, rounded at base. Involucral bracts porrect in

bud, 7–12 × 1.0–1.2 mm, dark greyish-green, the inner with paler margins, all linear-lanceolate, gradually drawn out to a narrow, acute apex, with numerous to dense stellate hairs especially on the margin, numerous, short to medium, pale, dark-based simple eglandular hairs and a few, short, dark glandular hairs. Ligules pale yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3–4 mm, reddish-black. Flowers 6–7. Apomictic. Native. On shaly, broken acidic rocks at Craigmichen Scars on the Selcoth Burn near Moffat (but growing with plants normally found on calcareous ground) and cliff ledges at the Devil’s Beef-tub in Dumfries-shire; cliff ledges at Craig an Dail Bheag (Little Craigandal) in Aberdeenshire; the Lairig Ghru in Inverness-shire; and rocks near Gallanach in Argyllshire. Endemic. Named after the brothers Edward Francis Linton (1848–1928) and William Richardson Linton (1850–1908). 253. H. britannicum F. Hanb. British Hawkweed H. wiesbaurianum subsp. britannicum (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–50(−60) cm, pale yellowish-green, sometimes tinted purplish below, more or less robust, erect, striate, with scattered, short to medium, pale simple eglandular hairs and occasional, short, dark glandular hairs and scattered stellate hairs in the upper part. Leaves usually rather flaccid, glaucous-green on upper surface, paler and more glaucous beneath, sometimes spotted, sometimes suffused purplish, basal numerous, with lamina 3–11 × 2–7 cm, the outer ovate, elliptical or subrotund, rounded to subacute and mucronate at apex, denticulate to dentate in the lower half, the teeth sharp, subtruncate at the base and sometimes with retrorse teeth, the inner ovate to broadly ovate or ovate-oblong, mostly acute-mucronate at apex, shallowly to deeply dentate, the teeth mammiform to aquiline-mammiform and often cusped, increasing in size downwards, unequal and the basal pair sometimes retrorse, narrow teeth sometimes disjunct on the petiole, and subtruncate at base; petioles up to 50 mm, with numerous to dense, medium to long, curly, pale simple eglandular hairs; cauline usually absent, when present linear or bract-like; all glabrous or with an occasional, medium simple eglandular hair on the upper surface, few to fairly numerous, short to long, pale simple eglandular hairs beneath, especially on the midrib and numerous to dense, pale, slightly rigid simple eglandular hairs on and near the margin. Inflorescence with 2–7(−28) capitula in fairly open cymosecorymbose, sometimes with a long, lower branch, with a rather long acladium; peduncles 15–60 mm, with numerous to dense stellate hairs, few to numerous, short, dark glandular hairs and few, short to medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at the base. Involucral bracts porrect in bud, 8–12 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linearlanceolate, long attenuate to an acute apex, with numerous to dense, short to long, pale, dark-based simple eglandular hairs, intermixed with few to numerous, short, dark glandular hairs, and with numerous stellate hairs on the margins. Ligules pale yellow, glabrous-tipped. Styles yellowish. Receptacular pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic.

39. Hieracium Native. Cliff ledges, rocks, quarries, rubble and grassy banks. The most abundant hawkweed in the Derbyshire and Staffordshire Dales in early summer. Endemic. 254. H. britannicoides P. D. Sell Confused Hawkweed H. britannicum auct.; H. britanniciforme auct.; H. stenolepis auct. Phyllopodous perennial herb with a branched stock. Stem 15–35 cm, pale yellowish-green, often tinted purplish below, slender to rather robust, erect, striate, with occasional, pale, short to medium simple eglandular hairs, and numerous stellate hairs and few to numerous, short, dark glandular hairs in the upper part. Leaves usually rather flaccid, more or less glaucous-green above, paler and more glaucous beneath and sometimes suffused purplish; basal numerous, with lamina 2–7(−9) × 1–4 cm, the outer ovate or lanceolate, rounded-mucronulate to more or less acute at apex, denticulate throughout or dentate in the lower half, the teeth mammiform, subtruncate and sometimes with retrorse teeth at base, the inner all ovate, acute and often twisted at apex, denticulate throughout or dentate in the lower half, the teeth mammiform, the base subtruncate and often with retrorse teeth; petioles up to 5(−7) cm, with dense, medium to long, curly, pale simple eglandular hairs; cauline often absent, sometimes with a linear lamina, sometimes like inner basal and petiolate; all glabrous or with an occasional, pale, medium simple eglandular hair on upper surface, few to fairly numerous, short to long, pale simple eglandular hairs beneath, especially on the midrib, and numerous to dense, pale, slightly rigid simple eglandular hairs on and near the margin. Inflorescence with 2–6 capitula, usually rather compactly cymosecorymbose, sometimes with a lower branch, the acladium usually rather short; peduncles 5–35 mm, with dense stellate hairs, numerous short, dark glandular hairs and sometimes an occasional simple eglandular hair. Capitula 20–35 mm in diameter, rounded at the base. Involucral bracts porrect in bud, 8–16 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, abruptly narrowed halfway up to a long, slender, sometimes filamentous, acute apex, with numerous, short to long, pale, usually dark-based simple eglandular hairs intermixed with rather few, short, dark glandular hairs, and few stellate hairs. Ligules pale yellow, glabrous-tipped. Styles yellowish. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. H. britannicoides is intermediate between H. britannicum and H. britanniciforme and has at different times been referred to both of them. It has the softer leaves and hair clothing of the former and the compact inflorescence, hairier sometimes filamentous involucral bracts, and more glandular peduncles of the latter. Native. Ledges and cracks on limestone cliffs. Great Ormes Head and Little Ormes Head in Caernarvonshire and Bryn Euryn at Colwyn Bay in Denbighshire. Endemic. 255. H. britanniciforme Pugsley Ormes Head Hawkweed H. britannicum auct.; H. britannicum var. subbritannicum auct.; H. caesium auct.; H. caesium var. smithii Baker

351

Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, sometimes tinted purplish below, more or less robust, erect, striate, with very few to fairly numerous, short to long, pale, curly simple eglandular hairs, few to numerous stellate hairs especially above and a few, short, dark glandular hairs above. Leaves thick and rigid, glaucous-green and usually spotted and blotched brownish-purple on the upper surface, paler, more glaucous and often suffused purplish beneath; basal numerous, often more uniform than in allied species, the outer or all with lamina 3–11 × 2–6 cm, ovate or subrotund, rounded at apex, with 2, large, reflexed basal teeth and sometimes other shallower ones above, subtruncate or subcordate at base, the inner all with lamina ovate, acute-mucronate at apex, with mammiform or aquiline-mammiform, unequal teeth in the lower half, the 2 lower often reflexed, occasionally only denticulate, subtruncate at base; petioles up to 30(−50) mm, often purplish, with dense, long, pale, shaggy hairs; cauline usually absent, sometimes with lamina lanceolate, acuminate at apex, entire or with a few, sharp teeth and subpetiolate; all glabrous or with few to fairly numerous, pale, rigid hairs on the upper surface, few to numerous, softer, pale simple eglandular hairs beneath, and numerous to dense, medium to long, pale, rigid simple eglandular hairs on and near the margins. Inflorescence with 1–5(−11) capitula usually rather compactly cymosecorymbose, rarely with a lower branch, the acladium of medium length; peduncles 10–40 mm, with dense stellate hairs, numerous, short, dark glandular hairs and sometimes an occasional, rarely few simple eglandular hairs. Capitula 40–50 mm in diameter, rounded at base. Involucral bracts porrect in bud, 8–14 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, abruptly narrowed halfway up to a long slender, sometimes filamentous, acute apex, with numerous, short to long, pale, often darkbased simple eglandular and numerous, short, dark glandular hairs intermixed, and stellate hairs numerous especially on the margins. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3–4 mm, reddish-black. Flowers 5–6. Apomictic. Native. Limestone cliffs by the sea at Great Ormes Head in Caernarvonshire and Ballygroggan in Kintyre, and steep grassy and stony slopes by the sea at Burnmouth and a sandstone scar of the River Whiteadder both in Berwickshire. Endemic. 256. H. vagicola P. D. Sell Tutshill Hawkweed H. britannicum var. subbritannicum auct.; H. subbritannicum auct. Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale green, sometimes tinted brownish-purple at base, striate, more or less robust, erect, glabrous or with an occasional, pale simple eglandular hair and a few stellate hairs above. Leaves usually rather rigid, slightly glaucousgreen on upper surface, paler and glaucous beneath, sometimes suffused with brownish-purple; basal numerous, with lamina 2–11 × 2–6 cm, the outer ovate, broadly ovate, broadly elliptical, broadly oblong or subrotund, broadly rounded-mucronulate at apex, deeply and irregularly dentate in all except the uppermost quarter, the teeth narrowly to broadly mammiform and often curved, and usually with

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250. H. dicella P. D. Sell & C. West (Stirlingshire)

251. H. sarcophylloides Dahlst. (Orkney)

252. H. fratrum Pugsley (Dumfries-shire)

253. H. britannicum F. Hanb. (Derbyshire)

254. H. britannicoides P. D. Sell (Caernarvonshire)

255. H. britanniciforme Pugsley (Caernarvonshire)

Outer and inner leaves of species of the Section Stelligera Zahn

39. Hieracium

256. H. vagicola P. D. Sell (Gloucestershire)

353

257. H. naviense J. N. Mills (Derbyshire)

258. H. subbritannicum (Ley) P. D. Sell & C. West (Breconshire) 259. H. stenolepiforme (Pugsley) (Somerset) P. D. Sell & C. West

260. H. stenopholidium (Dahlst.) Omang (Breconshire)

261. H. eustomon (E. F. Linton) Roffey (Somerset)

Outer and inner leaves of species of the Section Stelligera Zahn

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2 reflexed teeth at the truncate base, the inner with lamina ovate or lanceolate, acute or acuminate at the often twisted apex, usually deeply and very irregularly dentate, the teeth narrower, longer and more irregular than those of the outer, the petioles up to 60 mm, pale green, often tinted brownish-purple and with dense, very long, curly, pale simple eglandular hairs; cauline usually absent, when present linear and bract-like; all glabrous or with a very occasional simple eglandular hair on the upper surface, glabrous or with few to numerous, short to long, pale simple eglandular hairs beneath; especially on the midrib and with numerous, medium to long, pale, subrigid simple eglandular hairs on or near the margin. Inflorescence with 2–4(−7) capitula, usually fairly compactly cymose-corymbose, sometimes with a long lower branch, with short acladium; peduncles 15– 30(−60) mm, with dense stellate hairs, with few to fairly numerous, short, more or less equal, dark glandular hairs and sometimes with a few, medium simple eglandular hairs. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 8–16 × 1.0–1.5 mm, blackishgreen, at least the inner paler on the margin, all narrowly linear-lanceolate, with a long, narrow, almost filamentous, acute apex, with few to numerous, short to medium, pale, dark-based simple eglandular hairs, few to numerous, short, dark glandular hairs, and dense stellate hairs on the margin. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–6(−7). Apomictic. Native. Limestone cliff ledges and rocks and quarry debris. Above the River Wye, near Tutshill, Gloucestershire. Endemic. 257. H. naviense J. N. Mills Derby Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–35 cm, yellowish-green, usually tinted purplish below, rather slender, striate, with stellate hairs throughout, but particularly above, and a few, short simple eglandular and/or short, fine glandular hairs in the upper part. Leaves bluish-green on upper surface, paler beneath, sometimes purplish-tinted; basal few to numerous, with lamina 2– 5(−9 cult.) × 1.5–3.0(−4.0 cult.) cm, the outer elliptical, oblong or ovate, rounded-mucronate at apex, subentire to dentate and rounded or truncate at base, the inner with lamina ovate or oblong-ovate or obovate, more or less acute at apex, dentate, becoming deeply so, and often retrorse and descending the petiole in cultivation, the teeth narrowly mammiform or aquiline-mammiform, subtruncate at base; cauline often absent, when present usually linear or lanceolate and sometimes dentate; all glabrous or with occasional, pale, subrigid simple eglandular hairs on the upper surface, with sparse, medium, pale simple eglandular hairs beneath and numerous to dense, short and medium, more or less rigid, often curved simple eglandular hairs on and near the margin; petioles up to 30(−80 cult.) mm, often purplish, with dense, medium and long, pale, curly simple eglandular hairs. Inflorescence compactly cymose, with a short acladium; peduncles 30–40 mm, with dense stellate hairs and numerous, short, brownish-black glandular hairs and occasional short, pale simple eglandular hairs. Capitula 25–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 10–11 × 1.0–1.2 mm, rather pale

blackish-green with broad pale margins, all rather broad linear-lanceolate, abruptly narrowed two-thirds of the way up to an acute apex, with numerous stellate hairs, numerous, short, dark glandular hairs and usually very few, short or medium, pale simple eglandular hairs. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, purplish-black. Flowers 5–6. Apomictic. Native. Turfy ledges and cracks in rocks on limestone cliffs. Two colonies in Winnats Pass, Derbyshire. Endemic. 258. H. subbritannicum (Ley) P. D. Sell & C. West Limestone Hawkweed H. stenolepis var. subbritannicum Ley; H. britannicum var. subbritannicum (Ley) Pugsley; H. stenolepis auct. Phyllopodous perennial herb with a branched stock. Stem 20–42 cm, pale green, often tinted reddish below, striate, more or less robust, erect, with few, scattered, pale, short to long simple eglandular hairs, few stellate hairs below becoming numerous above and an occasional, short, dark glandular hair above. Leaves usually rather rigid, bright green to slightly glaucous above, more or less glaucous and paler beneath, often flushed or tinted reddish-purplish; basal numerous, with lamina 2–10 × 2–5 cm, the outer elliptical, ovate or subrotund, rounded-mucronulate at apex, denticulate to dentate mainly in the lower half and often with 2 reflexed teeth at the base and round, truncate or subcordate at base, the inner with lamina lanceolate, ovate or oblongovate, acute or acuminate or caudate at the often twisted apex, shallowly to deeply dentate, the teeth irregular, flattened mammiform, broadly mammiform or narrowly mammiform and sometimes cusped, the 2 basal usually large and reflexed and sometimes with a few teeth on the petiole, more or less truncate at base, the petioles up to 50 mm, pale green, often tinted reddish-purple, with numerous to dense, medium to long, curly simple eglandular hairs; cauline usually absent, when present the lamina usually linear or bract-like, occasionally like inner basal and petiolate; all glabrous or nearly so on the upper surface, glabrous or with short to long, pale simple eglandular hairs beneath, with numerous, pale, subrigid simple eglandular hairs on or near the margin. Inflorescence usually fairly compactly cymosecorymbose, sometimes with a long, lower branch; peduncles 10–20(−50) mm, with short acladium, with dense stellate hairs, numerous short, fairly equal, dark glandular hairs and few, medium to long, pale, dark-based simple eglandular hairs. Capitula 35–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 8–15 × 1.0– 1.5 mm, olive green to blackish-green, at least the inner paler on the margins, all narrowly linear-lanceolate with a long, narrow, filamentous, acute apex, with numerous, short to medium, occasionally long, pale, dark-based simple eglandular hairs, fewer, short, dark glandular hairs and sparse stellate hairs. Ligules pale yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–7. Apomictic. Native. Limestone cliff and quarry ledges, scree and rubble. Local in Gloucestershire, Herefordshire, Breconshire and Carmarthenshire. Endemic.

39. Hieracium 259. H. stenolepiforme (Pugsley) P. D. Sell & C. West Cheddar Hawkweed H. britannicum var. stenolepiforme Pugsley; H. stenolepis auct. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale green, often suffused purplish at the base, usually rather slender, flexuous, erect, striate, with scattered, short, medium and long, pale simple eglandular hairs throughout, usually a few, dark, short glandular hairs in the upper part and few to numerous stellate hairs above. Leaves flaccid, dark, slightly glaucous green, often paler between the veins, paler beneath, sometimes slightly tinted purplish; basal numerous, with lamina 3–12 × 2.0–5.0 cm, the outer lanceolate to ovate, rounded-mucronate to acute at apex, entire to denticulate or with a few basal teeth and more or less truncate at base, the inner with lamina lanceolate to oblong-lanceolate, acute to long-acuminate at apex, denticulate to deeply dentate, the teeth mammiform to aquiline-mammiform, often cusped and the basal ones retrorse, and usually truncate at base, all with petioles 20– 90 mm, slender and with numerous to dense, medium to long, pale, curly simple eglandular hairs; cauline usually absent, rarely 1, lanceolate, long-acute at apex, denticulate, truncate at base and shortly petiolate; all glabrous or with scattered, medium simple eglandular hairs on the upper surface especially near the margin, with few to numerous, short to long, pale simple eglandular hairs on the lower surface, and numerous, short to medium, sometimes subrigid, pale, curled simple eglandular hairs on the margins. Inflorescence with 1–6 capitula, laxly cymose; peduncles 10–60 mm, the acladium fairly short, with numerous to dense stellate hairs, numerous very short to short, fine, dark glandular hairs and few to numerous, short to medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts porrect in bud, 6–12 × 0.5–1.5 mm, olivegreen to blackish-green, the inner pale or with paler margins, all narrowly linear-lanceolate, the inner very narrow, gradually long-acute at apex, the inner very finely tapered, with numerous, short to long, pale, curly simple eglandular hairs, few to numerous, very short to short, dark glandular hairs, and few stellate hairs mainly towards the base. Ligules pale yellow, glabrous or with a few short hairs at tip. Styles pure yellow. Receptacle pits with margins subulatedentate. Achenes 2.5–3.0 mm, reddish black. Flowers 5–6. Apomictic. Native. Limestone cliff ledges, scree and rubble. Rare, endemic to Cheddar Gorge, Somerset. 260. H. stenopholidium (Dahlst.) Omang Western Hawkweed H. integrilaterum var. stenopholidium Dahlst. Phyllopodous perennial herb with a branched stock. Stem 25–40 cm, pale yellowish-green, sometimes reddish-purple towards the base, slender to slightly robust, striate, glabrous or with an occasional, short to medium simple eglandular hair, and a few stellate hairs or occasional short glandular hairs in the upper part. Leaves flaccid, bluish-green, paler beneath, rarely tinted reddish; basal often numerous, the outer with lamina 2–9 × 1.5–4.0 cm, elliptical, sub-

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rotund, oblong-elliptical or ovate-elliptical, roundedobtuse-mucronate, sinuate-dentate in the lower half, sometimes with a few mammiform teeth, and abruptly contracted or subtruncate at base to a fairly short petiole, the inner with lamina 4–10 × 1–3 cm, oblong, oblong-lanceolate, lanceolate, narrowly elliptical, more or less acute to acuminate at apex, dentate, with narrow to broad mammiform teeth in the lower half and abruptly contracted to subtruncate at base with rather long, long-hairy petioles, sometimes all the leaves are like the outer and sometimes all like the inner; cauline usually absent, sometimes 1, small, with lamina lanceolate, acute at apex and dentate; all with more or less numerous stellate hairs beneath, glabrous or nearly so on upper surface, with few to numerous, medium to long, pale simple eglandular hairs on the lower surface, especially the midrib, and on the margin where they are stiff to subrigid. Inflorescence with 1–6 capitula, furcatecorymbose, sometimes with a long lower branch; peduncles fairly long, suberect, with numerous stellate hairs and numerous, short, fine glandular hairs and sometimes an occasional, short simple eglandular hair. Capitula 30–50 mm in diameter, rounded at base. Involucral bracts porrect in bud, 3–10 × 0.7–1.0 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, acute at apex, with stellate hairs on the margin and at the base, numerous, pale, dark-based, short to medium simple eglandular hairs and numerous, very short and short, unequal, dark glandular hairs. Ligules yellow, glabrous. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–7. Apomictic. Native. Rocks and cliff ledges often on limestone. It has a very disjunct distribution over the western half of Great Britain from Devonshire to the Hebrides. It is also in Iceland. 261. H. eustomon (E. F. Linton) Roffey Bristol Channel Hawkweed H. schmidtii var. eustomon E. F. Linton; H. pallidum subsp. eustomon (E. F. Linton) Zahn; H. schmidtii var. devoniense F. Hanb.; H. pallidum subsp. devoniense (F. Hanb.) Zahn; H. devoniense (F. Hanb.) Roffey; H. eustomon var. devoniense (E. F. Linton) Pugsley Phyllopodous perennial herb with a branched stock. Stem 15–55 cm, yellowish-green, often reddish towards the base, more or less robust, striate, with numerous, very long, pale, wavy simple eglandular hairs below, which become absent above and few to numerous stellate hairs in the upper part. Leaves thick, bluish-green, paler beneath, often suffused reddish or purplish; basal few to numerous, the outer with lamina 4–10 × 1.5–5.0 cm, lanceolate to broadly ovate or subrotund, rounded-obtuse-mucronate at apex, entire, denticulate or with a few widely spaced teeth which are narrowly mammiform and sometimes pointed downwards, and rounded, cuneate or subtruncate at base to fairly short petioles, the inner with lamina 4–11 × 2–4 cm, lanceolate, elliptical or narrowly to broadly ovate, acute to acuminate at apex, dentate to serrate-dentate, the teeth narrow, cuneate at base to short, long-hairy petioles; cauline 0–2, the lower often large, with lamina lanceolate to broadly ovate, acute or more often acuminate with a long narrow apex, with

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widely spaced teeth which are widely to narrowly mammiform, rarely deeply laciniate, and cuneate or subtruncate at base to a short petiole, the upper sometimes also fairly large but narrow, usually only bract-like; all with medium to long, pale, rigid simple eglandular hairs on the margin and sometimes also in various amounts on the upper surface, with few to numerous, subrigid ones beneath, particularly on the midrib. Inflorescence with 1–6 capitula, furcate-corymbose, sometimes with long lower branches almost to base; peduncles fairly long, suberect, with numerous to dense stellate hairs, few to numerous, short to medium, pale simple eglandular hairs and few, short dark glandular hairs. Capitula 30–50 mm in diameter, rounded to subtruncate at base. Involucral bracts porrect in bud, 4–14 × 1.0–1.5 mm, dark green, the inner with pale margins, all linear-lanceolate, long drawn out to a narrow, subacute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs and few stellate hairs near the base. Ligules pale yellow, glabrous-tipped, sometimes stylose. Styles yellow. Receptacle pits with margins deeply subulate-dentate. Achenes 3.5–4.5 mm, blackish. Flowers 5–6. Apomictic. Closely allied to H. schmidtii from which it is distinguished by its large, thick leaves with irregular teeth and longer involucral bracts with fewer glandular hairs. As to whether the plants from Devonshire and Somerset are different from those in Wales, it is difficult to tell. Populations of both have been examined in the field and plants are very variable. Plants in Devonshire which are not found in Wales and vice versa occur, but considering all the plants it is difficult to separate them into two taxa. It seems also to be very close to H. veterascens from north Scotland and the Faeroes, but almost certainly must have had a different origin. Native. Cliff ledges and dry rocky slopes. Coastal areas of north Devonshire and Somerset, the Gower and Pembrokeshire and inland areas in the Brecon Beacons and Carmarthenshire. Endemic. Section 16. Vulgata (Griseb.) Willk. & Lange Taxon Vulgata Griseb.; Subsection Vulgata (Griseb.) P. D. Sell & C. West comb. inval.; Taxon Pleiophylla Peter; Section Pleiophylla (Peter) Zahn; Taxon Caesia Almq.; Subsection Caesia (Almq.) Dahlst.; Section Caesia (Almq.) A. R. Clapham; Subsection Euvulgata F. N. Williams; Section Euvulgata (F. N. Williams) A. R. Clapham; Subsection Pseudoreadea F. N. Williams; Subsection Subvulgata F. N. Williams, non (Almq.) Dahlst.; Subsection Vulgatina F. N. Williams Phyllopodous or hypophyllopodous perennial herbs. Stem usually hairy. Leaves mainly green, sometimes glaucous, sometimes suffused with purple, sometimes spotted; basal few to numerous, sometimes almost absent at flowering; cauline 2–14. Inflorescence paniculate-corymbose, with few to many capitula; peduncles and branches straight or curved. Capitula small to medium. Involucral bracts usually incumbent in bud, usually narrow, obtuse or acute at apex, variably clothed. Ligules mostly glabrous-tipped. Styles yellow or discoloured. Receptacle pits with margins usually dentate. Achenes dark.

Species of this section can usually be recognised by their rather few basal leaves, several cauline leaves, and large panicle of small to medium capitula. Species of the Section Oreadea can usually be distinguished by their often glaucous foliage on which are rigid or stiff hairs and fewer, larger capitula. Section Hieracium can usually be easily recognised by its 0–1 cauline leaves, though a few species hover on the boundary between the two sections. Species which grow in rocky places in mountains and particularly by streams, are common in Wales, northern England and Scotland, but are rather scarce in Ireland. Other native species occur on open heaths, hedgebanks and open woodland. By far the greater number of species appear to have been introduced into southern England and have been spread by road and rail to all parts of Great Britain. These species can become most abundant on disturbed ground. The species of the Section Vulgata are widespread over central and northern Europe where they are often common plants. 262. H. triviale (Norrl.) Norrl. Grey-headed Hawkweed H. vulgatum subsp. triviale Norrl.; H. laevicaule subsp. triviale (Norrl.) Zahn; H. vulgatum var. triviale (Norrl.) Pugsley; H. vulgatum var. nemorosum auct.; H. pinnatifidum auct.; H. adlerzii F. Hanb. Phyllopodous perennial herb with a branched stock. Stem 30–80 cm, pale yellowish-green, sometimes reddish-purple towards the base, usually robust, striate, with numerous, long, pale simple eglandular hairs at the base, becoming fewer upwards, and rather few stellate hairs above. Leaves pale yellowish-green on upper surface, paler and often washed reddish-purple beneath; basal rather few, the lamina 3–14 × 2.0–4.5 cm, broadly elliptical or ovate-lanceolate, obtuse-mucronate to acute at apex, denticulate to dentate, often incise-dentate with long, narrow, mucronate teeth, cuneate at base, the petioles medium with long simple eglandular hairs; cauline (2–)3–5(−6), the lower large, similar to basal, acute at apex and petiolate, the upper smaller and narrower; all glabrous or nearly so on upper surface or with a few short, stiff simple eglandular hairs, with more numerous, similar hairs beneath and on the margin. Inflorescence with 1–20 capitula, paniculate-cymose; peduncles rather short, erecto-patent, with numerous to dense stellate hairs and numerous, short or medium, pale simple eglandular hairs, without glandular hairs. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts 4– 11 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with few stellate hairs, numerous, short and medium, pale simple eglandular hairs and an occasional, short glandular hair. Ligules deep yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–8. Apomictic. The first valid description of H. adlerzii we can find is by F. J. Hanbury in 1894. It is based on specimens from the Menai Straits which are a mixture of H. triviale and H. anglorum. The brief description best fits H. triviale. Native. Open woods, roadside rocks, slate quarries and walls. Caernarvonshire, where it appears to be most frequent, and scattered records in Durham and Scotland. It

39. Hieracium may have been bypassed for H. vulgatum, and really be more common than the records indicate. Fennoscandia, Russia, Poland, Czechoslovakia and Austria. 263. H. vulgatum Fr. Common Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–80 cm, pale yellowish-green, often tinted reddish-purple in the lower half, slender to robust, striate, with numerous, short to long, pale simple eglandular hairs in the lower part becoming shorter and fewer in number upwards and numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves dull, pale or medium green on the upper surface, paler and often suffused reddishpurple beneath; basal usually few, the lamina 4–15 × 1–5 cm, oblong, narrowly to broadly elliptical, elliptic-oblong, lanceolate or lanceolate-oblong, obtuse-mucronate to acute at apex, minutely denticulate to dentate, the teeth sometimes large and mammiform, cuneate to attenuate at base, the petioles short to long and long-hairy; cauline 2–8, similar to basal and gradually smaller upwards, subentire to dentate, the teeth sometimes narrowly aquiline-mammiform, the lower usually shortly petiolate, the upper sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins, or the upper surface glabrous or nearly so. Inflorescence with 1–30 capitula, paniculatecorymbose, the upper part often subumbellate, often with a long lower branch; peduncles erect-spreading, straight, with dense stellate hairs, few to numerous, short to medium, pale simple eglandular hairs and usually a few, short, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–13 × 0.8–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, more or less acute at apex, with numerous, short to medium (not more than 1.0 mm), darkbased simple eglandular hairs, few to fairly numerous, short, dark glandular hairs and numerous stellate hairs especially on the margins. Ligules deep yellow, glabrous. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–8. Apomictic. The following forms may be quite distinct, but plants can often only be more or less referred to them. This widespread species shows much variation and the forms may not have become completely distinct when the plants became apomictic. Sometimes all the plants in a colony belong to one form, sometimes there is more than one form in a colony. (1) Forma vulgatum H. laevicaule subsp. triviale var. baltorum Zahn; H. vulgatum var. rosulatum Syme; H. acroleucum auct.; H. acroleucum var. daedalolepium auct. Plant usually slender. Basal leaves 4–10 × 2.0–2.5 cm, ascending, denticulate to dentate; cauline 2–4, usually smaller than basal and dentate. Capitula 1–20. Involucral bracts up to 10 mm. (2) Forma subfasciculare (W. R. Linton) P. D. Sell H. vulgatum var. subfasciculare W. R. Linton; H. subfasciculare (W. R. Linton) Roffey Plant usually robust. Basal leaves 6–11 × 1.0–2.5 cm, forming a spreading rosette, irregularly serrate-dentate;

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cauline 2–3, with coarse teeth. Capitula up to 30. Involucral bracts up to 12 mm. (3) Forma pseudosubramosum (Pugsley) P. D. Sell H. vulgatum var. pseudosubramosum Pugsley; H. subramosum auct. Plant usually robust. Basal leaves 6–10 × 2–3 cm, forming a spreading rosette, with narrow aquiline-mammiform teeth; cauline 4–8, smaller, narrower and less toothed than basal. Capitula up to 30. Involucral bracts up to 12 mm. (4) Forma subravusculum (W. R. Linton) P. D. Sell H. vulgatum var. subravusculum W. R. Linton; H. orarium var. ravusculum auct.; H. subravusculum (W. R. Linton) Roffey Plant usually robust. Basal leaves 4–8 × 1.5–3.0 cm, forming a spreading rosette, with spreading mammiform teeth; cauline 2–3, large, with coarse, spreading teeth. Capitula 6–20. Involucral bracts up to 10 mm. (5) Forma sejunctum (W. R. Linton) P. D. Sell H. vulgatum var. sejunctum W. R. Linton; H. laevicaule subsp. sejunctum (W. R. Linton) Zahn; H. sejunctum (W. R. Linton) Roffey Plant usually robust. Basal leaves 7–15 × 2.5–5.0 cm, often suffused reddish-purple, ascending, with spreading mammiform teeth; cauline 2–4 lower as large as basal, often with long spreading teeth. Capitula up to 25. Involucral bracts up to 13 mm. Native. Cliff ledges, grassy places and streamsides in upland areas, where it is native. By railways, roadsides, walls and waste places, where it is probably introduced. Common in Scotland, central and northern England and north Wales with a few records, probably introduced, further south. In Ireland it occurs in the north-east with a few records elsewhere. Abundant over large areas of Scandinavia and central Europe. An analysis of the 600-odd specimens of this species in the Cambridge herbarium (CGE) shows the following. Forma vulgatum is the commonest form found throughout the range of the species. This is borne out by observation in the field. Forma sejunctum is the next most common being frequent in northern England and Scotland, but seen only from Merionethshire in Wales. Impression in the field is that it is not as common as one would expect from the number of specimens. Forma subfasciculare is scattered through Wales, northern England and Scotland and may be more common than collecting suggests. Forma subravusculum, which is a difficult plant to recognise, is intermediate between forma subfasciculare and forma sejunctum, and is of scattered occurrence in Wales, northern England and Scotland. Forma pseudosubramosum is recorded only from Fifeshire, Perthshire, Argyllshire and Kintyre. 264. H. cravoniense (F. Hanb.) Roffey Craven Hawkweed H. duriceps var. cravoniense F. Hanb.; H. caesium subsp. cravoniense (F. Hanb.) Zahn; H. cravoniense var. pseudoduriceps Pugsley Phyllopodous perennial herb with a branched stock. Stem 25–60 cm, pale yellowish-green, often brownish- or

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reddish-purple towards the base, slender to robust, often flexuous, striate, with long, pale simple eglandular hairs throughout, and numerous stellate hairs and sometimes an occasional, short, dark glandular hair in the upper part. Leaves pale green on upper surface, even paler beneath and sometimes flushed reddish or purplish; basal few to fairly numerous, the lamina 3–10(−17) × 1–4(−5) cm, lanceolate, oblong, elliptical or elliptic-oblong, obtuse-mucronate to acute at apex, undulate-denticulate with larger, mammiform teeth towards the base or sometimes more deeply dentate with narrower teeth, cuneate or attenuate at base, the petioles up to 7 cm and with dense, long, pale, wavy simple eglandular hairs; cauline 2–4, the lower often like basal, sometimes even larger, acute at apex, incise-dentate with long, sharp teeth and petiolate, the upper smaller and narrower, acute at apex, dentate and sessile; all glabrous or with short, stiff simple eglandular hairs on the upper surface and numerous, short to long simple eglandular hairs beneath and on the margin. Inflorescence with 3–25 capitula, paniculate-corymbose, often with long lower branches; peduncles mostly rather short and slender, with dense stellate hairs, numerous, short to medium (−1.2 mm), pale, dark-based simple eglandular hairs and sometimes an occasional, rarely few, dark glandular hairs. Capitula 15–30 mm in diameter, usually with the flowers poorly developed, rounded at base. Involucral bracts incumbent in bud, 4– 10 × 1.0–1.2 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an obtuse apex, or with the inner acute, with dense, medium to long (mostly to 1.7, sometimes 2.0 mm), pale, dark-based simple eglandular hairs, numerous stellate hairs along the margin and at the apex, rarely with an occasional, short glandular hair. Ligules glabrous or with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with the margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Very similar to H. vulgatum with which it sometimes grows and to which it shows similar leaf variation, but no uniformity within a colony. It can be distinguished by its usually shorter involucral bracts, which are more obtuse at the apex, and by its longer simple eglandular hairs of the involucral bracts. It frequently has undeveloped ligules, a state which only rarely occurs in H. vulgatum. Native. Cliff ledges, grassy slopes, rocky woods and streamsides. Widespread and frequent in northern England and Scotland. Endemic. 265. H. coniops Norrl. Grey-haired Hawkweed H. coniops var. hebescens Brenner; H. subramosum subsp. coniops (Norrl.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale green, sometimes suffused purplish, sometimes flexuous, striate, with long, pale simple eglandular hairs below and few or none above, stellate hairs throughout and dense above, glandular hairs absent. Leaves medium yellowish-green on upper surface, paler beneath, sometimes flushed brownish-red; basal more or less numerous, the lamina 5–9 × 1.5–2.5 cm, oblong or narrowly oblongelliptical, obtuse-mucronate to acute at apex, denticulate to dentate, the teeth broad to narrow, cuneate to attenuate at base, the petioles short to rather long, densely hairy;

cauline 2–3, the lower similar to basal and shortly petiolate, the upper narrow or bract-like and sessile, entire to dentate; all with numerous, short to medium, pale simple eglandular hairs on both surfaces and the margins and numerous stellate hairs at least beneath, but often on both surfaces, especially on the cauline. Inflorescence with 2–20 capitula, paniculate-corymbose, the upper part sometimes subumbellate, sometimes with a long lower branch; peduncles with dense stellate hairs, few to fairly numerous simple eglandular hairs and occasional glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 0.8–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, narrowed to a fine, acute apex, with numerous to dense stellate hairs, numerous short or medium, pale simple eglandular hairs and a few, short, dark glandular hairs. Ligules medium yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 2.5–3.0 mm, reddishblack. Flowers 6–7. Apomictic. Native. Solitary records from Argyllshire, Arran and Kintyre. May be more common as it has probably been mistaken for H. vulgatum which it looks like in general appearance but has dense stellate hairs on the leaves. Described from Finland. 266. H. acroleucum (Stenstr.) Dahlst. Pale-headed Hawkweed H. diaphanum subsp. acroleucum Stenstr.; H. laevicaule subsp. acroleucum (Stenstr.) Zahn; H. acroleucum var. daedalolepium Dahlst. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes purplish below, slender and flexuous, with few to fairly numerous, long, pale simple eglandular hairs below, becoming fewer and shorter above, and numerous stellate hairs and a very occasional short, dark glandular hair in the upper part. Leaves bright medium yellowish-green on upper surface, paler beneath, sometimes stained purplish; basal few to numerous, with lamina 1.5–9.0 × 1–2 cm, narrowly lanceolate, narrowly elliptical or oblong-lanceolate, more or less acute at apex, subentire or minutely and shortly denticulate, attenuate at base, the petioles up to 5 cm, with fairly numerous, long, pale simple eglandular hairs; cauline 2–4, similar to basal, but getting smaller up the stem and the uppermost bract-like; all with short, pale simple eglandular hairs on the upper surface to nearly glabrous, and more or less numerous, short to long, pale simple eglandular hairs beneath and on the margin and sometimes some stellate hairs beneath. Inflorescence with 1–9 capitula, furcate-corymbose or sometimes subumbellate; peduncles long and straight, with dense stellate hairs and scattered, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–11 × 1.0–1.5 mm, dull pale green, the inner with paler margins, all narrow linear-lanceolate, gradually drawn out to a more or less acute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs and numerous stellate hairs particularly along the margin. Ligules yellow, glabrous-tipped. Styles yellowish or discoloured. Receptacle pits with margins obscurely dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic.

39. Hieracium The plants included here match specimens called H. acroleucum from Scandinavia, but are not very distinct from H. vulgatum forma vulgatum. In the field their narrow, paler green leaves and paler capitula seem to distinguish them. Our plants may best fit the var. daedalolepium. Native. Rocky banks of streams. Scattered localities in northern England and western Scotland and recorded from Llanberis in Caernarvonshire. Scandinavia. 267. H. caesionigrescens (Fr. ex Stenstr.) Omang Linton Falls Hawkweed H. silvaticum subsp. caesionigrescens Fr. ex Stenstr.; H. bifidum subsp. caesionigrescens (Fr. ex Stenstr.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–40(−80 cult.) cm, pale yellowish-green, often slightly brownish-purple at base, slender, striate, with a few, medium to long, pale simple eglandular hairs, particularly near the base, and few to fairly numerous stellate hairs above or nearly glabrous. Leaves medium green and lightly spotted brownish-purple on upper surface, paler and slightly bluish beneath; basal few to fairly numerous, the lamina 2–7(−11 cult.) × 1–2(−4 cult.) cm, lanceolate or elliptical, obtuse-mucronate or acute at apex, denticulate to shortly dentate, producing large, narrow teeth in cultivation, cuneate at base, the petioles fairly long and long-hairy; cauline 1–3, the lower similar to the basal and petiolate, the upper usually linear and bract-like; all with short to medium, subrigid, pale simple eglandular hairs on both surfaces and the margins or the upper surface nearly glabrous. Inflorescence with 2–8 (numerous in cultivation) capitula, cymosecorymbose; peduncles rather short and often curved, with dense stellate hairs and sometimes few to numerous, short, dark glandular hairs or short, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–11 × 0.8–1.2 mm, greyishgreen, the inner with paler margins, all linear-lanceolate, narrowed to an obtuse apex, with numerous stellate hairs, numerous, short to medium, dark-based simple eglandular hairs and a few, short, dark glandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 6–7. Apomictic. Native. Known only from boulders in the river at Linton Falls, slate cliffs on the top of Ingleborough and limestone at Skirethorns (which may be the same as Linton Falls) all in Yorkshire and a low, basalt cliff by the road to Loch Humphrey in Dunbartonshire. Southern Scandinavia. 268. H. pollichiae Sch. Bip. Roffey’s Hawkweed H. praecox var. oxyodontoides (Sch. Bip.) Sch. Bip.; H. pallescens var. pollichiae (Sch. Bip.) Arv.-Touv.; H. roffeyanum Pugsley Phyllopodous perennial herb with a branched stock. Stem 30–80 cm, pale yellowish-green, often flushed reddishpurple at the base, slender to robust, striate, with numerous, medium to long, pale simple eglandular hairs below, becoming less numerous upwards, and numerous stellate hairs and an occasional, short, dark glandular hair above. Leaves pale yellowish-green with brownish spots and blotches on upper surface, paler beneath and sometimes tinted purplish; basal few, the lamina 5–12 × 2–5 cm, elliptical to

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oblong-elliptical or oblong-lanceolate, obtuse-mucronate to acute at apex, denticulate to dentate, occasionally the teeth large and mammiform, cuneate or abruptly contracted at base, the petioles long and long-hairy; cauline 2–4, the lower usually large and similar to basal, acute at apex and often with large teeth, petiolate; all with short to long, stiff, pale simple eglandular hairs on both surfaces and the margins or nearly glabrous on upper surface. Inflorescence with 2–8 capitula, paniculate-corymbose; peduncles rather short, suberect or curved, with dense stellate hairs, numerous, short to medium, pale simple eglandular hairs and occasional glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4– 11 × 1.0–1.5 mm, dark green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, medium to long, pale simple eglandular hairs, few, short, dark glandular hairs, and a few stellate hairs towards the base. Ligules yellow, glabrous-tipped. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Grassy, shady banks, old walls and hedgerows. Kent, Sussex, Surrey and Hampshire. France, Germany, Switzerland, Austria, Hungary and Bulgaria. Named after Johann Adam Pollich (1740–80) and the synonym after John Roffey (1860–1927). 269. H. peccense (W. R. Linton) P. D. Sell Peak Hawkweed H. rubiginosum var. peccense W. R. Linton; H. sagittatum subsp. rubiginosum var. peccense (W. R. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 40–80 cm, pale yellowish-green, often purplish below, more or less robust, striate, with more or less numerous, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards, and with numerous stellate hairs in the upper part. Leaves bluish-green on upper surface, paler beneath and often suffused purplish; basal few, the lamina large, 3–12 × 2–7 cm, elliptical, ovate or oblong-elliptical, obtuse-mucronate to acute at apex, subentire to sinuatedenticulate or shallowly dentate, and cuneate at base, the petioles up to 8 cm; cauline 1–3, similar to basal and petiolate, the upper narrower and sessile and sometimes bractlike; all glabrous on upper surface, with few to fairly numerous, short to medium, pale simple eglandular hairs beneath, especially on the midrib, and on the margin. Inflorescence with 5–18 capitula, paniculate-corymbose, often with a long lower branch; peduncles rather short and slender, with dense stellate hairs, numerous, short, dark glandular hairs and few to numerous, short to medium, pale simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.5 mm, blackishgreen, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, short, dark glandular hairs, few to numerous, short to medium, pale, darkbased simple eglandular hairs, and stellate hairs along the margin and near the base. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and scree. Derbyshire and Staffordshire Dales. Endemic.

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270. H. caesiopilosum Pugsley Caesious Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–55 cm, pale yellowish-green, often suffused with brownish-purple below, more or less robust, striate, with numerous, long, pale, wavy simple eglandular hairs throughout and numerous stellate hairs in upper part. Leaves bluish- or greyish-green on upper surface, paler beneath; basal few to rather numerous, the lamina 5–12 × 2.5–5.0 cm, narrowly to broadly elliptical to oblong-elliptical or more or less ovate, obtuse-mucronulate to acute at apex, sinuate-dentate, the teeth sharply mammiform, and cuneate at base, the petioles up to 6 cm and with dense, long, wavy, pale simple eglandular hairs; cauline 2–4, gradually decreasing in size upwards, lower like basal and shortly petiolate, the median with lamina ovate, acute to cuspidate at apex, sharply dentate, sessile or shortly petiolate, the upper usually bract-like; all with scattered, medium to long, pale simple eglandular hairs on both surfaces and more numerous on the margins. Inflorescence with 2–10(−20) capitula, furcate-corymbose, with long, erect branches; peduncles long and slender, with dense stellate hairs, numerous, short to medium, pale, spreading simple eglandular hairs and few, short, dark glandular hairs. Capitula 30–40 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4.0–10.5 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all linear-lanceolate, obtuse to subacute at apex, with dense, short to medium, pale simple eglandular hairs, few, very short, dark glandular hairs, and stellate hairs along the margins and at the apex. Ligules pale yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.5–3.7 mm, reddishblack. Flowers 6–7. Apomictic. Native. Streamsides and rocky places. Upper Teesdale, Poldores Burn in Kirkcudbrightshire, near Dalmellington and Straiton in Ayrshire, and near Fintry and in the Campsie Hills in Stirlingshire. Endemic. 271. H. robertsii P. D. Sell Roberts’s Hawkweed H. rubiginosum var. glabrescens Pugsley, non H. glabrescens Murr Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale green, often suffused with purple below, rather slender and flexuous, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter above, and few to numerous stellate hairs and an occasional, short, dark glandular hair in the upper part. Leaves dark, dull green on upper surface, paler beneath; basal few, with lamina 5.5–13.0 × 2.0–4.5 cm, elliptical, elliptic-ovate or elliptic-oblong, obtuse-mucronulate to acute at apex, denticulate to very shallowly sinuate-dentate, cuneate at base, the petioles up to 6 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 2–3, the lowest usually large, petiolate and like the basal, the upper linear or bract-like; all glabrous or nearly so on upper surface, with a few, scattered, short or medium simple eglandular hairs beneath or on the margin. Inflorescence with 3–9 capitula, paniculate-corymbose, often with a long lower branch; peduncles rather short, straight or curved, with dense stellate hairs, numerous, short to medium, dark-based simple eglandular hairs and numerous, very short and short

(0.2–0.7 mm), dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed but subobtuse at apex, with numerous, short and medium, darkbased simple eglandular hairs, numerous, very short and short (0.2–0.7 mm), dark glandular hairs and a few stellate hairs along the margins and towards the base. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 6–7. Apomictic. Native. Known only from the cliffs of Craig Dulyn in Caernarvonshire. Endemic. Named after Evan Roberts (1906–91). 272. H. kintyricum P. D. Sell Kintyre Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, suffused brownish-purple in the lower half, slender to more or less robust, striate, with numerous, long, pale simple eglandular hairs below, becoming very scattered and much shorter upwards or absent, with few to numerous stellate hairs in the upper part. Leaves medium glaucous green on upper surface, paler beneath; basal few, the lamina 4–10 × 2.0–4.5 cm, broadly ovate, broadly elliptical or elliptical, narrowed above, but roundedobtuse at apex, subentire, remotely denticulate or with a few, small teeth, and abruptly contracted or cuneate at base, the petioles up to 4 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 3–5, the lowest, and often the first 2, like basal but sometimes with more numerous teeth and petiolate, the upper with lamina smaller, linear-lanceolate, long acute, dentate and sessile, or bract-like; all glabrous or nearly so on upper surface, with numerous, short to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 2–7 capitula, paniculate-corymbose; peduncles rather slender, with dense stellate hairs, numerous short to medium, pale simple eglandular hairs and a very occasional, short, dark glandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.5 mm, mostly pale green with little darkening in the centre, linear-lanceolate, obtuse at apex, with numerous, short to medium, pale simple eglandular hairs, few to fairly numerous, short, dark glandular hairs obscured by the simple hairs and few stellate hairs towards the base. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits and achenes not seen. Flowers 6–7. Apomictic. Native. Known only from a ravine on the Largybaan in Kintyre. Endemic. 273. H. aequiserratum P. D. Sell Equal-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–55 cm, pale yellowish-green, often purplish-red below, more or less slender, sometimes flexuous, striate, with numerous, long, pale simple eglandular hairs below, becoming gradually fewer and shorter upwards and sometimes very sparse above, and dense stellate hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal numerous, the lamina 3–8 × 1.5–3.5 cm, usually elliptical, sometimes ovate or lanceolate, obtuse to acute at apex, equally sinuate-serrate, the teeth small

39. Hieracium and mostly forward-pointing with a small apiculus at apex, cuneate to attenuate at base, the petioles mostly rather short, but up to 4 cm and with dense, long, pale simple eglandular hairs; cauline 2–3, similar to basal but smaller, narrower, sharply acute at apex and with a short, winged petiole, the uppermost usually bract-like; all with short, stiff, pale simple eglandular hairs on the upper surface and numerous, short to medium, pale simple eglandular hairs beneath and on the margin. Inflorescence with 2–10 capitula, compactly paniculate-corymbose; peduncles rather short, with dense stellate hairs, numerous, short to medium, pale simple eglandular hairs and few to numerous, short, dark glandular hairs. Capitula 25–30 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–10 × 1.0– 1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs, and numerous stellate hairs along the margins and at the apex. Ligules yellow, with occasional very short hairs at apex. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Grassy embankment of disused railway at Barras and walls of Shap Abbey, both in Westmorland. Endemic. 274. H. vinifolium P. D. Sell Claret-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale green often suffused with purplish-red, usually slender, rarely robust, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming much fewer and shorter upwards, and with few stellate hairs in the upper part. Leaves bluish-grey on upper surface, paler and often tinted reddish-purple beneath, whole leaf sometimes suffused reddish-purple; basal few to numerous, with the lamina 2–10(−14) × 2–5(−6) cm, elliptical, ovate or lanceolate, rounded-obtuse to subacute at apex, denticulate to dentate, the upper third sometimes entire, mostly cuneate, sometimes attenuate or subtruncate at base, the petioles up to 5 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 2–8, the lower and median like basal, shortly petiolate or sessile, the upper with lamina lanceolate, acute at apex, dentate and sessile or bract-like; all glabrous or with short, pale simple eglandular hairs on upper surface, with numerous, medium to long, pale simple eglandular hairs beneath and on the margins. Inflorescence with 2–16 capitula, corymbose, sometimes with a long lower branch; peduncles short, sometimes very short and often curved, with dense stellate, numerous, short and medium, pale, dark-based simple eglandular hairs and numerous, short, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–10 × 0.8–1.0 mm, dark olive green, the inner with paler margins, all linear-lanceolate, gradually tapered to an obtuse or subacute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs and an occasional stellate hair. Ligules deep yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins sharply incise-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic.

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Native. Rock-ledges on coastal cliffs. On and near the Mull of Kintyre and further north at Barrahormid in the same county. Endemic. 275. H. rubiginosum F. Hanb. Rusty-red Hawkweed H. vulgatum var. rubescens F. Hanb.; H. sagittatum subsp. rubiginosum (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often flushed reddish below, slender to robust, striate, with numerous, medium to long, pale, wavy simple eglandular hairs below, becoming less numerous and shorter upwards, and numerous stellate hairs in the upper part. Leaves dull medium green on upper surface, paler beneath and often suffused reddish; basal few to numerous, the lamina 4–12 × 2–5 cm, narrowly to broadly elliptical or lanceolate-elliptical, obtuse-mucronate to acute at apex, sharply dentate, the teeth narrowly mammiform, irregular and sometimes very narrow and cusped, cuneate at base, the petiole up to 6 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 1–3, the lower like the basal, shortly petiolate or sessile, the upper lanceolate, acute at apex, dentate and sessile or bract-like; all with few to numerous, short to long, pale simple eglandular hairs on both surfaces and the margin. Inflorescence with 3–9 capitula, paniculate-corymbose; peduncles short and often curved, with dense stellate hairs, numerous, medium, pale, dark-based simple eglandular hairs and few, short, dark glandular hairs. Capitula 30–35 mm in diameter, rounded or subtruncate at base. Involucral bracts incumbent in bud, 4–12 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse to more or less acute at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, some short, dark glandular hairs, and some stellate hairs along the margin. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and scree on limestone. Common in Yorkshire and Durham. Endemic. 276. H. fulvocaesium Pugsley Orange-flowered Hawkweed H. orarium var. fulvum F. Hanb. Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, often purplish below, usually slender, with numerous, medium to long, wavy, pale simple eglandular hairs below becoming fewer or absent upwards, and stellate hairs in the upper part. Leaves pale bluish-green on upper surface, paler beneath; basal few, the lamina 2.5–10.0 × 1.5–3.5 cm, elliptical, ovate or lanceolate, rarely subrotund, rounded-mucronulate to acute at apex, sharply denticulate to serrate with sharply ascending teeth which are sometimes cusped especially in cultivation, and cuneate at base, the petioles up to 7 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 2–4, the lowest with lamina elliptic-lanceolate, acute or acuminate at apex, coarsely and sharply serrate, the teeth sharp and sometimes cusped, in cultivation sometimes becoming laciniate with long, narrow teeth and shortly petiolate, the upper small and often laciniate, the teeth long and narrow, the uppermost bract-like; all glabrous or nearly so on upper surface,

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with numerous, short and medium, pale simple eglandular hairs beneath and long ones on the midrib beneath and the margin. Inflorescence with 1–numerous capitula, paniculate-corymbose; peduncles mostly short, with dense stellate hairs, numerous, short or medium, pale simple eglandular hairs and fairly numerous, short glandular hairs. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, dark greyishgreen the inner with paler margins, all linear-lanceolate, obtuse to subacute at apex, with dense stellate hairs on the margin and at the apex, numerous short and medium simple eglandular hairs and very short, dark glandular hairs. Ligules orange-yellow, short, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only from low cliffs on the east bank of the River Naver, below Bettyhill in Sutherland. Endemic. 277. H. amnicola P. D. Sell Riverside Hawkweed H. rubiginosum var. rivulare Pugsley, non H. rivulare Krocker Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale green, sometimes reddish below, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer and rather shorter above, and numerous stellate hairs in the upper part. Leaves dark green on upper surface, paler beneath; basal few, the lamina 3– 8 × 1.5–4.0 cm, ovate or ovate-elliptical, obtuse-mucronate to acute at apex, subentire or denticulate, sometimes with an occasional tooth, cuneate or abruptly contracted at base, the petioles up to 5 cm and densely long-hairy; cauline 1–4, the lower similar to basal and petiolate, the upper smaller, with lamina linear-lanceolate or lanceolate, entire or with an occasional tooth; all with numerous, short to long, stiff, curved simple eglandular hairs on both surfaces and the margins. Inflorescence with 2–12 capitula, paniculatecorymbose; peduncles long and more or less straight, with dense stellate hairs, numerous short to medium, pale simple eglandular hairs and fairly numerous glandular hairs. Capitula 30–42 mm in diameter, rounded to subtruncate at base. Involucral bracts incumbent in bud, 6–13 × 1.0–1.2 mm, blackish-green, with paler margins, all linearlanceolate, acute at apex, with numerous to dense, medium to long, pale, dark-based simple eglandular hairs, more or less numerous, short, dark glandular hairs, and few stellate hairs towards the base. Ligules yellow, with a few, very short hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Limestone bank of the River Wharfe at Linton Falls, near Grassington, Ingleborough Scars above Chapelle-dale and Twistleton Scaurs all in Yorkshire. Endemic. 278. H. stenophyes W. R. Linton Waterfall Hawkweed H. euprepes var. stenophyes (W. R. Linton) F. N. Williams; H. duplicatum var. stenophyes (W. R. Linton) W. R. Linton Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale green, striate, rather slender, with few simple eglandular hairs below and nearly glabrous above. Leaves dull medium green on upper surface, paler beneath;

basal few, with lamina 4.5–9.0 × 1.5–3.5 cm, lanceolate to oblong, obtuse-mucronate to acute at apex, distantly denticulate to sinuate-dentate, with broad, spreading, unequal, apiculate teeth, more or less abruptly narrowed below, the petioles up to 5 cm and slender; cauline 0–2, the lower with lamina lanceolate, acuminate at apex, with sharp, spreading teeth and petiolate, the upper similar but smaller and sessile; all clothed with scattered, short to medium, pale simple eglandular hairs on both surfaces and the margins, the petioles more densely hairy. Inflorescence with (3–)4–8(−30 cult.) capitula, with straight, erect branches exceeding the acladium; peduncles with dense stellate hairs, scattered simple eglandular hairs and sometimes a few, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts 6–13 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all broad linear-lanceolate, mostly obtuse at apex, with numerous, medium, dark simple eglandular hairs, few, short, dark glandular hairs, and some stellate hairs towards the base. Ligules yellow, glabrous-tipped. Styles yellowish. Receptacle pits with margins fimbriate-dentate. Achenes 3.0–3.5 mm, blackish. Flowers 6–7. Apomictic. Native. Rock-ledges and streamside rocks. Known only from Dumfries-shire and Peebles-shire. Endemic. 279. H. rhomboides (Stenstr.) Johanss. Rhombic-leaved Hawkweed H. gravestellum var. rhomboides Stenstr.; H. caesium var. rhomboides (Stenstr.) W. R. Linton; H. subramosum subsp. gravestellum var. rhomboides (Stenstr.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often tinted purplish below, nearly glabrous or with an occasional medium, pale simple eglandular hair and a few stellate hairs above. Leaves bluish to dark green on upper surface, paler beneath; basal few, the lamina 4.5–8.0 × 2.5–4.0 cm, ovate, elliptic-ovate, rhombic, lanceolate or elliptical, rounded-mucronulate to acute at apex, denticulate or dentate, with a few, sharp, spreading teeth, cuneate at base; cauline 1–3, similar to basal or with more, larger, sharp, spreading teeth, with petioles up to 15 mm or sessile; all with short to medium, pale simple eglandular hairs on both surfaces and the margins to nearly glabrous on upper surface. Inflorescence with 2–8 capitula, paniculate-corymbose; peduncles short to medium, slender, more or less straight, with more or less numerous stellate hairs and occasional short or medium simple eglandular hairs and/or short, dark glandular hairs. Capitula 26–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.5 mm, dark olivegreen, the inner with paler margins, all linear-lanceolate, narrowed to an obtuse or subacute apex, with sparse stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and few, short, slender, dark glandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rocks by streams, probably always on basic soils. Bethesda in Caernarvonshire, Ribbleshead in Yorkshire, the Campsie Hills in Stirlingshire, Allt Dubhchlair in Perthshire and the Bealach and near Kilmelfort in Argyllshire. Sweden.

39. Hieracium 280. H. subramosum L¨onnr. Pettycur Hawkweed Phyllopodous perennial herb with a branched stock. Stem 40–70 cm, pale yellowish-green, more or less robust, striate, with numerous, medium to long, pale simple eglandular hairs below, becoming fewer or absent upwards, and some stellate hairs in the upper part. Leaves medium yellowishgreen on upper surface, paler beneath; basal few, the lamina 4–13 × 1.5–5.0 cm, narrowly to broadly elliptical, lanceolate or oblong-lanceolate, acute at apex, dentate towards the base with a few, rather narrow, sharp, curved teeth and cuneate to attenuate at base, the petioles up to 11 cm, pale green, sometimes tinged purple, with long, pale, wavy simple eglandular hairs; cauline 2–4(−5), decreasing upwards, the lamina lanceolate or oblong-lanceolate, acute at apex, more or less dentate and narrowed to a sessile base, the upper usually narrower or bract-like; all with numerous, short to long simple eglandular hairs on both surfaces and the margins or glabrous on the upper surface. Inflorescence with 5–10 capitula, laxly paniculate-corymbose, often with a long lower branch; peduncles long and rather thick, with dense stellate hairs and more or less numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all rather broad linear-lanceolate, more or less acute apex, with numerous, short to medium, dark simple eglandular hairs, few, short, dark glandular hairs, and stellate hairs at the base and along the margins. Ligules yellow, with sparse, very short hairs at apex. Styles discoloured. Receptacle pits with margins incisedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Probably native. Known only from specimens collected on the shore between Burntisland and Pettycur in Fifeshire in 1876 and 1878. Southern Scandinavia. 281. H. praesigne (Zahn) Roffey Distinguished Hawkweed H. diaphanum var. praestans W. R. Linton; H. vulgatum subsp. praesigne Zahn; H. praesigne var. subdiaphanum Pugsley Phyllopodous perennial herb with a branched stock. Stem 25–70 cm, pale yellowish-green, often flushed reddishpurple, slender but rigid to more or less robust, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate and occasional, short, dark glandular hairs in the upper part. Leaves dull, dark green on upper surface, paler beneath and often flushed purplish, thick; basal few to fairly numerous, the lamina 4–12 × 1.5–3.0 cm, mostly narrowly elliptical, lanceolate or oblong-elliptical, sometimes oblong or elliptical, obtuse-mucronulate to acute at apex, subentire, denticulate or dentate, the teeth usually shallow and mammiform, sometimes larger and cusped, cuneate or attenuate at base, the petioles up to 8 cm, often purple, with numerous, long, pale, wavy simple eglandular hairs; cauline (2–)3–5 (−6), the lower with lamina narrowly oblong-elliptical, more or less acute at apex, usually denticulate or with small teeth, rarely with larger, longer teeth, cuneate or attenuate at base to long petioles, the upper with lamina lanceolate

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or linear, long-acute at apex, entire to dentate, sessile or shortly petiolate; all glabrous or nearly so on upper surface, with more or less numerous, short to medium, pale simple eglandular hairs beneath and on the margin, and longer ones on the midrib. Inflorescence with 6–20 capitula, paniculatecorymbose, often with a long lower branch; peduncles slender and erect, with dense stellate hairs, numerous short, dark glandular hairs and few to fairly numerous, short to medium, pale simple eglandular hairs. Capitula 25–35 mm, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.0– 1.2 mm, olive- to dark green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short and medium (0.4–1.0 mm), black, glandular hairs and occasional to few, short and medium, pale, dark-based simple eglandular hairs, without stellate hairs. Ligules yellow, glabrous-tipped. Styles yellow to discoloured. Receptacle pits with margins shortly dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges, rubble, grassy slopes and riverside rocks. Common on the limestone of England from Derbyshire northwards and in Kirkcudbrightshire and the Isle of Raasay. Endemic. 282. H. perscissum Jord. ex Boreau Long-toothed Hawkweed H. vulgatum subsp. perscissum (Jord. ex Boreau) Zahn; H. vulgatum subsp. deductum microgen. aurulentum var. perscissum (Jord. ex Boreau) Sudre; H. argutidens Uechtr.; H. pinnatifidum L¨onnr. ex Dahlst., non Willd.; H. vulgatum subsp. pinnatifidum Zahn; H. neopinnatifidum Pugsley Phyllopodous perennial herb with a branched stock. Stems 30–60 cm, pale green, often suffused with brownish-purple, robust but flexuous, markedly striate, with numerous to dense, short to long, pale simple eglandular hairs and numerous stellate hairs in the upper part. Leaves pale yellowishgreen on upper surface, paler and sometimes suffused purple beneath; basal numerous, the lamina 7–12 × 2–5 cm, broadly elliptical or ovate, obtuse-mucronate to more or less acute at apex, laciniate-dentate to incise-dentate, the teeth very unequal, from mammiform to long and narrow with an apiculate tip, sometimes free from the cuneate base, the long-hairy petiole up to 5 cm; cauline 3–8, gradually decreasing in size upwards, elliptical, oblong-elliptical, lanceolate or ovate, deeply incise-dentate to lobate, the teeth mostly long and narrow, apiculate at apex and very unequal, cuneate to attenuate at base; all with short to long, pale simple eglandular hairs on both surfaces and the margins, but often few or none on the upper surface, the upper cauline with some stellate hairs beneath. Inflorescence with few to numerous capitula, often with long lower branches, sometimes from near the base, the upper part corymbose; peduncles rather short, with dense stellate and numerous, short or medium, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud 4–10 × 0.8–1.2 mm, blackish-green, the inner with paler margins, all narrow linear-lanceolate, obtuse at apex, with numerous, unequal, short and very short, dark glandular hairs, few to numerous, short to medium, dark-based simple eglandular hairs, and few stellate hairs mainly towards the base. Ligules yellow, glabrous-tipped.

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Styles yellow to slightly discoloured. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, blackish. Flowers 5–6. Apomictic. Introduced. The inclusion of this species is based on material collected on grassy banks and cracks in concrete around Walthamstow Reservoirs in 1953, where it was common. Scandinavia and central Europe. It is difficult to find the correct name of this plant without seeing the types, but it fits very closely the original description of H. perscissum Jord. ex Boreau. It also fits the description of H. pinnatifidum L¨onnr. ex Dahlst. Dahlstedt cites Lindeberg’s material of H. vulgatum var. nemorosum, but Zahn states the material is mixed and all the sheets we have seen are referable to H. triviale Norrl. Pugsley’s H. neopinnatifidum was a new name for H. pinnatidum L¨onnr. ex Dahlst., non Willd., but the specimens he cites are not this plant.

283. H. subirriguum Dahlst. ex Porat Double-toothed Hawkweed H. vulgatum subsp. subirriguum (Dahlst. ex Porat) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale green, suffused brownish-purple, robust, flexuous, striate, with numerous, long, pale simple eglandular hairs below and often throughout, and numerous stellate hairs and an occasional, short, dark glandular hair above. Leaves dull medium green on upper surface, paler, slightly bluish and sometimes tinted purplish beneath; basal few, the lamina 5–9 × 3–5 cm, elliptical or ovate, mostly acute at apex, irregularly dentate, the teeth fairly large and mammiform or narrowly mammiform, interspersed with denticulations, abruptly contracted at base, the petioles up to 4 cm; cauline 4–5, gradually decreasing in size upwards, the lower like basal, similar in size and shortly petiolate, the upper with lamina lanceolate, long-acute at apex, dentate in the lower half and sessile; all with few, short to medium, pale simple eglandular hairs on both surfaces and more numerous ones on the margins, more or less numerous stellate hairs on the lower surface of the upper cauline and few to numerous, long simple eglandular hairs on the petioles. Inflorescence with 10–30 capitula, cymose-corymbose, with long lower branches from the leaf axils; peduncles short and suberect, with dense stellate hairs, numerous, short and medium, pale, dark-based simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–10 × 1.0–1.5 mm, blackish-green, inner with paler margins, all linearlanceolate, narrowed at apex, but obtuse to subacute, with numerous, unequal, very short to medium, dark glandular hairs, more or less numerous, short to medium, pale, darkbased simple eglandular hairs and some stellate hairs along the margins and at the base. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5– 7. Apomictic. Introduced. Known only from grassy banks at Walthamstow Reservoirs, Essex. Sweden, Germany, Austria, Czechoslovakia and Switzerland.

284. H. lepidulum (Stenstr.) Omang Irregular-toothed Hawkweed H. murorum subsp. lepidulum Stenstr.; H. vulgatum subsp. lepidulum (Stenstr.) Zahn; H. irriguum subsp. lepidulum (Stenstr.) Dahlst. Phyllopodous perennial herb with a branched stock. Stems 20–80 cm, pale yellowish-green, often suffused brownishpurple below, striate, slender to more or less robust, flexuous, with numerous, long, pale simple eglandular hairs below, which become shorter and sparser upwards, and numerous stellate hairs and an occasional, short, dark glandular hair in the upper part. Leaves dark green on upper surface, paler and rarely slightly suffused reddish-purple beneath; basal usually few, rarely more numerous, the lamina 2.5–12.0 × 1.5–4.0 cm, elliptical to lanceolate or oblong, narrowed to a rounded-mucronulate to acute apex, subentire, denticulate or irregularly dentate, the teeth mammiform and at variable intervals, cuneate to attenuate at base, the petioles up to 7 cm; cauline 2–5, gradually decreasing in size upwards, the lower sometimes larger than basal, elliptical, oblong or lanceolate, acute at apex, irregularly dentate, the teeth often sharply mammiform or aquilinemammiform, the lower petiolate, the upper sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins or glabrous or nearly so on upper surface, the uppermost cauline leaves sometimes with stellate hairs beneath. Inflorescence with 3–25 capitula, cymosecorymbose; peduncles slender and suberect, sometimes with a long lower branch, with dense stellate hairs, few to fairly numerous, short to medium, pale simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–9 × 0.8–1.2 mm, olivegreen, the inner with paler margins, all narrowly linearlanceolate, gradually narrowed to a narrow, more or less obtuse apex, with numerous, unequal, short, dark glandular hairs, few to fairly numerous, short to medium, dark-based simple eglandular hairs, and stellate hairs on the margin and near the base. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 2.5–3.0 mm, purplish-black. Flowers 5–7 and often again in autumn. Apomictic. Introduced. First recorded on a railway bank in Surrey in 1922 and may have been introduced in the First World War. It also occurs on grassy banks round Walthamstow Reservoirs in Essex, and on a disused railway line in that area; on a roadside bank near Royston, just in Cambridgeshire; and on a railway at Wymington in Bedfordshire. Scandinavia, Germany, Austria, Switzerland, Czechoslovakia and Hungary. 285. H. subviolascens P. D. Sell Violet-leaved Hawkweed H. irriguum subsp. violascens Almq. ex Dahlst.; H. vulgatum subsp. violascens (Almq. ex Dahlst.) Zahn; H. violascens (Almq. ex Dahlst.) Dahlst., non Borbas Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale green, suffused brownish-purple in lower half, flexuous but rigid, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter

39. Hieracium upwards, and numerous stellate and occasional, short, fine, dark glandular hairs above. Leaves bright green, often reddish round the margin on upper surface, paler and often suffused reddish-purple beneath; basal few to numerous, the lamina 3.5–6.5 × 1.5–3.5 cm, narrowly elliptical to elliptical or oblong-elliptical, rounded-mucronulate at apex, denticulate near the apex and incise-dentate in the lower two-thirds, the teeth mammiform or narrowly mammiform, interspersed with denticulations and often curved and redtipped, cuneate to shortly attenuate at base, the petioles up to 4 cm; cauline 3–4, gradually decreasing in size upwards, the lower like the basal and shortly petiolate, the upper with lamina lanceolate, linear-lanceolate or linear, sharply dentate to incise-dentate and sessile; all with numerous, short to medium, pale simple eglandular hairs on both surfaces and the margins, numerous stellate hairs on the undersurface of the upper cauline leaves and long, dense simple eglandular hairs on the petioles. Inflorescence with 5–16 capitula, cymose-corymbose; peduncles short, slender, with dense stellate, numerous, short to medium, pale simple eglandular hairs and numerous, very short, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–9 × 0.8–1.0 mm, olivegreen, the inner with paler margins, all narrowly linearlanceolate, gradually narrowed to a subacute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous short and very short, unequal, dark glandular hairs, and few stellate hairs along the margins. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Known only from grassy banks round Walthamstow Reservoirs in Essex, a disused railway track in the same area, and a railway embankment at Swanley Junction in Kent. Scandinavia and central Europe. 286. H. surrejanum F. Hanb. Surrey Hawkweed H. subramosum subsp. surrejanum (F. Hanb.) Zahn; H. surrejanum var. megalodon E. F. Linton ex F. Hanb.; H. subramosum subsp. surrejanum var. megalodon (E. F. Linton ex F. Hanb.) Zahn. Phyllopodous perennial herb with a branched stock. Stem 15–55 cm, pale green, usually brownish-purple in the lower half, slender, flexuous, striate, with numerous, long, pale simple eglandular hairs in the lower half, becoming shorter and less numerous upwards, and numerous stellate hairs and an occasional, short, fine, dark glandular hair above. Leaves rather pale green on upper surface and even paler beneath; basal usually few, rarely more numerous, the lamina 3–7 × 1.5–4.0 cm, narrowly to broadly elliptical or ovate, or subrotund, mostly rounded-mucronulate at apex, rarely subacute, subentire to shallowly dentate, the teeth patent and mammiform, those near the base sometimes narrowly mammiform, rounded or shortly cuneate at base, the petioles up to 7 cm; cauline 2–4, gradually decreasing in size upwards, the lower like basal and petiolate, the upper lanceolate or linear, more or less acute at apex and sessile; all with more or less numerous, short to medium, pale simple eglandular hairs on both surfaces and the margins and dense, long ones on the petioles. Inflorescence with 1–8 capitula, cymose-

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corymbose, rarely with a long lower branch; peduncles slender and suberect with dense stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and an occasional, short, dark glandular hair. Capitula 30– 35 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–10 × 1.5–1.7 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few, short, dark glandular hairs, and a few stellate hairs along the margin and at the base. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–6. Apomictic. Var. megalodon was distinguished from var. surrejanum by its more deeply dentate leaves and more numerous simple eglandular hairs on the involucral bracts and peduncles, but a range of plants occurs between the two varieties in both Sussex and Surrey. Native. Between Stedham and Woolbeding, Petworth and Bedham in Sussex and at Hindhead and Witley in Surrey. Endemic. 287. H. erubescens Jord. ex Boreau Red-flushed Hawkweed H. irriguum subsp. lepidulum var. haematophyllum Dahlst.; H. purpurascens Dahlst. nom. nud., non Scheele Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale green, although in most plants suffused brownish-purple in lower half and sometimes throughout, slender but rigid to more or less robust, flexuous, striate, with more or less numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and more or less numerous stellate hairs and occasional, short, fine glandular hairs in the upper part. Leaves dull, light to medium green, often with a reddish margin on the upper surface, paler and often suffused reddish or reddish-purple beneath; basal usually few, rarely more numerous, the lamina 2.5–9.0 × 1.5–4.0 cm, elliptical, obovate-elliptical or lanceolate, sometimes narrowed, but rounded mucronulate at apex, subentire to regularly and shallowly sinuatedentate through their whole length, the teeth more or less mammiform, sometimes flatly so, sometimes more pointed, cuneate to shortly attenuate at base, the petioles up to 6 cm; cauline 2–5, gradually decreasing in size upwards, the lower like basal and petiolate, with lamina lanceolate or linear, acute at apex, denticulate or sharply dentate and sessile; all with more or less numerous, short to long, pale simple eglandular hairs on both surfaces and the margins and long, dense ones on the petioles. Inflorescence with 3–15 capitula, cymose-corymbose, sometimes with a long lower branch; peduncles rather short, slender and suberect, with dense stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and few to numerous, short, dark glandular hairs. Involucral bracts incumbent in bud, 3–10 × 0.6–0.8 mm, dark to blackish-green, inner with paler margins, all narrow linear-lanceolate, gradually narrowed to a more or less acute apex, with numerous, very short to medium, dark glandular hairs, few to fairly numerous, short to medium, pale, dark-based simple eglandular hairs, and some stellate hairs mainly on the margin and at

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the base. Ligules yellow, glabrous-tipped. Styles yellowish to discoloured. Receptacle pits with margins incise-dentate. Achenes 2.5–3.0 mm, blackish-red. Flowers 5–7 and often again in the autumn. Apomictic. Introduced. Abundant on a disused railway station at Hurlingbarrows, St Agnes in Cornwall; railway banks between Clearbrook and Yelverton in Devonshire and grassy banks round Walthamstow Reservoirs and a nearby disused railway in Essex. There is a solitary specimen from a roadside at Settle in Yorkshire. France, Switzerland, Germany, Austria, Czechoslovakia, Hungary, Poland, Bulgaria, Denmark and Sweden. 288. H. commixtum Jord. Goring Gap Hawkweed H. maculatum subsp. commixtum (Jord.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–30 cm, pale green, flushed reddish-purple at least in the lower half, slender, striate, with numerous, long, pale simple eglandular hairs at least in the lower half and numerous stellate hairs with an occasional very short, dark glandular hair above. Leaves pale yellowish-green, faintly spotted and blotched reddish on upper surface, paler beneath; basal few, the lamina 4–5 × 1.0–1.5 cm, narrowly elliptical or lanceolate, acute at apex, more or less denticulate, cuneate to attenuate at base, the petioles long, slender and longhairy; cauline 2–3, lower similar to basal and petiolate, the upper linear or lanceolate and sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 1–9 capitula, furcate; peduncles with numerous to dense stellate hairs, few to numerous, very short, dark glandular hairs and an occasional simple eglandular hair. Capitula 15–20 mm in diameter, rounded below. Involucral bracts incumbent in bud, 5–10 × 0.8–1.0 mm, dark olive-green, the inner with pale green margins, all narrow linear-lanceolate, the inner very narrow, acute at apex, with numerous, unequal, short and very short, dark glandular hairs, few to fairly numerous simple eglandular hairs and few stellate hairs. Ligules yellow, glabroustipped. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 2.8–3.0 mm, reddish-black. Flowers 5–6. Apomictic. Possibly native. Known only from the Goring Gap in Oxfordshire. France, Switzerland, Germany, Austria, Hungary and Poland. 289. H. glanduliceps P. D. Sell & C. West Keld Head Hawkweed H. maculatum auct. Phyllopodous perennial herb with a branched stock. Stem 15–40 cm, pale yellowish-green, often reddish towards the base, slender, strongly striate, flexuous, with numerous, medium to long, pale simple eglandular hairs, scattered stellate hairs and without glandular hairs. Leaves dark green with brownish-purple spots and blotches on upper surface, paler and often flushed purplish beneath; basal few to numerous, the lamina 3–9 × 1.5–4.0 cm, subrotund, ovate, ovate-lanceolate, lanceolate or narrowly elliptical, rarely oblong, obtuse-mucronulate to acute, denticulate to dentate, the teeth small, mammiform or unguiculate-mammiform

and abruptly contracted or subtruncate at base, the petioles short and with numerous, medium to long, pale simple eglandular hairs; cauline 1–3, the lower similar to basal and petiolate, the upper with lamina linear or lanceolate, acute at apex, dentate at the base and sessile; all glabrous or with pale simple eglandular hairs near the margin on the upper surface, beneath and on the margin with few to numerous medium, pale simple eglandular hairs. Inflorescence with 1–8(−10) capitula, narrowly cymose, sometimes with a long lower branch from the axil of the upper cauline leaf; peduncles medium to long, suberect, with numerous stellate hairs, numerous, very short, dark glandular hairs and few, medium, dark simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all narrow linear-lanceolate, more or less acute at apex, with scattered stellate hairs, numerous very short and short, dark glandular hairs and few, medium, dark simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Ledges on limestone cliffs at Keld Head Scar, Twistleton Scars and north of Ingleton in Yorkshire and several sites in Westmorland. Endemic. 290. H. inquinatum Jord. ex Boreau Stained-leaf Hawkweed H. maculatum subsp. maculatum var. inquinatum (Jord. ex Boreau) Zahn Phyllopodous perennial herb with a branched stock. Stem 40–60 cm, pale yellowish-green, often tinted brownish-red towards the base, slender to slightly robust, striate, with numerous, short to long, pale simple eglandular hairs especially in the lower half, with fairly numerous stellate hairs above. Leaves medium yellowish-green with brownishpurple to almost black spots and blotches on the upper surface, paler often tinted reddish-purple beneath; basal rather few, the lamina 4–9 × 2–3 cm, elliptical, oblong-elliptical or lanceolate, acute at apex, subentire to remotely denticulate, cuneate at base, with medium, long-hairy petioles; cauline 3–5, lower like basal and more or less petiolate, the upper smaller, narrower and sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 2–15 capitula, cymose-corymbose; peduncles short, with dense stellate hairs, numerous, very short and short glandular hairs and occasional, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0– 1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less obtuse at apex, with numerous, very short, short and medium, unequal, dark glandular hairs, rarely a simple eglandular hair, and a few stellate hairs near the base and on the margin. Ligules yellow, glabroustipped. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 5–7. Apomictic. Possibly introduced. Known only from Horsea Island, Portsmouth, Hampshire. France, Switzerland, Germany and Austria.

39. Hieracium 291. H. naevuliferum Jord. ex Boreau Baldock Hawkweed H. spurcatum Jord. ex Boreau; H. thuringorum Sag.; H. maculatum subsp. naevuliferum (Jord. ex Boreau) Zahn Phyllopodous perennial herb with a branched stock. Stem 15–50 cm, pale yellowish-green, often with purple towards the base, rather slender, striate, with few to numerous, medium to long, pale simple eglandular hairs, and numerous stellate hairs and occasional, short, dark glandular hairs in the upper part. Leaves yellowish-green and heavily spotted and blotched brownish-purple on the upper surface, paler and often suffused purplish beneath; basal few to fairly numerous, the lamina 2–9 × 2–4 cm, ovate to broadly ovate or elliptical, more or less acute or obtuse at apex, subentire to denticulate or with a few teeth, abruptly contracted or rounded at base, with short, long-hairy petioles; cauline 2–4, like basal and petiolate or lanceolate and linear and sessile; all with short to long hairs on both surfaces and the margins. Inflorescence with 2–8 capitula; peduncles with dense stellate hairs, numerous short, dark glandular hairs and an occasional, medium, pale simple eglandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, greyishgreen, the inner with paler margins, all linear-lanceolate, acute at apex, with fairly numerous, very short, dark glandular hairs, and numerous stellate hairs especially on the margin. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 2.5– 3.0 mm, reddish-black. Flowers 7–9. Apomictic. Introduced. Known only from Weston Hills, near Baldock, Hertfordshire where it was collected in 1925 and an old railway near Kielder, Northumberland in 1963. Germany, Switzerland, France, Holland, Austria and Hungary. 292. H. asperatum Jord. ex Boreau Rough Hawkweed H. maculatum subsp. asperatum (Jord. ex Boreau) Zahn Phyllopodous perennial herb with a branched stock. Stem 50–70 cm, pale yellowish-green, often flushed reddishpurple towards the base, more or less robust, with few to numerous, medium to long, pale simple eglandular hairs especially towards the base, and numerous stellate hairs above. Leaves yellowish-green, spotted and blotched with brownish-purple on upper surface, paler beneath; basal few to fairly numerous, the lamina 6–10 × 3–5 cm, elliptical or ovate-lanceolate, acute at apex, dentate, the teeth in the lower half often narrowly aquiline-mammiform, cuneate at base, with medium, long-hairy petioles; cauline 2–5, the lower as large as and similar to the basal and petiolate, the upper with lamina lanceolate or ovate, acute at apex, dentate, petiolate or sessile; all with short to medium, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with numerous capitula, paniculate-corymbose, often with a long lower branch; peduncles suberect, with dense stellate hairs and sometimes an occasional, short, dark glandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.8– 1.0 mm, dark olive-green, the inner with paler margins, all narrow linear-lanceolate, more or less obtuse at apex, with numerous, very short glandular hairs, an occasional

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short to medium, pale simple eglandular hair and numerous stellate hairs particularly on the margins. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits sharply dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 5–7. Apomictic. Possibly introduced. Recorded from Devonshire, Surrey and Merionethshire. France, Switzerland, Germany and Austria. 293. H. fictum Jord. ex Boreau Lacerate-leaved Hawkweed H. incisifolium Jord. ex Boreau; H. maculatum subsp. fictum (Jord. ex Boreau) Zahn; H. maculatum subsp. fictum forma incisifolium (Jord. ex Boreau) Zahn Phyllopodous perennial herb with a branched stock. Stem 40–90 cm, pale yellowish-green, often suffused with brownish-purple in the lower half, robust, striate, with few, medium or long, pale simple eglandular hairs, sometimes fairly numerous below, usually very sparse above, with numerous to dense stellate hairs above, rarely with a very short glandular hair above. Leaves medium yellowish-green and heavily spotted and blotched with brownish-purple on the upper surface, paler beneath and sometimes tinted purple; basal usually few, the lamina 5–8 × 2.5–3.5 cm, elliptical, lanceolate or elliptic-lanceolate, obtuse-mucronulate to acute at apex, dentate or incise-dentate, the teeth long, narrow and acute, cuneate to attenuate at base with long petioles clothed with dense, long, curly simple eglandular hairs; cauline 3–5, similar to basal, usually acute at apex, usually incise-dentate with long, narrow, acute teeth, the lower petiolate, the upper sessile; all with numerous, short to long, pale simple eglandular hairs on both surfaces and the margins or glabrous or nearly so on the upper surface. Inflorescence with 6–17 capitula, cymose-corymbose; peduncles rather short, with dense stellate hairs, few to numerous, short, dark glandular hairs and occasional simple eglandular hairs. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 2–8 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all linearlanceolate, obtuse at apex, with numerous, short, dark glandular hairs, an occasional, dark-based simple eglandular hair and numerous stellate hairs especially on the margins. Ligules yellow, sometimes with an occasional, simple eglandular hair at apex. Styles yellow or slightly discoloured. Receptacle pits with the margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Probably introduced. Railway banks and roadsides. A few scattered localities in southern England, Derbyshire and Argyllshire. South and central France. 294. H. spilophaeum Jord. ex Boreau Spotted Hawkweed H. maculatum subsp. maculatum var. spilophaeum (Jord. ex Boreau) Zahn; H. maculatum Sm., non Schrank; H. maculatum Teesdale ex Turner & Dillwyn nom. nud.; H. vulgatum forma maculatum Backh.; H. vulgatum var. maculatum F. Hanb. Phyllopodous perennial herb with a branched stock. Stem 30–80 cm, pale yellowish-green often suffused brownishpurple especially in the lower half, slender to robust, striate,

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with numerous, medium to long, pale simple eglandular hairs especially in the lower half and numerous stellate hairs and an occasional, short, dark glandular hair in the upper part. Leaves pale yellowish-green, spotted and blotched with brownish-purple on the upper surface but sometimes clear green in shade, paler and sometimes flushed purple beneath; basal few, the lamina 4–9 × 2–4 cm, elliptical, ovate-lanceolate or lanceolate-oblong, obtuse to acute at apex, entire to shallowly dentate, abruptly contracted to cuneate at base, the long-hairy petioles usually long; cauline 2–5, the lower with lamina lanceolate or ovate, acute at apex, shallowly dentate, cuneate and petiolate, the upper similar, but smaller and the uppermost linear; all with numerous, medium to long, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 6– 20 capitula, cymose-corymbose; peduncles rather short and suberect, with numerous to dense stellate hairs, few to numerous, short, dark glandular hairs and sometimes an occasional, medium, pale simple eglandular hair. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all linear-lanceolate, much narrowed at apex to a subacute tip, with numerous, short, dark glandular hairs, an occasional, short, pale simple eglandular hair and numerous stellate hairs especially on the margins. Ligules yellow, glabrous-tipped. Styles discoloured or yellowish. Receptacle pits with margins incise-dentate. Achenes 2.5–3.0 mm, purplish-black. Flowers 5–7. Apomictic. Probably introduced. Grassy and waste places, waysides, railway banks, open woodland and walls. Widespread in southern England, rarer further north, and a few records for Ireland. Probably widespread in Continental Europe, but its native distribution is not known. 295. H. subamplifolium (Zahn) Roffey Balloon-leaved Hawkweed H. vulgatum var. amplifolium Ley; H. sciaphilum var. amplifolium (Ley) W. R. Linton; H. vulgatum subsp. subamplifolium Zahn; H. maculatum subsp. asperatum forma immaculatum Zahn Phyllopodous perennial herb with a branched stock. Stem 50–100 cm, pale yellowish-green, sometimes pinkish or purple below, slender to robust, striate, with long, wavy, pale simple eglandular hairs, numerous below becoming fewer upwards, numerous to dense stellate hairs in the upper half and sometimes an occasional, short glandular hair above. Leaves yellowish-green on upper surface, paler beneath and often suffused with purple; basal numerous, the outer with lamina 5–11 × 3–6 cm, broadly elliptical, subrotund, obovate, rounded-mucronulate at apex, entire to shallowly sinuate-dentate with shallow mammiform teeth or rarely with larger mammiform teeth, and rounded, subtruncate or abruptly contracted at base, the inner with lamina 7–12 × 3– 5 cm, elliptical, lanceolate-elliptical or ovate, narrowed to an obtuse or subacute apex, serrate-dentate, with curved mammiform teeth and cuneate or abruptly contracted at base, the petioles short to medium and with dense, long, pale simple eglandular hairs; cauline 2–4, similar to inner basal and becoming gradually smaller upwards, all except the uppermost petiolate; all with numerous, short to medium,

pale, subrigid hairs on both surfaces and the margins. Inflorescence with 3–45 capitula, paniculate-corymbose, often with a long branch from the upper leaf axils; peduncles suberect or erecto-patent, with dense stellate hairs, few to numerous, short, dark glandular hairs and few to numerous, pale, short to medium simple eglandular hairs. Capitula 30– 35 mm in diameter. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, more or less acute at apex, with numerous, very short and short, dark glandular hairs, numerous short and medium, dark-based simple eglandular hairs, and a few stellate hairs mostly at the base and on the margins. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–8. Apomictic. Native. Rocky banks, low cliffs, quarries, roadside and railway banks, often on limestone. South Wales and the west of England border country. Endemic. 296. H. mammidens P. D. Sell Breast-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–85 cm, pale yellowish-green, often flushed brownishpurple at base, slender to robust, striate, with numerous to dense, long, pale, wavy simple eglandular hairs below, becoming gradually fewer and shorter upwards, and numerous to dense stellate and occasional, very short, dark glandular hairs in the upper part. Leaves yellowish-green on upper surface, paler beneath and often suffused with purple; basal few to numerous, the lamina 4–10 × 2–4 cm, elliptical to broadly elliptical, obovate or nearly subrotund, rounded or obtuse at apex, usually dentate, rarely subentire or denticulate, the teeth mammiform, sometimes narrowly so, and cuneate or subtruncate at base, the petioles up to 9 cm, with dense, long, pale simple eglandular hairs; cauline 3–5, gradually decreasing in size upwards, the lower like basal and petiolate, the upper sometimes ovate or lanceolate and often more toothed; all with numerous, short, pale simple eglandular hairs on the upper surface and longer ones beneath especially on the midrib, the cauline with varying amounts of stellate hairs beneath. Inflorescence with 3–20 capitula, paniculate-corymbose, with long branches; peduncles rather short and slender, erect, with dense stellate hairs, numerous short, dark glandular hairs and numerous medium, pale simple eglandular hairs. Capitula 30– 40 mm in diameter. Involucral bracts incumbent in bud, 5.0–10.5 × 0.8–1.0 mm, olive-green, the inner with a paler margin, all narrow linear-lanceolate, obtuse at apex, with numerous very short and short, fine, dark glandular hairs, an occasional medium, pale, dark-based simple eglandular hair, and a few stellate hairs on the margin. Ligules yellow, with very short simple eglandular hairs at the tips. Styles yellow. Receptacle pits shortly dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 5–6. Apomictic. ?Native. Sandy banks near Fleet in Hampshire, clay banks by Fittleworth Station in Sussex and shady hedgebanks around Pembury and between Capel and Matfield in Kent. ?Endemic.

39. Hieracium 297. H. submutabile (Zahn) Pugsley Variable Hawkweed H. vulgatum subsp. argillaceum var. submutabile Zahn; H. vulgatum var. mutabile Ley; H. acroleucum var. mutabile (Ley) W. R. Linton; H. mutabile (Ley) Ley, non F. W. Schultz; H. reclinatum auct. Phyllopodous perennial herb with a branched stock. Stems 20–70 cm, pale yellowish-green, often brownish-purple towards the base, slender to robust, striate, with numerous, long, pale, wavy simple eglandular hairs below becoming fewer and shorter upwards and with numerous stellate hairs and a few, short, dark glandular hairs in the upper part. Leaves pale, rather greyish-green on upper surface, paler beneath; basal few to numerous, the lamina 2–9 × 1.5–4.0 cm, elliptical, oblong-elliptical or ovate, obtuse-mucronulate to acute at apex, subentire, denticulate or shallowly dentate, rarely with a few larger teeth, rounded or cuneate at base, the petioles usually short, but up to 4 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 1–4, often 2, with a large one very near the base, petiolate and similar to basal, the rest when present very small and sessile, so that there usually appears to be a tall stem almost without leaves; all with short to medium, rather stiff, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 1–16 capitula, paniculatecorymbose, sometimes with a long lower branch; peduncles slender and erect, with dense stellate hairs, numerous, short, fine dark glandular hairs and sometimes an occasional, pale simple eglandular hair. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, dark green, the inner with paler margins, all linear-subulate much narrowed at apex, obtuse except the acute innermost, with short (0.4–0.7 mm), fine, dark glandular hairs and a few stellate hairs, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 2.5–3.0 mm, purplish-black. Flowers 6–7. Apomictic. Native. Rocky banks and streamsides and grassy and waste places. Fairly common in south and central Wales extending into Herefordshire and Gloucestershire. Endemic. 298. H. diaphanum Fr. Dark-leaved Hawkweed H. vulgatum subsp. diaphanum (Fr.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, sometimes purplish towards the base, usually slender, with numerous, medium to long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards, with fairly numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves dark, bluish-green on upper surface, paler beneath; basal few, the lamina 4–6 × 1.5–2.5 cm, narrowly elliptical, lanceolate or ovate, mostly more or less acute at apex, sometimes obtuse, denticulate or shallowly dentate, with small, sharp teeth, cuneate at base, the petioles up to 4 cm, with dense, long, wavy simple eglandular hairs; cauline 2–5, lower like basal and shortly petiolate, the upper lanceolate, acute at apex, entire to dentate and sessile or bract-like;

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all with short, stiff, pale simple eglandular hairs on upper surface and long, softer ones beneath and on the margin. Inflorescence with 2–16 capitula, paniculate-corymbose; peduncles straight, slender and more or less erect, with numerous stellate hairs, numerous, short or very short, dark glandular hairs and sometimes an occasional, medium simple eglandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–9 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, short, black glandular hairs and occasional slightly longer, simple eglandular hairs, stellate hairs absent. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. Probably introduced. Roadsides, railway banks and a chalkpit. A few scattered records in England, Wales and southern Scotland. Sweden and central Europe. 299. H. daedalolepioides (Zahn) Roffey Petite-leaved Hawkweed H. vulgatum subsp. daedalolepioides Zahn; H. acroleucum var. daedalolepium auct.; H. vulgatum var. daedalolepium auct.; H. adlerzii auct.; H. diaphanoides var. ornatum auct. Phyllopodous perennial herb with a branched stock. Stem 10–40 cm, pale yellowish-green, often purplish in the lower half, slender, flexuous, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards and numerous stellate hairs and few, short, dark glandular hairs in the upper part. Leaves bright or bluish-green, sometimes reddish round the margin on upper surface, paler and often suffused purple beneath, rather thick and rigid; basal few to fairly numerous, the lamina 2–7 × 1–3 cm, elliptical or lanceolate, obtuse to acute at apex, subentire or with a few, sharp teeth, abruptly contracted or cuneate at base, the petioles up to 5 cm, but usually much shorter and with numerous, long, pale, wavy simple eglandular hairs; cauline 2–5, lower like basal or sometimes ovate and petiolate, the upper lanceolate or linearlanceolate, acute at apex, sharply toothed and sessile; all with short, rather stiff simple eglandular hairs on upper surface and numerous, softer, medium to long, pale simple eglandular hairs beneath. Inflorescence with 3–10 capitula, paniculate-corymbose, rarely with a long lower branch; peduncles long, slender, straight and erect, with dense stellate hairs, numerous, short, dark glandular hairs and few, short to medium, dark-based simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.4 mm, blackishgreen, the inner with paler margins, all linear-lanceolate, gradually narrowed to an acute apex, with numerous, very unequal, short to medium (0.4–1.0 mm), slender, black glandular hairs and occasional simple eglandular hairs, without stellate hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 2.0–2.5 mm, reddish-black. Flowers 6–7. Apomictic. This species can be difficult to distinguish from H. anglorum, with which it often grows, especially when pressed on herbarium sheets. In the field its bright green, often suffused

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reddish, thickish, small, little-toothed leaves are diagnostic, but in the herbarium small, little-toothed specimens of H. anglorum are probably mistaken for it. Native. Cliff ledges, scree and riverside rocks. Fairly common in the Welsh mountains; typical specimens have been seen from the Staffordshire and Yorkshire limestone and from Little Langdale in Westmorland and good material from Kirkcudbrightshire. A few scattered records from roadsides and railway banks may be small forms of H. anglorum. 300. H. lepidiceps (Dahlst.) Prain Skipton Hawkweed H. irriguum subsp. lepidiceps Dahlst.; H. vulgatum subsp. lepidiceps (Dahlst.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often suffused purplish in the lower half, slender, flexuous, striate, with dense, long, wavy, pale simple eglandular hairs below, becoming fewer and shorter upwards and dense stellate and numerous, short, dark glandular hairs in the upper part. Leaves medium to dark green on upper surface, paler and often suffused purplish beneath, sometimes suffused purplish or reddish-purple throughout; basal few, the lamina 3–9 × 1– 4 cm, lanceolate or ovate, sometimes narrowly elliptical or oblong-elliptical, rounded-obtuse to acute at apex, denticulate to dentate, the teeth narrow and sharp to rather sharply mammiform, cuneate to slightly attenuate at base, the petioles up to 9 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 2–4, lower like the basal, with a petiole which is sometimes very long, the upper lanceolate or ovate, acute at apex, shortly dentate, cuneate at base, often sessile, or bract-like; all with short to medium, stiff, pale simple eglandular hairs on the upper surface and longer ones beneath and on the margin. Inflorescence with 2–10 capitula, corymbose or with a narrow panicle; peduncles short and straight, with dense stellate hairs and numerous, very short and short, dark glandular hairs. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.8–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an obtuse tip, with dense, very unequal, very short to medium (0.2–1.5 mm), black glandular hairs, some of them very slender, without stellate or simple eglandular hairs. Ligules deep yellow, minutely hairy at apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Probably introduced. Abundant on a railway embankment about 2 km from Skipton in Yorkshire, near Chatsworth in Derbyshire and on walls north of Rhu in Dunbartonshire. Sweden and central Europe. 301. H. scanicum (Dahlst.) Ley Fang-toothed Hawkweed H. pinnatifidum subsp. scanicum Dahlst.; H. vulgatum subsp. scanicum (Dahlst.) Zahn Phyllopodous perennial herb with a branched stock. Stem 25–60 cm, pale yellowish-green, sometimes flushed brownish-purple, slender to robust, striate, with dense, long,

pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards and numerous stellate hairs and fairly numerous, short, dark glandular hairs in the upper part. Leaves rather pale to medium green on upper surface, paler beneath; basal few, the lamina 4–10 × 1.5–3.0 cm, narrowly elliptical, oblong-elliptical or ovate-oblong, more or less obtuse to acute at apex, incise-dentate with longpointed teeth, attenuate at base, the petioles up to 4 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 3–5, lower like the basal and petiolate, the upper oblong or oblong-lanceolate, acute at apex, deeply incise-dentate in the lower half and sessile; all glabrous or with short, pale simple eglandular hairs on the upper surface particularly near the margin and numerous, medium to long ones beneath and on the margin. Inflorescence with 10–25 capitula, paniculate-corymbose, sometimes forming an umbel near the top, often with a long lower branch; peduncles long, slender and suberect, with dense stellate hairs, numerous, short, black glandular hairs and a few, short to medium simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4– 11 × 0.8–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, narrowed to an acute apex, with numerous, unequal, very short to medium (0.2–1.2 mm), dark, slender glandular hairs and an occasional, medium, dark-based simple eglandular hair, without stellate hairs. Ligules yellow, with few, very short hairs at apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Native or possibly introduced. Cliff ledges, roadsides, walls, sand-dunes and quarry. Scattered localities in Wales and Gloucestershire and at Little Langdale in Westmorland. Scandinavia and central Europe. 302. H. subminutidens (Zahn) Pugsley Llanwrtyd Hawkweed H. caesium subsp. adlerzii var. subminutidens Zahn; H. adlerzii auct. Phyllopodous or hypophyllopodous perennial herb with a branched stock. Stem 30–80 cm, pale yellowish-green, often reddish-purple in the lower part, more or less robust, striate, with numerous, long, pale simple eglandular hairs towards the base and fewer upwards, and numerous stellate hairs in the upper part. Leaves deep green, sometimes tinted reddishpurple beneath; basal few, the lamina 5–8 × 2–3 cm, elliptical or elliptic-lanceolate, obtuse-mucronate to acute at apex, denticulate or remotely dentate, with short, long-hairy petioles; cauline 5–9, decreasing in size upwards, the lower elliptical, elliptic-lanceolate or lanceolate, acute at apex, denticulate to dentate and shortly petiolate or sessile, the upper lanceolate, acute at apex, dentate or incise-dentate, the teeth narrow and sessile; all with scattered, short or medium, pale simple eglandular hairs on both surfaces and more numerous on the margins, sometimes the upper with stellate hairs beneath. Inflorescence with 3–20 capitula, paniculatecorymbose, sometimes with a lower branch; peduncles long and suberect, with dense stellate hairs, more or less numerous, short to medium, dark-based simple eglandular hairs

39. Hieracium and more or less numerous, short, dark glandular hairs. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts more or less porrect in bud, 4–11 × 1.0–1.2 mm, dark green, the inner with paler margins, all narrowly linearoblong, obtuse at apex, with numerous, very unequal, very short, short and medium, fine, dark glandular hairs, an occasional, medium, dark-based simple eglandular hair, and a few stellate hairs towards the base. Ligules yellow, glabroustipped. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, blackish. Flowers 6–7. Apomictic. Native. Riverside rocks at Llanwrtyd and Abergwesyn in Breconshire. Endemic. 303. H. anglorum (Ley) Pugsley Anglian Hawkweed H. maculatum subsp. divisum var. genuinum auct.; H. scytophyllum auct.; H. scanicum var. anglorum Ley; H. adlerzii auct.; H. diaphanoides auct.; H. irriguum auct. Phyllopodous perennial herb with a branched stock. Stem 25–80 cm, pale yellowish-green, occasionally purplish towards the base, more or less robust, striate, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer upwards and with dense stellate hairs and a few, short, dark glandular hairs above. Leaves rather pale green on upper surface and paler beneath, rarely tinted reddish, usually rather flaccid; basal usually few, the lamina 3–10 × 1.5–4.5 cm, elliptical, ovate, oblong-elliptical or rarely subrotund, rounded-obtuse to acute at apex, dentate to deeply incise-dentate, the teeth often mammiform, sometimes much narrower and sharper, abruptly contracted or cuneate at base, the petioles up to 5 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 3– 5(−9), the lower elliptic-oblong, elliptic or ovate, acute at apex, dentate, often deeply so, the teeth long, narrow and unequal, abruptly contracted at base, and usually petiolate, the upper ovate or lanceolate, acute at apex, usually deeply laciniate-dentate, the teeth long and sharp, sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 10– 20(−40) capitula, paniculate-corymbose; peduncles slender and erect-spreading, with dense stellate hairs, numerous, short, dark glandular hairs and numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an acute apex, with numerous, short to medium (0.4–1.0 mm), slender, black glandular hairs and sometimes with an occasional solitary simple eglandular hair, without stellate hairs. Ligules yellow, glabrous-tipped. Styles slightly discoloured. Receptacle pits with margins shortly subulate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. In Wales and northern England where it occurs in natural habitats on cliffs and scree and by streams it is probably native. Elsewhere where it occurs on roadsides, open woodland and railway banks it is probably introduced. It is common in Wales and adjacent England and in northern England. In Scotland it has only been seen in Kirkcudbright-

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shire and there are records for Co. Leix, Co. Kildare and Co. Down in Ireland. Endemic. 304. H. diaphanoides Lindeb. Diaphanous Hawkweed H. medium Lindeb., non Jord.; H. diaphanoides var. apiculatum E. F. Linton; H. apiculatum (E. F. Linton) Druce Phyllopodous perennial herb with a branched stock. Stem 35–60 cm, pale yellowish-green, occasionally purplish towards the base, slender to robust, striate, with numerous, long, pale, wavy simple eglandular hairs towards the base becoming fewer and shorter upwards, and with numerous stellate hairs and a few, fine, dark glandular hairs in the upper part. Leaves medium green on upper surface, paler and often bluish-green beneath; basal few, the lamina 3–13 × 1.5–5.0 cm, lanceolate, elliptical, oblong-elliptical, oblong-lanceolate, oblong or ovate, obtuse-mucronulate to acute at apex, denticulate to dentate, the teeth small and sharp to large and mammiform, cuneate to attenuate at base, the petioles up to 6 cm and with dense, long, pale simple eglandular hairs; cauline 1–3, the lower often like basal but more or less sessile, most frequently linear or linearlanceolate, acute at apex, dentate and sessile; all usually with numerous, short to long, pale simple eglandular hairs on both surfaces and the margins, but sometimes glabrous or nearly so on the upper surface. Inflorescence with 3– 15 capitula, paniculate-corymbose, often with a long lower branch; peduncles long and slender, straight or curved, with dense stellate hairs, numerous, short, dark glandular hairs and an occasional, short or medium, dark-based simple eglandular hair. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0– 1.4 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an obtuse apex, with dense, unequal, short to medium (0.4–1.0 mm), slender, black glandular hairs and sometimes an occasional medium simple eglandular hair, without stellate hairs or with a few near the base. Ligules yellow, glabrous-tipped. Styles more or less discoloured. Receptacle pits with margins shortly dentate. Achenes 2.5–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and rocky streamsides. Herefordshire and scattered localities in Wales, northern England and Scotland and on stabilised dunes and walls in and near Dublin in Ireland. Scandinavia and central Europe. 305. H. megapodium Dahlst. Fine-bracted Hawkweed Phyllopodous perennial herb with a branched stock. Stem 45–80 cm, pale yellowish-green, sometimes flushed purplish towards the base, slender to robust, striate, with rather few, long, pale simple eglandular hairs and numerous stellate hairs in the upper part. Leaves dull, rather pale green on upper surface, paler beneath; basal usually rather few, the lamina 6–11 × 2–4 cm, oblong, elliptic-oblong, oblanceolate-oblong or elliptical, rounded-mucronulate to subacute at apex, denticulate to dentate, the teeth shallowly mammiform, rarely more deeply dentate with narrower, more sharp teeth, cuneate to attenuate at base, the petioles up to 10 cm, slender, with few to fairly numerous, long,

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pale simple eglandular hairs; cauline (2–)3–4, lower sometimes similar to basal but often lanceolate, ovate or elliptical, acute at apex, dentate, the teeth fairly sharp, cuneate at base and petiolate, the upper lanceolate, acute at apex, dentate or entire, cuneate and sessile; all with short, stiff simple eglandular hairs on upper surface to nearly glabrous and rather few, similar but sometimes longer hairs beneath and on the margin. Inflorescence with 10–30 capitula, paniculatecorymbose, often with a long lower branch exceeding the acladium; peduncles often long, slender, erect, with dense stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and sometimes a few, dark glandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, olive-green, the inner with broad, pale margins, all oblonglanceolate, narrowed to an obtuse apex, with numerous short (0.4–0.7 mm), dark glandular, and few to numerous, short to medium, dark-based simple eglandular hairs, without or with very occasional stellate hairs. Ligules yellow, glabroustipped. Styles yellow to slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. Native or possibly introduced, but not known elsewhere. Grassy banks. Ashdown Forest in Sussex, Langton, Ashurst Park and Sundridge Park in Kent and Walthamstow Reservoirs in Essex. Endemic. 306. H. lagganense P. D. Sell Laggan Hawkweed H. diaphanum var. glaucovirens auct. Phyllopodous perennial herb with a branched stock. Stem 40–60 cm, pale yellowish-green, often purplish towards the base, slender but rigid, striate, with numerous, long, pale, wavy simple eglandular hairs towards the base, fewer or absent above, and numerous stellate and short, dark glandular hairs in the upper part. Leaves medium yellowish-green, often with a purple mark round the rim on the upper surface, paler and often suffused purple beneath; basal few to fairly numerous, with the lamina 5.0–7.5 × 2–3 cm, narrowly elliptical or ovate, subacute at apex, subentire or remotely denticulate, cuneate to attenuate at base, the petioles up to 5 cm, often purple, with numerous, long, pale, wavy simple eglandular hairs; cauline (1–)2(−3), lowermost usually very low on stem, the lamina ovate, more or less acute, denticulate and shortly petiolate, the upper with lamina lanceolate, acute at apex, entire and sessile, or bract-like; all with short, stiff simple eglandular hairs on both surfaces and the margins, the cauline with numerous stellate hairs beneath. Inflorescence with 3–12 capitula, paniculate-corymbose, often with a long lower branch; peduncles long and slender, with dense stellate hairs and numerous, short, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.8–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, very narrow at apex and acute, with numerous, short and very short, dark, slender glandular hairs and a few stellate hairs near the base, without simple eglandular hairs. Ligules yellow, glabroustipped. Styles yellow. Receptacle pits with margins dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 7–8. Apomictic.

Native. River-banks at Crathie and Laggan Bridge in Inverness-shire. A plant from a wood at Balnakeilly, Pitlochry, in Perthshire is similar. Endemic. 307. H. acuminatum Jord. Tall Hawkweed H. vulgatum var. acuminatum (Jord.) Arv.-Touv.; H. vulgatum subsp. acuminatum (Jord.) Sudre; H. barbareifolium auct.; H. cacuminatum var. barbareifolium auct.; H. diaphanum var. glaucovirens auct. Phyllopodous perennial herb with a branched stock. Stems 50–110 cm, pale green, often suffused brownish-purple, robust, markedly striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and few to numerous, short, dark glandular hairs above. Leaves medium yellowish-green, sometimes reddish round the margins on the upper surface, paler beneath and sometimes tinted purplish; basal usually few, sometimes numerous, the lamina 6–15 × 2–6 cm, mostly oblong-elliptical, sometimes lanceolate to ovate or oblong-lanceolate, obtuse-mucronate to acute or acuminate at apex, dentate to incise-dentate, the teeth narrow and apiculate, cusped and up to 15 mm, sometimes free on the petiole, cuneate to attenuate at base, the petioles up to 11 cm; cauline 6–14, gradually decreasing in size upwards, the lower like basal with petioles up to 10 cm, the upper ovate to lanceolate, long-acute to acuminate at apex, deeply incise-dentate with narrow, curved teeth up to 20 mm and sessile or shortly petiolate; all with short to long, pale simple eglandular hairs on both surfaces and the margins, or nearly glabrous on the upper surface, the petioles with dense, long simple eglandular hairs. Inflorescence with 5–50 capitula, paniculate-corymbose, often with several long branches from the leaf axils, each branch bearing a corymb at its apex; peduncles short to long, erectspreading, with dense stellate hairs, numerous, short, dark glandular hairs and rarely an occasional, medium, pale simple eglandular hair. Capitula 25–30 in diameter, rounded at base. Involucral bracts incumbent in bud, 3–11 × 1.0–1.2 mm, dull greyish-green the inner with paler margins, all linear-lanceolate, acute at apex, with numerous very short to medium, dark glandular hairs, an occasional, medium, dark-based simple eglandular hair and numerous stellate hairs particularly on the margins and towards the base. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–8. Apomictic. Easily recognised by its numerous, deeply cut cauline leaves with teeth up to 20 mm and a large panicle of glandular-hairy capitula with numerous stellate hairs along the margin of the involucral bracts. Native. Rocky limestone woods and streamsides in the England/Wales border country where it is presumably indigenous, although it also occurs on roadsides and railway banks in that area. It is the plant which A. Ley called H. barbareifolium. A modern assessment of its distribution there is badly needed. It is also recorded for Baildon in Yorkshire and walls of St John’s College at Oxford where it is presumably introduced. A specimen from rocks at Rowallan, near Saintfield, Co. Down seems also to be it. Central, southwest, south and south-east Europe.

39. Hieracium 308. H. consociatum Jord. ex Boreau Sociable Hawkweed H. vulgatum subsp. deductum microgen. aurulentum var. consociatum (Jord. ex Boreau) Sudre; H. vulgatum subsp. consociatum (Jord. ex Boreau) Zahn; H. fastigiatum Fr., non Tausch; H. vulgatum subsp. lachenalii var. fastigiatum Zahn; H. cacuminatum auct.; H. scanicum auct. Phyllopodous perennial herb with a branched stock. Stems 30–90 cm, pale green, usually suffused brownish-purple at least in the lower half, slender to robust, often flexuous, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and few to numerous, short glandular hairs above. Leaves medium yellowish-green, often reddish round the margin, paler and sometimes suffused purplish beneath; basal few to numerous, the lamina 4–12 × 2–5 cm, lanceolate to ovate or elliptical-ovate, rounded-mucronulate to more or less acute or acuminate at apex, more or less dentate, the teeth mammiform or narrowly mammiform, often cusped and some more than 6 mm, cuneate to attenuate at base, the petioles up to 8 cm; cauline 3–7, gradually decreasing in size upwards, the lower similar to basal, often with some teeth up to 10 mm and petiolate, the upper lanceolate to linear, long-acute or acuminate at apex, dentate to incise-dentate with teeth narrowly mammiform, curved and up to 10 mm, sessile or shortly petiolate; all with numerous, short to long simple eglandular hairs on both surfaces and the margins, or sparse on the upper surface, the petioles with long simple eglandular hairs. Inflorescence with 5–30 capitula, paniculate-corymbose, often with long lower branches; peduncles erect-spreading to suberect, with dense stellate hairs, numerous, short, dark glandular and an occasional pale simple eglandular hair. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.2–1.5 mm, olive green to blackish-green, inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous to dense, unequal, very short to medium, dark glandular hairs, an occasional, pale, darkbased simple eglandular hair and numerous stellate hairs which are often dense along the margins. Ligules yellow, with very short, pale simple eglandular hairs at the apex. Styles yellow to slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, blackish. Flowers 6–8. Apomictic. Native. Banks of streams, rocky slopes and scree in upland areas where it is presumably native. Also occurs in quarries, railway and roadside banks, walls and waste land where it has been introduced. Widespread in Wales and in scattered localities throughout England. France, Switzerland, Germany, Czechoslovakia, Austria, Hungary and Bulgaria. The epithet consociatum is very apposite as it often grows with either H. acuminatum or H. argillaceum with which it forms a connecting link, causing Sell and West to put all three taxa into one variable species. These three species do, however, occur uniformly in some populations and it seems best to follow Jordan and treat them as separate species. H. consociatum was the plant A. Ley consistently called H. cacuminatum in his herbarium, but appears to have published it (1909) as H. scanicum Dahlst.

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309. H. radyrense (Pugsley) P. D. Sell & C. West Radyr Hawkweed H. lachenalii var. radyrense Pugsley; H. sciaphilum forma paucifolia Ley ex Riddelsd. Phyllopodous perennial herb with a branched stock. Stem 25–80 cm, pale yellowish-green, usually purplish or reddish towards the base, robust, striate, with dense, long, pale, tuberculate-based simple eglandular hairs below, becoming much fewer and shorter upwards, with numerous stellate hairs and few, short, dark glandular hairs above. Leaves yellowish-green on upper surface, paler and often suffused pinkish-red beneath; basal usually rather few, the outer with lamina 4–11 × 2–6 cm, elliptical, broadly ovate or nearly subrotund, rounded-obtuse at apex, dentate, especially in lower half, with large, broad to narrow mammiform teeth to denticulate and subtruncate and sometimes with two reflexed teeth at base, the inner with lamina broadly ovate or oblong-ovate, obtuse to subacute at apex, dentate, the teeth large and mammiform, and subtruncate at base with large spreading or deflexed teeth sometimes descending on the medium to very long, densely long-hairy petioles; cauline 2–3, with lamina 4–12 × 2–7 cm, like inner basal, shortly petiolate; all with short to long, pale simple eglandular hairs on both surfaces and the margin, those on the upper surface rather stiff, sometimes with a few stellate hairs on the lower surface. Inflorescence with 2–14 capitula, cymosesubcorymbose; peduncles slender, curved, with dense stellate hairs and numerous, more or less equal, short, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–11 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linearlanceolate, obtuse at apex, with numerous, short to medium unequal, dark glandular hairs, a very occasional, medium, dark-based simple eglandular hair, and dense stellate hairs on the margins and scattered ones elsewhere. Ligules yellow, with numerous, very short simple eglandular hairs at apex. Styles yellow. Receptacle pits shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–7. Apomictic. Native. Shady banks and ledges. Known only from the region of Radyr in Glamorganshire. Endemic. 310. H. argillaceum Jord. Southern Hawkweed H. vulgatum subsp. argillaceum (Jord.) Sudre; H. vulgatum var. argillaceum (Jord.) Arv.-Touv.; H. vulgatum var. sciaphilum Uechtr.; H. sciaphilum (Uechtr.) F. Hanb.; H. lachenalii auct.; H. vulgatum subsp. lachenalii auct.; H. lachenalii var. transiens Ley; H. deductum auct.; H. brevidentatum Jord. var. multiflorum Zahn; H. aspernatum var. anglicum Zahn; H. arrectarium auct.; H. alfvengrenii var. plurifolium Zahn; H. chlorophyllum auct.; H. vulgatum var. genuinum Syme; H. vulgatum var. medium auct.; H. cacuminatum auct.; H. scanicum auct. Phyllopodous perennial herb with a branched stock. Stems 20–90 cm, in shade pale green, in exposed situations deep brownish-purple, and in large populations showing every degree of brownish-purple colouring in between, slender to robust, often flexuous, striate, with numerous, long, pale simple eglandular hairs below, becoming shorter and less numerous upwards, and numerous stellate hairs and few to

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fairly numerous, short, dark glandular hairs above. Leaves dull, medium green, paler beneath, sometimes tinted purplish especially if growing on walls or in dry situations; basal usually numerous, sometimes few, the lamina 5– 11 × 2–4 cm, elliptical to broadly elliptical, ovate, lanceolate or oblong-elliptical, mostly more or less acute at apex but sometimes obtuse-mucronulate, denticulate to shortly dentate, the teeth mammiform to narrowly mammiform and not more than 6 mm, cuneate to attenuate at base, the petioles up to 9 cm; cauline 3–7, gradually decreasing in size upwards, the lower similar to basal, the teeth short and petiolate, the upper with lamina lanceolate to linear, entire, denticulate or shortly dentate, the teeth not exceeding 5 mm, and sessile; all with numerous, short to long, pale simple eglandular hairs on both surfaces and the margins or sometimes nearly glabrous on the upper surface and the petioles with dense, long simple eglandular hairs. Inflorescence with 3–50 capitula, in a paniculate-corymbose cyme, sometimes with a branch from the upper leaf axils; peduncles slender to rather robust, suberect to spreading, straight to curved, with dense stellate hairs, numerous to fairly dense, fine to quite strong, dark glandular hairs and rarely with a solitary simple eglandular hair. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–13 × 1.0– 1.5 mm, olive-green to blackish-green, inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous to dense, unequal, very short to medium, dark glandular hairs, sometimes with occasional to few, dark-based, medium simple eglandular hairs, and numerous to dense stellate hairs especially along the margins. Ligules yellow, glabrous or with very short simple eglandular hairs at apex. Styles yellow to discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–8. Apomictic. Very variable. Plants in the open, especially if on walls, have thick leaves and dark glandular hairs. Plants in shade tend to have thinner, more flaccid leaves and paler glands, with variable intermediates if there is a large colony. Possibly introduced. Grassy banks, quarries, hedge sides, roadsides, waste places, railway banks, tracksides and rocky places. Throughout most of England and Wales and has been recorded for Co. Kerry in Ireland. Central Europe from Spain and France to Hungary. It is possible that H. consociatum and H. nemophilum are its native relatives, and its introduction has confused the taxonomy. 311. H. chlorophyllum Jord. ex Boreau Green-leaved Hawkweed H. vulgatum subsp. chlorophyllum (Jord. ex Boreau) Zahn; H. vulgatum subsp. argillaceum var. chlorophyllum (Jord. ex Boreau) Sudre Phyllopodous perennial herb with a branched stock. Stems 50–65 cm, pale yellowish-green, sometimes brownishpurple towards the base, rather slender, flexuous, striate, with numerous, pale, long simple eglandular hairs below, becoming fewer and shorter upwards, with numerous stellate hairs above. Leaves medium yellowish-green on upper surface, paler beneath; basal few, the lamina 5–11 × 2–5 cm, elliptical to broadly elliptical or obovate-elliptical, mostly rounded-mucronulate, rarely acute at apex, subentire to den-

ticulate or shortly dentate, the teeth short and sharp or mammiform, cuneate to attenuate at base, the long-hairy petioles up to 5 cm; cauline 2–4, gradually decreasing in size upwards, the lower like the basal and usually as large, acute and petiolate, the upper with lamina ovate or lanceolate, gradually narrowed to an acute apex, dentate in the lower half, sessile; all with numerous, short to long, pale simple eglandular hairs on both surfaces and the margins, particularly on the midrib beneath, or with very few on the upper surface. Inflorescence with up to 12 capitula in an irregular cyme, sometimes with a long lower branch; peduncles long, slender and suberect, with dense stellate hairs, few to fairly numerous, short to medium, pale simple eglandular hairs, and few to fairly numerous, very short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 0.8–1.2 mm, rather pale olive-green, the inner with broad paler margins, all narrow linear-lanceolate, more or less acute, with numerous, unequal, slender, short and very short, dark glandular hairs and occasional, medium, dark simple eglandular hairs, and very few stellate hairs on the margins. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins incise-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Possibly native. Known only from Stouting in Kent. Widespread in central Europe from France to Hungary. 312. H. nemophilum Jord. ex Boreau Grassland Hawkweed H. vulgatum subsp. argillaceum var. asperatum Sudre; H. cantianum auct. Phyllopodous perennial herb with a branched stock. Stems 25–70 cm, pale green, usually suffused brownish-purple in lower half, slender but rigid, flexuous, striate, with numerous, long, pale simple hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and few to fairly numerous, short, dark glandular hairs above. Leaves dull, medium green on upper surface, paler beneath and sometimes tinted purplish beneath; basal few to numerous, the lamina 2.5–6.0(−10) × 1.5–3.0 cm, narrowly elliptical, lanceolate to ovate, oblong-lanceolate or oblong-elliptical, mostly acute, rarely rounded-mucronulate at apex, denticulate to dentate, the teeth short and sharp or sharply mammiform, cuneate to attenuate at base, the slender petioles up to 5 cm; cauline 3–6, often rapidly decreasing in size upwards, the lower similar to basal and shortly petiolate, the upper with lamina lanceolate or linear, acute at apex, entire to dentate and sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins and with dense long ones on the petioles. Inflorescence with 5–30 capitula, cymose-corymbose, sometimes with a long lower branch; peduncles with dense stellate hairs, numerous, very short and short, fine, dark glandular hairs and occasional, short or medium, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, greyish olive-green, the inner with paler margins, all narrowly linear-lanceolate, more or less acute at apex, with numerous, very short to medium, fine, dark glandular hairs, sometimes with an occasional, medium, dark-based simple eglandular hair and numerous

39. Hieracium stellate hairs particularly along the margins and at the base. Ligules yellow, with numerous very short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Chalk and limestone cliffs, scree and grassland where it seems to be indigenous in western and northern England and Wales, with a few scattered records elsewhere in England where it may be introduced. A solitary specimen has been seen from Blairgowrie in Perthshire. Central Europe. 313. H. aviicola Jord. ex Boreau Many-toothed Hawkweed H. vulgatum subsp. deductum microgen. aurulentum var. aviicola (Jord. ex Boreau) Sudre: H. vulgatum subsp. aviicola (Jord. ex Boreau) Zahn; H. sciaphilum var. strumosum Ley; H. strumosum (Ley) Ley Phyllopodous perennial herb with a branched stock. Stems 40–80 cm, pale green, usually purplish in lower half, slender and flexuous but rigid, striate, with numerous, long, pale simple eglandular hairs below, which are fewer and shorter upwards, and numerous stellate hairs and a few, short glandular hairs above. Leaves dull, medium yellowish-green on upper surface, paler beneath; basal few to numerous, the lamina 6–10 × 2–4 cm, narrowly to broadly elliptical or lanceolate, usually more or less acute at apex, sometimes rounded-mucronulate, sharply denticulate to regularly and sharply serrate-dentate, the teeth small and sharp to narrowly and sharply mammiform, cuneate to attenuate at base, the petioles up to 7 cm; cauline 4–6, gradually decreasing in size upwards, most similar to basal, or uppermost linear to lanceolate, acute or acuminate at apex, entire or with a few teeth, usually sessile; all with numerous, short to long, pale simple eglandular hairs on both surfaces and the margins and dense, long ones on the petioles. Inflorescence with 5– 30 capitula, paniculate-corymbose, usually with long lower branches from the leaf axils; peduncles slender and suberect or erecto-patent, with numerous to fairly dense stellate hairs, numerous, short, fine, dark glandular hairs and sometimes with an occasional, pale, medium simple eglandular hair. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–11 × 1.0–1.2 mm, olive to blackish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to a more or less acute apex, with numerous, unequal, very short to short, sometimes medium, dark glandular hairs, occasional to few, medium, dark-based simple eglandular hairs and numerous stellate ones along the margins. Ligules yellow, glabroustipped or with very short simple eglandular hairs at apex. Receptacle pits with margins sharply dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 6–8. Apomictic. The group of plants brought together here are characterised by having their leaves narrowed at both ends with sharp regular teeth and a large panicle of capitula which are glandular-hairy with stellate margins to the involucral bracts. The type of glandular hair is, however, rather variable. Probably introduced. Ranmore in Surrey; railway near Royston in Cambridgeshire; St Vincent’s Rocks, Bristol

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and a limestone bank near Stroud, Gloucestershire; wood by the Grwyne Valley, Herefordshire; roadside bank south of Abergwesyn and the Upper Tawe Valley in Breconshire; wood at Pont Esgob in Monmouthshire; Leigh in Worcestershire and Eglwyseg in Denbighshire. The lectotype of H. strumosum (Ley) Ley is this plant, but some specimens named as that by Ley are other species. It is possible it is native in some of the Welsh localities. France, Switzerland, Germany, Austria and Hungary. 314. H. latebrosum Jord. ex Boreau Walthamstow Hawkweed H. deductum Sudre; H. festinum subsp. deductum (Sudre) Sudre; H. vulgatum subsp. deductum subvar. latebrosum (Jord. ex Bor.) Sudre; H. vulgatum subsp. jaccardii var. deductum forma latebrosum (Jord. ex Bor.) Zahn Phyllopodous perennial herb with a branched stock. Stems 30–60 cm, pale yellowish-green, often suffused with brownish-purple below, more or less robust, flexuous, striate, with dense, medium to long, pale simple eglandular hairs below, becoming much fewer and shorter upwards, and numerous stellate hairs and a few, short, dark glandular hairs in the upper part. Leaves dull medium green on upper surface, paler and sometimes tinted reddish beneath; basal few, the lamina 4–8 × 2–4 cm, ovate or elliptical, more or less acute at apex, subentire to dentate with rather small mammiform or narrow mammiform teeth, cuneate at base, with, long-hairy petioles up to 4 cm; cauline 3–6, gradually decreasing in size upwards, ovate, acute at apex, dentate, the teeth sometimes narrowly mammiform, cuneate at base, the lowest with a short petiole, the upper sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins, or nearly absent from the upper surface, the upper cauline with a few stellate hairs beneath. Inflorescence with up to 15 capitula, cymose-corymbose, often with a long lower branch; peduncles very slender, with dense stellate hairs, numerous, short, dark glandular hairs and few to numerous, short and medium, pale, darkbased simple eglandular hairs. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–10 × 1.0–1.2 mm, dark olive-green, the inner with pale margins, all linear-lanceolate, obtuse at apex, with numerous, unequal, slender, short and very short, dark glandular hairs, a few, pale, dark-based, medium simple eglandular hairs and numerous stellate hairs especially on the margins. Ligules yellow, glabrous or puberulous at apex. Styles dark orange. Receptacle pits with margins sharply dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Known only from grassy banks and cracks in concrete round Walthamstow Reservoirs. A specimen from scree on Breidden Hill, Montgomeryshire and an old specimen from near Studley in Yorkshire may be it, but need confirmation. Central Europe. 315. H. pulchrius (Ley) W. R. Linton Beautiful Hawkweed H. sciaphilum var. pulchrius Ley; H. vulgatum subsp. pulchrius (Ley) Zahn

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Phyllopodous perennial herb with a branched stock. Stem 30–85 cm, pale yellowish-green, often suffused reddishpurple especially towards the base, slender to robust, striate, with numerous, medium to long, pale simple eglandular hairs below, becoming fewer upwards and numerous stellate hairs and few to numerous, short, black glandular hairs above. Leaves bright green on upper surface, paler beneath and often tinted purplish; basal few to numerous, the lamina 3–13 × 2–7 cm, subrotund, elliptical, obovate or oblong-obovate, obtuse-mucronate to acute at apex, denticulate to sinuate-dentate the teeth often apiculate, abruptly contracted to cuneate at base, the long-hairy petioles usually rather short; cauline 2–4, the lower with lamina elliptical or ovate, acute at apex, dentate, cuneate at base and petiolate, the upper smaller, the lamina ovate, lanceolate or linear, acute at apex, subentire to dentate, mostly sessile; all with few to numerous, short to long, pale simple eglandular hairs on both surfaces or glabrous on the upper surface. Inflorescence with 4–20 capitula, cymose-corymbose, sometimes with a long, lower branch; peduncles rather short, with dense stellate hairs, numerous, unequal, very short and short, black glandular hairs and an occasional dark simple eglandular hair. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–13 × 1.2–1.5 mm, almost black, linear-oblong, obtuse at apex, with numerous, unequal, very short, short and medium, black glandular hairs, without simple eglandular hairs and with few stellate hairs towards the base. Ligules golden yellow, glabroustipped. Styles discoloured. Receptacle pits with margins sharply subulate-dentate. Achenes 3.0–3.5 mm, blackish. Flowers 6–8. mm. Apomictic. Native. Ledges on limestone and sandstone cliffs. Craig Du, Craig Cerrig-gleisiad, Fan Nedd, Craig-y-cilau (Craig Cille) and the central cliff of the Beacons in Breconshire and Llyn-y-fan-fach in Carmarthenshire. Endemic. 316. H. lortetiae Balbis Lortet’s Hawkweed H. vulgatum subsp. deductum var. lortetiae (Balbis) Sudre; H. vulgatum subsp. jaccardii var. lortetiae (Balbis) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale green, suffused brownish-purple at the base, slender, flexuous, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and an occasional, short, dark glandular hair above. Leaves pale green on upper surface, even paler and slightly greyish beneath; basal few, the lamina 6–9 × 2.0–3.5 cm, narrowly elliptical to lanceolate, acute at apex, entire or denticulate or with an occasional tooth, cuneate to attenuate at base, the petioles up to 5 cm; cauline 3–5, gradually decreasing in size upwards, the lower like basal and petiolate, the upper lanceolate or linear, entire or nearly so and sessile; all with few to numerous, pale, short to medium simple eglandular hairs on both surfaces and the margins, with longer ones on the petioles. Inflorescence with 2–27 capitula, cymose-corymbose, often with a long lower branch; peduncles rather short, erect-spreading, with numerous to fairly dense stellate hairs, numerous, yellowish to dark, short glandular hairs and sometimes with occasional, medium, pale simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts

incumbent in bud, 5–13 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, more or less acute at apex, with numerous, unequal, very short to short, dark glandular hairs, very rarely with a solitary, pale simple eglandular hair and numerous stellate hairs particularly on the margins and near the base. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 6–7. Apomictic. Native. Torpantau and a road bridge over the River Usk, near Brecon in Breconshire, Whitbourne in Herefordshire, rocks by the river in a wood below Pen-y-craig in Denbighshire and a wet, rocky gorge, Cwm Rhiwiau on the boundary of Montgomeryshire and Denbighshire. France. Named after Cl´emence Lortet (1772–1835). 317. H. festinum Jord. ex Boreau Hairy-leaved Hawkweed H. vulgatum subsp. argillaceum microgen. festinum (Jord. ex Boreau) Sudre; H. vulgatum subsp. festinum (Jord. ex Boreau) Sudre Phyllopodous perennial herb with a branched stock. Stems 50–70 cm, pale green, sometimes purplish in lower half, more or less robust but flexuous, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter above, and numerous stellate hairs and occasional, dark, short glandular hairs above. Leaves dull, pale yellowish-green on upper surface, even paler beneath and sometimes tinted purplish; basal few to numerous, the lamina 6–12 × 2–4 cm, oblong, narrowly elliptical-oblong or obovate-oblong, sometimes narrowed, but rounded-mucronulate at apex, undulate-denticulate to shortly undulate-dentate, the teeth short and sharp or sharply mammiform, cuneate to attenuate at base, the petiole up to 5 cm; cauline 3–7, gradually decreasing in size upwards, the lamina narrowly elliptical, oblong-elliptical or oblong, obtuse to acute at apex, entire or dentate in lower half and shortly petiolate, the upper smaller, long-acute or acuminate, entire or with a few teeth and shortly petiolate or sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins, or glabrous or nearly so on upper surface, stellate hairs present on under surface of upper cauline leaves. Inflorescence with 14–35 capitula, laxly paniculate-corymbose, with long, slender branches from the upper leaf axils and the uppermost part often subumbellate; peduncles slender, suberect or spreading, with dense stellate hairs, few, short, dark glandular hairs and few to fairly numerous, short to medium, dark-based simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, olive-green to slightly blackish-green, the inner with paler margins, all linear-lanceolate, drawn out to a narrow but subobtuse apex, with numerous unequal, very short and short, dark glandular hairs, occasional, medium, dark-based simple eglandular hairs, and some stellate hairs towards the base. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 6–8. Apomictic. Probably introduced. There are colonies at Bitchett and Pembury in Kent and at Gray’s Chalk Pit in Essex. A

39. Hieracium specimen from rocks by a stream at Cwm Irfon in Breconshire appears to be it and could be native. Central Europe from France and Holland to the Caucasus. 318. H. cheriense Jord. ex Boreau Cher Hawkweed H. vulgatum subsp. argillaceum var. cheriense (Jord. ex Boreau) Zahn; H. vulgatum subsp. argillaceum var. sublaeve Sudre; H. tunbridgense Pugsley; H. lachenalii var. pseudoporrigens Pugsley; H. porrigens auct.; H. lachenalii forma nemorale Pugsley Phyllopodous perennial herb with a branched stock. Stems 30–70 cm, pale green, often brownish-purple in lower half, usually slender and flexuous, sometimes slightly robust, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter above, and numerous stellate hairs and an occasional, very short, fine, dark glandular hair above. Leaves pale to medium green on upper surface, paler and sometimes tinted purple beneath; basal few to numerous, the lamina 2–9 × 1–4 cm, lanceolate, ovate or elliptical, mostly more or less acute at apex, sometimes rounded-mucronulate, denticulate to dentate, the teeth small and sharp to mammiform or narrowly mammiform, rounded to cuneate at base, the petiole up to 7 cm; cauline 2–6, gradually decreasing in size upwards, the lower like basal and petiolate, the upper lanceolate or linear, long-acute at apex, entire to dentate and sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins, sometimes sparse on the upper surface and with dense, long ones on the petiole. Inflorescence with 2–20 capitula, laxly paniculate-corymbose, usually with 1 or more long, slender branches from the leaf axils; peduncles long and slender, with numerous to fairly dense stellate hairs, a few, scattered, very short, dark glandular hairs and sometimes an occasional simple eglandular hair. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–11 × 1.0–1.2 mm, olive-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, unequal, short or very short, fine, dark glandular hairs, a few stellate hairs towards the base and sometimes with occasional simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles yellow to slightly discoloured. Receptacle pits with margins incise-dentate. Achenes 3.0– 3.5 mm, purplish-black. Flowers 6–8. Apomictic. Probably introduced. Scattered records across southern England on roadsides and wood margins, particularly in Kent, with isolated records in Hertfordshire, Cambridgeshire, Monmouthshire, Breconshire, and Argyllshire. Central Europe from France to Hungary. 319. H. rectulum Ley Dyfed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–50(−80) cm, pale yellowish-green, often reddish-purple in the lower half, usually rather slender, rarely robust, striate, with numerous to dense, long, whitish simple eglandular hairs below, becoming fewer, shorter and darkbased upwards, and numerous stellate hairs and an occasional, short, dark glandular hair above. Leaves medium green on upper surface, paler and often suffused purple beneath; basal few, the lamina 2.5–8.0(−10.0) × 1.5– 3.5(−4.0) cm, lanceolate, elliptical or oblong-lanceolate,

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obtuse-mucronate to acute at apex, entire or denticulate, rounded to cuneate at base, the petioles short and densely long-hairy; cauline 1–4, like the basal and petiolate or lanceolate or linear and sessile; all with short to long, pale simple eglandular hairs on both surfaces and the margins, or nearly glabrous above. Inflorescence with 3–10(−20) capitula, cymose-corymbose, sometimes with a long lower branch; peduncles mostly short, suberect, with dense stellate hairs, numerous, very short, dark glandular hairs and an occasional, short or medium, pale simple eglandular hair. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, blackishgreen, the inner with paler margins, all linear-lanceolate, subobtuse to acute at apex, with numerous, very short and short, unequal, dark glandular hairs and very rarely a solitary, dark simple eglandular hair, stellate hairs absent or a few near the base. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rocky slopes and ledges, roadside banks and a cemetery. Known only from near Llangadoc in Carmarthenshire, several localities in Cardiganshire and Llanfairtalhaiarn in Denbighshire. Section 17. Hieracium Section Hieraciotypus Dumort. nom. inval.; Section Pulmonaria Monnier; Series Pulmonaria (Monnier) Fries; Taxon Aurelloidea Arv.-Touv.; Section Aurelloidea (Arv.-Touv.) Gremli; Taxon Bifida Arv.-Touv.; Subsection Bifida (Arv.-Touv.) Pugsley; Series Bifida (Arv.-Touv.) P. D. Sell & C. West comb. inval.; Section Bifida (Arv.-Touv.) A. R. Clapham; Taxon Subvulgata Almq.; Subsection Subvulgata (Almq.) Dahlst., non F. N. Williams; Taxon Subcaesia Almq.; Subsection Subcaesia (Almq.) Dahlst.; Subsection Aggregata F. N. Williams; Subsection Pictorum F. N. Williams; Subsection Silvatica F. N. Williams; Subsection Trichopetala F. N. Williams; Subsection Communia Rouy; Subsection Glandulosa Pugsley; Section Glandulosa (Pugsley) A. R. Clapham; Subsection Sagittata Pugsley; Section Sagittata (Pugsley) A. R. Clapham; Subsection Stellatifolia Pugsley Phyllopodous perennial herbs. Stems hairy or not. Leaves green, rarely glaucous, sometimes spotted; basal usually numerous, the outer often different from the inner; cauline 0–1(−3). Inflorescence paniculate-corymbose, usually with many capitula; peduncles and branches often curved. Capitula small to medium. Involucral bracts usually incumbent in bud, usually narrow, variably clothed. Ligules glabrous or hairy at apex. Styles yellow or discoloured. Receptacle pits dentate or dentate-fimbriate. Achenes dark. Species of this section occur throughout Great Britain, but are more common in the west and north, and are scarce in Ireland. Native species occur in rocky places and by streams in the mountains and possibly in open woods and on heaths in the lowlands. Many species are introduced and are abundant on disturbed railway banks and roadsides. In these localities a half dozen species can grow intermixed and one or two plants can soon become hundreds. In this situation there is a greater chance of sports appearing and

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reproducing themselves, although a population can soon disappear again if the area becomes overgrown. 320. H. aggregatum Backh. fil. Aggregate-headed Hawkweed H. bifidum subsp. aggregatum (Backh. fil.) Zahn Phyllopodous perennial herb with a branched stock. Stems 20–50 cm, pale yellowish-green, sometimes reddish tinted below, strictly erect, striate, more or less floccose throughout, but particularly so above, sometimes with an occasional, pale simple eglandular hair. Leaves deep green on upper surface, paler beneath; basal few to numerous, with lamina 2–14 × 1–6 cm, ovate, lanceolate or elliptical, obtuse-mucronate to acute at apex, nearly entire to undulatedenticulate to irregularly dentate, the teeth large and mammiform, the base cuneate or subtruncate, often asymmetrical and sometimes with teeth on the rather long petiole which has numerous, long, pale simple eglandular hairs; cauline often absent, when present with lamina lanceolate, acute at apex, more or less dentate especially below, subtruncate at base and petiolate; hairiness very variable, sometimes glabrous on both surfaces, sometimes with scattered, short and medium, pale simple eglandular hairs, sometimes with dense stellate hairs on the lower surface, usually with short to medium, pale simple eglandular hairs on the margin. Inflorescence corymbose-subumbellate, the acladium rather short to long, peduncles with dense stellate hairs, without or with occasional simple eglandular or glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4.0–11.5 × 1.0–1.3 mm, dark green, the inner with paler margins, all linear-lanceolate, mostly obtuse at apex, with numerous stellate hairs particularly on the margins and at the base, with scattered very short to short, dark glandular hairs and a few solitary, short or medium, pale, dark-based simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins obscurely dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 7–8. Apomictic. Despite the great variation in the leaves, this species is easily recognised by its subumbellate inflorescence, peduncles with stellate but without glandular or simple eglandular hairs, and the sparsely clothed involucral bracts. Very large plants have been collected on Cairntoul, Aberdeenshire. Native. A very local, alpine species of rocky ledges and streamsides. Mountains of Clova in Forfarshire, south Aberdeenshire, Argyllshire and the Cairngorms. Endemic. There are very few modern records. 321. H. caesiomurorum Lindeb. Long-stalked Hawkweed H. caesium subsp. caesiomurorum (Lindeb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–80 cm, pale yellowish-green, often reddish or purplish below, slender to robust, striate, with few to fairly numerous, pale, long simple eglandular hairs below, sometimes some shorter ones above, but often absent, and few to numerous stellate hairs in upper part. Leaves bright or bluishgreen on upper surface, paler beneath and often tinted purplish; basal few, the lamina 2.5–13.0 × 2.0–6.0 cm, ovate, elliptical, lanceolate, lanceolate-oblong or ellipticoblong, obtuse-mucronate to acute or acuminate at apex,

denticulate to dentate or incise-dentate, the teeth mostly mammiform or narrowly mammiform and sometimes those near the cuneate base long and narrow, the petioles long, up to 10 cm, and with numerous, long, pale simple eglandular hairs; cauline (0–)1–2(−3), the lower like the basal, sometimes with long teeth and often with long petioles, the upper with lamina lanceolate, acute or acuminate at apex, toothed and sessile, or bract-like; all glabrous on upper surface, with numerous, short to long, pale simple eglandular hairs beneath, especially on the midrib, and on the margins, and sometimes with stellate hairs on the underside of the cauline. Inflorescence with 2–10(−15) capitula, paniculate-corymbose, sometimes with a long, lower branch; peduncles rather short and slender, with dense stellate hairs, numerous, short, dark glandular hairs and few to numerous, short to medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 0.8–1.2 mm, dark olive green, the inner with paler margins, all linear-lanceolate, gradually narrowed, but mostly obtuse at apex, with dense, short and medium, pale, dark-based simple eglandular hairs and few, short, dark glandular hairs, and numerous stellate hairs along the margin and at the apex. Ligules yellow, with very short simple eglandular hairs at apex. Styles yellowish. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Our plants fit well the Scandinavian plant, but seem to have larger capitula. Plants with long teeth at the base of the leaves may be referable to var. umbraticum K. Johans. ex Dahlst. Native. Riverside rocks and cliff ledges. Widespread in the Scottish mountains. Scandinavia. 322. H. irregularidens P. D. Sell Blunt-toothed Hawkweed H. dissimile var. poliaenum auct. Phyllopodous perennial herb with a branched stock. Stem 30–65 cm, pale yellowish-green, often reddish-purple below, slender to robust, striate, with fairly numerous, long, pale simple eglandular hairs below, becoming fewer and shorter or absent upwards, and numerous stellate hairs in the upper part. Leaves more or less bluish-green on upper surface, paler beneath; basal few to numerous, with the lamina often large, 4–14 × 3.5–6.0 cm, ovate-oblong, ovate or lanceolate-oblong, mostly acute but occasionally obtuse at apex, irregularly dentate, often deeply so, with the lower teeth often long and narrow but most teeth obtuse though apiculate, abruptly contracted or subtruncate at base and with the lower teeth sometimes extending onto the petiole, the petioles usually rather short but up to 9 cm and with numerous, long, pale, wavy simple eglandular hairs; cauline 0–2, one often large like the basal and petiolate, the upper when present narrow, sometimes with long, narrow teeth; all glabrous or nearly so on upper surface, with numerous, short to long simple eglandular hairs beneath and on the margin. Inflorescence with 2–12 capitula, paniculatecorymbose, often with a long lower branch; peduncles rather slender, erect, with dense stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral

39. Hieracium bracts incumbent in bud, 4–12 × 1.2–1.5 mm, dark oliveto blackish-green, the inner with paler margins, all linearlanceolate, gradually narrowed but obtuse at apex, with very numerous, short to medium, pale, dark-based simple eglandular hairs, fairly numerous, short, dark glandular hairs, and numerous stellate hairs along the margins. Ligules yellow, with short simple eglandular hairs at apex. Styles yellow or slightly discoloured. Receptacle pits with margins subulatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. This species differs from H. caesiomurorum in its more numerous, blunt leaf-teeth and usually shorter petioles, and from H. cuspidens in its blunter leaf-teeth and more contracted base to the leaf. Native. Grassy banks, rocks and shingle by streams and on cliff ledges. Common in the mountains of central Scotland. Endemic. 323. H. armadalense P. D. Sell Armadale Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–30 cm, pale yellowish-green, brownish-purple towards the base, more or less robust, with a few, long, pale simple eglandular hairs below and few or none above, and numerous stellate hairs and sometimes an occasional glandular hair in the upper part. Leaves medium, slightly bluish-green on upper surface, paler beneath; basal numerous, with the lamina 2.5–10.0 × 1–6 cm, elliptical to broadly elliptical or ovate, usually more or less acute, sometimes roundedmucronulate at apex, denticulate to shallowly dentate or serrate-dentate and with the teeth sharp, rounded or cuneate at base, the petioles short, up to 2 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1, when present with the lamina linear or linear-lanceolate, acute at apex, dentate, sessile; all glabrous or with a few, stiff simple eglandular hairs near the margin on the upper surface, with few, short to medium simple eglandular hairs beneath and more numerous ones on the margins. Inflorescence with 5– 12 capitula, condensed paniculate-corymbose, often with a long lower branch; peduncles mostly short and rigid, with dense stellate hairs, numerous, short to medium, pale simple eglandular hairs and an occasional, short, dark glandular hair. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all broadly linear-lanceolate, narrowed to a subacute apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, occasional, short, dark glandular hairs and some stellate hairs on the margins and towards the base. Ligules golden yellow, with some very short simple eglandular hairs towards the apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 6–7. Apomictic. Native. Known only on sandhills at Armadale in Sutherland. Endemic. 324. H. clivicola (F. Hanb.) Pugsley Hill Hawkweed H. euprepes var. clivicola F. Hanb.; H. sagittatum subsp. euprepes var. clivicola (F. Hanb.) Zahn; H. euprepes F. Hanb., non Peter; H. sagittatum subsp. euprepes Zahn

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Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, sometimes slightly tinted below, slender to robust, with numerous, long, pale, wavy simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate and an occasional short, dark glandular hair in the upper part. Leaves dark green on upper surface, paler and often suffused purplish beneath; basal few to numerous, with lamina 6–12 × 2.5–4.5 cm, mostly elliptical or ovate-elliptical, sometimes oblongelliptical, obtuse-mucronate to acute at apex, sharply denticulate to shortly dentate and the teeth more or less mammiform, cuneate at base, the petioles up to 6 cm and with numerous, pale, long, wavy simple eglandular hairs; cauline usually 1, with lamina linear to lanceolate, acute at apex, sharply dentate, cuneate at the sessile base; all with short to medium, pale, stiff hairs on both surfaces, longer on the midrib beneath and on the margins. Inflorescence with 3–9 capitula, paniculate-corymbose, sometimes with a long lower branch; peduncles short, with dense stellate hairs, numerous, pale, dark-based, short to medium simple eglandular hairs and occasional, short, dark glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.2– 1.5 mm, blackish-green, the inner with paler margins, all broad linear-lanceolate, obtuse at apex, with dense, short to medium, dark-based simple eglandular hairs, few, short, dark glandular hairs, and some stellate hairs on the margins and towards the base. Ligules deep yellow, glabrous-tipped. Styles yellowish. Receptacle pits with margins fimbriatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and scree. Wales, particularly Breconshire and Caernarvonshire, and north Ireland. Endemic. 325. H. orcadense W. R. Linton Orkney Hawkweed H. subramosum subsp. orcadense (W. R. Linton) Zahn; H. sagittatum subsp. sagittatum var. abrasum Dahlst.; H. oistophyllum var. abrasum (Dahlst.) Pugsley; H. subexpallescens Dahlst.; H. sagittaticeps Dahlst.; H. kalsoense subsp. burnense Druce & Zahn; H. paraliaeforme Dahlst.; H. subalpestrifrons Dahlst.; H. holopleurum auct.; H. ardisodon Dahlst. Phyllopodous perennial herb with a branched stock. Stem 25–55 cm, pale yellowish-green, often purplish towards the base, slender, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, numerous stellate hairs throughout and an occasional, short, dark glandular hair in the upper part. Leaves deep green on upper surface, paler beneath; basal few to numerous, with lamina 3–8 × 2–4 cm, elliptical, ovate, lanceolate or oblanceolate, rounded-mucronulate to acute at apex, subentire, denticulate or dentate, the teeth sometimes sharp and sometimes mammiform, the petioles up to 5 cm, with numerous, long, pale simple eglandular hairs; cauline 0–1(−3), when present with lamina lanceolate, acute at apex, subentire to dentate, cuneate at base and petiolate; all with short to long, stiff, pale simple eglandular hairs on both surfaces and the margins or nearly glabrous on the upper surface. Inflorescence with 2–12 capitula, paniculatecorymbose; peduncles often curved, those of the acladium and adjacent capitulum often very short, with numerous

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stellate hairs, numerous, short to medium, dark-based simple eglandular hairs and few, short, dark glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, blackishgreen, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short to medium, pale, darkbased simple eglandular hairs and few, short, dark glandular hairs, and stellate hairs at apex and towards the base. Ligules golden yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins fimbriate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and streamsides. Orkney Islands and scattered localities on the mainland of Scotland. Faeroes. 326. H. subtenue (W. R. Linton) Roffey Dark-bracted Hawkweed H. silvaticum var. subtenue W. R. Linton; H. bifidum subsp. subtenue (W. R. Linton) Zahn; H. subtenue var. canispense Pugsley Phyllopodous perennial herb with a branched stock. Stem 25–45 cm, pale yellowish-green, often suffused brownishpurple in the lower half, slender, flexuous, striate, with few to fairly numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate and an occasional, short, dark glandular hair in the upper part. Leaves medium dull green on upper surface, paler beneath; basal usually few, the outer with lamina 3– 6 × 1–3 cm, elliptical, rounded-mucronulate at apex, subentire or minutely denticulate, and rounded or shortly cuneate at base, the inner with lamina 4–10 × 2.0–3.5 cm, oblong, lanceolate or oblong-lanceolate, obtuse-mucronate to more or less acute at apex, subentire to denticulate or shallowly sinuate-dentate and cuneate to attenuate at base, the petioles up to 8 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline usually 1, large, with lamina 5– 10 cm, lanceolate or oblong-lanceolate, long-acute at apex, subentire, denticulate or with occasional large teeth towards the cuneate or attenuate base and petiolate; all glabrous or nearly so, or with scattered, short, pale simple eglandular hairs on upper surface, with more or less numerous, medium to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 2–9(−15 cult.) capitula, paniculatecorymbose, sometimes with a long lower branch; peduncles rather long and slender, straight or curved, with numerous to dense stellate hairs, few to fairly numerous, medium, pale, dark-based simple eglandular hairs and short (0.2–0.4 mm) glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–11 × 1.0– 1.4 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, obtuse to acute at apex, with numerous, short to medium, pale, dark-based simple eglandular hairs, few to numerous, very short to short (0.2–0.4 mm), dark glandular hairs, and few to numerous stellate hairs on the margins. Ligules lemon yellow, glabrous or with few, very short simple eglandular hairs at the apex. Styles yellow to slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Often mixed with H. duriceps on herbarium sheets, with which species it often grows. The glandular hairs of the involucre are quite different in the two species.

Native. Cliff ledges, rocky slopes, screes and streamsides. A common species in Ross-shire and Sutherland and occurring in a few localities further south into central Scotland. Endemic. 327. H. dipteroides Dahlst. Aberfeldy Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, sometimes tinted reddish towards the base, rather slender but rigid, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate and short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal few to fairly numerous, the outer with lamina 2– 7 × 2.0–3.5 cm, ovate or elliptical, rounded-mucronulate at apex, subentire, denticulate or with a few shallow teeth and rounded or abruptly contracted at base, the inner with lamina 7–13 × 2.0–4.5 cm, oblong, narrowly elliptical, oblonglanceolate, or lanceolate, subobtuse to acute at apex, subentire especially in the upper two-thirds, denticulate or with a few shallow teeth, the basal pair narrow and hooked or spreading, and abruptly contracted or cuneate at base, the petioles up to 9 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 0–2, when present with lamina linear-lanceolate, gradually narrowed to an acute apex, entire or with an occasional tooth, sessile or petiolate; all with few to numerous, short to medium, pale simple eglandular hairs on both surfaces and longer ones on the midrib beneath and the margins. Inflorescence with 4–10 (numerous in cultivation) capitula, paniculate-corymbose, sometimes with a long lower branch; peduncles curved and quite long, with dense stellate hairs, numerous, short or very short (0.2–0.5 mm), dark glandular hairs and few to numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.3 mm, olive green, the inner with paler margins, all linear-lanceolate, obtuse to acute at apex, with numerous, short (0.3–0.6 mm), dark glandular hairs, numerous, short to medium, pale, darkbased simple eglandular hairs, and numerous stellate hairs along the margins and at the apex. Ligules yellow, glabroustipped. Styles yellow to slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 6–7. Apomictic. In the field the narrow, nearly entire leaves of this species are diagnostic, but in cultivation they become broad and deeply toothed so as to look like a different species, but not one that is known. Native. Cliff ledges and rocky streamsides. In several localities in Perthshire, especially in its type locality at the Birks of Aberfeldy where there are at least 500 plants, near Braemar in Aberdeenshire, near Moy in Lochaber and north of Cluny Castle in Inverness-shire. Endemic, but closely related to the Scandinavian H. dipterum Dahlst. 328. H. inspissatum P. D. Sell Crass-leaved Hawkweed H. silvaticum var. crassum Ley, non H. crassum Almq. Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often purplish at the base, usually rather slender, sometimes robust, striate, with

39. Hieracium numerous, medium to long, pale, wavy simple eglandular hairs below, becoming less so or almost absent above and numerous stellate hairs in the upper part. Leaves deep green with purple spots and blotches on upper surface, paler and often suffused purplish beneath; basal rather few, the outer with lamina 3–9(−13) × 2–4(−5) cm, ovate, elliptical or oblong-elliptical, rounded-mucronulate at apex and denticulate with a few, large, mammiform teeth towards the cuneate base, the inner with lamina ovate or oblongelliptical, acute at apex, denticulate or with a few, large, mammiform teeth towards the base, rarely with very large teeth, in cultivation with numerous teeth, the base cuneate and sometimes with 1–2 of the teeth extending onto the petioles, the petioles up to 4 cm, with numerous, long, pale simple eglandular hairs; cauline 0–1, when present with lamina lanceolate, acute at apex, with a few teeth near the base and petiolate, or bract-like; all with short to medium, pale simple eglandular hairs on both surfaces and the margins and the cauline sometimes with stellate hairs beneath. Inflorescence with 2–5 capitula, furcate-corymbose; peduncles rather long, usually straight, with dense stellate hairs and a few, short or medium, pale simple eglandular hairs. Capitula 35–40 mm in diameter, broad, rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.5–1.7 mm, blackish-green, the inner with paler margins, all broad linear-lanceolate, the outer obtuse, the inner acute at apex, with dense, short to long, dark-based simple eglandular hairs, covered with stellate hairs particularly on the margins and with occasional, very short, dark glandular hairs. Ligules yellow, glabrous or with very few, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 5–7. Apomictic. Native. Limestone cliffs. Pwll Byffre and Y Fan Nedd in the Breconshire Beacons. Endemic. 329. H. subhirtum (F. Hanb.) Pugsley Hairy-headed Hawkweed H. rivale var. subhirtum F. Hanb.; H. sagittatum var. subhirtum (F. Hanb.) W. R. Linton; H. silvaticum var. rivale forma subhirtum (F. Hanb.) F. N. Williams; H. sagittatum subsp. sagittatum var. subhirtum (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, often tinted reddish or purplish below, usually slender, striate, with few to fairly numerous, medium to long, pale simple eglandular hairs below and fewer and shorter ones above, and numerous stellate hairs in the upper part. Leaves bright or yellowish-green on upper surface, paler beneath; basal few to numerous, the outer with lamina 2–7 × 1.0–3.5 cm, elliptical, ovate or subrotund, rounded-mucronulate at apex, denticulate to sinuate-dentate and cuneate, rounded or truncate at base, the inner with lamina 4–10 × 1–3 cm, lanceolate to ovatelanceolate, acute at apex, denticulate to dentate, the lowest teeth sometimes narrowly mammiform and spreading, the petioles up to 5 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1, when present with lamina linear or linear-lanceolate, long-acute at apex, sharply dentate, narrowed at base, petiolate; all with short to medium, rough, pale simple eglandular hairs on the upper surface,

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softer ones beneath and long ones on the midrib beneath and on the margins. Inflorescence with 2–6 capitula, narrowly corymbose; peduncles long, straight to arcuate, with numerous stellate, numerous short to medium, pale simple eglandular hairs and usually a few, minute, dark glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.8–1.2 mm, greyish-green, the inner with paler margins, all linearlanceolate, more or less acute, with dense, short to medium, dark-based simple eglandular hairs, usually a few, minute glandular hairs and a few, scattered stellate hairs. Ligules yellow, usually with a few, very short simple eglandular hairs at apex. Styles yellow to discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddishblack. Flowers 6–7. Apomictic. Native. By rocky streams, on cliff ledges and in other rocky places. Common in central Scotland and Kintyre. Endemic. 330. H. neocoracinum Pugsley Craig Gleisiad Hawkweed H. caesium var. coracinum Ley, non H. coracinum (Dahlst.) Durand & B. D. Jackson Phyllopodous perennial herb with a branched stock. Stem 20–45 cm, pale yellowish-green, sometimes purplish towards the base, striate, flexuous, with few to numerous stellate hairs above, with scattered, short to medium, pale simple eglandular hairs especially above, without glandular hairs. Leaves rather pale yellowish-green on upper surface, paler and sometimes tinted purplish beneath, glabrous or with scattered, short, pale simple eglandular hairs on the upper surface especially near the margin and few to numerous, short to medium, pale simple eglandular hairs especially on the midrib and medium to long simple eglandular hairs on the margins; basal with lamina 2–11 × 1–5 cm, lanceolate, elliptical, ovate or oblong-lanceolate, obtuse-mucronate to acute and sometimes cuspidate at apex, undulate-denticulate to undulate-dentate, sometimes with large aquiline-mammiform teeth at the base which sometimes extend on to the petiole, cuneate to subtruncate at the often asymmetrical base, the petioles short to long, with dense, long, pale, wavy simple eglandular hairs; cauline usually absent, when present with lamina small and linear to lanceolate, acute at apex and more or less entire. Inflorescence compactly cymose-corymbose with a short acladium; peduncles short, with dense stellate hairs, few to numerous, short to medium, pale, dark-based simple eglandular hairs and few, very short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded-based. Involucral bracts more or less porrect in bud, 5–12 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an acute apex, the inner very narrow, with stellate hairs at the base and sometimes along the margins, with numerous, pale, dark-based, short to medium simple eglandular hairs and a few, short, dark glandular hairs almost completely obscured by the simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 3.0–3.5 mm, blackish. Flowers 5–7. Apomictic. Native. Ledges and rocks on old red sandstone. Endemic to Craig Cerrig-gleisiad in the Brecon Beacons.

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331. H. oistophyllum Pugsley Sagittate-leaved Hawkweed H. murorum var. sagittatum Lindeb.; H. sagittatum (Lindeb.) Dahlst., non Hoffmans. & Link; H. silvaticum subsp. sagittatum (Lindeb.) Stenstr¨om; H. sagittatum var. philanthrax auct.; H. philanthrax auct. Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, sometimes suffused reddish towards the base, usually slender, but robust in large plants, flexuous, striate, with numerous, medium to long, pale simple eglandular hairs below and sometimes throughout, but often getting shorter and fewer upwards, and dense stellate and few to numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler and sometimes tinted reddish beneath; basal few to numerous, the outer with lamina 2.5–8.0 × 1–5 cm, broadly elliptical, broadly ovate or subrotund, broadly rounded-mucronulate at apex, denticulate or shallowly dentate in the lower two-thirds and subtruncate at base, the inner with lamina 4–10 × 2–5 cm, ovate, elliptic-ovate or oblong-ovate, usually more or less acute at apex, denticulate to shallowly dentate, the teeth shallowly mammiform with a prominent apiculus and very characteristic, and subtruncate at base, the 2 basal teeth often reflexed, the petioles up to 7 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 0–1(−2), when present ovate, acute or acuminate, denticulate to shallowly dentate, shortly petiolate; all with numerous, short or medium, rather stiff simple eglandular hairs on the upper surface and more numerous, longer, softer hairs beneath and on the margins, the cauline also with stellate hairs beneath. Inflorescence with 2–20 capitula, broadly paniculate-corymbose; peduncles short and usually curved, with dense stellate hairs, numerous, short and medium, pale simple eglandular hairs and few, short, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3.0–8.5 × 1.0–1.2 mm, olivegreen, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short and medium, pale simple eglandular hairs, a few, short, dark glandular hairs, and numerous stellate hairs along the margins. Ligules yellow, glabrous or with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins obscurely dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Grassy slopes, limestone pavement, cliffs and streamsides especially on limestone. Fairly common in northern England with thinly scattered records in Scotland from Dumfries-shire to Ross-shire. Scandinavia and central Europe. 332. H. lintonii Ley Linton’s Hawkweed H. sagittatum var. maculigerum W. R. Linton; H. sagittatum subsp. loennrothianum var. maculigerum (W. R. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 25–60 cm, pale yellowish-green, often brownish-purple towards the base, slender to robust, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards and often deciduous except for the

bases, and few to fairly numerous stellate hairs in the upper part. Leaves deep green, often speckled with brownishpurple on upper surface, paler and often suffused purple beneath; basal few, the lamina 3–12 × 1.5–5.5 cm, ovate, ovate-lanceolate or broadly elliptical, obtuse-mucronate to acute at apex, irregularly dentate with small to large teeth particularly in the lower half, the teeth narrowly mammiform, abruptly contracted or cuneate at base, the teeth sometimes occurring on the petioles, the petioles up to 7 cm, often purplish, with numerous, long, pale simple eglandular hairs; cauline 0–1(−2), the lamina ovate, acute at apex, dentate, sometimes deeply so with narrow teeth, truncate or cuneate at base and petiolate; all with numerous, rigid, pale, short to medium simple eglandular hairs on both surfaces and the margins. Inflorescence with 2–8 capitula, paniculatecorymbose; peduncles rather short and often curved, with numerous stellate, numerous short to medium, pale, darkbased simple eglandular hairs and numerous short, dark glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all rather broad linear-lanceolate, the outer obtuse, the inner acute at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, numerous, short, dark glandular hairs, and some stellate along the margin towards the base. Ligules golden yellow, glabrous or with some very short simple eglandular hairs at apex. Styles yellowish. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. This species is very closely allied to H. rubiginosum of the Section Vulgata, but usually has 0–1 cauline leaf, brighter green spotted leaves and more glandular involucral bracts. Native. Cliff ledges and scree. Frequent on the limestone of Breconshire, Carmarthenshire and Yorkshire. Endemic. Named after William Richardson Linton (1850–1908). 333. H. orithales E. F. Linton Short-stemmed Hawkweed H. lintonianum Druce nom. illegit. Phyllopodous perennial herb with a branched stock. Stem 25–45 cm, pale yellowish-green, sometimes reddish below, slender but rigid, flexuous, striate, with numerous, long, wavy, pale simple eglandular hairs below, becoming shorter and fewer upwards, and numerous stellate and few, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath, sometimes flushed reddish; basal usually few, sometimes 1–2 outer, with lamina 2–6 × 2–4 cm, broadly ovate, roundedmucronate at apex, shallowly dentate and truncate at base, the inner, or sometimes all similar, with the lamina 5– 7 × 2.5–4.0 cm, ovate to broadly ovate, more or less acute at apex, denticulate or shallowly dentate and more or less truncate at base, the petioles up to 5 cm, with dense, long pale simple eglandular hairs; cauline usually 1, ovate, acute or acuminate at apex and dentate towards the subtruncate base; all with short to medium, pale simple eglandular hairs on upper surface to nearly glabrous, with numerous, medium to long, pale simple eglandular hairs beneath, especially on the midrib, and on the margin. Inflorescence with 2–5

39. Hieracium capitula, paniculate-corymbose; peduncles often fairly long and slender, with dense stellate hairs, numerous, short or medium, pale, dark-based simple eglandular hairs and few to fairly numerous, short, dark glandular hairs. Capitula 25–35 mm in diameter, rounded to subtruncate at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linearlanceolate, gradually narrowed to an obtuse apex, with dense, short to medium, pale, dark-based simple eglandular hairs, few, dark, short glandular hairs, and stellate hairs along the margin and towards the base. Ligules yellow, glabrous or with a few, very short simple eglandular hairs at tip. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 4.0–4.5 mm, reddish-black. Flowers 6–7. Apomictic. This species is closely allied to H. oistophyllum, but is distinguished by its fewer, obviously larger heads and larger achenes. Native. Cliffs, rocky slopes and streamsides. In a few localities in Perthshire, Aberdeenshire, Inverness-shire and Argyllshire. Endemic. 334. H. maculoides P. D. Sell & C. West Small-spotted Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, yellowish-green, sometimes purplish at base, glabrous or with a few, short to long, pale simple eglandular hairs, particularly below, and numerous stellate hairs in the upper part. Leaves bright medium yellowish-green on the upper surface with numerous brownish-purple spots and blotches, paler beneath and often purple tinted; basal numerous, the lamina 2.5–7.0 × 1–3 cm, ovate, lanceolate, elliptical or broadly elliptical, obtuse or acute at apex, denticulate or dentate, the teeth often apiculate and sometimes very narrow, cuneate at base, shortly petiolate; cauline 0–1(−3), similar to basal but the lamina of the uppermost linear or filiform; all on both surfaces and the margins with few to numerous, short to medium simple eglandular hairs. Inflorescence with 1–8 capitula, paniculate-corymbose; peduncles short and slender, with dense stellate hairs, few to numerous, pale, sometimes dark-based, short or medium simple eglandular hairs and few, very short or short, dark glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0– 1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with scattered stellate hairs, numerous, medium or long, pale, darkbased simple eglandular hairs and rarely a few, very short, dark glandular hairs. Ligules yellow, with a few, very short hairs at apex. Styles dark. Receptacle pits with margins long-dentate. Achenes about 4 mm, dark. Flowers 5–6. Apomictic. Native. Cliff ledges and scree. Scattered over the Yorkshire limestone and at Arnside Knott in Westmorland. Endemic. 335. H. uisticola Pugsley Uist Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, purplish towards the base, slender to robust, striate, with numerous, long, pale sim-

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ple eglandular hairs below, becoming fewer and shorter or absent upwards and some stellate hairs in the upper part. Leaves bright green on upper surface, paler beneath and often tinted reddish; basal numerous, the lamina 2–8 × 1– 4 cm, ovate, elliptical or subrotund, obtuse-mucronate to acute at apex, denticulate to sinuate-serrate or sinuatedentate and abruptly contracted to subtruncate at base, the petioles up to 7 cm and with numerous, long, pale, wavy simple eglandular hairs; cauline 0–2, when present lower like basal and petiolate, the upper linear-lanceolate, acute at apex, dentate and sessile, or bract-like; all glabrous or nearly so on upper surface, with few to fairly numerous, short to medium, pale simple eglandular hairs beneath and more numerous ones on the margin and midrib. Inflorescence with 2–7 capitula, paniculate-corymbose, often with a long lower branch; peduncles fairly long and slender, with dense stellate hairs, numerous, short and medium, pale simple eglandular hairs and a few, very short, dark glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.2– 1.5 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with dense, short to medium, pale, dark-based simple eglandular hairs, few, very short, dark glandular hairs and a few stellate hairs towards the base. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and rocky streamsides. Described from North Uist and Lewis in the Outer Hebrides and also occurring in a few localities in Perthshire and Argyllshire. Endemic. 336. H. sinuolatum P. D. Sell Sinuate-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, often brownish-purple in the lower half, slender to fairly robust, with numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and a few, short, dark glandular hairs in the upper part. Leaves medium yellowish-green and speckled with brownish-purple on the upper surface, pale and often suffused purple beneath; basal numerous, with the lamina 2–11 × 1.5–4.5 cm, subrotund, broadly elliptical, oblong-elliptical, elliptical or ovate, rounded-obtuse to acute at apex, regularly sinuate-dentate in the lower two-thirds with short, apiculate teeth, sometimes with the lowest teeth long, narrow and curved, sometimes some leaves with the margins only denticulate, rounded or subtruncate at base, the petioles up to 5 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1, even in large plants often absent, when present usually with the lamina linear and bract-like; all with short to medium, rather stiff, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 2–11 capitula, paniculatecorymbose; peduncles short and straight, with dense stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and numerous, short, fine, dark glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all narrowly

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linear-lanceolate, gradually narrowed at apex to an obtuse to acute tip, with numerous, short to medium, pale, dark-based simple eglandular hairs, numerous, short, dark glandular hairs, and numerous stellate hairs along the margins, particularly towards the base. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5– 7. Apomictic. Differs from H. lintonii, with which it grows, by its regularly sinuate-dentate basal leaves and lack of a large cauline leaf. ?Native. Grassy banks of old disused railway at Threshfield and at Linton in Yorkshire. Endemic. Despite its close relationship to H. lintonii this species could be introduced. A single specimen from rocks above Byre Bank Wood, near Arncliffe, which seems to be the same species may confirm that it is native to the area. 337. H. silvaticoides Pugsley Wood Hawkweed H. silvaticum auct.; H. scandinaviorum auct.; H. murorum var. pachyllum auct.; H. praetenerum auct.; H. murorum var. phaeotrichum auct. Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, sometimes purplish below, slender to more or less robust, striate, with few to fairly numerous, short to medium, pale simple eglandular hairs and numerous stellate hairs. Leaves deep green on upper surface, pale and often suffused purple beneath; basal usually few, the outer with lamina 3–8 × 2.0–5.5 cm, ovate, broadly elliptical or subrotund, rounded-mucronulate at apex, entire to shallowly dentate in the lower half and often with 2 large teeth at the subtruncate or subcordate base, the inner with lamina 5–10 × 2.5–5.5 cm, ovate or broadly ovate, usually more or less acute at apex, entire to shallowly dentate in lower half with 2 large teeth at the subcordate to subtruncate base, the petioles medium to long, with numerous, long, pale simple eglandular hairs; cauline 0–1, ovate, acute at apex, entire to dentate and petiolate; all glabrous or with a few simple hairs on upper surface and numerous, short to long simple eglandular hairs on the lower surface. Inflorescence with 2–8 capitula, cymosecorymbose; peduncles usually curved, with numerous to dense stellate hairs, few to numerous, short, dark glandular hairs and very rarely an occasional simple eglandular hair. Capitula 40–50 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–12 × 1.0–1.2 mm, olive-green, at least the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, short to medium, darkbased simple eglandular hairs, a few, short, dark glandular hairs, and numerous stellate hairs particularly towards the base and on the margins. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins slightly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Rocky streamsides and grassy banks. Fairly frequent on the limestone in the north of England and solitary localities at Fealar in Perthshire, Bridge of Avon near Tomintoul in Banffshire, Torr Head in Co. Antrim and Benevenagh in Co. Londonderry Endemic.

338. H. breadalbanense F. Hanb. Breadalbane Hawkweed H. pictorum var. breadalbanense (F. Hanb.) F. N. Williams; H. sagittatum subsp. breadalbanense (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–55 cm, pale yellowish-green, sometimes purplish below, more or less robust, striate, with few to fairly numerous, long, pale simple eglandular hairs below and shorter ones higher up, and scattered stellate hairs above. Leaves medium yellowish-green on upper surface, paler beneath and sometimes tinted purplish; basal usually numerous, the outer with lamina 3–7 × 2.0–4.5 cm, broadly elliptical to subrotund, rounded at apex, entire or remotely denticulate and rounded or abruptly contracted at base, the inner with lamina 5– 11 × 2.5–4.5 cm, elliptical, broadly elliptical or oblongelliptical, narrowed to an obtuse apex or acute at apex, entire to very shallowly sinuate-denticulate and abruptly contracted or rounded at base, the petioles up to 6 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1(−2), when present sometimes large like inner basal, usually lanceolate, acute or acuminate at apex, entire or denticulate and petiolate; all with rather stiff, pale, medium simple eglandular hairs on the upper surface and numerous longer ones beneath, particularly on the midrib, and on the margins. Inflorescence with 2–10 capitula, paniculatecorymbose, sometimes with a long, lower branch; peduncles usually rather short, straight or curved, with dense stellate hairs and numerous, short and medium, dark-based simple eglandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts more or less porrect in bud, 4– 11 × 1.0–1.2 mm, dark green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous to dense, short to medium, dark-based simple eglandular hairs, and numerous stellate hairs on the margins, glandular hairs not obvious but a few can usually be found hidden among the simple eglandular hairs. Ligules yellow, glabrous or with occasional very short hairs at apex. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and rocky streamsides. Breadalbane Hills of Perthshire and a few localities in Aberdeenshire, Banffshire, Inverness-shire and Argyllshire. Endemic. 339. H. auratiflorum Pugsley Dark-flowered Hawkweed H. crebridens auct. Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, sometimes slightly purplish towards the base, slender to robust, striate, with fairly numerous, long, pale simple eglandular hairs below, becoming much sparser upwards and few to fairly numerous stellate hairs above. Leaves dark green on upper surface, paler beneath and sometimes slightly tinted reddish; basal usually few, the outer with lamina 2.5–10 × 2.5–8.0 cm, subrotund, broadly elliptical or broadly ovate, rounded mucronulate at apex, denticulate to sinuate-dentate and subtruncate to slightly cordate at base, the inner with lamina 6–14 × 4.0–8.5 cm, broadly ovate, acute to acuminate at apex,

39. Hieracium sinuate-dentate, the teeth broadly mammiform, mostly cordate at base and often with a down-turned tooth at each side, the petioles medium to rather long and with long, pale simple eglandular hairs; cauline often absent, occasionally 1 like inner basal but much smaller and petiolate; all with short to medium, pale simple eglandular hairs on both surfaces and the margins or nearly glabrous on upper surface. Inflorescence with 2–12 capitula, cymose-corymbose; peduncles spreading, with dense stellate hairs, few to numerous, short, dark glandular hairs and some, medium, pale simple eglandular hairs. Capitula 40–50 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–15 × 1.0–1.6 mm, greyish-green with paler margins, all long linear-lanceolate, obtuse at apex, with numerous, long, dark-based simple eglandular hairs, more or less numerous, short, dark glandular hairs and often dense stellate hairs on the margin and apex. Ligules golden yellow, with very short simple eglandular hairs at apex. Styles dull yellow to slightly discoloured. Receptacle pits with margins subulatedentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–7. Apomictic. Native. Limestone rocks and ledges and grassy banks. Yorkshire Dales and Teesdale. Endemic. 340. H. triangularifolium P. D. Sell Triangular-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 35–45 cm, pale yellowish-green, sometimes tinted purplish towards the base, with few to numerous, long, pale simple eglandular hairs below, becoming much shorter upwards, and with numerous stellate hairs in the upper part. Leaves pale green on upper surface, even paler and often tinted or suffused purplish beneath; basal few, the lamina 6–9 × 3.0– 5.5 cm, broadly triangular-ovate, rounded-obtuse to acute at apex, dentate or serrate-dentate, the teeth sharp and apiculate, truncate or subcordate at the base, the petioles long, up to 7 cm and with dense, long, wavy, pale simple eglandular hairs; cauline 0–1, when present lamina small, ovate, acute, sharply dentate and petiolate; all glabrous or with an occasional simple eglandular hair on upper surface, with scattered short to medium, pale simple eglandular hairs on the lower surface and more numerous ones on the margin. Inflorescence with 2–4 capitula, furcate-corymbose; peduncles long, with numerous stellate hairs, numerous, short to medium, pale, dark-based simple eglandular hairs and a few, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs, few to fairly numerous, short, dark glandular hairs, and numerous stellate hairs along the margin and near the apex. Ligules yellow, with few, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits and achenes not seen. Flowers 7–8. Apomictic. Native. A very distinct species known only by the River Falloch at the head of Glen Falloch, Perthshire, and from schistose cliff ledges on Meall Buidhe near the Bridge of Orchy, Beinn a’ Chaisteil (Ben Chaisteil) near Tyndrum and Ben Laoigh (Ben Lui), all in Argyllshire. Endemic.

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341. H. crebridens F. N. Williams Chapel-le-dale Hawkweed H. bifidum subsp. crebridens (F. N. Williams) Zahn; H. sagittatum var. lanuginosum auct. Phyllopodous perennial herb with a branched stock. Stem 18–55 cm, pale yellowish-green, often purplish towards the base, usually more or less slender, striate, with a few, long, pale simple eglandular hairs below and an occasional one upwards, few to numerous stellate hairs in the upper part and sometimes an occasional short, dark glandular hair. Leaves medium yellowish-green on upper surface, paler and sometimes suffused violet-purple beneath; basal usually numerous, the outer with lamina 3–8 × 2.5–4.5 cm, broadly elliptical to oblong-elliptical, rounded-obtuse at apex, regularly sinuate-denticulate or dentate and abruptly contracted or subtruncate at base, the inner with lamina 6–13 × 3–5 cm, elliptical, oblong-elliptical or ovate-lanceolate, acute at apex, regularly sinuate-dentate, the teeth mammiform, abruptly contracted or subtruncate at base, the petioles medium to long, pale simple eglandular hairs; cauline 0–1, often absent, when present the lamina linear or lanceolate, entire or dentate, sessile or petiolate; all glabrous or nearly so on the upper surface, with short to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 2–8 capitula, furcate-corymbose; peduncles suberect, with numerous to dense stellate hairs, fairly numerous short to medium, pale simple eglandular hairs and few, short, dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 0.7– 1.0 mm, blackish-green, the inner with paler margins, all narrow linear-lanceolate, obtuse at apex, with numerous, medium to long, dark-based simple eglandular hairs, fewer, short, dark glandular hairs, and dense stellate hairs especially on the margins. Ligules yellow, with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins sharply subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. H. Dahlstedt never validated the name H. crebridens, which he used on herbarium sheets. The first person to apply a description to the name was F. N. Williams (1902). Although Dahlstedt’s specimens were cited by Williams, the description does not fit them. Of the specimens cited the nearest to fitting the description is from Chapel-le-dale in Yorkshire. Many of the plants referred to H. sagittatum var. lanuginosum by the brothers Linton belong here. Native. Limestone cliffs, pavements and river banks, and roadside and railway embankments. Hawnby, Linton, Chapel-le-dale, Buckden, Ribbleshead, Clapham and High Force in Yorkshire and by the Tyne in Northumberland. Endemic. 342. H. crebridentiforme Pugsley Close-toothed Hawkweed H. crebridens auct.; H. bifidum subsp. crebidens auct.; H. ciliatum auct.; H. ciliatiflorum auct. Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often reddish towards the base, more or less robust, striate, with rather few, medium to long, pale simple eglandular hairs, and numerous stellate

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hairs and a few, short, dark glandular hairs in the upper part. Leaves bright green on upper surface, pale beneath and often suffused brownish-purple, the outer with lamina 3–8 × 1.5– 3.5 cm, ovate or elliptical, obtuse-mucronate to more or less acute at apex, entire to denticulate or dentate and abruptly contracted or subtruncate at base, the inner with lamina 6– 14 × 2–7 cm, elliptical, elliptical-oblong, ovate-oblong or ovate, short to long acute or acuminate at apex, denticulate to dentate with a few, large mammiform teeth and abruptly contracted or subtruncate at base, the petioles medium to rather long, with numerous, long, pale simple eglandular hairs; cauline 0–1, with lamina linear and entire or ovatelanceolate, long-acute at apex, dentate and with a short petiole; all with numerous, short to medium, rather stiff, pale simple eglandular hairs on both surfaces and the margins or becoming almost glabrous on the upper surface. Inflorescence with 2–10 capitula, cymose-corymbose but often furcately branched; peduncles often rather long and suberect, with more or less dense stellate hairs, numerous, short, dark glandular hairs and few to fairly numerous, medium, pale simple eglandular hairs. Capitula 30–40 mm in diameter, narrowed and scarcely rounded at base. Involucral bracts porrect in bud, 4–12 × 1.0–1.5 mm, dark green with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, short, dark glandular hairs, numerous, short to medium, dark-based simple eglandular hairs, and some stellate hairs mainly towards the base and on the margins. Ligules yellow, with few to numerous, very short simple eglandular hairs at apex. Styles more or less discoloured. Receptacle pits with margins sharply subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and scree. Common on the Yorkshire limestone where it often grows with H. crebridens. Endemic. 343. H. pseudosarcophyllum Pugsley Dumfries Hawkweed H. sarcophyllum auct. Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, often purplish towards the base, slender, flexuous, striate, with a few, long simple eglandular hairs near the base and some stellate hairs above to nearly glabrous. Leaves glaucous green on upper surface, sometimes blotched brownish-purple, paler beneath; basal numerous, the outer with lamina 2–6 × 1.5–3.0 cm, elliptical or subrotund, rounded-mucronulate at apex, entire, denticulate or shallowly dentate, the teeth mammiform, and abruptly contracted or rounded at base, the inner with lamina 3–9 × 1.3–3.0 cm, oblong-elliptical or narrowly elliptical, more or less acute at apex, denticulate to shallowly dentate, the basal teeth narrowly mammiform and often curved, and cuneate or subtruncate at base, the petioles up to 7 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1, when present with lamina lanceolate, acute at apex, entire to sharply dentate, the lower teeth often mammiform, cuneate to subtruncate at base and petiolate; all glabrous or nearly so on upper surface, with scattered short and medium simple eglandular hairs beneath and on the margin and with numerous stellate hairs on the underside of the cauline. Inflorescence with 2–8 capitula, rather

compactly paniculate-corymbose; peduncles slender and curved, with dense stellate hairs, numerous very short and short (0.2–0.5 mm), dark glandular hairs and an occasional, short to medium simple eglandular hair. Capitula 25– 30 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–12 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, more or less acute at apex, with numerous, very short to short (0.2– 0.5 mm), dark glandular hairs and few to fairly numerous, short to medium, dark simple eglandular hairs, and numerous stellate hairs on the margin. Ligules yellow, glabroustipped. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and rocky sides of streams. Moffat Hills of Dumfries-shire, about Talla, Peebles-shire and Culter Fell, Lanarkshire. Endemic. 344. H. cymbifolium Purchas Boat-leaved Hawkweed H. bifidum subsp. cymbifolium (Purchas) Zahn Phyllopodous perennial herb with a branched stock. Stem 25–55 cm, pale yellowish-green, usually purple-tinted below, slender to robust, often flexuous, striate, with few to fairly numerous, medium to long, pale simple eglandular hairs below, becoming shorter upwards and with numerous stellate hairs and an occasional short glandular hair in the upper part. Leaves thick, dull medium green on upper surface, paler beneath; basal usually numerous, often laterally incurved and boat-shaped, the outer with lamina 3–10 × 1–4 cm, elliptical, oblong-elliptical, oblong-ovate or oblong, rounded-mucronulate at apex, subentire, denticulate or shallowly sinuate-dentate, the teeth mammiform or narrowly mammiform, and more or less truncate at base, the inner with lamina 5–10 × 2–4 cm, ovate or oblong-ovate, more or less acute at apex, subentire, denticulate or shallowly sinuate-dentate, the lower teeth sometimes large and deflexed at the more or less truncate base, the petioles up to 12 cm, with numerous, long, wavy, pale simple eglandular hairs; cauline 0–1, when present with lamina ovate, acute at apex, usually entire, sometimes with shallow teeth, subtruncate at base and petiolate; all with few, short, pale simple eglandular hairs on upper surface, and more numerous, longer ones beneath and on the margins, and with stellate hairs on both surfaces, often densely so beneath. Inflorescence with 3–14 capitula, paniculate-corymbose, often with a long lower branch; peduncles rather long and slender, spreading and curved, with dense stellate hairs, numerous very short to short (0.2–0.5 mm), dark glandular hairs and a few, medium, dark-based simple eglandular hairs. Capitula 30–35 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, dark olivegreen, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, unequal, short and very short (0.2–0.7 mm), dark glandular hairs, numerous short to medium, dark-based simple eglandular hairs, and numerous stellate hairs along the margins and at the base. Ligules yellow, with a few, very short, pale simple eglandular hairs at the tips. Styles yellow to slightly discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic.

39. Hieracium Native. Cliff ledges, rocks and streamsides on limestone and other basic rocks. Common in the Derbyshire and Yorkshire Dales and extending into Staffordshire and Westmorland; also on limestone outcrops in Lanarkshire, Inverness-shire, Argyllshire and the Isle of Raasay in the North Ebudes. Endemic. 345. H. pictorum E. F. Linton Picts Hawkweed H. murorum subsp. pictorum (E. F. Linton) Zahn; H. semicrassiceps Pugsley; H. murorum subsp. crassiceps auct.; H. serratifrons var. crassiceps auct. Phyllopodous perennial herb with a branched stock. Stems 20–50 cm, pale yellowish-green, sometimes purplish towards the base, slender to fairly robust, striate, with few to numerous, short to long, pale simple eglandular hairs especially below and few to numerous stellate and few to numerous, very short to short (0.2–0.5 mm), dark glandular hairs in the upper part. Leaves bright medium green on the upper surface, paler and sometimes flushed purplish beneath; basal few to fairly numerous, with lamina 2–10 × 1.5–4.5 cm, the outer elliptical or oblong-elliptical, rounded-mucronulate at apex, denticulate to shallowly dentate and rounded or subtruncate at base, the inner with lamina lanceolate, ovate, elliptical or oblong-lanceolate, more or less acute at apex, dentate and sometimes incise-dentate towards the base, the teeth often narrowly aquiline-mammiform, and sometimes an isolated one on the petiole which is short to rather long, often purplish and with long, wavy, pale simple eglandular hairs; cauline usually absent, when present the lamina lanceolate or ovate, acute at apex, dentate and petiolate, or bractlike; all usually glabrous on the upper surface, with few to numerous, short to long, pale simple eglandular hairs on the lower surface and margin and especially on the midrib, sometimes also a few on the upper surface near the margin. Inflorescence with 2–6(–15) capitula, cymose-corymbose, with a rather short acladium and sometimes with a long lower branch; peduncles arcuate, with numerous to dense stellate hairs, few, scattered, very short to short (0.1–0.5 mm), dark glandular hairs and sometimes an occasional, short simple eglandular hair. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts more or less porrect in bud, 4–13 × 1.0–1.4 mm, blackish-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an obtuse apex, mostly with few stellate hairs but sometimes with them more numerous along the margins and at the apex, numerous, very short to short (0.2–0.7 mm), dark glandular hairs and few to numerous, short to medium, pale, dark-based simple eglandular hairs. Ligules glabrous or slightly hairy at tip. Styles yellowish or slightly livid, darkening when dry. Receptacle pits with margins incisedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Although a variable species H. pictorum is usually recognisable by its ovate, lanceolate or ovate-oblong, mainly acute leaves which are dentate towards the base and glabrous or nearly so on the upper surface, an inflorescence of few to fairly numerous, rather small capitula, which have narrow, but more or less obtuse involucral bracts clothed with numerous glandular and few to numerous simple eglandular hairs. Stellate hairs on the involucral bracts

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are usually few, but some plants from Clova, which were named H. semicrassiceps, have them more numerous. The ligules are usually glabrous tipped but can be more or less hairy at tip. The only confusion could be in Clova where H. pruinale is similar in appearance, but the upper surface of its leaves is hairy and the involucral bracts have the hairs all glandular or with a very occasional simple eglandular hair. The type specimens of H. pictorum, H. semicrassiceps and H. pruinale all come from Clova. Native. Rocks, cliff ledges and streamsides mostly between 550 and 950 m. All specimen labels which record the type of rock have put granite, with one exception which gives sandstone. A common species of the central Scottish Highlands extending in the north-west to the coast of Sutherland and south to Peebles-shire and Dumfries-shire. Endemic. 346. H. elongatifolium P. D. Sell Elongate-leaved Hawkweed Phyllopodous perennial herb with a branched stock. Stem 35–50 cm, pale yellowish-green, sometimes reddish-tinted below, rather slender and flexuous, striate, with few, medium to long, pale simple eglandular hairs, few stellate hairs and few, short, dark glandular hairs in the upper part. Leaves medium green on upper surface with the veins sometimes paler, paler and slightly glaucous beneath; basal usually numerous, the outer with lamina 2–9 × 2–4 cm, elliptical, oblong-elliptical or oblong, rounded-mucronulate at apex, subentire, denticulate or dentate in the lower half, with the lower teeth sometimes mammiform and curved, and abruptly contracted or subtruncate at base, the inner with lamina 8–14 × 3.0–5.5 cm, oblong-lanceolate to oblong, acute at apex, denticulate or shallowly dentate in upper half, dentate in lower part, with the teeth bluntly to acutely mammiform, usually narrow and often curved, abruptly contracted or subtruncate, sometimes asymmetrically, at base, sometimes with very narrow teeth extending on to the petiole, and the petioles up to 8 cm, with numerous, long, pale simple eglandular hairs; cauline absent, or with the lamina linear and bract-like; all glabrous or nearly so on upper surface, but with few, medium to long, pale simple eglandular hairs scattered over the lower surface and more numerous ones on the midrib and margin. Inflorescence with 2–9 capitula, rather openly paniculate-corymbose; peduncles rather long and curved, slender, with dense stellate hairs, numerous, very short to short (0.2–0.5 mm), dark glandular hairs, and occasional short to medium, pale simple eglandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–11 × 1.0–1.2 mm, dull greyish-green, the inner with paler margins, all linearlanceolate, narrowly acute at apex, with numerous, short and medium, pale, dark-based simple eglandular hairs intermixed with very short to short (0.2–0.5 mm), dark glandular hairs, and numerous stellate hairs along the margins and at the apex. Ligules yellow, sometimes poorly developed, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Closely allied to H. pictorum, from which it differs in its larger, more oblong leaves and acute involucral bracts.

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Native. Streamside rocks and low cliffs. Perthshire, Aberdeenshire, Banffshire, Inverness-shire, Argyllshire, Ross-shire and Sutherland. Endemic. 347. H. mariae P. D. Sell McCallum Webster’s Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, sometimes tinted purplish below, slender to more or less robust, striate, with fairly numerous, long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and a few, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal few to numerous, the outer with lamina 2.5– 9.0 × 1.5–4.5 cm, broadly elliptical, nearly subrotund or obovate, rounded-mucronulate at apex, subentire to sinuatedenticulate, and rounded or subtruncate at base, the inner with lamina 5–11 × 3.0–4.0 cm, narrowly to broadly elliptical, elliptic-oblong or oblong, more or less acute at apex, denticulate or denticulate with a few shallow teeth towards the base which is rounded or subtruncate, the petioles up to 8 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, when present with lamina linearlanceolate, long-acute, denticulate, cuneate at base and petiolate, or bract-like; all with numerous, short to medium, pale simple eglandular hairs on the upper surface or sometimes nearly glabrous and with numerous, medium to long, pale simple eglandular hairs beneath, particularly on the midrib, and on the margin. Inflorescence with 4–numerous capitula, paniculate-corymbose; peduncles variable in length and slender, with dense stellate hairs, numerous very short to medium, dark glandular hairs and few, short to medium, pale, dark-based simple eglandular hairs. Capitula 25– 35 mm in diameter, narrowed to a rounded base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, dark olivegreen, the inner with paler margins, all linear-lanceolate, obtuse at apex, with numerous very short and short, dark glandular hairs, numerous, short and medium, pale, darkbased simple eglandular hairs, and some stellate hairs at the apex and towards the base. Ligules yellow, sometimes with a few, very short hairs at the apex. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 3.0– 3.5 mm, blackish-red. Flowers 6–7. Apomictic. Native. Sandstone cliffs north of Rosemarkie and by the Eathie Burn, both in the Black Isle, above Glassnock, Kishorn, and near Loch Duich all in Ross-shire and Creag na h’ Uidhe in Sutherland. Endemic. Named after Mary McCallum Webster (1906–85) for her great contribution to Scottish botany. 348. H. rivale F. Hanb. Burnside Hawkweed H. caniceps F. Hanb., non Norrlin; H. silvaticum var. rivale (F. Hanb.) F. N. Williams; H. sagittatum subsp. rivale (F. Hanb.) Zahn; H. ciliatum auct.; H. ciliatiflorum auct.; H. praecox subsp. ciliatum auct. Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, often purplish towards the base, more or less robust, striate, with scattered, long, pale simple eglandular hairs towards the base and dense stellate hairs and numerous, short, dark glandular hairs in the upper

part. Leaves bright green on upper surface, paler and slightly bluish beneath; basal usually numerous, the outer with lamina 2.5–9.0 × 1.5–4.0 cm, ovate or broadly elliptical, rounded-mucronulate at apex, regularly denticulate or very shallowly dentate and rounded or subtruncate at base, the inner with lamina 5–9.0 × 2–4 cm, mostly elliptical, sometimes ovate, more or less acute at apex, regularly and shallowly sinuate-denticulate or sinuate-dentate and abruptly contracted at base, the petioles up to 8 cm, with dense, long, wavy simple eglandular hairs; cauline 0–1, ovate, acute at apex, shallowly dentate, petiolate; all with few to numerous, short to medium, pale simple eglandular hairs on both surfaces and the margins and some longer ones on the midrib beneath. Inflorescence with 3–10 capitula, paniculatecorymbose, sometimes with a long lower branch; peduncles straight or arcuate, with dense stellate hairs, numerous, short or very short (0.2–0.5 mm), dark glandular hairs and an occasional, short or medium, dark-based simple eglandular hair. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts 5–12 × 1.2–1.5 mm, greyish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, short (0.3–0.7 mm), dark glandular hairs, numerous, short to medium, dark-based simple eglandular hairs, and dense stellate hairs on the margins. Ligules yellow, with numerous, very short simple eglandular hairs at apex. Styles yellowish to slightly discoloured. Receptacle pits with margins incise-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges and rocky streamsides. Sutherland. Endemic. When first described as H. caniceps the citation included a mixture of specimens. The original description best fits the plants which grow commonly in Sutherland. This plant has also been commonly called H. ciliatum, i.e. H. ciliatiflorum. H. ciliatiflorum Pugsley is technically a new name for H. ciliatum Almq., non Willd., which is not a British plant. 349. H. sanguineum (Ley) W. R. Linton Bloody Hawkweed H. murorum var. sanguineum Ley; H. silvaticum var. sanguineum (Ley) F. N. Williams; H. murorum subsp. subsanguineum Zahn: H. subsanguineum (Zahn) Roffey Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, sometimes reddish below, robust, striate, glabrous or with a few, medium or long simple eglandular hairs below and numerous stellate, numerous short, dark glandular and a few, short to medium, dark-based simple eglandular hairs in the upper part. Leaves rather pale green on upper surface, paler and often suffused reddish beneath; basal usually numerous, the outer with lamina 4–7 × 2.0–4.5 cm, ovate, elliptical, oblong-elliptical or ovate-oblong, rounded-mucronulate at apex and entire, denticulate or with a few shallow teeth near the subtruncate base, the inner with lamina 6–12 × 2.5–5.0 cm, ovate, elliptical, oblong-elliptical or oblong, more or less acute at apex, denticulate or shallowly sinuate-dentate and with a few larger teeth near the subtruncate base, the petioles up to 8 cm, with dense, long, pale, wavy simple eglandular hairs; cauline usually absent, sometimes one, linear and bractlike, larger ones produced in cultivation; all glabrous or nearly

39. Hieracium so on the upper surface, with scattered, short to medium, pale simple eglandular hairs beneath and longer and more numerous ones on the midrib and margins. Inflorescence with 2–6 capitula, furcate-corymbose, sometimes with a long, lower branch; peduncles thick, straight to arcuate, with dense stellate, numerous, very short to short (0.2–0.7 mm), dark glandular hairs and few, short to medium, dark-based simple eglandular hairs. Capitula 30–40 mm in diameter, broad and subtruncate at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, dark green, the inner with paler margins, all linear-lanceolate, mostly obtuse at apex, with numerous, unequal, very short to medium (0.2–1.0 mm), dark glandular hairs, short to medium, dark-based simple eglandular hairs, and dense stellate hairs on the margins and at the apex. Ligules yellow, often with very short simple eglandular hairs at apex. Styles yellowish to discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 5–7. Apomictic. Native. Sandstone cliff ledges. Dyffryn Crawnon, Breconshire. Endemic. 350. H. boreoanglicum P. D. Sell North England Hawkweed H. pseudostenstroemii Pugsley (1948), non Pugsley (1941); H. serratifrons var. stenstroemii auct.; H. serratifrons subsp. stenstroemii auct. Phyllopodous perennial herb with a branched stock. Stem 30–55 cm, pale yellowish-green, sometimes tinted purple at the base, slender to robust, striate, with few, long, pale simple eglandular hairs below, becoming more numerous, but shorter above, and numerous stellate hairs and an occasional short, dark glandular hair in the upper part. Leaves dull caesious-green on upper surface, paler beneath and often tinted reddish, rather thick; basal few to numerous, the outer with lamina 2.5–9.0 × 2.0–3.5 cm, ovate, elliptical or oblong-elliptical, rounded-mucronulate at apex, subentire, denticulate or dentate, with the teeth shallowly mammiform, and subtruncate at base, the inner with lamina 7–12 × 2.5–4.0 cm, mostly ovate-oblong or oblong, sometimes lanceolate, acute to shortly caudate at apex, dentate, often with large mammiform or aquiline-mammiform teeth below which are occasionally decurrent on the petiole, and subtruncate at base, the petioles up to 12 cm, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1, when present with lamina lanceolate, acute at apex, dentate, petiolate; all with scattered, short or medium, pale simple eglandular hairs on the upper surface or nearly glabrous, with more numerous simple eglandular hairs beneath and on the margins and often with numerous stellate hairs beneath. Inflorescence with 3–10 capitula, furcate-corymbose, sometimes with a long lower branch; peduncles rather thick, straight or curved, with dense stellate hairs, numerous, very short or short (0.2–0.5 mm), dark glandular hairs and numerous, short or medium, dark-based simple eglandular hairs. Capitula 30–35 mm in diameter, rounded to subtruncate at base. Involucral bracts porrect in bud, 5–12 × 0.8–1.0 mm, dark green, the inner with paler margins, all lanceolate-oblong, more or less acute at apex, with numerous, very unequal, very short to medium (0.2–1.2 mm), dark glandular hairs, few to numerous, short to medium, pale,

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dark-based simple eglandular hairs, and numerous stellate hairs on the margins. Ligules yellow, with numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins sharply dentate. Achenes 3.5– 4.0 mm, reddish-black. Flowers 6–7. Apomictic. When H. W. Pugsley first described H. pseudostenstoemii in 1941 he did not give it a Latin description and a type. He did, however, refer to a Latin description of H. silvaticum var. stenstroemii of F. N. Williams which validated it. Unfortunately Williams’s description is of the true H. stenstroemii and so H. pseudostenstroemii becomes a synonym of it. Pugsley’s 1948 description of H. pseudostenstroemii is of H. boreoanglicum. Native. Cliff ledges, rocky slopes, grykes and streamsides. Common on the Yorkshire limestone and on the Durham border at Winch Bridge. A specimen from the North Ebudes on the Isle of Raasay seems indistinguishable. Endemic. 351. H. pachyphylloides (Zahn) Roffey Carboniferous Hawkweed H. murorum subsp. pachyphylloides Zahn; H. murorum var. pachyphyllum Purchas; H. pachyphyllum (Purchas) F. N. Williams, non Brenner Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often flushed purple below, slender to robust, striate, with few, long, pale simple eglandular hairs below and numerous stellate, few to numerous, short, dark glandular and few, short to medium, dark-based simple eglandular hairs in the upper part. Leaves thick, bluish-green on upper surface, paler and often suffused purplish beneath; basal usually numerous, the outer with lamina 2.5–7.0 × 1.5–5.0 cm, subrotund, elliptical or ovate, rounded-mucronulate at apex and denticulate or with larger teeth near the subtruncate base, the inner with lamina 6–9 × 3.0–4.5 cm, mostly ovate or slightly oblong-ovate, roundedmucronulate to subacute or slightly cuspidate at apex, denticulate to dentate, the lower teeth often mammiform and sometimes reflexed at the subtruncate or subcordate base, sometimes the outer and inner leaves hardly different, the petioles up to 9 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, sometimes large, ovate, cuspidate at apex, dentate, subtruncate at base and petiolate; all glabrous or nearly so on the upper surface, with numerous, short to long, pale simple eglandular hairs below, especially on the midrib, and on the margins. Inflorescence with 2–12 capitula, furcate-corymbose, sometimes with a long, lower branch; peduncles long and slender, straight, with dense stellate hairs, numerous, very short to short (0.2–0.5 mm), dark glandular hairs and an occasional short, dark-based simple eglandular hair. Capitula 30–38 mm in diameter, broad and subtruncate based. Involucral bracts incumbent in bud, 5–12 × 1.0–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to a long, thin, but obtuse apex, with numerous short and medium, dark-based simple eglandular hairs, fewer, very short or short, dark glandular hairs, and dense stellate hairs on the margins. Ligules deep yellow, glabroustipped. Styles yellowish to slightly discoloured. Receptacle

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pits with margins dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 5–7. Apomictic. Native. Carboniferous limestone cliffs and rubble on both sides of the Wye Valley between Ross and Monmouth. Endemic. 352. H. pauculidens P. D. Sell & C. West Skerray Hawkweed H. serratifrons var. caliginosum auct. Phyllopodous perennial herb with a branched stock. Stem 20–55 cm, pale yellowish-green, sometimes slightly purplish below, markedly striate, with few to numerous stellate hairs particularly in the upper part, with few, short or medium, pale simple eglandular hairs especially below and a few, short, dark glandular hairs above. Leaves dull medium green on upper surface, paler beneath and often tinted purplish, with more or less numerous, short, pale simple eglandular hairs on both surfaces and long ones on the margin and midrib beneath; basal more or less numerous, 1–9 × 1–4 cm, the outer with lamina subrotund, elliptical or ovate, rounded-mucronulate at apex, subentire to denticulate and rounded, subtruncate or subcordate at base, the inner with lamina elliptical, ovate or lanceolate, more or less acute at apex, denticulate or sometimes with 1–2 mammiform teeth near the base and subtruncate, often unequally at the base, with rather short petioles which have numerous to dense, medium to long, pale simple eglandular hairs; cauline 0–1, when present with lamina lanceolate, longacute at apex, subentire, cuneate at base and petiolate. Inflorescence with 2–8 capitula, compactly cymose with a short acladium; peduncles short or medium, more or less arcuate, with numerous stellate hairs, numerous, short (0.2–0.5 mm), dark glandular hairs and fewer, short to medium, pale, dark-based simple eglandular hairs. Capitula 30–45 mm in diameter, rounded at base. Involucral bracts more or less incumbent in bud, 3.0–10.5 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse to acute at apex, with scattered stellate hairs at the base and on the margins, numerous short and very short (0.2– 0.7 mm), dark glandular hairs and numerous, short and medium, pale, dark-based simple eglandular hairs. Ligules pale yellow, glabrous or slightly hairy at tip. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0– 3.5 mm, blackish. Flowers 6–7. Apomictic. Native. Abundant on grassy slopes by the sea at Skerray and rocks above the Tongue road at Coldbackie, Sutherland; also on the rocky slope of An Teallach, Ross-shire. Endemic. 353. H. varifolium P. D. Sell & C. West Variable-leaved Hawkweed H. silvaticum var. asymmetricum auct.; H. sarcophyllum var. expallidiforme auct. Phyllopodous perennial herb with a branched stock. Stems 20–45 cm, pale yellowish-green, often tinted purplish towards the base, markedly striate, with few to numerous stellate hairs especially in the upper part, few, medium to long, pale simple eglandular hairs especially below and few, very short, dark glandular hairs in the upper part. Leaves medium green often blotched and spotted on the upper

surface, paler and often suffused purplish beneath; basal few to numerous, variable, 2–9 × 1–4 cm, the outer with lamina subrotund, ovate or elliptical, rounded-mucronulate to subacute at apex, dentate with shallow, mammiform teeth and cuneate to truncate at base, the inner with lamina lanceolate, ovate or oblong-ovate, acute and often caudate at apex, dentate, with mammiform or aquiline-mammiform teeth and sometimes with a reflexed pair at the subtruncate base, occasionally with 1–2 on the short to rather long petioles which are clothed with dense, long, pale simple eglandular hairs; cauline absent or rarely 1, with the lamina lanceolate, long-acute at apex, slightly dentate and petiolate; all more or less glabrous on the upper surface, with numerous, short to medium, pale simple eglandular hairs beneath and long ones on the midrib and margins. Inflorescence narrowly cymose, with short acladium; peduncles arcuate, with numerous to dense stellate hairs, numerous, short (0.2–0.5 mm), dark glandular hairs and numerous, short to medium, pale, dark-based simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–15 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, with a long-drawn-out, acute apex, the inner almost filamentous, with few stellate hairs mainly on the margins near the base, numerous very short or short (0.2–0.5 mm), dark glandular hairs and numerous, short or medium, dark or pale, dark-based simple eglandular hairs. Ligules deep yellow, puberulous at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, blackish. Flowers 5–8. Apomictic. Native. Sandy, dolomitic and other cliffs and rocks up to 1,800 m on Ben Hope, Coldbackie, Foinaven, Heilan Ferry and Ben Loyal in Sutherland. Endemic. 354. H. anguinum (W. R. Linton) Roffey Snake-headed Hawkweed H. stenolepis var. anguinum W. R. Linton; H. bifidum var. anguinum (W. R. Linton) F. N. Williams; H. bifidum subsp. anguinum (W. R. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stems 20–40 cm, pale yellowish-green, often tinted purplish towards the base, erect, slender, flexuous, striate, usually glabrous except for some minute stellate hairs in the uppermost part. Leaves medium green and sometimes with a few small spots on the upper surface, paler and often suffused purplish beneath; basal with lamina 2–9 × 1–4 cm, few to fairly numerous, subrotund, lanceolate, lanceolateoblong or ovate, the outer obtuse-mucronulate, the inner more or less acute at apex, subentire to denticulate, sometimes with a few larger teeth towards the base which are sometimes reflexed; the petioles often rather long, sometimes reddish, with few to numerous, medium to long, pale simple eglandular hairs; cauline usually absent, rarely 1 resembling the inner basal and petiolate, sometimes bractlike; all glabrous on upper surface, glabrous or with scattered to fairly numerous, short, pale simple eglandular hairs beneath, usually with short, pale simple eglandular hairs on the margin. Inflorescence with 2–6 capitula, cymosesubcorymbose, often with long acladium but sometimes geminate; peduncles rather long, with more or less dense

39. Hieracium stellate hairs, few, scattered, very short to short, dark glandular hairs and occasionally a short or medium, pale simple eglandular hair. Capitula 30–40 mm in diameter, narrowed to a rounded base. Involucral bracts porrect in bud, 4–14 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually drawn out to an acute apex, the inner almost filamentous, with few stellate hairs near the base, with numerous very short and short (0.2–0.5 mm), dark glandular hairs and numerous, short and medium, pale, dark-based simple eglandular hairs. Ligules glabrous-tipped. Styles yellow to slightly discoloured. Receptacle pits with margins more or less dentate. Achenes 3.0–3.5 mm, blackish. Flowers 6–7. Apomictic. Native. Rocky streamsides. By the Grey Mare’s Tail and elsewhere in the Moffat valley, Dumfries-shire and Talla in Peebles-shire. Probably endemic, but a single plant seen from Norway seems identical with it. 355. H. prolixum Norrl. Ben Hope Hawkweed H. silvaticum subsp. prolixum (Norrl.) Dahlst.; H. silvaticum var. prolixum (Norrl.) F. N. Williams; H. bifidum subsp. prolixum (Norrl.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–35 cm, pale yellowish-green, often tinted brownishpurple below, slender, flexuous, striate, with a few, scattered, medium to long, pale simple eglandular hairs below and numerous stellate hairs and occasional to few, short, dark glandular hairs above. Leaves thick, dull, medium or dark green on upper surface, sometimes with a few dark blotches, paler and slightly bluish-green beneath; basal few to fairly numerous, the outer with lamina 2–4 × 1.5– 3.0 cm, ovate, rounded-mucronulate at apex, serrate-dentate in the lower half and rounded or subtruncate at base, the inner with lamina 4–6 × 2–3 cm, lanceolate to ovate, subacute to narrowly and sharply acute at apex, dentate or serrate-dentate in the lower two-thirds, the teeth mammiform or narrowly mammiform, often cusped, subtruncate at base, the petioles up to 4 cm, with numerous, long, wavy, pale simple eglandular hairs; cauline 0–1, lanceolate or ovate, caudate at apex, dentate, shortly petiolate; all glabrous or nearly so on both surfaces but with medium to long, pale, wavy simple eglandular hairs round the margin, or with scattered, short to medium hairs on one or both surfaces especially on the midrib beneath. Inflorescence with 2–14 capitula, paniculate-corymbose, sometimes with a long lower branch; peduncles straight or curved, with dense stellate hairs, few to numerous, short, dark glandular hairs and sometimes an occasional, medium simple eglandular hair. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts porrect in bud, 4–13 × 1.0– 1.3 mm, olive-green, the inner with a pale margin, all narrowly linear-lanceolate, gradually tapered to an acute apex, with numerous very short to medium (0.2–1.0 mm), dark glandular hairs and numerous stellate hairs, and an occasional, medium, dark-based simple eglandular hair. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 7–8. Apomictic. Native. The headquarters of this species are the cliffs of Ben Hope in Sutherland. It also occurs on sandhills at

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Auchinver, below Coldbackie, by Tongue Bay in the same county. Scandinavia. 356. H. mucronellum P. D. Sell & C. West Fleck-leaved Hawkweed H. silvaticum var. asymmetricum auct. Phyllopodous perennial herb with a branched stock. Stem 15–55 cm, pale yellowish-green, often purplish towards the base, slender, flexuous, strongly striate, glabrous or with an occasional simple eglandular hair below and numerous stellate and a very occasional, short, dark glandular hair above. Leaves medium to dark green, flecked and blotched with brownish-purple on upper surface, paler and often flushed with purple or reddish-purple beneath; basal usually numerous, the outer with lamina 2–7 × 2–4 cm, elliptical, ovate or subrotund, rounded-mucronulate at apex, subentire, denticulate or dentate and subtruncate at base, the inner with lamina 3–11 × 1–5 cm, very variable, oblong, elliptic-oblong, lanceolate-oblong, ovate or lanceolate, mostly acute but some obtuse-mucronate at apex, denticulate to grossly dentate, the teeth mammiform and very irregular, the basal sometimes reflexed and subtruncate at base; cauline 0–1, usually in the lower half and divaricate, the lamina lanceolate, long acute at apex, dentate, petiolate; all glabrous or nearly so on upper surface, with scattered, medium to long, pale simple eglandular hairs beneath and more numerous on the midrib beneath and the margins. Inflorescence with 2–8 capitula, compactly corymbose, sometimes with a lower branch; peduncles short, slender and curved, with dense stellate and numerous, very short and short (0.2–0.4 mm), dark glandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 0.8–1.0 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, drawn out to a long, narrow, acute apex, with numerous, very short to short (0.2–0.5 mm), dark glandular hairs, numerous short to medium, dark-based simple eglandular hairs, and scattered stellate hairs particularly on the margins. Ligules pale yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Low cliffs on the west side of the River Naver, near Bettyhill and limestone rocks by the sea at Keoldale, Durness, both in Sutherland. Endemic. 357. H. caesitium Norrl. Ribbleshead Hawkweed H. silvaticum subsp. maculosum (Dahlst. ex Stenstr¨om) Omang; H. maculosum (Dahlst. ex Stenstr¨om) Omang; H. bifidum subsp. maculosum (Dahlst. ex Stenstr¨om) Zahn; H. silvaticum var. maculosum (Dahlst. ex Stenstr¨om) F. N. Williams Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, often reddish-purple towards the base, slender, striate, with few to fairly numerous, medium to long, pale simple eglandular hairs towards the base and few upwards, and few to numerous stellate hairs throughout. Leaves medium green with brownishpurple flecks and small blotches on upper surface, paler and often flushed purple beneath; basal few, with lamina 3–7 × 1.5–2.5(–3.5) cm, lanceolate, ovate-lanceolate,

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oblong-lanceolate or linear-lanceolate, obtuse-mucronate to acute at apex, subentire, denticulate or dentate towards the base where a few teeth are often more or less mammiform, subtruncate or abruptly contracted at base, the petioles medium to long, slender and with numerous, long, pale simple eglandular hairs; cauline 0–1, with lamina linear and acute at apex; all with short, medium or long, pale simple eglandular hairs on both surfaces and the margin and particularly the midrib beneath. Inflorescence with 3–15 capitula, paniculate-corymbose with the upper part sometimes subumbellate; peduncles slender, with dense stellate hairs, few to rather numerous, short to medium, dark-based simple eglandular hairs and an occasional, very short glandular hair. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts more or less porrect in bud, 4–8 × 0.6– 1.0 mm, olive-green, the inner with paler margins, all narrow linear-lanceolate, very narrow towards the acute apex, with numerous, short to medium, dark-based simple eglandular hairs, few, very short, dark glandular hairs, and stellate hairs along the margins. Ligules orange-yellow, with a few, short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Ledges, grassy slopes and grykes on the limestone. Frequent near Ribbleshead in Yorkshire and an old record from above the Dale Beck on Twistleton Scars. A specimen from Keld Head Scars may be a good record, but the spotted-leaved plant which grows there is H. glanduliceps and there may be a mixture of specimens. The species looks native, but in the area where it is most common it is associated with a quarry and the railway. South and middle Scandinavia, Finland and Austria. 358. H. piligerum (Pugsley) P. D. Sell & C. West Brindled Hawkweed H. variicolor var. piligerum Pugsley; H. variicolor auct.; H. integratum auct.; H. murorum subsp. variicolor auct.; H. orbicans auct.; H. rotundatum auct. Phyllopodous perennial herb with a branched stock. Stem 30–70 cm, pale yellowish-green, sometimes reddish towards the base, slender to more or less robust, striate, with few, long, pale simple eglandular hairs to glabrous below and dense stellate hairs and few to fairly numerous, short, dark glandular hairs in the upper part. Leaves bright, medium green, usually with brownish-purple spots and blotches on the upper surface and paler and often bluish beneath and sometimes suffused purple; basal usually numerous, the outer with lamina 3–9 × 1.5–4.5 cm, elliptical, subrotund or oblong-elliptical, rounded-obtuse at apex, subentire, denticulate or shallowly sinuate-dentate and rounded or subtruncate at base, the inner with lamina 7–11 × 2.5–6.0 cm, ovate, elliptical or elliptical-oblong, rounded-mucronulate to acute at apex, subentire, denticulate or shallowly sinuate-dentate and abruptly contracted or subtruncate at base, the petioles up to 8 cm and with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, the lamina lanceolate to ovate, acute at apex, shallowly dentate in lower half, abruptly contracted at base and petiolate; all with numerous, short to medium, pale, rather stiff simple eglandular hairs on the upper surface to

nearly glabrous, with numerous, short to long, pale simple eglandular hairs beneath, particularly on the midrib, and on the margin, sometimes with stellate hairs on the underside of the cauline. Inflorescence with 3–7(–20 cult.) capitula, paniculate-corymbose; peduncles straight or curved, with dense stellate hairs, numerous, very short to short (0.2–0.5 mm), dark glandular hairs and occasional, short or medium, dark-based simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts more or less porrect in bud, 4–12 × 1.0–1.4 mm, blackish-green, the inner with paler margins, all linearlanceolate, obtuse at apex, with numerous, very short to medium (0.2–1.0 mm) glandular hairs, few to numerous, short and medium, dark-based simple eglandular hairs and dense stellate hairs on the margins and near the apex. Ligules yellow, with very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins subulatedentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. This species differs from H. variicolor Dahlst. ex Stenstr¨om in its more rounded leaves and simple eglandular hairs as well as glandular hairs on the peduncles and involucral bracts. Even E. S. Marshall no. 3844 from the Keltney Burn, Perthshire, placed by H. W. Pugsley under typical H. variicolor, has these characters. Native. Rocky streamsides and cliff ledges. Widespread in central Scotland. Endemic. 359. H. pollinarioides Pugsley Calcareous Hawkweed H. candelabrae auct.; H. murorum var. crassiusculum auct. Phyllopodous perennial herb with a branched stock. Stem 20–70 cm, pale yellowish-green, often purplish below, slender to robust, flexuous, striate, with an occasional, pale simple eglandular hair below and with dense stellate hairs and a few, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, sometimes with brownish-purple flecks and blotches, paler beneath and often flushed reddish-purple; basal usually numerous, the outer with lamina 3–9 × 2–7 cm, very broadly elliptical, subrotund or very broadly ovate, rounded-mucronulate at apex, denticulate to shallowly sinuate-dentate and truncate to cordate at base, the inner with lamina 5–12 × 3–7 cm, very broadly ovate, rounded-subacute to acute at apex, shallowly sinuate-dentate with large mammiform teeth near the base, the petioles up to 6 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, when present lanceolate, long-acute at apex, dentate, petiolate; all glabrous or nearly so on upper surface, with numerous, short to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 3–15 capitula, compactly paniculate-corymbose, sometimes with a long lower branch; peduncles rather short, straight or curved, with dense stellate hairs and numerous, very short to short (0.2– 0.5 mm), dark glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.0–1.2 mm, dark olive-green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, short and very short (0.2–0.5 mm), dark glandular hairs, numerous, short to medium, dark-based simple eglandular hairs, and numerous stellate hairs on the margins and towards the base. Ligules yellow, glabrous-tipped. Styles

39. Hieracium yellow or slightly discoloured. Receptacle pits with margins slightly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Limestone cliffs on the north coast of Sutherland and on the Isle of Raasay in the North Ebudes. Endemic. 360. H. duriceps F. Hanb. Hard-headed Hawkweed H. bifidum subsp. duriceps (F. Hanb.) Zahn; H. murorum var. pulcherrimum F. Hanb.; H. pulcherrimum (F. Hanb.) Roffey; H. murorum subsp. pulcherrimum (F. Hanb.) Zahn; H. pellucidum var. pulcherrimum (F. Hanb.) W. R. Linton; H. killinense auct.; H. angustatum auct. Phyllopodous perennial herb with a branched stock. Stem 20–55 cm, pale yellowish-green, sometimes tinted reddishpurple towards the base, slender to robust, striate, with few to numerous, medium to long, pale simple eglandular hairs below, becoming fewer upwards, and dense stellate hairs and few, short, dark glandular hairs above. Leaves rather pale yellowish-green and sometimes spotted brownish-purple on upper surface, paler beneath; basal usually numerous, the outer with lamina 2–9 × 1.5–3.5 cm, broadly ovate or elliptical, rounded-mucronulate at apex, subentire, denticulate or shallowly dentate in the lower half and rounded, subtruncate or abruptly contracted at base, the inner with lamina 4–10 × 1.5–3.5 cm, elliptical, lanceolate, ovate, oblong-elliptical or oblong, obtuse to acute at apex, subentire to denticulate, usually with a few teeth towards the base, usually shallow but sometimes narrowly mammiform and usually subtruncate at base, the petioles up to 4 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 0–1(–2), lanceolate, linear-lanceolate or bract-like, acute at apex, entire or dentate in the lower half, narrowed or subtruncate at base, petiolate; all with numerous, short to medium, pale simple eglandular hairs on the upper surface to nearly glabrous, and with numerous, medium to long, pale simple eglandular hairs beneath and on the margins. Inflorescence with few to numerous capitula, irregularly paniculate or subcorymbose, often with a long lower branch; peduncles slender and curved, with dense stellate hairs, numerous, very short to medium (0.2–1.0 mm), dark glandular hairs mixed with fairly numerous longer dark simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4– 12 × 1.0–1.4 mm, blackish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, unequal, very short to medium (0.2–1.2 mm), black glandular hairs, numerous, medium to long, dark simple eglandular hairs, and few to numerous stellate hairs along the margins. Ligules yellow, with numerous, very short simple eglandular hairs along the apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges, grassy slopes and streamsides; perhaps always on basic rock. One of the commonest hawkweeds in Scotland and northern England south to Lancashire and Yorkshire; recorded from Co. Leix, Co. Down, Co. Antrim and Co. Londonderry in Ireland. Endemic. Note on the nomenclature of H. duriceps F. Hanb. and associated species. H. stenophyes was described from Black’s Hope, Moffat in 1893 by W. R. Linton. In 1902 F. N. Williams made it

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a variety of H. euprepes and in 1905 W. R. Linton himself made it a variety of the Scandinavian H. duplicatum Almq. Plants from the Moffat area collected and grown by W. R. Linton which he called H. sarcophyllum seem to be the same thing. A cultivated plant from Moffat sent out in the Lintons’ Set of British Hieracia no. 93 as H. orarium var. fulvum is also H. stenophyes. In 1896 W. R. Linton described H. stenophyes var. oxyodus from the Midlaw Burn, Moffat as no. 149 of the Lintons’ Set of British Hieracia. As this was a distributed set and had a printed description it is validly published. As pointed out by Pugsley (1948) no. 149 is a mixture of H. stenophyes and H. longilobum. The original description, however, is clearly referable to H. longilobum. H. oxyodus was raised to the rank of species by W. R. Linton in 1905. H. longilobum was sent out by H. Dahlstedt as an exsiccata in 1893, but was not validly published. E. S. Marshall discussed it in 1908, but he also did not validly publish it. The first valid publication which can be traced is that of Zahn in 1921 who called it H. murorum subsp. longilobum. It was left to J. Roffey (1925) to raise it to the rank of species. H. oxyodus (W. R. Linton) W. R. Linton is thus the oldest name for H. longilobum (Zahn) Roffey. In 1894, F. J. Hanbury described H. aggregatum var. prolongatum from Glen Lochay in Perthshire. In making H. oxyodus a species in 1905, W. R. Linton included plants from Perthshire as well as Dumfries-shire. After the account he says ‘This is the plant named H. aggregatum var. prolongatum in JB 1894, 228,’ but does not give Glen Lochay as a locality. Pugsley (1948), realising the Moffat plant was H. longilobum, used the name H. oxyodus for the Perthshire plant. Unfortunately, H. oxyodus must be used for the Moffat plant and, as the Perthshire plant does not have a specific name, it is called H. protentum. When raising var. oxyodus to specific rank in 1905 W. R. Linton described a new variety of it, var. delicatulum from Coire Dubhchlair (Corrie Dubh Ghalair) and Glen Lochay in Perthshire. This is a distinct species. Pugsley (1948) made var. delicatulum a variety of H. duriceps, to which it is closely allied. In 1891, H. Dahlstedt sent out exsiccatae of H. micracladium from Norway, but did not validly publish it. In 1902, F. N. Williams identified Scottish plants with it and validly published the name as H. silvaticum var. micracladium. Although he cites Dahlstedt’s plant, Williams clearly states that his description is based on the Lintons’ Set of British Hieracia no. 17 from Killin, which is different from Dahlstedt’s plant and is identical with H. oxyodus var. delicatulum. Var. micracladium was raised to specific rank by A. Ley in 1905. The first description of the Scandinavian plant which can be traced is H. murorum subsp. micracladium Zahn in 1921. S. O. F. Omang made it a species in 1924. The Scandinavian plant should therefore be called H. neomicracladium P. D. Sell. Zahn (1921), realising that the var. micracladium of Williams and the Lintons was not the Scandinavian plant, called the Scottish plant H. murorum subsp. killinense, which was raised to specific rank by Roffey in 1925. Pugsley’s (1948) description of H. killinense is not referable to H. micracladium, but to H. duriceps. The specimens of H. prolongatum and H. oxyodus var. delicatulum (= micracladium), whose types come from the

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same area, are hopelessly muddled in herbaria, despite being easily distinguishable. H. duriceps was described by F. J. Hanbury in 1892 from Oykell Bridge in Sutherland. It is a widespread, characteristic species of Scotland and northern England. In 1892, F. J. Hanbury published a new variety, H. murorum var. pulcherrimum, from Catrigg (Catterick) Force, near Settle in Yorkshire and Carnedd Dafydd in Caernarvonshire, and it was raised to specific rank by J. Roffey in 1925. The description is of the Yorkshire plant, which is H. duriceps. These Yorkshire plants are usually very large, have long, pointed leaves and involucral bracts with glandular hairs longer than most, but not all Scottish plants, but it has been impossible to clearly distinguish them. The Carnedd Dafydd plant has been named H. snowdoniense P. D. Sell & C. West (1955). In 1894 F. J. Hanbury published H. murorum var. camptopetalum and in 1948 H. W. Pugsley transferred it as a variety to H. duriceps. In 1893, H. Dahlstedt named a Scandinavian plant H. serratifrons subsp. caliginosum. British botanists identified some of our plants with it, and Roffey (1925) raised it to the rank of species. None of our plants is identical with the Scandinavian one and the Sutherland plants so identified differ from H. murorum var. camptopetalum only in having fully developed ligules. H. murorum var. camptopetalum was therefore raised to specific rank by P. D. Sell & C. West (1955) to cover both these plants. 361. H. protentum P. D. Sell Meall Garbh Hawkweed H. aggregatum var. prolongatum F. Hanb. Phyllopodous perennial herb with a branched stock. Stem 30–55 cm, pale yellowish-green, sometimes purplish at the very base, with numerous, long, pale simple eglandular hairs throughout and numerous stellate hairs in the upper part. Leaves medium yellowish-green on upper surface, paler and greyish beneath; basal numerous, the outer with lamina 2.5– 7.0 × 1.5–2.5 cm, narrowly elliptical or oblong, roundedmucronulate at apex, shallowly undulate-dentate or with a few, larger, cusped teeth towards the cuneate base, the inner with lamina oblong or lanceolate-oblong, more or less acute at apex, shallowly dentate or with large cusped teeth below and cuneate to attenuate at base, the petioles up to 5 cm, with numerous, medium to long, pale simple eglandular hairs; cauline 0–2(–3 cult.), when present similar to inner basal and shortly petiolate or bract-like; all with short simple eglandular hairs on upper surface and longer, more numerous ones beneath, especially on the midrib, and on the margins. Inflorescence with 2–7(–26 cult.) capitula, furcatecymose, becoming subumbellate in cultivation and sometimes with a long, lower branch; peduncles short and slender, with dense stellate hairs, numerous, short and medium pale simple eglandular hairs, and an occasional, short glandular hair. Capitula 20–30 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–10 × 1.0– 1.2 mm, olive-green, the inner with broad pale margins, all linear-lanceolate, obtuse at apex, with dense, short and medium, dark-based simple eglandular hairs, and a few stellate hairs towards the base, without or with a very occasional, short, dark glandular hair. Ligules yellow, with very short, simple eglandular hairs at apex. Styles discoloured.

Receptacle pits with margins shortly dentate. Achenes 2.5– 3.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only from Allt Coire Chaorach and Allt Tir Artair, near Killin in Perthshire. Endemic. 362. H. micracladium (F. N. Williams) Ley Small-headed Hawkweed H. silvaticum var. micracladium F. N. Williams; H. murorum subsp. killinense Zahn; H. killinense (Zahn) Roffey; H. oxyodus var. delicatulum W. R. Linton; H. duriceps var. delicatulum (W. R. Linton) Pugsley; H. bifidum subsp. oxyodus var. delicatulum (W. R. Linton) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, often purplish towards the base, erect, rather slender, flexuous, striate, with an occasional, medium to long, pale simple eglandular hair particularly towards the base and dense stellate hairs and few to numerous, short, dark glandular hairs in the upper part. Leaves deep green on upper surface, paler beneath and sometimes tinted purplish; basal numerous, the outer with lamina 4–6 × 2.0–2.5 cm, elliptical, rounded-obtusemucronulate at apex, subentire, denticulate or shallowly dentate and rounded at base, the inner with lamina elliptical to oblong-elliptical, acute or acuminate at apex, dentate especially in the lower half, the teeth small and often mammiform, cuneate and often asymmetrical at base, the petioles 3–7 cm, long and slender, with numerous, long, pale, wavy simple eglandular hairs; cauline 0–1, like inner basal but smaller and petiolate, or bract-like; all with few to numerous, short to medium, pale simple eglandular hairs on both surfaces and the margin. Inflorescence with 2–10 (–15 cult.) capitula, cymose-paniculate and often subumbellate, sometimes with a long lower branch; peduncles rather short, slender, with dense stellate hairs, numerous, short, dark glandular hairs and numerous, short to medium, dark-based simple eglandular hairs. Capitula 20– 30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, unequal, short to medium, black glandular hairs and numerous, medium, dark simple eglandular hairs, stellate hairs few towards the base or absent. Ligules rather pale yellow, glabrous-tipped. Styles yellow or slightly livid. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. For an explanation of the complicated nomenclature of this species see under H. duriceps. Native. The only certain locality for this plant is Meall na Samhna (Meall-na-Saone), Glen Lochay in Perthshire. Endemic. 363. H. subprasinifolium Pugsley Leek-coloured Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–60 cm, pale yellowish-green, usually reddish-purple near the base, slender to robust, striate, with a few, long, pale simple eglandular hairs near the base and a fair number of shorter ones upwards, and numerous stellate hairs and short, dark glandular hairs in the upper part. Leaves deep bluish-green on upper surface and bluish, often suffused

39. Hieracium purple beneath, thick; basal usually numerous, the outer with lamina 3–6 × 1.5–3.0 cm, elliptical or ovate-elliptical, obtuse-mucronulate at apex, entire, remotely denticulate or with a few, small teeth near the base and rounded, truncate or cordate at base, the inner 5–9 × 1.5–4.0 cm, ovate, oblongovate, lanceolate or rarely oblong, acute or narrowed and cuspidate at apex, subentire, denticulate or sinuate-dentate, the teeth sometimes narrowly mammiform, and truncate at base, the petioles up to 5 cm, with numerous to dense, long, pale, wavy simple eglandular hairs; cauline 0–1, lanceolate to ovate, acuminate at apex, denticulate or with a few teeth, cuneate to truncate at base and petiolate; all glabrous or nearly so on both surfaces or with scattered, medium, pale simple eglandular hairs beneath and numerous ones on the margin. Inflorescence with 2–10 capitula, paniculatecorymbose, sometimes with a long lower branch; peduncles with dense stellate hairs, numerous, very short or short (0.2– 0.7 mm), dark glandular hairs and an occasional, medium, dark-based simple eglandular hair. Capitula 25–35 mm in diameter, rounded or subtruncate at base. Involucral bracts incumbent in bud, 4–9 × 0.7–1.0 mm, dark olive-green, the inner with paler margins, all narrow linear-lanceolate, acute at apex, with dense, very short or short (0.2–0.5 mm), black glandular hairs, few to numerous, short or medium, darkbased simple eglandular hairs, and numerous stellate hairs along the margin at the apex. Ligules yellow, glabroustipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. This species is common on ledges, rock scree and banks throughout the Derbyshire Dales and extending into Staffordshire in the Manifold Valley. It appears to be native, but is often associated with the railway and tracksides and may be introduced. Endemic. 364. H. discophyllum P. D. Sell & C. West Disc-leaved Hawkweed H. pellucidum var. lucidulum auct.; H. murorum subsp. subsanguineum auct. Phyllopodous perennial herb with a branched stock. Stem 20–40(–60) cm, pale yellowish-green, usually purplish in lower half, slender to robust, striate, with few to numerous, medium to long, pale simple eglandular hairs below, numerous stellate hairs above and sometimes a minute, dark glandular hair in the upper part. Leaves yellowish-green with small brownish-purple spots and flecks on the upper surface, paler and often purple-tinted beneath; basal usually rather few, the outer with lamina 2.5–5.0 × 2–3 cm, subrotund or ovate, rounded-mucronulate at apex, subentire to denticulate or slightly dentate and rounded or subtruncate at base, the inner with lamina 4–8 × 3–5 cm, mostly ovate, sometimes elliptical-ovate, mostly obtuse, sometimes more or less acute at apex and denticulate to shallowly dentate with a few, larger, often mammiform teeth near the base, the petioles rather short, with numerous, long, pale simple eglandular hairs; cauline 0–1, often large, petiolate and similar to inner basal; all with the surfaces glabrous or with scattered, medium, pale simple eglandular hairs with longer ones on the midrib beneath and on the margin. Inflorescence with 3–6 capitula, numerous in

395

cultivation, furcate-corymbose; peduncles medium to long, suberect, with dense stellate hairs, numerous, short, dark glandular hairs and occasional, medium, dark-based simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.2– 1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, abruptly narrowed to an obtuse or subacute apex, with numerous, short to medium, dark-based simple eglandular hairs, numerous short and very short, dark glandular hairs, and dense stellate hairs at the base and on the margin. Ligules deep yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins long dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. K. H. Zahn (1921) created the name H. murorum subsp. subsanguineum as a replacement name for H. sanguineum (W. R. Linton) Ley, and it must be placed as a synonym of that species. His description, however, based on E. F. Linton no. 140 is of this species. Native. Limestone cliff ledges and scree. Pwll Byffre near Penwyllt, Dyffryn Crawnon, Craig Maillard in Cefn Coed, Taren-yr-esgob, Y Fan Beg in the Beacons and Taren Lewyd all in Breconshire and Fan Fechan in Carmarthenshire. Endemic. 365. H. scotostictum Hyl. Dappled Hawkweed H. praecox auct. Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, yellowish-green, sometimes purplish at base, striate, slender, with few to numerous, long, wavy, pale simple eglandular hairs particularly towards the base and numerous stellate hairs and very short or short, fine, dark glandular hairs above. Leaves glaucous-green, heavily spotted and marbled brownish-purple on the upper surface, paler and usually tinted purplish beneath; basal with lamina 3– 9 × 2–4 cm, mainly ovate or lanceolate, sometimes oblong or oblong-lanceolate, mainly acute, sometimes obtuse at apex, subentire or denticulate in the lower half or dentate in the lower half with sharp teeth, subtruncate or subcordate at base, with long petioles up to 15 cm clothed with numerous, long, wavy, pale simple eglandular hairs; cauline 0–1, when present the lamina lanceolate or ovate, acute at apex, sharply toothed, petiolate; more or less glabrous on the upper surface, with numerous, medium to long, pale, wavy simple eglandular hairs beneath (particularly on the midrib) and on the margin. Inflorescence cymose-corymbose, with a rather short acladium; peduncles with dense stellate hairs, numerous, very short and short (0.2–0.7 mm), dark glandular hairs and sometimes an occasional, medium, dark simple eglandular hair. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.7–1.0 mm, blackish-green, the inner with pale margins, all narrow linear-lanceolate, gradually narrowed to a long, fine more or less acute apex, with few stellate hairs mainly on the margin and towards the base, numerous, very short and short (0.2– 0.6 mm), dark glandular hairs and few to numerous, short to medium, rather dark simple eglandular hairs. Ligules medium yellow, glabrous-tipped. Styles orange. Receptacle pits with subulate-dentate margins. Achenes 2.5–3.0 mm, reddish-black. Flowers 5–7. Apomictic.

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Introduced. First recorded from a railway cutting at Smitham Bottom in Surrey in 1920. It soon spread in the area around London and has been recorded in scattered localities on road and railway banks north to Westmorland and west to Cornwall. In Cambridge it is a weed in various gardens and grows on the walls of Pembroke College. It was one of several species formerly growing on grassy banks round Walthamstow Reservoirs in Essex. In Continental Europe it occurs in Sweden, Denmark and Germany and has frequently been called H. praecox. 366. H. baliophyllum Dahlst. ex Hyl. Oxford Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, sometimes tinted purplish towards the base, striate, robust, with numerous, pale, medium to long simple eglandular hairs below becoming fewer upwards, and numerous stellate hairs and few, short glandular hairs in the upper part. Leaves medium yellowgreen, with scattered spots and blotches on the upper surface, paler and tinted reddish beneath; basal numerous, the lamina 5–11(–13) × 2.4–5.0 cm, elliptical or oblongelliptical, rounded-mucronulate to acute at apex, sinuatedenticulate or toothed towards the base, the teeth mammiform or narrowly mammiform, the basal pair often curved outwards at the truncate base, the petioles up to 7 cm, with dense, long, pale simple eglandular hairs; cauline 0–1, ovate, acute at apex, sinuate-dentate, petiolate; all with scattered, short to medium, pale simple eglandular hairs on the upper surface and numerous, medium to long similar ones beneath and on the margins. Inflorescence with 5–15 capitula, paniculate-corymbose, sometimes with a long lower branch; peduncles with dense stellate, numerous, short to long, pale simple eglandular hairs and few, short (0.3–0.5 mm) dark glandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.2–1.5 mm, dull blackish-green, the inner with paler margins, all with numerous, very short to medium, pale simple eglandular hairs, numerous, very short (0.2–0.5 mm), dark glandular hairs and stellate hairs, along the margins and towards the base. Ligules yellow, glabroustipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. This species was collected by G. C. Druce at Oxford in July 1918 and distributed through the Botanical Exchange Club without a name. It is known elsewhere only in two localities in Sweden. 367. H. zygophorum Hyl. Suffolk Hawkweed Phyllopodous perennial herb with a branched stock. Stems 40–90 cm, yellowish-green, often purplish towards the base, robust, striate, with more or less numerous, medium to long, pale simple eglandular hairs throughout, few to numerous stellate hairs and a few, very short, dark glandular hairs in the upper part. Leaves yellowish- to bluish-green on upper surface, often tinted purplish on the lower surface; basal few to numerous 4–15 × 1.0–5.5 cm, the outer with lamina oblong or lanceolate-oblong, narrowed to an obtuse apex, subentire or with a few teeth near the base, the inner or sometimes all of them with lamina oblong-lanceolate,

oblong, lanceolate or ovate, narrowed to a more or less acute apex, entire or denticulate in the upper third, dentate below with large mammiform, aquiline-mammiform or narrowly mammiform teeth, sometimes with small teeth between the large ones, sometimes with teeth on the petiole, rounded or subtruncate at base, the petioles up to 15 cm, with dense, long, pale simple eglandular hairs; cauline 0–1, when present with lamina lanceolate or ovate, acute to long acuminate at apex, usually deeply and irregularly dentate, the teeth very narrowly mammiform or long and sharp, petiolate; all with few to fairly numerous, short to medium, pale simple eglandular hairs on the upper surface, more numerous ones on the lower surface and numerous long ones on the margin and the midrib beneath, with scattered stellate hairs beneath the cauline. Inflorescence cymosecorymbose and often subumbellate, frequently with a long lower branch, with a fairly short acladium; peduncles arcuate, with dense stellate hairs, numerous very short to short (0.2–0.6 mm), fine, dark glandular hairs and few to numerous, short to medium, pale simple eglandular hairs. Capitula 40–48 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 0.7–1.0 mm, greyish-green with paler margins, very narrowly linear-lanceolate, mostly suddenly narrowed about halfway up, then gradually narrowed to a long, narrow, acute apex, the inner ones almost filamentous, with numerous stellate hairs on the margin and base, with numerous very short and short (0.2–0.5 mm, rarely to up to 0.7 mm), fine, dark glandular hairs and numerous, short and medium, pale, dark-based simple eglandular hairs. Ligules medium yellow, shortly pilose-tipped at least when young. Styles pure yellow or slightly discoloured (Hylander gives dark, but his specimens from the type locality still have them yellow). Receptacle pits with subulate-dentate margins. Achenes 3.0–3.5 mm, blackish. Flowers 5–7. Apomictic. Introduced. Calcareous, rabbit-grazed grassland and margin of railway tracks. Known only to the north of Weston Crossing, Suffolk. It was described from Karlskrona, Blekinge, Sweden where also it may not be native. Either its native locality is unknown or it evolved as a grassland weed. When grown in the Botanic Garden at Cambridge it spread as a weed in the beds and along paths and the leaves became more bluish-green and spotted and blotched as in Swedish plants. 368. H. gougetanum Gren. & Godr. Gouget’s Hawkweed H. praecox subsp. gougetanum (Gren. & Godr.) Zahn; H. prasiophaeum Arv.-Touv. & Gaut.; H. maculosum auct. Phyllopodous perennial herb with a branched stock. Stems 30–60 cm, pale yellowish-green, sometimes tinted purplish towards the base, stout, striate, with a few, long, pale, wavy simple eglandular hairs and a few stellate and short, fine, dark glandular hairs above. Leaves glaucous-green, speckled and spotted with brownish-purple on upper surface, paler and sometimes tinted purplish beneath; basal numerous with lamina 4–10 × 2–5 cm, oblong, ovate or ovate-oblong, mostly acute at apex, deeply and grossly dentate or serrate-dentate, the teeth long, cusped and narrowly mammiform or aquiline-mammiform, often with some teeth extending on to the petiole, subtruncate at base, some leaves

39. Hieracium sometimes less dentate and cuneate at base, the petioles up to 8 cm, with long, shaggy simple eglandular hairs; cauline 0–1, when present with lamina ovate or lanceolate, acute at apex, deeply toothed like the basal and petiolate; all glabrous on upper surface and with numerous, medium to long, pale, wavy simple eglandular hairs beneath (especially on the midrib) and margins. Inflorescence cymosecorymbose, with a rather long acladium and sometimes a long lower branch; peduncles arcuate, with dense stellate hairs, very short to short (0.2–0.5 mm), dark glandular hairs and sometimes an occasional, medium, dark simple eglandular hair. Capitula 25–45 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–13 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linearlanceolate, gradually narrowed to a long, thin, more or less acute apex, the inner almost filamentous, with few stellate hairs mainly on the margins and near the base, numerous, unequal, very short to medium (0.2–0.8 mm), black glandular hairs and numerous, medium to long, dark simple eglandular hairs. Ligules medium yellow, glabrous-tipped. Styles yellow. Receptacle pits with shortly dentate margins. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–6. Apomictic. Although unmistakable in all its forms, it is very variable in toothing, base of leaf, spotting and degree of hairiness much of which seems to be due to the time of the year. Introduced. Railway banks, spoil heaps and by the Royal Canal near Liffey Junction, Dublin, where it has been known since at least 1903. It then appears to have spread to railway sidings at North Wall and more recently has been recorded at Astagob sand-pits, between Liffey Junction and Cabra and walls between the Royal Canal and the railway at Drumcondra. It also occurs on railway sidings at Louise Bridge Station, Leixlip, in Co. Kildare and on old colliery spoil tips near Flemington, Lanarkshire. Named after ?Edouard Gouget. 369. H. liljeholmii Dahlst. Liljeholm’s Hawkweed Phyllopodous perennial herb with a branched stock. Stems 30–70 cm, pale yellowish-green, often flushed purplish towards the base, markedly striate, fairly robust, with a few, scattered, pale, medium to long simple eglandular hairs throughout and few to numerous stellate and short, dark glandular hairs in the upper part. Leaves yellowish-green or slightly glaucous-green, spotted and blotched brownishpurple on the upper surface, paler and flushed purplish beneath; basal few to numerous, with lamina 5–9 × 2–4 cm, ovate to oblong-ovate or oblong-lanceolate, obtusemucronate to acute at apex, denticulate in the upper part becoming dentate or rarely serrate-dentate in the lower part, the teeth narrowly to broadly mammiform or sometimes aquiline-mammiform, subtruncate at base, on long petioles up to 10 cm with very long, pale, wavy simple eglandular hairs; cauline nearly always 1 large, ovate or lanceolate, long acute or acuminate at apex, serrate-dentate, truncate at base, petiolate; all with numerous, short to medium, pale simple eglandular hairs on both surfaces and long ones on the margin. Inflorescence cymose-corymbose, with a rather long acladium and sometimes a long, lower branch; peduncles arcuate, with dense stellate hairs and numerous, short (0.4– 0.5 mm), subequal, dark glandular hairs. Capitula 13–30

397

mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.5–1.0 mm, blackish-green with paler margins, narrowly linear-lanceolate, gradually narrowed to an obtuse or subacute apex, with few stellate hairs mainly on the margin and at the base, numerous very short and short (0.2–0.5 mm), dark glandular hairs and few, short to medium, dark simple eglandular hairs. Ligules medium yellow, glabrous-tipped. Styles pure yellow. Receptacle pits with shortly dentate margins. Achenes 3.0–3.5 mm, blackish. Flowers 5–6. Apomictic. Introduced. Clearing in Joyden’s Wood, Dartford, Kent and grassy banks at Walthamstow Reservoirs in Essex. Described from G¨oteborg, Sweden where also it was considered to be introduced. Named after A. F. Liljeholm (fl. 1896–1912). 370. H. cuneifrons (Ley) Pugsley Wedge-leaved Hawkweed H. subulatidens var. cuneifrons Ley; H. murorum subsp. subulatidens var. cuneifrons (Ley) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–80 cm, pale yellowish-green, sometimes purplish towards the base, more or less robust, striate, with a few, scattered, medium to long, pale simple eglandular hairs, and numerous stellate hairs and a few, short, black glandular hairs in the upper part. Leaves bright green on upper surface, paler beneath and sometimes suffused purplish; basal usually few, similar except for size, the lamina 2.5– 17.0 × 1–6 cm, elliptical, ovate or oblong, mostly acute at apex but sometimes obtuse, shallowly to deeply dentate, the teeth sometimes large and mammiform or longmammiform, cuneate at base, the petioles up to 7 cm, with numerous, long, pale, wavy simple eglandular hairs; cauline (0–)1–2, like the basal and petiolate, or bractlike; all with short to long, pale simple eglandular hairs on both surfaces and the margins or nearly glabrous on upper surface. Inflorescence with 3–15(–40 cult.) capitula, paniculate-corymbose, with a long lower branch; peduncles often incurved, with dense stellate hairs and numerous, short (0.4–0.5 mm), black glandular hairs, without simple eglandular hairs. Capitula 30–40 mm in diameter, rounded or subtruncate at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.2 mm, blackish-green, the inner with paler margins, linear-lanceolate, obtuse at apex, with dense, very unequal, very short to medium (0.2–1.5 mm), black glandular hairs, and fairly numerous stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Native. Cliff ledges, quarries and lanesides, probably always on limestone. Monmouthshire, Glamorganshire, Carmarthenshire and particularly Breconshire. Endemic. 371. H. uistense (Pugsley) P. D. Sell & C. West Island Hawkweed H. clovense var. uistense Pugsley Phyllopodous perennial herb with a branched stock. Stem 8–30 cm, pale yellowish-green, often suffused purplish, slender and flexuous, striate, with numerous, long to very long, pale simple eglandular hairs below, becoming fewer

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364. H. discophyllum P. D. Sell & C. West

366. H. baliophyllum Hyl.

368. H. gougetanum Gren. & Godr.

365. H. scotostictum Hyl.

367. H. zygophorum Hyl.

369. H. liljeholmii Dahlst.

Inner and outer basal leaves of species of the Section Hieracium

39. Hieracium

399

371. H. uistense (Pugsley) P. D. Sell 370. H. cuneifrons (Ley) Pugsley

374. H. sylvularum Jord. ex Boreau 372. H. ronasii P. D. Sell

376. H. firmiramum Hyl.

377. H. onychodontum Hyl.

Inner and outer basal leaves of species of the Section Hieracium

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upwards, and numerous stellate hairs and numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler and sometimes suffused reddish beneath; basal few to fairly numerous, the outer with lamina 1.5–5.0 × 1.5–3.0 cm, subrotund to elliptical, rounded-mucronate at apex, denticulate to sharply dentate and rounded at base, the inner with lamina 4–6 × 2–3 cm, ovate, lanceolate or elliptical, more or less acute at apex, with sharp teeth and a cuneate or abruptly contracted base, the petioles short, up to 3 cm, with numerous, very long, pale simple eglandular hairs; all with few to numerous, short to medium, stiff simple eglandular hairs on the upper surface, with more numerous and longer ones beneath and on the margin. Inflorescence with 2–4 capitula, furcate-corymbose; peduncles short, straight or curved, with dense stellate hairs, numerous, very short to short (0.2– 1.0 mm), dark glandular hairs and an occasional simple eglandular hair. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.2–1.5 mm, blackish-green, the inner with paler margins, all linearlanceolate, obtuse at apex, with dense, very short to medium (0.2–1.0 mm), fine, dark glandular hairs, few stellate hairs on the margins and few, medium, dark-based simple eglandular hairs. Ligules deep yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins long-dentate. Achenes 2.5–3.0 mm, blackish. Flowers 6–7. Apomictic. Native. Cliff ledges. Ben Eaval on Uist, Uig on Skye, the Merrick and Poldores Burn, Carsphairn in Kirkcudbrightshire and Beinn na Cille, Morvern in Argyllshire. Endemic. 372. H. ronasii P. D. Sell Ronas Voe Hawkweed Phyllopodous perennial herb with a branched stock. Stem 40–60 cm, pale yellowish-green, sometimes purplish towards the base, more or less robust, with numerous, long, pale, wavy simple eglandular hairs at the very base and few or none upwards, and a few stellate hairs and few, short, dark glandular hairs in the upper part. Leaves rather bluishgreen on upper surface, paler beneath; basal numerous, with the lamina 4–10 × 2.5–4.0 cm, the outer ovate or elliptical, rounded-obtuse at apex, denticulate or with a few, shallow teeth and cuneate at base, the inner elliptical, obovateoblong or oblong, obtuse to acute at apex, denticulate and usually with a few teeth towards the base, and cuneate at base, the petioles up to 6 cm and with numerous, long, pale simple eglandular hairs; cauline 1–2, lower similar to basal and petiolate, the upper when present bract-like; all with more or less numerous, short to medium, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with numerous capitula, paniculate-corymbose; peduncles mostly short, with numerous stellate hairs and numerous, short, dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, rather pale green, the inner with paler margins, all linear-lanceolate, more or less obtuse, with numerous, fine, very short to short (0.2–0.5 mm), dark glandular hairs, and few stellate hairs along the margin, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic.

Native. Precipitous crags and dry rocky stream banks on granite. Very scarce on stream banks at Heylor to the south of the Ronas Voe, and among crags west of Feal near the mouth of the Grud Burn on the north side of the Mainland, Shetland Islands. A specimen in BM labelled Booner’s Hill, Unst may be the same thing, but needs confirmation. Endemic. 373. H. grandidens Dahlst. Grand-toothed Hawkweed H. murorum subsp. grandidens (Dahlst.) Zahn; H. exotericum forma grandidens (Dahlst.) Pugsley; H. macrodon Sudre, non N¨ageli & Peter; H. murorum subsp. murorum microgen. macrodon Sudre Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often suffused brownishpurple in the lower half, slender to robust, striate, with numerous, medium to long, pale simple eglandular hairs in the lower part, and numerous stellate hairs and numerous, short, dark glandular hairs above. Leaves dull, medium yellowish-green on upper surface, paler and slightly bluish beneath, often suffused reddish-purple; basal usually numerous, the outer with lamina 3–7 × 2–5 cm, ovate or elliptical, rounded-mucronulate at apex, grossly dentate, the teeth large and rounded-mammiform to narrowly mammiform, the lower pair often deflexed and the base subtruncate, the inner with lamina 7–10 × 2.5–4.0 cm, lanceolateoblong, elliptic-oblong or oblong, obtuse-mucronate to acute, markedly dentate, the teeth pointed mammiform and getting larger towards the base, interspersed with occasional smaller teeth or denticulations, the basal pair often retrorse, and the base subtruncate, the petioles up to 7 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 0–1(–2), with lamina ovate, long-acute at apex, toothed like basal and petiolate, a second if present bract-like or with lamina linear-lanceolate with long narrow apex, narrow teeth and sessile; all with scattered, short to medium, stiff, pale simple eglandular hairs on the upper surface to nearly glabrous, with numerous, medium to long, pale simple eglandular hairs beneath, particularly on the midrib, and on the margin. Inflorescence with few to numerous capitula, paniculate-corymbose, often with a long lower branch; peduncles rather short, curved or divaricate, with dense stellate hairs, numerous, very short to medium(0.2– 1.0 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.7–1.0 mm, blackish-green, the inner with paler margins, all narrow linear-lanceolate, the inner very narrow, acute at apex, with dense, unequal, very short to medium (0.2–1.0 mm), black glandular hairs, and stellate hairs along the margin, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Roadside and railway banks and other disturbed waste places. Scattered records over Great Britain. The only Irish record is a wall by Millpark House, Co. Offaly. Even in its restricted sense, N. Hylander (1943) found it abundant in Sweden, and in Norway, Finland, Germany, France, Austria, Hungary and Poland. It is

39. Hieracium probably one of the most common hawkweeds in Europe, having spread by road and railway. 374. H. sylvularum Jord. ex Boreau Ample-toothed Hawkweed H. murorum subsp. murorum microgen. macrodon var. sylvularum (Jord. ex Boreau) Sudre; H. murorum subsp. sylvularum (Jord. ex Boreau) Zahn Phyllopodous perennial herb with a branched stock. Stem 35–55 cm, slender to robust, pale yellowish-green, sometimes purplish towards the base, striate, with numerous, medium to long, pale simple eglandular hairs below, becoming fewer upwards, and numerous stellate and numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath, sometimes suffused reddish-purple; basal few to fairly numerous, the outer with lamina 2.5–6.0 × 2–4 cm, ovate or elliptic-ovate, rounded-mucronulate at apex, with large, round mammiform teeth, the basal pair sometimes narrower and retrorse, and the base subtruncate, the inner with lamina 5–10 × 2.5– 5.0 cm, ovate, elliptical or ovate-oblong, obtuse to acute at apex, dentate, with large, rounded mammiform teeth, gradually getting larger towards the base, the basal pair often narrower and retrorse, and truncate at base, the petioles up to 7 cm, with dense, long, pale simple eglandular hairs; cauline 0–1(–2), with lamina usually large, 6–9 × 3–5 cm, ovate, acute or acuminate at apex and often drawn out to a very narrow tip, with large, rounded mammiform teeth, truncate at base and petiolate; all with medium to rather long, pale simple eglandular hairs on both surfaces, particularly on the margin and midrib beneath. Inflorescence with 4–20 capitula, paniculate-corymbose, often with a long lower branch; peduncles rather short and slender, curved or divaricate, with dense stellate hairs and numerous, very short or short, dark glandular hairs, without simple eglandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–12 × 1.0–1.2 mm, green, the inner with paler margins, all linear-lanceolate, acute at apex, with dense, unequal, very short to medium (0.2–1.5 mm), black glandular hairs, and some stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 5–7. Apomictic. Closely allied to H. grandidens from which it is distinguished by its more ovate leaves and large, rounded teeth on the leaves. The teeth on the leaves of H. grandidens are narrower and more pointed and the leaves tend to be more oblong. Introduced. Roadside and railway banks, an old quarry and a wall. Recorded from Devonshire, Cambridgeshire, Shropshire, Westmorland, Lanarkshire, Perthshire and Aberdeenshire, all in the twentieth century. Widely recorded from France to the Caucasus. 375. H. adelphicum P. D. Sell Glamorganshire Hawkweed H. praecox subsp. recensitum microgen. fraternum Sudre; H. praecox subsp. fraternum (Sudre) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–55 cm, pale yellowish-green, sometimes slightly

401

purplish towards the base, rather slender, striate, with few to numerous, long, pale simple eglandular hairs below, and numerous stellate hairs and a few, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath and sometimes tinted purplish; basal few to numerous, not clearly divided into outer and inner, the lamina 2–9 × 1.5–4.0 cm, lanceolate, ovate or ovate-oblong, pointed but mostly obtuse at apex, markedly dentate, the teeth unequal and mostly broadly mammiform, truncate at base, often with 2 retrorse teeth, the petioles up to 8 cm, with dense, long, pale simple eglandular hairs; cauline 0–1, with lamina lanceolate or ovate, pointed or caudate at apex, markedly dentate with large mammiform teeth, subtruncate at base, petiolate; all with short to medium, rather stiff, pale simple eglandular hairs on the upper surface and longer ones beneath, especially on the midrib, and on the margin. Inflorescence with 4–15 capitula, openly paniculate-corymbose, mostly with a long lower branch; peduncles long and slender, with dense stellate hairs and numerous, very short and short (0.2–0.5 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–9 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with dense, unequal, very short to short (0.2–0.7 mm), dark glandular hairs, and dense stellate hairs on the margin and scattered elsewhere, without simple eglandular hairs. Ligules yellow, with numerous, very short simple eglandular hairs at the apex. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Included by A. Ley in his H. cinderella, but different from that species in its markedly different involucral bracts. Collected in some quantity by A. Ley in the neighbourhood of Caerphilly, Glamorganshire in 1906. Could be introduced. Central Europe from south-west France to Hungary. 376. H. firmiramum Hyl. Dense-branched Hawkweed Phyllopodous perennial herb with a branched stock. Stem 45–90 cm, pale green, purplish below, robust, striate, with numerous, pale, long simple eglandular hairs below, becoming shorter and fewer upwards, and numerous stellate and numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath and sometimes flushed purplish; basal numerous, the outer with lamina 6–8 × 3–4 cm, ovate or elliptical, rounded at apex, entire or denticulate in upper half, dentate below with large mammiform teeth and subtruncate, often asymmetrically at base, the inner with lamina 8–11 × 3.0– 4.5 cm, broadly oblong, rounded or obtuse-mucronulate at apex, dentate in the lower two-thirds with large mammiform, sometimes acute-mammiform teeth, and truncate at base, the petioles 5–13 cm, with numerous, long, pale simple eglandular hairs; cauline 0–1, ovate, long acuminate at apex, dentate, with large, acute-mammiform teeth, truncate at base, usually petiolate; all with short to long simple eglandular hairs on both surfaces and the margins, especially the midrib beneath. Inflorescence with numerous capitula, paniculate-corymbose; peduncles rather short and curved,

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with dense stellate hairs, numerous very short to short (0.2–0.6 mm), fine, dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–9 × 0.7–1.1 mm, blackish-green, the inner with paler margins, all narrow linear-lanceolate, acute at apex, with dense, very short to short (0.2–0.7 mm), fine, dark glandular hairs, without stellate or simple eglandular hairs. Ligules yellow, glabroustipped. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-brown. Flowers 5–6. Apomictic. Introduced. Common on chalk under trees, near Royston, Cambridgeshire. Described from Sweden where it is probably not native. 377. H. onychodontum Hyl. Giant-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 40–75 cm, pale green, often purplish near the base, robust, striate, with numerous, long, pale simple eglandular hairs at the base becoming fewer and shorter upwards, and with few to numerous stellate hairs and numerous, short to medium, dark glandular hairs in the upper part. Leaves medium yellowish-green and often mottled on upper surface, paler beneath; basal all similar, few, the lamina 6–12 × 2–4 cm, broadly oblong, rounded-mucronulate to subacute at apex, deeply dentate except sometimes in the upper third, the teeth large and mammiform, sometimes narrowly and sharply mammiform, subtruncate at base, often asymmetrically, and the basal teeth sometimes retrorse, the petioles 6–12 cm, often purplish, with dense, long, pale, wavy simple eglandular hairs; cauline 0–1, when present like basal or with lamina ovate, acute at apex, toothed like basal, subtruncate and petiolate; all with numerous, short or medium, pale simple eglandular hairs on both surfaces and the margin and particularly on the midrib beneath. Inflorescence with numerous capitula, paniculate-corymbose; peduncles short and curved, with numerous to dense stellate hairs, numerous very short to medium, fine, dark glandular hairs and without simple eglandular hairs. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3–9 × 0.8–1.0 mm, olive-green, the inner with paler margins, all linear-lanceolate, gradually narrowed to an acute apex, with dense, very unequal, very short to medium (0.2– 1.1 mm), slender, dark glandular hairs, and few stellate hairs on the margin, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 5–6. Apomictic. Introduced. Frequent on chalk at Royston in Cambridgeshire, near Kielder Station in Northumberland and Lochawe village in Argyllshire. Described from Sweden where it is probably not native either. 378. H. quadridentatum Hyl. Four-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 40–70 cm, pale yellowish-green, suffused purplish towards the base, robust, striate, with few to fairly numerous, long, pale simple eglandular hairs below, and fairly numerous stellate hairs and short and medium, dark glandular hairs in the upper part. Leaves rather pale yellowish-green on upper

surface, paler beneath and heavily flushed with reddishpurple; basal few to fairly numerous, the outer with lamina 5–8 × 3–4 cm, ovate or oblong-ovate, rounded-mucronate at apex, entire in the upper part, with narrow to broad mammiform teeth below and the basal pair sometimes retrorse, the inner with lamina 5–11 × 3.5–4.5 cm, ovate or oblongovate, sometimes almost oblong, acute at apex, with large, rounded or narrowly mammiform teeth and mostly truncate at base, the petioles up to 7 cm, with numerous, long, pale simple eglandular hairs; cauline 1, large, 9–10 × 4–6 cm, ovate, with a long-acute apex, dentate with large mammiform teeth, truncate at base, petiolate; all with rather short to medium, pale simple eglandular hairs on both surfaces and with longer ones on the margin and midrib beneath. Inflorescence characteristic, paniculate-corymbose, large and open with numerous capitula; peduncles rather short, rigid and curved or divaricate, with dense stellate hairs and numerous, very short to medium (0.2–1.0 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25– 30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, greyish-green, the inner with paler margins, the outer narrow linear-lanceolate, the inner very narrow, all acute at apex, with dense, unequal, very short to medium (0.2–1.0 mm), dark glandular hairs, without stellate or simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 5–7. Apomictic. Introduced. Known only from two specimens collected at Buckland in the Moor in Devonshire in 1959. Described from S¨odermanland, Sweden, where also it may not be native. 379. H. seriflorum Hyl. Gigantic-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–70 cm, pale yellowish-green, robust, striate, with numerous, long, pale simple eglandular hairs below, becoming fewer upwards, and numerous stellate and short, dark glandular hairs in the upper part. Leaves medium yellowishgreen on upper surface, paler and bluish beneath; basal numerous, the outer with lamina 2.5–10 × 2.0–6.0 cm, subrotund to ovate or elliptical, rounded mucronulate at apex, with large mammiform teeth at least in the lower half and truncate at base, the inner large, with lamina 9–16 × 4–7 cm, oblong or narrowly elliptical, obtuse to more or less acute at apex, with large, irregular, rounded, mammiform teeth in all but the upper third, the largest near the base and the lowest pair sometimes retrorse, subtruncate at base, the petioles up to 12 cm, with numerous, long, pale simple eglandular hairs; cauline 1, large, 6–12 × 3–7 cm, broadly ovate, acuminate at apex to a long caudate tip, deeply and irregularly mammiform-dentate, the teeth large and pointed, truncate at base, petiolate; all with scattered to fairly numerous, short to medium, pale simple eglandular hairs on the upper surface and longer and more numerous ones beneath and on the margins, especially on the midrib, sometimes with stellate hairs beneath. Inflorescence with 8–20 capitula, paniculatecorymbose, sometimes with a long lower branch; peduncles short, slender and curved, with dense stellate hairs and numerous, very short to short, dark glandular hairs,

39. Hieracium without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts 4–11 × 0.8– 1.2 mm, pale to medium green, the inner with even paler margins, all narrow linear-lanceolate, acute at apex, with dense, very unequal, very short to medium (0.2–1.5 mm), blackish glandular hairs and a few stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 5–7. Apomictic. Introduced. Recorded only from Cirencester in Gloucestershire, Witley in Surrey, a chalk cutting near Royston in Cambridgeshire, wood above Kilmuir in Ross-shire and Altamont House, Tullow, Co. Carlow in Ireland. Described from Sweden where also it is probably not native. 380. H. candelabrae W. R. Linton Candelabra Hawkweed H. murorum subsp. candelabrae (W. R. Linton) Zahn; H. exotericum var. candelabrae (W. R. Linton) Pugsley Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often suffused purplish below, robust, striate, with numerous, medium and long, pale simple eglandular hairs below, becoming fewer upwards, and numerous stellate hairs and numerous, short and medium, dark glandular hairs in the upper part. Leaves rather thick and leathery, medium yellowish-green on upper surface, paler and bluish beneath and often suffused with reddish-purple; basal numerous, the outer with lamina 3– 6 × 2–5 cm, broadly ovate or broadly elliptical, obtusemucronate at apex, dentate, especially in the lower half, the teeth more or less mammiform, and truncate at base, the inner with lamina broadly elliptical or oblong-ovate, obtuse to acute at apex, dentate in the lower half with mammiform teeth and cordate to truncate at base or the inner sometimes similar to outer but often the lamina narrower and more acute at apex, sometimes all leaves like the outer, the petioles up to 6 cm, with dense, long, pale simple eglandular hairs; cauline 0–1(–2), the lamina often large, ovate or ovate-oblong, acute at apex, dentate in the lower half, petiolate; all glabrous or nearly so on upper surface, with numerous, short to long, pale simple eglandular hairs beneath, particularly on the midrib, and on the margin. Inflorescence of 2–20 capitula in a broad corymb, often with a long, lower branch; peduncles rather long, slightly curved or straight and divaricate, with dense stellate hairs and numerous, very short to medium (0.2–1.0 mm), black glandular hairs, without simple eglandular hairs. Capitula 35–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–14 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse to acute at apex, with dense, very short to medium (0.2– 1.5 mm), black glandular hairs, and numerous stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 5–7. Apomictic. Native. Known only from the limestone cliffs and pavement around Ingleborough in Yorkshire where it is rare and has not been seen recently. Endemic.

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381. H. subcrassum (Almq. ex Dahlst.) Johanss. Trackway Hawkweed H. serratifrons subsp. subcrassum Almq. ex Dahlst.; H. murorum subsp. subcrassum (Almq. ex Dahlst.) Zahn Phyllopodous perennial herb with a branched stock. Stems 45–70 cm, pale yellowish-green, sometimes brownishpurple towards the base, slender, striate, with numerous, short to long, pale simple eglandular hairs throughout, and few to numerous stellate and unequal, very short to medium, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath, sometimes with few dark blotches; basal few to numerous, inner and outer not particularly distinct, 5–15 × 2–6 cm, ovate, sometimes broadly so, oblong-ovate or elliptical, rounded-obtuse to subacute at apex, denticulate to dentate especially in the lower half, the teeth shallowly to narrowly mammiform, sometimes with 2 retrorse ones at base, more or less subtruncate, the petioles of medium length, with dense, long, pale simple eglandular hairs; cauline usually 1, large, 3–12 × 1– 6 cm, ovate, gradually narrowed to an acute or acuminate apex, serrate-dentate, with narrow, mammiform teeth, subtruncate at base, petiolate; all with few to numerous, short to medium, pale simple eglandular hairs on the upper surface and more numerous, medium to long ones on the lower surface (especially the midrib) and margin. Inflorescence paniculate-corymbose, with 4–14 capitula; the acladium fairly short; branches and peduncles slender and arcuate, with dense stellate hairs and numerous, very short to short (0.2–0.7 mm), slender, dark glandular hairs, without simple eglandular hairs. Capitula 30–40 mm in diameter, rounded to slightly cuneate at base. Involucral bracts incumbent in bud, 5–12 × 0.7–1.0 mm, pale blackish-green, the inner with paler margins, all narrowly linear-lanceolate, acute at apex, with few to numerous stellate hairs along the margins, with numerous, unequal, very short to medium (0.2– 1.2 mm), slender, dark glandular hairs, without simple eglandular hairs. Ligules yellow, very shortly pilose-tipped. Styles discoloured. Receptacle pits shortly dentate. Achenes 2.5–3.0 mm, blackish. Flowers 6–7. Apomictic. H. subcrassum is characterised by its broad, basal leaves, mammiform-dentate in the lower half, large, petiolate cauline leaf, discoloured styles and pale narrow involucral bracts with slender, unequal glandular hairs. Probably introduced. Banks by roads or tracks. Ashwood Dale in Derbyshire and Grisedale in Westmorland. Sweden. 382. H. cinderella (Ley) Ley Long-bracted Hawkweed H. serratifrons var. cinderella Ley; H. exotericum var. cinderella (Ley) Pugsley Phyllopodous perennial herb with a branched stock. Stem 18–80 cm, pale yellowish-green, sometimes purplish near the base, striate, usually rather slender, with few to numerous, short to long, pale simple eglandular hairs near the base, becoming fewer upwards and usually absent in the upper part, with few to numerous, unequal (0.2–0.8 mm), slender, dark glandular hairs in the upper parts, and few to numerous stellate hairs mostly in the upper part. Leaves rather pale to medium yellowish-green, paler and sometimes slightly bluish beneath; basal numerous, the outer

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375. H. adelphicum P. D. Sell 373. H. grandidens Dahlst.

379. H. seriflorum Hyl.

378. H. quadridentatum Hyl.

380. H. candelabrae W. R. Linton

381. H. subcrassum (Almq. ex Dahlst.) Johanss.

Inner and outer basal leaves of species of the Section Hieracium

39. Hieracium

405

382. H. cinderella (Ley) Ley 383. H. koehleri Dahlst.

385. H. kentii P. D. Sell

384. H. aterrimum Hyl.

387. H. elevatum P. D. Sell 386. H. stenstroemii (Dahlst.) Johanss.

Inner and outer basal leaves of species of the Section Hieracium

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with lamina 2–12 × 1.5–6.0 cm, elliptical to broadly ovate or oblong, rounded to obtuse at apex, often entire towards the apex and denticulate to dentate below, the teeth sharp and apiculate and rounded, truncate or subcordate at base, the inner with lamina ovate or oblong, usually more or less acute at apex, more deeply dentate, the teeth usually narrowly mammiform or aquiline-mammiform and the basal pair sometimes long and retrorse, the petioles usually long, up to 12 cm, pale and slender and with long, pale, shaggy simple eglandular hairs; cauline 0–1, with lamina lanceolate or ovate, acute at apex, very irregularly dentate with narrow, mammiform teeth, subtruncate to cuneate-based and obviously petiolate, all with few to numerous, short to medium, pale simple eglandular hairs on both surfaces, medium to long ones on the margin and midrib beneath. Inflorescence paniculate-corymbose, with 2–14 capitula, sometimes with a long lower branch; peduncles short to fairly long, arcuate, slender, the acladium usually rather long, with dense stellate hairs and numerous, slightly unequal ((0.4–)0.5(–0.6) mm), slender, dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded to cuneate at base. Involucral bracts porrect in bud, 6–12 × 0.9– 1.2 mm, pale to rather dark green with paler margins, long and narrowly linear-lanceolate, more or less acute at apex, with stellate hairs particularly on the margins and near the apex and numerous to dense, very unequal (0.2–1.0 mm), slender, dark glandular hairs, without simple eglandular hairs but the stellate ones near the apex are rather tufted. Ligules rather pale yellow, ragged, with short eglandular hairs at tip. Styles yellow to slightly discoloured. Achenes 3.0–3.5 mm, dark reddish. Flowers 5–7(–8). Apomictic. Native. Riverside rocks, roadsides and wood banks. South and central Wales and the border country in Breconshire, Glamorganshire, Montgomeryshire, Gloucestershire, Worcestershire, Herefordshire and Shropshire. There is an isolated locality at Bodnant Gardens in Denbighshire where it is presumably introduced. Endemic. 383. H. koehleri Dahlst. Koehler’s Hawkweed Phyllopodous perennial herb with a branched stock. Stems 40–60 cm, pale yellowish-green, often tinted reddish-purple towards the base, usually robust, striate, with numerous, medium to long, pale simple eglandular hairs below, becoming fewer upwards, and numerous stellate hairs and numerous, short, dark glandular hairs in the upper part. Leaves pale yellowish-green, sometimes with darker veins, paler beneath and rarely tinted reddish; basal numerous, the outer with lamina 5–7 × 2–4 cm, ovate or elliptical, roundedmucronulate at apex, dentate, with numerous, shallow, mammiform teeth in the lower two-thirds and a truncate base, the inner with lamina oblong, oblong-elliptical or elliptical, acute at apex, dentate in the lower two-thirds, the teeth narrowly mammiform and the lower pair often retrorse, the petioles up to 8(–15) cm, with dense, long, pale simple eglandular hairs; cauline (0–)1–2, ovate or lanceolate, acute or acuminate at apex, sharply dentate, with curved, mammiform teeth, petiolate; all with short, pale simple eglandular hairs on the upper surface to nearly glabrous and with numerous longer hairs beneath and on the margins. Inflorescence with 8–15 capitula, paniculate-

corymbose, sometimes subumbellate, often with a long lower branch; peduncles rather short and slender, with dense stellate hairs and numerous, short (0.3–0.6 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, acute at apex, with dense, very short to medium (0.2–1.0 mm) dark glandular hairs, and few stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabroustipped. Styles yellow, sometimes slightly discoloured when dry. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Roadside banks, walls and waste places. Surrey, Cambridgeshire–Hertfordshire border, Monmouthshire, Cumberland and Inverness-shire. Sweden. Named after Oscar Eug`ene K¨ohler (1851–1911). 384. H. aterrimum Hyl. Patent-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 50–65 cm, pale yellowish-green, often tinted purplish towards the base, robust, striate, with numerous, long, pale simple eglandular hairs below, becoming much fewer upwards, and numerous to dense stellate hairs and numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on the upper surface, paler beneath and sometimes tinted reddish-purple; basal few to numerous, all fairly similar except in size, the lamina 3– 12 × 2–5 cm, lanceolate, lanceolate-oblong, ovate or ovateoblong, most more or less acute at apex, some narrowed to an obtuse apex, irregularly dentate, the teeth of various sizes, mammiform or narrowly mammiform and spreading, the basal pair sometimes retrorse, the teeth interspersed with a few denticulations, mostly subtruncate at base, the petioles up to 8 cm, with dense, long, pale simple eglandular hairs; cauline 1–2, the lower like the basal and as large, petiolate, the upper a small version of the lower or bract-like; all with numerous, short to medium, pale simple eglandular hairs on the upper surface and numerous, medium to long simple eglandular hairs beneath and on the margin. Inflorescence with numerous capitula, rather open paniculatecorymbose, usually with a long, lower branch; peduncles straight or curved, slender, with dense stellate hairs and numerous, very short to short (0.2–0.7 mm), dark glandular hairs, without simple eglandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–9 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, acute at apex, with dense, very short to short (0.2–1.0 mm), dark glandular hairs, and numerous stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, with very short simple eglandular hairs at apex. Styles yellow, sometimes turning dark when dry. Receptacle pits with margins sharply dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. This species differs from H. grandidens in its spreading leaf-teeth and usually yellow styles. Introduced. Roadside and railway banks and waste places. Cheddar Gorge in Somerset, Surrey, Cambridgeshire–Hertfordshire border near Royston, a pine plantation

39. Hieracium near Brandon in Norfolk, Selsley in Gloucestershire, Machen in Monmouthshire, Harton in Shropshire, and a wall near Grantown-on-Spey Station in Morayshire. Described from Sweden where it may not have been native. 385. H. kentii P. D. Sell Kent’s Hawkweed Phyllopodous perennial herb with a branched stock. Stem 40–70 cm, pale yellowish-green, often purplish towards the base, strictly erect, striate, with numerous, medium to long, pale simple eglandular hairs below which become gradually fewer up the stem, and numerous stellate and numerous, very short to short, yellowish-black glandular hairs in the upper part. Leaves medium yellowish-green on the upper surface, paler beneath and not purplish; basal few to fairly numerous, the outer with lamina 4–10 × 2.5–6.0 cm, subrotund, broadly ovate, broadly elliptical, broadly rounded and mucronulate to obtuse at apex, undulate-dentate, the teeth more or less mammiform and truncate at base with the basal pair of teeth often retrorse, the inner when present, with lamina 6–10 × 2.5–5.0 cm, elliptical or oblongelliptical, subobtuse to more or less acute at apex, toothed like the outer, the petioles often short, but can be up to 8 cm and with numerous, long, pale simple eglandular hairs; cauline 0–1(–2), when present lanceolate to ovate, acute to acuminate at apex, dentate, usually sharply, but sometimes with mammiform teeth, truncate at base, sessile or shortly petiolate; all with numerous, short to medium, subrigid, pale simple eglandular hairs on both surfaces and longer ones on the margin and midrib beneath, the cauline with numerous to dense stellate hairs beneath. Inflorescence corymbose, often with a long lower branch, the acladium fairly short; peduncles mostly rather short and slender, with dense stellate hairs, numerous, very short to short, unequal, yellowish-black glandular hairs and occasional short, pale simple eglandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–9 × 1.0–1.2 mm, olive-green, the inner with paler margins, all narrow linear-lanceolate, gradually narrowed to a more or less acute apex, with dense, very short to very unequal, yellowish-black glandular hairs, and numerous stellate along the margin and at the base, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–6. Apomictic. Perhaps native. Grassy places and banks. East Grinstead and near Lodsworth in Sussex. Ashurst Park, West Peckham and Tunbridge Wells in Kent and Godalming, Caterham, Witley and Hindhead in Surrey. Endemic. Named after Douglas Henry Kent (1920–98). 386. H. stenstroemii (Dahlst.) Johanss. Stenstr¨om’s Hawkweed H. serratifrons subsp. stenstroemii Dahlst.; H. serratifrons var. stenstroemii (Dahlst.) W. R. Linton; H. murorum subsp. stenstroemii (Dahlst.) Zahn; H. cuneifrons var. decipiens Pugsley; H. serratifrons var. triangulare auct.; H. serratifrons var. torticeps auct.; H. subulatidens auct.;

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H. serratifrons var. lepistoides auct.; H. pseudostenstroemii Pugsley Phyllopodous perennial herb with a branched stock. Stem 40–80 cm, pale yellowish-green, often brownish-purple towards the base, more or less robust, striate, with more or less numerous, medium to long, pale simple eglandular hairs, and numerous stellate and short, dark glandular hairs in the upper part. Leaves medium yellowishgreen on upper surface, paler beneath, sometimes suffused purplish; basal often numerous, the outer with lamina 2– 9 × 2–6 cm, subrotund, ovate, elliptical or oblong-elliptical, rounded-mucronulate at apex, subentire to shallowly and broadly sinuate-dentate and sometimes with larger teeth at the base which is rounded, truncate or subcordate, the inner with lamina 6–14 × 3–7 cm, mostly elliptic-oblong to ovate-oblong, more or less acute or acuminate at apex, dentate, often with large, mammiform teeth, sometimes with the teeth sharper, the lower pair often retrorse, truncate or subcordate at base, the petioles up to 10 cm, with dense, long, pale simple eglandular hairs; cauline 0–1, when present often large, up to 13 × 7 cm, ovate to ovate-oblong or elliptical-oblong, acute or acuminate at apex, usually with large, irregular teeth, often mammiform, the basal pair often retrorse, truncate or subcordate at base, petiolate; all with short to medium, pale, appressed simple eglandular hairs on the upper surface, with similar ones on the lower surface, and longer ones on the margin and the midrib beneath. Inflorescence usually with numerous capitula, rather open paniculate-corymbose, usually with a long lower branch; peduncles rather short and often curved, with dense stellate hairs and numerous, short (0.4–0.7 mm), dark glandular hairs, without simple eglandular hairs. Capitula 40–50 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–12 × 1.2–1.8 mm, olive-green, the inner with paler margins, all linear-lanceolate, obtuse at apex, with dense, very short to medium (0.2–1.5 mm), very unequal, dark glandular hairs, and stellate hairs along the margin and near the base, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles yellow or slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 4.0– 4.5 mm, reddish-black. Flowers 5–7. Apomictic. Native. Woods and rocky places on limestone in quarries, roadside banks and railway cuttings. Gloucestershire, Herefordshire, Monmouthshire, Breconshire and Yorkshire. Norway and Sweden. Named after Karl Otto Edvard Stenstr¨om (1858–1901). H. stenstroemii and the following three species were put into one species by Sell and West (1968). The group made a tight distribution in south-east Wales and adjacent England. In the 1950s they were common in the area; their present status in not known. H. stenstroemii has large leaves with large teeth, some of the leaves at least being oblong in their shape. H. glevense has similar shaped rather smaller leaves without or with shallow teeth only in the lower half. H. elevatum had large leaves with shallow teeth. H. dowardense has ovate or elliptical leaves one of which is often like a large balloon, the toothing is variable, the heads are darker than the other three. Three of these species occur on the Great Doward in Herefordshire, but apparently not together.

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387. H. elevatum P. D. Sell Taf-fechan Hawkweed H. sparsidens var. elatius Ley; H. elatius (Ley) Druce, non Rehmann Phyllopodous perennial herb with a branched stock. Stem 40–80 cm, pale yellowish-green, sometimes tinted brownish-purple below, more or less slender and flexuous, striate, with more or less numerous, medium to long, pale simple eglandular hairs throughout, and numerous stellate and few to numerous, short glandular hairs in the upper part. Leaves flaccid, medium to dark green on upper surface, paler beneath; basal usually few, the outer with lamina 7–10 × 4–5 cm, elliptical or ovate, narrowed, but obtuse at apex, subentire or sinuate-denticulate or sinuate-dentate and cuneate to rounded or subcordate at base, the inner with lamina 4–14 × 3–8 cm, broadly ovate, ovate-elliptical or oblong-elliptical, acute, shortly acuminate or cuspidate at apex, sinuate-dentate, the teeth mammiform or sharply mammiform, the lowest pair sometimes cusped or retrorse, cuneate, truncate or subcordate at base, the petioles 3– 10 cm, slender and with numerous medium to long, pale, spreading simple eglandular hairs; cauline 1–3, the lower large, up to 10 × 6 cm, ovate, acute at apex, irregularly dentate, with large mammiform teeth, cuneate or truncate at base with petioles up to 4 cm, the upper similar, but smaller and sometimes bract-like; all with numerous, short to medium, appressed, pale simple eglandular hairs on the upper surface, similar but long hairs beneath and on the margin, and especially on the midrib. Inflorescence with 2–20 capitula, fairly open cymose-corymbose, usually with a long lower branch; peduncles short to fairly long, erect or curved, with dense stellate hairs and numerous, short to medium (0.3–1.5 mm), fine, dark glandular hairs, without simple eglandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3– 10 × 1.3–1.5 mm, olive-green, the inner with paler margins, linear-lanceolate, obtuse at apex, with dense, very short to medium (0.2–1.7 mm), fine, dark, very unequal glandular hairs, and few stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins subulate-dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Known only in the wooded, rocky glen of the Taf-fechan, Breconshire where Augustin Ley found it in abundance in 1908 and where it has not been seen since, and in a hedgebank at Capel-y-ffin where it was re-found in 1953. It approaches H. radyrense in its broad, 1–3 cauline leaves, but that species has larger more stellately hairy involucres. Endemic. 388. H. dowardense P. D. Sell Doward Hawkweed Phyllopodous perennial herb with a branched stock. Stem 40–70 cm, pale yellowish-green, often brownish-purple towards the base, slender to robust, striate, with few to numerous, medium to long, pale simple eglandular hairs, and numerous stellate and short, dark glandular hairs above. Leaves medium yellowish-green on upper surface, paler beneath, sometimes tinted red; basal usually numerous, the outer with lamina 3–10 × 2–6 cm, at least one of them large and subrotund or very broadly elliptical, sometimes ovateelliptical, rounded-mucronulate at apex, sinuate-dentate in

the lower half, sometimes denticulate in the upper half and rounded, truncate or subcordate at base, the inner with lamina 6–9 × 2.5–6.0 cm, usually ovate or elliptical, sometimes elliptic-ovate, rounded-obtuse to more or less acute at apex, sinuate-dentate, with the teeth mammiform or sharply mammiform, and truncate or subcordate at base, the petioles up to 9 cm, with dense, long, pale simple eglandular hairs; cauline 0–1, when present with the lamina lanceolate or ovate, acute or acuminate at apex, more or less dentate, sometimes with the teeth large and mammiform, truncate at base, petiolate; all with short to medium, pale simple eglandular hairs on both surfaces and long ones on the margins and petiole. Inflorescence usually with numerous capitula and often with a long lower branch, paniculate-corymbose; peduncles mostly rather short, with dense stellate hairs and numerous, short (0.2–0.7 mm), dark glandular hairs, without simple eglandular hairs. Capitula 35–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–13 × 0.8–1.5 mm, blackish-green, the inner with paler margins, all linearlanceolate, obtuse to acute at apex, with dense, very unequal, very short to medium (0.2–1.5 mm), black glandular hairs, stellate hairs on the margins and near the base, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–7. Apomictic. This species sits between H. stenstroemii and H. glevense, differing from both in having subrotund, ovate or broadly elliptical leaves, one of which is often large and balloonlike, and the stem leaf, when present, usually large, ovate, narrowly acute at apex and with large teeth. The heads are darker than in both the other species. H. dowardense has long been known on the slopes of the Great Doward in Herefordshire and is probably native. It is also recorded from the Caplar slip in the same county, a wall and stream in Tintern village, Monmouthshire, and Capel-y-ffin in Breconshire. Endemic. 389. H. glevense (Pugsley) P. D. Sell & C. West Gloucester Hawkweed H. exotericum var. glevense Pugsley; H. stenstroemii var. subcordatum Pugsley; H. serratifrons var. crassiceps auct.; H. serratifrons var. lepistoides auct.; H. serratifrons var. torticeps auct. Phyllopodous perennial herb with a branched stock. Stem 25–70 cm, pale yellowish-green, often tinted brownishpurple below, striate, slender and flexuous to robust, with few, medium or long, pale simple eglandular hairs, mostly below, and few to numerous stellate hairs and short, dark glandular hairs above. Leaves medium yellowish-green on upper surface, paler beneath; basal often numerous, the outer with lamina 2–9 × 1.5–4.5 cm, subrotund, ovate, elliptical or oblong-elliptical, obtuse-mucronulate to subacute at apex, subentire or shallowly dentate towards the base and rounded or subcordate at base, the inner with lamina 4.0–10.5 × 1.5–4.0 cm, often oblong-elliptical, sometimes ovate, elliptical or lanceolate, more or less acute at apex, subentire, dentate or sinuate-dentate, the teeth usually shallow and sharply pointed and cuneate to subcordate at base, the petioles up to 8 cm, and with dense, long, pale

39. Hieracium simple eglandular hairs; cauline 0–1, ovate, shortly acuminate or cuspidate, subentire or toothed and petiolate, sometimes linear, acute at apex, subentire or toothed and sessile; all with numerous, short to medium, pale, appressed simple eglandular hairs on the upper surface and numerous similar ones beneath, with longer ones on the midrib and on the margin. Inflorescence with 2–numerous capitula, cymose-corymbose, often with a long lower branch; peduncles rather short, straight to curved, with dense stellate hairs and numerous, short (0.4–0.7 mm), fairly uniform on any one plant, black glandular hairs, without simple eglandular hairs. Capitula 35–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–13 × 0.8–1.5 mm, olive-green, the inner with paler margins, all narrow linearlanceolate, obtuse to more or less acute at apex, with numerous, very short to medium (0.2–1.5 mm), very unequal, dark glandular hairs, and some stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabroustipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.5–4.0 mm, reddish-black. Flowers 5–7. Apomictic. This species differs from H. stenstroemii in its leaves being subentire to shallowly toothed towards the base and its narrow involucral bracts. Woods, walls, quarries, roadsides and railway banks, and grassy cliffs, in some places looking native, in others introduced. Leigh Woods by Bristol in Somerset, Gloucestershire, Herefordshire, Breconshire and Aberedw Rocks in Radnorshire, extending east to Buckinghamshire and Berkshire. Endemic. 390. H. monstrosum Hyl. Nipple-toothed Hawkweed Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, sometimes flushed brownish-purple towards the base, slender to robust, striate, with few to numerous, medium to long, pale simple eglandular hairs towards the base and very few upwards, with numerous stellate hairs and numerous, short, rather slender, dark glandular hairs in the upper part. Leaves medium yellowish-green, spotted and blotched with purple on the upper surface, or without spotting or blotching in later flowering plants, paler beneath; basal numerous, hardly divided into outer and inner, the lamina 4–10 × 2.5–4.0 cm, oblongelliptical, oblong-lanceolate, oblong or ovate, narrowed to a rounded-mucronate point, entire in the upper third, dentate in the lower two-thirds, the teeth getting longer towards the base and mammiform or narrowly mammiform with a long mucro at tip, subtruncate at base, the petioles up to 7 cm, with dense, long, pale simple eglandular hairs; cauline 0–1, the lamina ovate, long-acute or acuminate at apex, sharply dentate, abruptly contracted at base, petiolate, or sometimes linear and bract-like; all with short to medium, rather stiff, pale simple eglandular hairs on the upper surface and longer, softer ones beneath especially on the midrib and margins. Inflorescence with 2–20 capitula, paniculatecorymbose, often with a long lower branch; peduncles with dense stellate hairs and numerous, short (0.3–0.5 mm), dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts 4–11 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or

409

less acute at apex, with numerous, very short to medium (0.2–1.0 mm), dark glandular hairs, and numerous stellate hairs along the margin, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins subulate-dentate. Achenes 3.0– 3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Common on roadside and railway banks at Tavistock in Devonshire. Described from S¨odermanland in Sweden, but may not be native there either. 391. H. medium Jord. Tavistock Hawkweed H. murorum var. medium (Jord.) Gren. & Godr.; H. praecox subsp. petiolare var. medium (Jord.) Sudre; H. praecox subsp. medium (Jord.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–45 cm, pale yellowish-green, often suffused brownishpurple, slender, flexuous, striate, with few to numerous, long, pale simple eglandular hairs below, slowly dwindling upwards, and numerous stellate hairs and short, dark glandular hairs in the upper part. Leaves medium yellowishgreen, spotted and blotched with brownish-purple on upper surface, paler beneath; basal few, not clearly differentiated into outer and inner, the lamina 5.5–7.0 cm, lanceolate, ovate or elliptic-oblong, narrowed and mucronulate to acute at apex, entire in the upper half to denticulate or shallowly serrate below, the teeth narrow and sharp, subtruncate at base, the petioles up to 7 cm, with dense, long, wavy, pale simple eglandular hairs; cauline 0–1, with lamina lanceolate, acute at apex, dentate towards the subtruncate base, petiolate; all with short or medium, pale simple eglandular hairs on the upper surface and long ones beneath and on the margins. Inflorescence with 2–9 capitula, rather compactly corymbose; peduncles short and slender, with dense stellate hairs and numerous, short (0.3–0.5 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.5 mm, blackish-green, the inner with paler margins, all linear-lanceolate, long drawn out to an acute apex, with numerous, short (0.4–0.7 mm), dark glandular hairs, and stellate hairs along the margin, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Roadsides and railway banks about Tavistock in Devonshire where it grows with H. monstrosum, which is also a spotted-leaved plant. It can be distinguished from that species by its shorter, less dentate leaves, yellow styles and shorter glandular hairs of the involucre. It occurs also in France and Spain. 392. H. hortense Hyl. Garden Hawkweed Phyllopodous perennial herb with a branched stock. Stem 20–30 cm, pale yellowish-green, sometimes purplish towards the base, slender, striate, with numerous, medium to long, pale simple eglandular hairs below, becoming fewer upwards and with numerous stellate hairs and few to numerous, short glandular hairs in the upper part. Leaves bluishgreen on upper surface, paler beneath; basal numerous, not divisible into 2 kinds, the lamina 2.5–6.0 × 1.5–2.5 cm, ovate or lanceolate, sometimes oblong-lanceolate, acute at

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388. H. dowardense P. D. Sell 389. H. glevense (Pugsley) P. D. Sell & C. West

391. H. medium Jord. 390. H. monstrosum Hyl.

392. H. hortense Hyl.

393. H. subaequialtum Hyl.

Inner and outer basal leaves of species of the Section Hieracium

39. Hieracium

394. H. sublepistoides (Zahn) Roffey

411

395. H. cardiophyllum Jord. ex Sudre

397. H. sylvivagum Jord. ex Bor. 396. H. severiceps Wiinst.

398. H. microspilum (Jord. ex Sudre) A. W. Hill 399. H. oblongum Jord. ex Bor. Inner and outer basal leaves of species of the Section Hieracium

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apex, dentate with regular small teeth in the lower half, truncate or cuneate at base, the petioles up to 5 cm, with long, dense, pale simple eglandular hairs; cauline 0–1, with lamina ovate, narrow acute or caudate at apex, dentate with small teeth, sessile or shortly petiolate; all with numerous short to medium, stiff, pale simple eglandular hairs on the upper surface and more numerous, longer ones below and on the margins. Inflorescence with 4–8 capitula, in a fairly tight corymb, sometimes with a lower branch; peduncles short and thin, with numerous, very short to short (0.2–0.5 mm), dark glandular hairs and occasional to few, short to medium, pale simple eglandular hairs. Capitula 15–20 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.7–1.0 mm, greyish-green, the inner with paler margins, all with numerous, very short to short (0.2–0.5 mm), dark glandular hairs, few stellate hairs and occasional to few, pale to dark, short to medium simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Abundant on grassy banks of Walthamstow Reservoirs in Essex in 1953. Described from a garden in G¨oteborg, Sweden and therefore not known anywhere in the wild. 393. H. subaequialtum Hyl. Black-bracted Hawkweed Phyllopodous perennial herb with a branched stock. Stem 25–60 cm, pale yellowish-green, sometimes brownishpurple below, more or less slender, with few to numerous, medium to long, pale simple eglandular hairs below becoming fewer upwards, and numerous stellate hairs and numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath and sometimes tinted purplish; basal few to numerous, not distinctly divisible into 2 kinds, with lamina 2–8 × 1.5–3.5 cm, ovate, lanceolate or oblong-lanceolate, obtuse-mucronate to acute at apex, subentire to denticulate or with a few sharp teeth at the base, subtruncate or rounded at base, the petioles up to 7 cm, with numerous to dense, long, pale simple eglandular hairs; cauline 0–1, ovate or lanceolate, acute or acuminate at apex, subentire or denticulate, rounded or subtruncate at base, sessile or petiolate; all with short to medium, stiff, appressed, pale simple eglandular hairs on the upper surface and longer ones beneath and on the margins. Inflorescence with 5–15 capitula, paniculate-corymbose, sometimes with a long lower branch; peduncles straight to curved, slender, with dense stellate hairs, numerous, very short to short (0.2–0.7 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 0.7–1.0 mm, blackish-green, the inner with paler margins, all narrow linear-lanceolate, the inner very narrow, acute at apex, with numerous to dense, unequal, very short to medium (0.2–1.5 mm), black glandular hairs and few stellate hairs, without simple eglandular hairs. Ligules yellow, with very short, simple eglandular hairs at apex when young. Styles slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic.

This plant has long been known as black-headed H. sublepistoides, being distinguished from true H. sublepistoides by its blackish-green involucral bracts, which are covered with black glandular hairs with hardly any stellate hairs. Probably always introduced. It is common in the Derbyshire Dales where it is mostly on the limestone, but in disturbed habitats. It also occurs on roadsides and railway banks in Monmouthshire, Glamorganshire, Radnorshire, Southport dunes in Lancashire, Invergarry in Invernessshire and St Peter Port in Guernsey. H. subaequialtum was described from Sweden where it may not be native either. Some small plants of H. glevense approach it, but that species has larger heads. H. hortense is very similar in appearance, but has some simple eglandular hairs on the involucral bracts. 394. H. sublepistoides (Zahn) Druce Grey-bracted Hawkweed H. murorum subsp. lepistoides var. sublepistoides Zahn; H. exotericum var. sublepistoides (Zahn) Pugsley Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, yellowish-green, sometimes purplish towards the base, slender and flexuous, striate, with few to numerous, medium to long, pale simple eglandular hairs below, becoming fewer upwards, and with numerous stellate hairs and numerous, short, dark glandular hairs in the upper part. Leaves yellowish-green, often with spots and blotches of brownish-purple on upper surface, paler beneath and often suffused purple; basal numerous, not obviously divided into outer and inner, the lamina 2.5–7.0(–9.0) × 1.5–3.5 cm, mostly ovate, sometimes subrotund, elliptical or lanceolate, rounded to more or less acute at apex, subentire or shallowly sinuate-dentate in the lower half, truncate at base, the petioles up to 4 cm, with numerous, long, pale simple eglandular hairs; cauline 0–1, the lamina ovate, acute at apex, subentire or with a few shallow teeth, sessile or petiolate; all with numerous, short, stiff, pale simple eglandular hairs on the upper surface and medium to long ones beneath and on the margin. Inflorescence with 2–10 capitula, paniculatecorymbose, sometimes subumbellate; peduncles short and curved, with dense stellate hairs and numerous, very short to short (0.2–0.5 mm), fine, dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4– 10 × 0.8–1.0 mm, greyish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with numerous, unequal, very short to medium (0.2–1.0 mm), slender, dark glandular hairs, and dense stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 5–7. Apomictic. Introduced. Woods, roadsides and walls. Scattered records in Great Britain north to Cumberland. Central Europe. 395. H. cardiophyllum (Jord. ex Sudre) Juxip Heart-leaved Hawkweed H. murorum subsp. exotericum microgen. cardiophyllum Jord. ex Sudre; H. murorum subsp. cardiophyllum (Jord.

39. Hieracium ex Sudre) Zahn; H. murorum var. cardiophyllum (Jord. ex Sudre) Rouy Phyllopodous perennial herb with a branched stock. Stem 50–70 cm, pale yellowish-green, sometimes purplish towards the base, more or less slender and flexuous, striate, with numerous, long, pale simple eglandular hairs at the base becoming fewer upwards, and few stellate hairs and few to fairly numerous, short, fine, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler, bluish and sometimes tinted reddish-purple beneath; basal usually numerous, all more or less similar except for size, the lamina 4–12 × 2–7 cm, broadly ovate or broadly elliptical, obtuse-mucronate to more or less acute at apex, regularly sinuate-denticulate or with a few shallow teeth towards the base, cordate or truncate at base, sometimes asymmetrically so, the petioles up to 10 cm, with few to numerous, long, pale simple eglandular hairs; cauline (0–)1(–2), with lamina ovate, acute at apex, regularly denticulate, cordate at base, petiolate; all with a few, scattered, short, pale simple eglandular hairs on the upper surface, more numerous longer hairs beneath especially on the midrib and numerous, short to long hairs on the margin. Inflorescence with numerous capitula, paniculatecorymbose, often with a long, lower branch; peduncles rather short and slender, with dense stellate hairs and numerous, more or less equal, short (0.4–0.5 mm), dark glandular hairs, without simple eglandular hairs. Capitula 20– 30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–9 × 0.7–1.2 mm, greyish-green, the inner with paler margins, all narrowly linear-lanceolate, obtuse at apex, with numerous, very short to medium (0.2– 1.0 mm), dark glandular hairs, and fairly numerous stellate hairs mainly on the margin, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles yellowish to discoloured. Receptacle pits with margins shortly dentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Abundant on a railway bank at Shilling Bridge near Dunton Green in Kent, near Royston in Cambridgeshire and on a streamside between Llangattock village and Llangattwg Quarries in Breconshire. A specimen from the border of a drive in Rhu, Dunbartonshire differs only in having stellate hairs on the under side of its cauline leaf. South-west and central Europe east to Russia. 396. H. severiceps Wiinst. Strict-headed Hawkweed Phyllopodous perennial herb with a branched stock. Stems 40–80 cm, yellowish-green, often purplish towards the base, rather robust, faintly striate, with few to numerous, short to long, pale simple eglandular hairs mainly in the lower half and few to numerous, very short to short, fine, dark glandular hairs and few to numerous, dense stellate hairs in the upper half. Leaves medium yellowish-green on upper surface, sometimes suffused purplish, paler and greyishgreen beneath, with few to numerous, short, pale simple eglandular hairs on the upper surface and numerous, medium to long ones on the under surface and margin, particularly the midrib; basal rather few, 3–10 × 2–4 cm, the outer with lamina broadly elliptical to ovate, rounded-

413

mucronulate at apex, subentire to undulate-denticulate or undulate-dentate and rounded or subtruncate at base, the inner with lamina lanceolate to ovate, more or less acute at apex, more or less dentate or serrate-dentate in the lower two-thirds, the teeth mammiform or sometimes aquilinemammiform and cuneate to truncate, often asymmetrical at base, the petioles up to 8 cm and with dense, long, pale simple eglandular hairs; cauline 0–1, when present with lamina usually lanceolate, rarely ovate, long-acute or acuminate at apex, toothed like the inner, cuneate or subtruncate at base and petiolate. Inflorescence cymose-corymbose, often with a long lower branch, with up to 22 capitula; peduncles with numerous to dense stellate hairs and numerous, short, fairly equal, dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 0.9–1.2 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, gradually narrowed to an obtuse or subacute apex, with numerous, short to medium, unequal, black glandular hairs, and very few stellate hairs on the margins, without simple eglandular hairs. Ligules medium yellow, shortly pilose-tipped. Styles yellow. Receptacle pits with subulate-dentate margins. Achenes 3.5–4.0 mm, reddishblack. Flowers 5–7. Apomictic. Introduced. Roadsides, railway banks and a quarry. Sussex, Derbyshire, Lancashire, Yorkshire, Durham, Fifeshire, Dunbartonshire, Argyllshire and Ross-shire. Denmark and Sweden. 397. H. sylvivagum Jord. ex Boreau Wandering Hawkweed H. murorum subsp. murorum microgen. gentile var. sylvivagum (Jord. ex Boreau) Sudre; H. murorum subsp. gentile var. sylvivagum (Jord. ex Boreau) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–50 cm, pale yellowish-green, sometimes purplish towards the base, slender, flexuous, striate, with numerous long, pale simple eglandular hairs below, becoming fewer and shorter upwards, and numerous stellate hairs and few to numerous, short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler and sometimes tinted reddish beneath; basal usually numerous, not clearly divided into outer and inner, the lamina 2–8 × 1.5–4.0 cm, mostly broadly triangular-ovate, sometimes lanceolate or oblong-ovate, mostly more or less acute at apex, but sometimes rounded-obtuse-mucronulate, shortly dentate in lower three-quarters, the teeth small and sharply apiculate, rounded or subtruncate at base, the petioles up to 8 cm and with dense, long, pale, wavy simple eglandular hairs; cauline 1–2, the lower usually large, like the basal and petiolate, the upper narrower or bract-like; all with short, rather stiff, pale simple eglandular hairs on the upper surface to nearly glabrous, and longer, softer ones beneath and on the margin. Inflorescence with 2–8 capitula, rather compactly corymbose; peduncles short and slender, with dense stellate hairs and dense, very short or short (0.2– 0.7 mm), black glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all narrowly

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linear-lanceolate, with a long, narrow, acute apex, with dense, very short to medium (0.2–1.0 mm), black glandular hairs, and numerous stellate hairs along the margin, without simple eglandular hairs. Ligules yellow, glabroustipped. Styles yellow to slightly discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Roadside and railway banks, old walls and riverside rocks. Caterham in Surrey; Symonds Yat and Llanthony Abbey in Monmouthshire; Buildwas in Shropshire; Glyn Neath and Castell Coch in Glamorganshire; riverside rocks at Abercraf (Abercrave) in Breconshire; and Grey Abbey in Co. Down. Central Europe from France to Russia. 398. H. microspilum (Jord. ex Sudre) A. W. Hill Minute-toothed Hawkweed H. murorum subsp. murorum microgen. gentile var. microspilum Jord. ex Sudre; H. murorum subsp. microspilum (Jord. ex Sudre) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–55 cm, pale yellowish-green, often tinted purplish towards the base, erect, slender, flexuous, striate, with numerous, medium to long, pale simple eglandular hairs below becoming less numerous upwards, and stellate and short, dark glandular hairs in the upper part. Leaves medium yellowish-green on the upper surface, paler beneath; basal usually numerous, the outer with lamina 3–7 × 1.5– 4.0 cm, ovate, oblong or oblong-ovate, rounded-obtuse at apex, entire or with minute denticulations and subtruncate, rounded or subcordate at base, the inner with lamina 6– 8 × 2.0–3.5 cm, narrowly elliptical, ovate or lanceolate, more or less acute at apex, entire or minutely denticulate, and rounded or subcordate at base, the petioles up to 6 cm, with dense, long, curly, pale simple eglandular hairs; cauline 0–1, when present with lamina ovate or lanceolate, acute or acuminate at apex to a slender point, entire or denticulate, shortly petiolate; all with rather stiff, short to medium, pale simple eglandular hairs on upper surface and numerous medium to long ones beneath, particularly on the midrib, and on the margins. Inflorescence with 2–7 capitula, fairly tightly corymbose; peduncles short and slender, with dense stellate hairs and numerous, more or less equal, short (0.3– 0.4 mm), dark glandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with numerous, very short to medium (0.2–1.0 mm), black glandular hairs, and numerous stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, with few to numerous, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced or possibly native. H. C. Watson cultivated this plant in his garden in Surrey in 1853 and said it was wild at Long Ditton in north Surrey. It has been more recently collected in Surrey on chalk rubble near Juniper Hall, Dorking and at Coulsden. It was collected in the last century

on limestone rocks between Tintern and the Wyndcliffe in Monmouthshire and more recently in that county on limestone cliffs in Lady Park Wood above the Biblin. It occurs in central Europe from France to Hungary. 399. H. oblongum Jord. Weedy Hawkweed H. murorum var. oblongum (Jord.) Gren. & Godr.; H. murorum subsp. oblongum (Jord.) Zahn Phyllopodous perennial herb with a branched stock. Stems 30–50 cm, pale yellowish-green, often purplish towards the base, striate, often flexuous, with numerous, short to long, pale simple eglandular hairs towards the base, usually much fewer in the upper part and sometimes none, and few to numerous stellate hairs and short, dark glandular hairs near the inflorescence. Leaves yellowish-green on upper surface, paler and rather greyish-green beneath, often tinted or flushed purplish beneath and sometimes near the margin of the upper surface; basal numerous, with lamina 2–10 × 2.0–3.5 cm, the outer elliptical or ovate, obtuse at apex, entire or denticulate and rounded at base, the inner with lamina oblong, elliptic-oblong or lanceolate, usually acute at apex, entire or denticulate and sometimes with a few, larger, more or less acute teeth near the base, the petioles medium or long, with numerous, long, pale simple eglandular hairs; cauline 0–1(–2), when present with lamina lanceolate, acute or acuminate at apex, entire, denticulate or dentate, petiolate; all with few to numerous, short to medium, stiff, pale simple eglandular hairs on the upper surface and numerous to dense, medium to long, softer, pale simple eglandular hairs beneath and particularly on the midrib and margins. Inflorescence furcatecorymbose, often with a long lower branch and rather long acladium; peduncles more or less arcuate, with dense stellate hairs and numerous, short (0.3–0.7 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25– 35 mm in diameter, more or less rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, blackish, the inner with paler margins, all narrowly linear-lanceolate, the inner almost filamentous, acute at apex, with dense, short to medium (0.3–1.0 mm), very unequal, black glandular hairs, without or with very few stellate hairs and without simple eglandular hairs. Ligules medium yellow, glabrous-tipped. Styles more or less yellow or slightly discoloured, mostly discoloured when dry. Receptacle pits shortly dentate. Achenes 3.0–3.5 mm, blackish. Flowers 5–7. Apomictic. Introduced. Grassy banks, by railways and on walls. Chalkpit at Farlington in Hampshire; Walthamstow Reservoirs in Essex; wall of Wadham College at Oxford; grounds of Newnham College in Cambridge; railway and mine workings at Kilton in Yorkshire; Hallington Reservoir in Northumberland and wall-tops at Monasterevan in Co. Kildare. Central and west Europe. 400. H. integratum (Dahlst. ex Stenstr.) Dahlst. Toothless Hawkweed H. sylvaticum subsp. integratum Dahlst. ex Stenstr.; H. murorum subsp. integratum (Dahlst. ex Stenstr.) Zahn Phyllopodous perennial herb with a branched stock. Stems 40–75 cm, pale yellowish-green, sometimes slightly

39. Hieracium purplish below, slender to more or less robust, striate, with few to numerous, medium to long, pale, wavy simple eglandular hairs particularly in the lower part, but sometimes almost completely absent, with few to numerous stellate hairs and occasional to few, short glandular hairs in the upper part. Leaves pale yellowish-green on upper surface, paler and rather greyish beneath, sometimes tinted purplish; basal usually rather few, the lamina ovate, elliptical, lanceolate or more rarely subrotund, rounded-obtuse to acute at apex, entire, remotely denticulate or with a pair of mammiform teeth at the base, rounded or subtruncate at base, with fairly long petioles; cauline 0–1, with lamina lanceolate, long-acute or acuminate at apex, denticulate or with 1–2 small teeth, petiolate; all usually more or less glabrous on upper surface, sometimes with a few simple eglandular hairs, with numerous, short to long, pale, wavy simple eglandular hairs beneath and on the margin and particularly on the midrib. Inflorescence fairly compactly cymosecorymbose, with a medium acladium, sometimes with a long lower branch; peduncles arcuate, with dense stellate hairs and few to numerous, short (0.3–0.5 mm), dark glandular hairs, without simple eglandular hairs. Capitula 30– 35 mm in diameter, rounded at base. Involucral bracts 5–10 × 1.0–1.2 mm, blackish-green, the inner with paler margins or pale green with only the base dark, all linearlanceolate, gradually narrowed at tip to a more or less obtuse apex, with very few stellate hairs at base or at apex and numerous, short (0.3–0.7 mm), slightly unequal, fine, dark glandular hairs, without simple eglandular hairs. Ligules medium yellow, glabrous-tipped. Styles discoloured. Receptacle pits with subulate-dentate margins. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Native. Limestone cliffs and rocks. Ingleborough Scaurs in Yorkshire and Arnside Knott in Westmorland. It also occurs on a sandstone cliff by the Castle Museum at Nottingham where it may be introduced. Scandinavia, central Europe and Greece. 401. H. pellucidum Laest. Pellucid-leaved Hawkweed H. murorum subsp. pellucidum (Laest.) Zahn; H. murorum var. lucidulum Ley; H. pellucidum var. lucidulum (Ley) W. R. Linton; H. murorum subsp. lucidulum (Ley) Zahn; H. lucidulum (Ley) Roffey; H. melanolepis Almq. ex Norrlin; H. nigriglandulosum L¨onnr.; H. murorum var. pellucidum (Laest.) F. Hanb.; H. silvaticum var. pellucidum (Laest.) Almq.; H. silvaticum var. lucidulum (Ley) F. N. Williams Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often purplish towards the base, slender and flexuous to robust, striate, with few to fairly numerous, medium to long, pale simple eglandular hairs or with them almost absent and few to numerous stellate hairs and the occasional, short, dark glandular hair above. Leaves rather dark, shining green on upper surface, sometimes with a few purplish spots, paler bluishgreen beneath and often purpurescent; basal numerous, the outer with lamina 2–8 × 2–4 cm, subrotund, ovate or elliptical, obtuse-mucronate to broadly rounded at apex, entire to faintly undulate-denticulate or sometimes with a few

415

obscure teeth, and truncate or cordate at base, the inner with lamina 6–11 × 3–5 cm, ovate or oblong-ovate, more or less acute at apex, entire or with a few shallow teeth and cordate or truncate at base, sometimes the leaves are all like the outer, sometimes all like the inner, the petioles up to 10 cm, with numerous, long simple eglandular hairs; cauline 0–1, when present with lamina ovate, acute or acuminate at apex, denticulate or sharply dentate, truncate or cordate at base, petiolate; all glabrous or nearly so on upper surface, with few to numerous, medium to long, pale simple eglandular hairs beneath, particularly on the midrib, and on the margin. Inflorescence with 3–20(–50 cult.) capitula, paniculate-corymbose and sometimes subumbellate, often with a long, lower branch; peduncles mostly short, straight and slender, with dense stellate and numerous fairly uniform, short (0.4–0.5 mm), black glandular hairs, without simple eglandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 3– 9 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with dense, unequal, very short to medium (0.2–1.0 mm), black glandular hairs, and a few stellate hairs towards the base, without simple eglandular hairs. Ligules yellow, glabrous or with a very few, short hairs at the apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0– 3.5 mm, reddish-black with a particularly white pappus. Flowers 5–7. Apomictic. A. Ley originally thought the plants with the leaves all obtuse (lucidulum) could be distinguished from those with the leaves all more or less acute (pellucidum), but all combinations occur both here and in Scandinavia. The type specimen of H. pellucidum has acute leaves. H. pellucidum is an easily recognised species with glabrous upper surface of dark, shining leaves, small black capitula and almost white pappus. Native. Cliff ledges, grassy banks and streamsides. Common in the limestone areas of south Wales, the Derbyshire Dales and northern England. Two specimens from the shingle of the River Dee near Glen Tanar in Aberdeenshire seem to be an exact match for this species. 402. H. exotericum Jord. ex Boreau Jordan’s Hawkweed H. murorum subsp. exotericum (Jord. ex Boreau) Sudre; H. syngenes Jord. ex Boreau; H. murorum var. exotericum (Jord. ex Boreau) Rouy Phyllopodous perennial herb with a branched stock. Stem 40–65 cm, pale yellowish-green, sometimes purplish towards the base, usually slender, striate, with numerous, medium to long, pale simple eglandular hairs towards the base, becoming fewer upwards, and numerous stellate hairs and short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath; basal few to fairly numerous, not obviously divided into 2 kinds except for size, the lamina 4–10 × 2–4 cm, mostly elliptical or lanceolate-elliptical, rounded-mucronulate to more or less acute at apex, subentire to denticulate or shallowly sinuate-dentate near the base, rounded or truncate at base, the petioles up to 11 cm, with dense, long, pale simple

416

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401. H. pellucidum Laest.

400. H. integratum (Dahlst. ex Stenstr.) Dahlst.

403. H. neosparsum (Zahn) P. D. Sell 402. H. exotericum Jord. ex Boreau

405. H. gentile Jord. ex Boreau 404. H. pruinale (Zahn) P. D. Sell & C. West Inner and outer basal leaves of species of the Section Hieracium

39. Hieracium

417

406. H. itunense Pugsley 407. H. hjeltii Norrl.

410. H. camptopetalum (F. Hanb.) P. D. Sell & C. West 409. H. pollinarium F. Hanb.

412. H. asteridiophyllum P. D. Sell & C. West 411. H. snowdoniense P. D. Sell & C. West Inner and outer basal leaves of species of the Section Hieracium

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eglandular hairs; cauline 0–1(–2), when present lamina ovate or ovate-oblong, acute at apex, subentire to sinuatedentate, petiolate; all with short to medium, pale, appressed simple eglandular hairs on the upper surface and numerous medium to long ones beneath and on the margin. Inflorescence with 10–20(–numerous) capitula, paniculatecorymbose, often with a long lower branch; peduncles rather slender, curved, with dense stellate hairs and numerous to dense, short to very short (0.2–0.7 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent to slightly porrect in bud, 4–11 × 0.8–1.0 mm, blackish-green, the inner with paler margins, all narrow linear-lanceolate, more or less acute at apex, with dense, unequal, very short to medium (0.2–1.2 mm), dark glandular hairs, and a few stellate hairs along the margins, without simple eglandular hairs. Ligules deep yellow, glabrous-tipped. Styles usually pure yellow, but sometimes discoloured. Receptacle pits with margins dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Probably introduced. Roadside and railway banks, walls and waste places. Widely scattered localities in Great Britain north to Inverness-shire. Possibly widespread in central Europe from Spain to Russia 403. H. neosparsum (Zahn) P. D. Sell Bank Hawkweed H. murorum subsp. neosparsum Zahn; H. sparsum Jord. ex Boreau, non Friv. Phyllopodous perennial herb with a branched stock. Stem 25–60 cm, pale yellowish-green, often tinted purplish towards the base, more or less robust, striate, with numerous, medium to long, pale simple eglandular hairs below, becoming fewer upwards, and dense stellate hairs and numerous, short, dark glandular hairs above. Leaves medium yellowish-green on upper surface, paler beneath, rarely tinted reddish; basal few to numerous, the outer with lamina 3–11 × 2–5 cm, more or less elliptical or ovateelliptical, rarely oblong-elliptical, rounded-mucronulate at apex, entire to denticulate or shallowly sinuate-dentate and subtruncate or subcordate at base, the inner with lamina 7–10 × 3–5 cm, mostly oblong to oblong-elliptical, obtusemucronate to more or less acute at apex, sinuate-denticulate or shallowly sinuate-dentate and subtruncate to subcordate at base, the petioles up to 9 cm, with numerous, long, pale simple eglandular hairs; cauline usually 1, the lamina often large, ovate, acute-caudate at apex, dentate, subtruncate at base, petiolate; all with more or less numerous, short or medium, pale simple eglandular hairs on the upper surface and numerous, medium to long, pale simple eglandular hairs beneath and on the margins. Inflorescence with 2–23 capitula, paniculate-corymbose, often with a long lower branch; peduncles rather short, straight or curved, with dense stellate hairs and numerous, very short to short (0.2–0.7 mm), black glandular hairs, without simple eglandular hairs. Capitula 25–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0– 1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, more or less acute at apex, with dense,

unequal, very short to medium (0.2–1.0 mm), black glandular hairs, and few to fairly numerous stellate hairs along the margin, without simple eglandular hairs. Ligules yellow, glabrous or with a few, very short simple eglandular hairs at apex. Styles yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Roadside and railway banks and waste places. Recorded from Sussex, Essex, Cambridgeshire, Bedfordshire, Gloucestershire, Glamorganshire, Breconshire and Inverness-shire. Widespread in Europe from southern France to Bulgaria. 404. H. pruinale (Zahn) P. D. Sell & C. West Pruinose Hawkweed H. euprepes var. pruiniferum W. R. Linton, non H. pruiniferum Norrlin; H. murorum subsp. pruinale Zahn; H. murorum var. subulatidens auct.; H. subulatidens auct. Phyllopodous perennial herb with a branched stock. Stems 25–40 cm, pale yellowish-green, striate, rather slender, with numerous, medium to long, pale simple eglandular hairs towards the base and occasional ones in the upper part, and more or less numerous stellate hairs and occasional to few, short, dark glandular hairs in the upper part. Leaves medium yellowish-green, paler beneath; basal few, 2–9 × 1–4 cm, the outer with lamina ovate, elliptical or subrotund, rounded-mucronulate to subacute at apex, denticulate or dentate, and rounded or subtruncate at base, the inner with lamina lanceolate, ovate or elliptical, acute to acuminate at apex, denticulate to deeply dentate with sharp teeth, and subtruncate or shortly cuneate at base, the petioles short to medium with dense, long, wavy, pale simple eglandular hairs; cauline usually absent, occasionally 1, the lamina lanceolate, acute at apex, dentate, cuneate at base, petiolate; all with few to numerous, short to medium, rather stiff simple eglandular hairs on the upper surface and numerous medium to long ones beneath and on the margin, particularly the midrib. Inflorescence with 2–4(–10 cult.) capitula, cymose-corymbose, with a short acladium; peduncles arcuate, with dense stellate hairs and few to fairly numerous, very short to short (0.2–0.7 mm), dark glandular hairs. Capitula 20–25 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, blackishgreen, the inner with paler margins, all linear-lanceolate, obtuse to subacute at apex, with dense stellate hairs at the base and on the margin, numerous very short to medium (0.2–0.8 mm), unequal, black glandular hairs and without or with an occasional, short to medium, dark simple eglandular hair. Ligules golden yellow, with short hairs at tip at least when young. Styles yellow to slightly discoloured. Receptacle pits with subulate-dentate margins. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. H. pruinale differs from the widespread H. pictorum in having a hairy upper surface to the leaf and in having no simple eglandular hairs or only an occasional one on the involucral bracts. Native. Cliff ledges and rocks by streams. Known only from Ben Lawers, Fin Glen and Ben y Vrackie in Perthshire and Glen Doll (Dole), Clova in Forfarshire. Endemic.

39. Hieracium 405. H. gentile Jord. ex Boreau Foreign Hawkweed H. murorum subsp. murorum microgen. gentile (Jord. ex Boreau) Sudre; H. murorum subsp. gentile (Jord. ex Boreau) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, pale yellowish-green, often reddish-purple towards the base, rather slender and flexuous, with numerous, medium to long, pale simple eglandular hairs below, becoming fewer upwards, and with numerous stellate and short, dark glandular hairs in the upper part. Leaves medium to dark yellowish green on upper surface, paler and occasionally tinted reddish beneath; basal numerous, more or less similar except for size, the lamina 1.5–8.0 × 1.0– 3.5 cm, lanceolate, ovate or elliptical, most more or less acute, sometimes obtuse at apex, shallowly dentate with mammiform teeth, the upper third entire or denticulate, abruptly contracted or subtruncate at base, the petioles up to 6 cm and with dense, long, pale simple eglandular hairs; cauline 0–1, when present lamina lanceolate, long-acute at apex, dentate, truncate-based and petiolate, or bract-like; all with numerous, short to long, pale simple eglandular hairs on both surfaces and the margins. Inflorescence with 4– 15 capitula, paniculate-corymbose, sometimes with a long lower branch; peduncles erect or curved, slender, with dense stellate hairs and numerous, very short to short (0.2–0.5 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, rounded at base. Involucral bracts 4–10 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linear-lanceolate, acute at apex, with dense, unequal, very short to medium (0.2–1.0 mm), slender, black glandular hairs, and some stellate hairs towards the base, without simple eglandular hairs. Ligules yellow, with a few, very short simple eglandular hairs at apex. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, blackish-red. Flowers 5–7. Apomictic. Introduced. Roadside and railway banks, quarry and waste places. Ashurst Park in Kent, Cambridgeshire– Hertfordshire border at Royston, Tintern and Symonds Yat in Monmouthshire, Ysgwd Gwladys in Glamorganshire, Capel Curig in Caernarvonshire, Oyne in Aberdeenshire and Lochawe village and Kilmun in Argyllshire. Said to be widespread in Continental Europe, but that may be as an aggregate. 406. H. itunense Pugsley Barras Hawkweed H. cumbriense auct.; H. atratum subsp. cumbriense auct. Phyllopodous perennial herb with a branched stock. Stem 40–60(–80 cult.), pale yellowish-green, sometimes slightly tinged reddish, rather slender, striate, with few, medium, pale, dark-based simple eglandular hairs, and numerous stellate hairs, and numerous short, dark glandular hairs in the upper part. Leaves medium yellowish-green on upper surface, paler beneath and sometimes tinted reddish; basal numerous, the lamina 4–7 × 2–3 cm, ovate, elliptical or elliptical-lanceolate, acute to acuminate or cuspidate at apex, denticulate to sinuate-serrate or sharply dentate with long, spreading teeth, subtruncate or shortly cuneate at base,

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the petioles up to 6 cm, slender, with numerous, long, pale simple eglandular hairs; cauline 0–1, when present lamina narrowly linear or bract-like; all with scattered, short or medium, pale simple eglandular hairs on both surfaces or nearly glabrous, more numerous hairs on the margins. Inflorescence with 6–12(–numerous cult.) capitula, paniculate-corymbose, with suberect branches; peduncles slender, curved or suberect, with dense stellate hairs and numerous short to medium (0.3–1.0 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25– 30 mm in diameter, rounded at base. Involucral bracts porrect in bud, 5–12 × 1.0–1.3 mm, blackish-green, the inner with paler margins, all linear-lanceolate, obtuse at apex except the innermost, with dense, unequal, very short to medium (0.2–1.0 mm), black glandular hairs, an occasional, medium, dark simple eglandular hair, and a very occasional stellate hair. Ligules golden yellow, glabrous or with occasional very short, simple eglandular hairs at the apex. Styles discoloured. Receptacle pits unknown. Achenes 3.0– 3.2 mm, reddish-black. Flowers 6–8. Apomictic. Native. Known only from limestone rocks near South Stainmore (near Barras, Westmorland) where the population is very small. Endemic. 407. H. hjeltii Norrl. Hjelt’s Hawkweed H. murorum subsp. hjeltii (Norrl.) Zahn Phyllopodous perennial herb with a branched stock. Stems 25–40 cm, yellowish-green, slender, striate, with few to numerous, short to long, pale simple eglandular hairs in the lower half and few to numerous stellate and few to numerous, short, dark glandular hairs above. Leaves medium yellowish-green, paler beneath; basal few, with lamina 3– 7 × 1.5–3.5 cm, lanceolate, ovate or elliptical, rounded or subacute at apex, subentire or denticulate near the base, rounded or subtruncate at base, the petioles short to medium and with dense, long, pale simple eglandular hairs; cauline usually 1, with lamina lanceolate or ovate, acute or acuminate at apex, subentire or denticulate in lower half, subtruncate at base, petiolate; all with occasional, short to medium, pale simple eglandular hairs on the upper surface and numerous short to long, pale simple eglandular hairs on the lower surface and margins. Inflorescence cymose-corymbose, with 2–7 capitula, the acladium fairly short; peduncles slender and rather straight, with dense stellate and very short to short (0.2–0.4 mm), dark glandular hairs. Capitula 30–40 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 5–10 × 0.8– 1.0 mm, blackish-green, the inner with paler margins, all narrowly linear-lanceolate, acute at apex, with numerous stellate hairs on the margins, numerous, short (up to 0.6 mm), dark glandular hairs, without simple eglandular hairs. Ligules yellow, slightly hairy or glabrous-tipped. Styles probably yellow. Receptacle pits unknown. Achenes (immature) about 3 mm, blackish-red. Flowers 5–7. Apomictic. Probably introduced. Specimens collected by the railway at Silverdale in Lancashire by J. Cryer in 1912 match exactly authentic specimens of H. hjeltii. Specimens collected at Storth, near Arnside, Westmorland are

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very similar in appearance but the upper surface of the leaves is hairy and the styles discoloured, but this may not be enough to distinguish them as a distinct species. Native of Fennoscandia. Named after Albert Hjalmar Hjelt (1851–1925). 408. H. patale Norrl. Dartmoor Hawkweed H. murorum subsp. patale (Norrl.) Zahn Phyllopodous perennial herb with a branched stock. Stem 30–60 cm, yellowish-green, sometimes purplish near the base, striate, usually rather slender, with a few, medium to long, pale simple eglandular hairs towards the base, and numerous to dense stellate and scattered, minute (0.3– 0.4 mm), dark glandular hairs in the upper part. Leaves yellowish-green, often suffused purplish beneath; basal usually few, with lamina 4–10 × 2.0–4.5 cm, the outer elliptical or with lamina elliptic-lanceolate, more or less rounded at apex, undulate-denticulate, with shallow, apiculate teeth and rounded or subtruncate at base, the inner with lamina elliptical to oblong or sometimes ovate, acute at apex, undulate-denticulate to shallowly dentate, with small mammiform teeth, and subtruncate to cuneate at base, the petioles long (up to 9 cm), often purplish, with numerous to dense, long, pale, wavy simple eglandular hairs; cauline 0–1, when present the lamina small, lanceolate, acute at apex, sharply dentate and petiolate, or bract-like; all glabrous or nearly so on upper surface, with scattered to numerous, medium to long, pale simple eglandular hairs on the lower surface and margin and dense along the midrib. Inflorescence rather compactly paniculate-corymbose, with 5–22 capitula, the acladium fairly long; peduncles rather short, arcuate, with dense stellate hairs and more or less uniform, short (0.4–0.5 mm), dark glandular hairs, without simple eglandular hairs. Capitula 20–30 mm in diameter, rounded or slightly narrowed at base. Involucral bracts incumbent in bud, 5–10 × 1.0–1.2 mm, blackish-green with paler margins, all narrow linear-lanceolate, more or less acute at apex, with stellate hairs on the margin and a tuft of hairs at apex and more or less numerous, more or less equal, short (0.4–0.5 mm), dark glandular hairs, without simple eglandular hairs. Ligules medium yellow, with short simple eglandular hairs at apex. Styles discoloured. Achenes about 3 mm, reddish-black. Flowers 5–7. Apomictic. Introduced. Occurs in considerable abundance on roadsides, railway banks and in woods of Dartmoor in Devonshire, near Newtown in Montgomeryshire and on a wall and rough ground in a quarry at Onich in Inverness-shire. It is native of Fennoscandia and has presumably been introduced although it looks native in some Devonshire localities. 409. H. pollinarium F. Hanb. Naver Hawkweed H. murorum subsp. pollinarium (F. Hanb.) Zahn Phyllopodous perennial herb with a branched stock. Stem 20–50 cm, pale yellowish-green, often brownish-purple below, more or less robust, striate, with a few, long, pale simple eglandular hairs below or nearly glabrous, and numerous stellate and numerous, short, dark glandular hairs in

the upper part. Leaves thick, glaucous-green on upper surface, paler beneath and sometimes tinted purplish; basal usually numerous, the outer with lamina 2.0–7.5 × 1.5–5.5 cm, broadly elliptical to subrotund, rounded-mucronulate at apex, subentire or with shallow teeth and rounded at base, the inner with lamina 6.5–10.0 × 2.0–5.0 cm, acute at apex, entire or denticulate, sometimes with a few larger teeth and truncate or rounded at the base, the petioles up to 5 cm, with numerous, long, pale simple eglandular hairs; cauline 0–1, when present with lamina lanceolate, acute at apex, entire or denticulate, shortly petiolate; all glabrous or nearly so on the upper surface, with few to numerous, short to medium, pale simple eglandular hairs beneath and on the margin. Inflorescence with 2–10(–many cult.) capitula, furcatecorymbose, usually with geminate acladium; peduncles short and stiff, with dense stellate hairs and numerous, short to medium (0.3–1.0 mm), dark glandular hairs, without simple eglandular hairs. Capitula 25–30 mm in diameter, subtruncate at base. Involucral bracts incumbent in bud, 4–11 × 1.5–2.0 mm, blackish, the inner with paler margins, all broadly linear-lanceolate, obtuse at apex, with numerous, very short to short (0.2–0.5 mm), black glandular hairs, and a few stellate hairs along the margins, without simple eglandular hairs. Ligules yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins incisedentate. Achenes 2.5–3.0 mm, reddish-black. Flowers 6–7. Apomictic. Native. Cliff ledges. Between Bettyhill and Farr Church and by the River Naver, Sutherland. Endemic. 410. H. camptopetalum (F. Hanb.) P. D. Sell & C. West Short-flowered Hawkweed H. murorum var. camptopetalum F. Hanb.; H. duriceps var. camptopetalum (F. Hanb.) Pugsley; H. caliginosum auct.; H. serratifrons var. caliginosum auct.; H. murorum subsp. caliginosum auct. Phyllopodous perennial herb with a branched stock. Stem 25–50 cm, pale yellowish-green, occasionally reddish towards the base, slender and flexuous, striate, sometimes with a few, medium to long, pale simple eglandular hairs towards the base, sometimes with them absent almost throughout, with numerous stellate and few to numerous, short, dark glandular hairs in the upper part. Leaves dull, deep green on upper surface, paler and bluish-green beneath, sometimes tinted reddish or purplish especially near the margin; basal numerous, hardly divided into outer and inner, the lamina 1.5–10.0 × 1.5–5.0 cm, subrotund, broadly elliptical or ovate, mostly rounded-mucronulate at apex, but sometimes more or less acute, subentire, denticulate or sharply and shallowly dentate, the teeth with mucros, rounded or cuneate at base, the petioles up to 8 cm, with numerous, long, pale simple eglandular hairs; cauline usually absent or bract-like, but sometimes with lamina ovate, acute at apex, dentate, truncate at base and petiolate; all glabrous or nearly so on upper surface, with few to numerous, medium to long, pale simple eglandular hairs beneath and on the margin. Inflorescence with 3– 10 capitula, paniculate-corymbose, sometimes with a lower branch; peduncles short, slender and spreading, with dense stellate hairs and numerous, short (0.4–0.5 mm), fine, dark

40. Arctotis glandular hairs, without simple eglandular hairs. Capitula 15–25(–30) mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0–1.2 mm, greyishgreen, the inner with paler margins, all linear-lanceolate, obtuse at apex, with very short to medium (0.2–1.0 mm), dark glandular hairs, and numerous to dense stellate hairs especially on the margins, without simple eglandular hairs. Ligules often poorly developed, yellow, glabrous-tipped. Styles greenish-yellow. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddish-black. Flowers 6–7. Apomictic. Native. Mountain and sea-cliffs and by rocky streams. In a few scattered localities in Ross-shire and Sutherland. Endemic.

411. H. snowdoniense P. D. Sell & C. West Snowdonia Hawkweed H. murorum var. pulcherrimum auct.; H. pellucidum var. pulcherrimum auct. Phyllopodous perennial herb with a branched stock. Stems 20–40 cm, pale yellowish-green, sometimes tinted purplish below, striate, slender, flexuous, with few to numerous, short to long, pale simple eglandular hairs below and occasional ones above, and numerous stellate hairs and few to numerous, short, fine, dark glandular hairs in the upper part. Leaves pale green on upper surface, very pale and often suffused with reddish-purple beneath; basal with lamina 1.5–7.0 × 1.5–3.5 cm, ovate, obovate, elliptical or obovateelliptical, rounded-mucronulate to broadly acute at apex, mostly denticulate or undulate-denticulate, sometimes with a few, small teeth at the subtruncate or shortly cuneate base, the petioles short to medium with dense, long, pale simple eglandular hairs; cauline 1(–3), the lower with lamina lanceolate, acute at apex, denticulate, shortly petiolate, the upper when present bract-like; all with few to fairly numerous, short to medium, subrigid, pale simple eglandular hairs on the upper surface and numerous, short to long ones beneath and on the margin and particularly on the midrib. Inflorescence compactly cymose-corymbose, with a fairly long acladium; peduncles arcuate, with dense stellate hairs and numerous, very short to medium (0.2–1.0 mm), very unequal, slender, dark glandular hairs, without simple eglandular hairs. Capitula 20–30 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–10 × 1.0– 1.2 mm, blackish, the inner with paler margins, all narrowly linear-lanceolate, the inner very narrow, acute at apex, with stellate hairs along the margin and sometimes at the apex and dense, very short to medium (0.2–1.0 mm), very unequal, black glandular hairs, without simple eglandular hairs although some medium glands with tiny heads may be mistaken for them. Ligules golden yellow, glabroustipped. Styles discoloured. Receptacle pits with margins incise-dentate. Achenes 2.5–3.0 mm, blackish. Flowers 6–7. Apomictic. Native. Cliff ledges. Cwm Glas Fach on Snowdon, the Devil’s Kitchen, Ysgolion Duon on Carnedd Dafydd and Nant Ffrancon all in Caernarvonshire. Only one plant found in a recent survey. Endemic.

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412. H. asteridiophyllum P. D. Sell & C. West Star-haired Hawkweed H. pellucidum var. lucidulum auct. Phyllopodous perennial herb with a branched stock. Stem 20–40 cm, pale yellowish-green, often suffused reddishpurple below, slender to fairly robust, flexuous, striate, with a few, long, pale simple eglandular hairs below, petering out upwards, and few to numerous, short, dark glandular hairs in the upper part. Leaves dark green on upper surface, paler and often suffused purple beneath, thick; basal usually numerous, the outer with lamina 2–7 × 1.5–4.0 cm, broadly ovate, subrotund or sometimes almost reniform, broadly rounded-mucronulate at apex, shallowly dentate in the lower two-thirds and truncate or cordate at base, the inner with lamina 4–9 × 2–4 cm, broadly ovate to lanceolate, rounded-mucronate to more or less acute at apex, denticulate to strongly dentate, with the teeth mammiform or aquiline-mammiform and the basal pair sometimes retrorse, subtruncate at base the petioles up to 5 cm, with long, pale simple eglandular hairs; cauline 0–1, the lamina ovate or lanceolate, acute or acuminate at apex, more or less dentate, petiolate; all with stellate hairs on both surfaces and medium, scattered, pale simple eglandular hairs beneath and on the margins. Inflorescence with 3–10 capitula, paniculate-corymbose, sometimes with a long lower branch; peduncles straight or curved, with dense stellate hairs and numerous, short (0.4–0.5 mm), black glandular hairs. Capitula 30–35 mm in diameter, rounded at base. Involucral bracts incumbent in bud, 4–11 × 1.0–1.2 mm, blackish-green, the inner with paler margins, all linearlanceolate, acute at apex, with numerous, very short to short (0.2–0.5 mm), dark glandular hairs, few to fairly numerous, short to medium, dark-based simple eglandular hairs, and numerous stellate hairs along the margins. Ligules medium yellow, glabrous-tipped. Styles discoloured. Receptacle pits with margins shortly dentate. Achenes 3.0–3.5 mm, reddishblack. Flowers 6–7. Apomictic. Native. Ledges on limestone cliffs at Craig-y-cilau (Craig Cille), Breconshire mainly on the main lower Craig-y-cilau cliff and at the west end of Darren Cilau quarry. Endemic. Tribe 2. Arctotideae Cass. Tribe Gorterieae Lindl.; Tribe Gundelieae DC. ex Lecoq & Juillet Plants not producing white latex. Capitula with flowers of 2 kinds, the ligulate flowers sterile, the ligules yellow to orange, 3- to 4-lobed at apex; tubular flowers 5-lobed. Anthers without or with very short tails, apical appendage usually separated from thecae; filaments usually smooth; pollen usually spiny. Style bilobed at apex or with short or long branches, mostly with longer hairs beneath the branches or swollen apical part. Contains 2 subtribes, 16 genera and about 200 species, mainly in South Africa with few species elsewhere in Africa, south-west Asia and Australia. 40. Arctotis L. Perennial herbs. Stems erect, branches spreading, leafy. Leaves alternate, entire or pinnately lobed, woolly. Capitula

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solitary at the ends of stems or branches. Involucral bracts in several rows, with broadly scarious margins. Flowers of 2 kinds, the outer ligulate and female with 3 lobes at apex, the inner tubular and bisexual. Corolla of ligulate flowers cream, yellow, orange, brick-red or purple, of tubular flowers dark purplish-violet. Anthers with tails short or absent, apical appendages distinctly demarcated from the thecae; filaments smooth. Style slender, with a conspicuously thickened, shortly bilobed apical part which has a ring of hairs at its base. Receptacle flat, pitted, the pits ciliate, without scales. Achenes broadly obconical, with 2 furrows and 3 ridges on one side; pappus of 2 rows of scarious, oblong, scales. About 50 species in South Africa, Namibia and Angola. A. breviscapa Thunb. has occurred as a casual garden escape. 1. A. stoechadifolia Bergius White Arctotis Perennial herb with a strong tap-root. Stems 30–75 cm, bluish-green, erect, robust and sometimes slightly woody at base, hollow, markedly striate, glabrous or slightly woolly especially in the upper part, diffusely branched, leafy. Leaves 10–25 × 2–10 cm, whitish- or greyish-green, obovate or oblong-obovate, obtuse at apex, entire or pinnately lobed, the lobes few, broad and obtuse at apex, undulatedentate, gradually attenuate at base, woolly all over or only on the lower surface; petiole long, striate and more or less woolly. Capitula 40–70 mm in diameter, solitary at the ends of stems or branches; peduncles long, striate and woolly. Involucral bracts in several rows, 3–16 × 2–4 mm, brownish or purplish-brown, with a broad scarious margin, narrowly to broadly oblong, rounded-obtuse at apex, more or less woolly. Flowers of 2 kinds, the outer 20–35 mm, ligulate and female, the ligules cream, yellow, orange, brick red or purple and shortly 3-lobed at apex, the inner 8–12 mm, tubular, bisexual and dark purplish-violet. Receptacle flat and pitted, the pits ciliate, without scales. Achenes 2.5–4.0 mm, brownish, broadly obconical, with 2 furrows and 3 ridges on 1 side, with dense, long, white to yellowish hairs; pappus 4–6 mm, of 2 rows of brown, scarious, oblong scales. Flowers 9–10. Introduced. An established garden escape naturalised since 1978 in a quarry in Sark in the Channel Islands. Native of South Africa. Very variable. Most of our garden plants belong to var. grandis (Thunb.) Less. 41. Arctotheca J. C. Wendl. Cryptostemma R. Br. Annual or perennial herbs. Stems decumbent to erect, leafy only near the base. Leaves alternate, mainly deeply pinnately divided, white tomentose beneath. Capitula solitary and axillary. Involucral bracts in several rows, free, with conspicuous scarious tips. Inner flowers tubular, greenish-black and bisexual, the outer ligulate, pale yellow above, greyish-green, purplish or brownish below and sterile, with 3 lobes at apex. Anthers with tails short or absent, apical appendages distinctly demarcated from the thecae; filaments papillose. Style slender, with a conspicuously thickened, shortly bilobed apical part which has a ring of hairs at its base. Receptacle flat, pitted, with-

out scales. Achenes linear to obovoid, ribbed, densely woolly-hairy; pappus of a single row of 4–8, minute scales. Four species in South Africa (mainly the Cape) and Mozambique. 1. A. calendula (L.) Levyns Plain Treasureflower Arctotis calendula L.; Arctotis calendulacea Hill; Cryptostemma calendulacea (Hill) R. Br.; Cryptostemma calendula (L.) Druce; A. calendulacea (Hill) Lewin Annual to perennial herb, very variable in habit and sometimes rooting at the stem nodes. Stems 20–40 cm, pale green, sometimes tinted brownish-purple, decumbent to erect, fleshy, striate, glabrous or with sparse arachnoid hair. Leaves mostly near the base, 5–25 × 2–6 cm, yellowishgreen on upper surface, greyish-white beneath, ovate, mucronate at apex, entire to denticulate, with a cuneate to truncate base, to lyrate-pinnatifid, the terminal lobe broadly ovate, rounded at apex, usually with small teeth, and truncate based, the lateral lobes 3–5 pairs, linear to oblong or lanceolate to ovate and entire to dentate, glabrous on upper surface, thinly white-felted beneath; petioles up to 12 cm, striate. Capitula solitary, 25–50 mm in diameter; peduncles 10–25 cm, striate, with a few, very short hairs, axillary. Involucral bracts in several rows, imbricate, 4– 9 × 2–4 mm, pale, dull green, with a narrow, black margin and conspicuous scarious apex, lanceolate to ovate, acute to obtuse at apex, sometimes with traces of white, woolly hair. Flowers of 2 kinds; the outer 15–20 mm, ligulate, the ligules lemon yellow on upper surface, partly yellow and washed greenish-grey, purplish or brownish beneath, 3-lobed at apex; the inner 10–15 mm, tubular, yellowish to blackish. Receptacle flat and pitted, without scales. Achenes 1.8–2.0 mm, linear to obovoid, ribbed, enveloped in copious, pinkish-brown, silky wool; pappus of a single row of 4–8, minute, delicate, acute scales. Flowers 7–8. 2n = 18. Introduced. Fairly frequent wool alien in arable fields and waste places. Scattered records in Great Britain. Native of South Africa. 42. Gazania Gaertn. Annual to perennial herbs. Stems decumbent to ascending, often woody at the base, leafy mainly below. Leaves alternate, entire to pinnately divided, white-felted beneath. Capitula solitary and axillary. Involucral bracts in 2–3 rows, the outer connate at the base so as to form a cup-like involucre. Inner flowers tubular, orange and bisexual, the outer ligulate, yellow to bright orange, sometimes with a black patch in which is a central white spot, 4 teeth at apex and sterile. Anthers without a tail, apical appendages more or less continuous with the thecae; filaments papillose. Style slender, with rather long branches, the shaft glabrous with a hairy apical part. Receptacle flat to convex, without scales. Achenes obovoid, ribbed, densely hairy; pappus of linearlanceolate scales. Seventeen species in South Africa, mostly in the Cape, Namibia and one species extending to tropical east Africa. Roessler, H. (1959). Revision der Arctotidae – Gorteriinae. Mitt. Bot. Staats. M¨unchen 3: 71–500.

43. Berkheya 1. Ligules yellow to orange with a black blotch in which is a 1(i). rigens var. rigens central white spot 2. 1. Ligules yellow 2. Upper surface of leaves and involucral bracts glabrous or 1(ii). rigens var. uniflora nearly so 2. Upper surface of leaves and involucral bracts 1(iii). rigens var. leucolaena white-woolly

1. G. rigens (L.) Gaertner Treasureflower Gorteria rigens L. Annual to perennial herb. Stems often more or less woody at base, 20–50 cm, pale green, sometimes tinted brownishpurple, decumbent to ascending, sometimes rooting at the nodes, striate, glabrous or slightly woolly. Leaves mainly in the lower part, 2–12 × 0.4–1.5 cm, yellowishgreen on upper surface, white beneath, mostly narrowly linear-oblanceolate, obtuse to subacute at apex, entire and long-attenuate at base, sometimes the basal deeply pinnately lobed with narrowly elliptical, obtuse, entire lobes, all glabrous or tomentose on upper surface, white-felted beneath except on the midrib and margin; petioles short to long, narrowly winged. Capitula 40–60 mm in diameter, solitary; peduncles 10–15 cm, axillary. Involucral bracts in 2–3 rows, the outer fused below in a cup-like structure 10– 15 mm, acute and scarious at apex, dark green and glabrous or white-woolly, the inner acute and scarious at apex and glabrous. Flowers of 2 kinds; outer 20–40 mm, ligulate, the ligules all yellow or with orange towards the base, or yellow to orange with a black blotch bearing a central white spot, with 4 lobes at apex; inner 10–12 mm, tubular, orange. Receptacle flat to convex, pitted, without scales. Achenes 6–7 mm, obovoid, ribbed, densely silky-hairy; pappus of narrow, linear-lanceolate, acuminate, delicate scales. Flowers 8–10. 2n = 10, 14. (i) Var. rigens Leaves glabrous above, white-felted beneath. Involucral bracts glabrous or sparingly hairy. Ligules yellow to orange with a black blotch bearing a central white spot. (ii) Var. uniflora (L. fil.) Roessler G. uniflora (L. fil.) Sims; Gorteria uniflora L. fil.; G. uniflora var. pinnata Harv.; Meridiana uniflora (L. fil.) Kuntze Leaves glabrous above, white-felted beneath. Involucral bracts glabrous or sparingly hairy. Ligules yellow. (iii) Var. leucolaena (DC.) Harv. G. leucolaena DC.; G. uniflora var. leucolaena (DC.) Harv. Leaves tomentose on upper surface, white-felted beneath. Involucral bracts white-woolly. Ligules yellow. Introduced. Grown in gardens and more or less naturalised on walls, rocks and cliffs near the sea. Isles of Scilly and Channel Islands. Native of South Africa. 43. Berkheya Ehrh. Crocodilodes Adans.; Stobaea Thunb. Perennial herbs sometimes woody at base. Stems erect, leafy. Leaves alternate, pinnately lobed or pinnatifid, often white-tomentose beneath. Capitula in a paniculate inflores-

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cence. Involucral bracts in several rows, the outer connate at base, spinose. Flowers all tubular and bisexual, with some behaving as male, or the outer ligulate, 4-lobed at apex and sterile, the inner tubular and bisexual. Anthers not tailed, apical appendages more or less continuous with the thecae; filaments smooth. Style slender, with rather long branches, the shaft glabrous, the apex hairy. Receptacle deeply pitted, with yellowish, rigid hairs. Achenes turbinate, ribbed, glabrous or hairy; pappus of numerous scales. About 75 species, mainly in South Africa, with a few in Tropical Africa north to Nigeria and Ethiopia. Roessler, H. (1959). Revision der Arctotidae – Gorteriinae. Mitt. Bot. Staats. M¨unchen 3: 71–500. 1. Leaves slightly tomentose to nearly glabrous; pappus 1. pinnatifida scales acute to subobtuse at apex, in 2 rows 1. Leaves white-tomentose beneath and sometimes above; 2. pappus scales obtuse at apex, in a single row 2. Lower leaves more or less obovate with broad triangular-spinose teeth; pappus scales 0.2–0.5 mm, 2. heterophylla oblong 2. Lower leaves linear to lanceolate with spinous margins; 3. rigida pappus scales about 0.2 mm, oblong-subrotund

1. B. pinnatifida (Thunb.) Thell. Lobed African Thistle Stobaea pinnatifida Thunb.; Stobaea apitrachys DC.; Crocodilodes pinnatifidum (Thunb.) Kuntze; Crocodilodes epitrachys (DC.) Kuntze Perennial herb sometimes with a more or less woody rhizome. Stem 20–50 cm, yellowish-green, sometimes slightly purplish, more or less erect, branched, more or less tomentose and sometimes with a few, sparse simple eglandular hairs. Leaves alternate, rigid; lower 3–8(–10) cm, linearlanceolate, spinose-acute at apex, pinnatifid or pinnately lobed, the lobes up to 10 pairs, triangular to linear, when pinnatifid the divisions broadly laciniate, narrowed at base, semiamplexicaul or shortly petiolate; upper gradually smaller but similar; all slightly tomentose or rarely nearly glabrous. Inflorescence paniculate. Capitula 15–50 mm in diameter. Involucral bracts in several rows, triangularlanceolate or linear-subulate, spiny on the margin, the spines 1–3 mm, the outer more or less tomentose and rigid-patent. Flowers all tubular and more or less the same size, yellow. Receptacle deeply pitted, with yellow rigid hairs 2–5 mm. Achenes 2–3 mm, turbinate, glabrous or rarely with short hairs; pappus about 2 mm, of 2 rows of oblong-lanceolate or lanceolate-subulate, acute to subobtuse scales which are denticulate at their apices. Flowers 6–9. Introduced. A wool casual. Native of South Africa. 2. B. heterophylla (Thunb.) O. Hoffm. Variable African Thistle Stobaea heterophylla Thunb.; Apuleje heterophylla (Thunb.) Less.; Stobaea biloba DC.; Crocodilodes heterophyllum (Thunb.) Kuntze; Crocodilodes bilobum (DC.) Kuntze Perennial herb often with a more or less woody rhizome. Stem 10–60 cm, pale green, sometimes flushed purplish, erect, branched, tomentose to more or less glabrous. Leaves alternate; basal 5–12(–20) × 1–4(–7) cm, sometimes in a

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rosette, more or less obovate, obtuse at apex, pinnately lobed or broadly triangular spinose-dentate, narrowed at base; cauline gradually smaller upwards, oblong, with small lobes or spinous-dentate; all glabrous or slightly tomentose above, white-tomentose beneath. Inflorescence paniculate. Capitula 20–30(–35) mm in diameter. Involucral bracts in several rows, 8–15 mm, ovate-lanceolate to linearlanceolate, spinous-acute at apex, the spine 1–3 mm, glabrous to slightly tomentose. Flowers of 2 kinds, the outer ligulate, the inner tubular, or all tubular, yellow. Receptacle deeply pitted with yellowish rigid hairs 1(–2) mm long. Achenes 1.5–2.0 mm, turbinate, with short hairs or nearly glabrous; pappus of 1 row of about 20, oblong, obtuse, denticulate scales 0.2–0.5 mm. Flowers 6–9. Introduced. A wool casual in cultivated and waste ground. Native of the Cape Province of South Africa. 3. B. rigida (Thunb.) Ewart, J. White & Rees Rigid African Thistle Stobaea rigida Thunb.; Apuleja zeyheri Less.; Stobaea zeyheri (Less.) DC.; Stobaea rubricaulis DC.; Stephanocoma rubricaulis (DC.) Sch. Bip.; Cocodilodes rigidum (Thunb.) Kuntze Perennial herb with a more or less woody rhizome. Stem 20– 60 cm, pale green, sometimes suffused brownish-purple, erect, branched in upper part, leafy, tomentose or becoming glabrous. Leaves alternate, rigid; lower 2–5(–8) cm, linear or lanceolate, spinose-acute at apex, pinnatifid, with spinose margins, semiamplexicaul; remainder similar but gradually smaller; all more or less glabrous or slightly tomentose above, usually white-tomentose beneath. Inflorescence paniculate, dense. Capitula 15–30(–35) mm in diameter. Involucral bracts in a several rows, 8–15(–20) mm, ovatetriangular or ovate-lanceolate, spinose-acute at apex, with spines 1–2 mm along the margin, the outer tomentose, the inner more or less glabrous. Flowers all tubular of more or less the same size. Receptacle deeply pitted, with yellowish, rigid hairs 1(–2) mm. Achenes 1.5–2.5 mm, turbinate with sparse, short hairs or glabrous; pappus about 0.2 mm, of 1 row of about 20, oblong-subrotund, obtuse, minutely denticulate scales. Flowers 6–9. Introduced. A wool casual in cultivated and waste ground. Native of South Africa. (Epaltes australis Less. has been recorded as a casual.)

Subfamily 2. Asteroideae (Cass.) Lindl. Subfamily Tubiflorae auct.; Subfamily Adenostyloideae (Cass.) Lindl.; Subfamily Ambrosioideae Raf.; Subfamily Anthemidoideae (Cass.) Lindl.; Subfamily Artemisioideae Burmeist.; Subfamily Baccharidoideae Burmeist.; Subfamily Buphthalmoideae Burmeist.; Subfamily Calenduloideae (Cass.) Lindl.; Subfamily Chrysanthoideae Burmeist.; Subfamily Ecliptoideae Burmeist.; Subfamily Eupatorioideae (Cass.) Lindl.; Subfamily Gnaphalioideae Burmeist.; Subfamily Helenioideae Burmeist.; Subfamily Inuloideae (Cass.) Lindl.; Subfamily Melampodioideae Burmeist.; Subfamily Othonnoideae Burmeist.

Plant not producing white latex. Stem leaves usually spiral, sometimes opposite or absent. Capitula mostly with ligulate and tubular flowers, sometimes with only tubular flowers. Tubular flowers usually with 5 short lobes or teeth, often yellow to orange. Filaments joining anthers at base. Style branches 2, each with 2 stigmatic surfaces, 1 near each margin of the inner face. Pollen grains spiny. Tribe 3. Gnaphalieae (Cass.) Lecoq & Juillet Taxon Gnaphalieae Cass.; Subtribe Gnaphaliinae (Cass.) Dumort. Annual to perennial herbs, rarely shrubs. Leaves alternate, simple. Capitula with all tubular flowers, or tubular and ligulate ones. Flowers yellow, brownish or white. Involucral bracts in several rows, more or less scarious. Receptacle without scales, or the outer achenes with scales doubtfully separable from the involucral bracts. Pappus of hairs. Contains 5 subtribes, 162 genera and more than 2,000 species. Worldwide but particularly in southern Africa and Australia. 44. Filago L. Annual herbs. Stems leafy, branched, woolly-hairy. Leaves alternate, narrow, entire, woolly-hairy. Capitula in terminal and axillary or basal subglobose clusters, rarely solitary. Involucral bracts in a few ill-defined rows, the outer herbaceous, the inner scarious. Flowers all tubular, shallowly 5-lobed, the outer and sometimes some of the inner female and filiform, some or all of the inner bisexual or sometimes functionally male. Corolla yellow. Anthers tailed, apical appendage ovate, flat. Style with oblong-linear branches. Receptacle from more or less flat to conical, obconical or cylindrical, naked except for scales subtending the outer flowers which are often doubtfully distinguishable from the involucral bracts. Achenes oblong to ovoid, slightly compressed, the outer usually larger and sometimes enclosed by the involucral bracts and shed with them; pappus of central flowers of many simple hairs, of outer flowers of few hairs or absent, sometimes absent on all achenes. About 50 species mainly in Europe, Asia, North Africa and North America. Rich, T. C. G. (1999). Conservation of Britain’s biodiversity: Filago lutescens Jordan (Asteraceae), Red-tipped cudweed. Watsonia 22: 251–260. Rich, T. C. G. (1999). Conservation of Britain’s biodiversity. IV. Filago pyramidata (Asteraceae), Broad-leaved cudweed. Edinb. Jour. Bot. 56: 61–73. Rich, T. C. G., Gibson, C. K. & Marsden, M. (1999). Reestablishment of the extinct native plant Filago gallica L. (Asteraceae), narrow-leaved cudweed, in Britain. Biol. Conserv. 91: 1–8. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [F. gallica, F. lutescens and F. pyramidata.] 1. Capitula (5–)10–40 in each cluster; outer involucral bracts aristate at apex, erect or divergent in fruit; marginal

44. Filago

1.

2.

2.

3.

3.

4.

4.

5. 5.

flowers subtended but not enclosed by involucral bracts 2. or receptacular scales Capitula 2–7(–14) in each cluster, sometimes solitary; outer involucral bracts obtuse to acute at apex, spreading star-like in fruit; marginal flowers more or less enclosed by involucral bracts or receptacular scales resembling 4. them Leaves widest below the middle, the most apical ones not overtopping the clusters of capitula; capitula in clusters of 1. vulgaris (15–)20–35(–40) Leaves widest above the middle, the most apical ones usually overtopping the clusters of capitula; capitula in 3. clusters of (5–)10–20(–25) Plant usually yellowish-woolly; clusters of capitula each overtopped by (0–)1–2 leaves; outer involucral bracts 2. lutescens with an erect, red-tinged apical arista Plant white-woolly; clusters of capitula each overtopped by 2–4(–5) leaves; outer involucral bracts with a 3. pyramidata recurved, yellowish apical arista Leaves 8–25 mm; cluster of capitula usually much overtopped by subtending leaves; marginal achenes 6. gallica completely enclosed by involucral bracts Leaves 4–20 mm; clusters of capitula usually not overtopped by subtending leaves; marginal achenes incompletely enclosed by involucral bracts or 5. receptacular scales Plant densely white-woolly; leaves erect but not closely 4. arvensis appressed to the stem Plant greyish with short, silky hairs; leaves erect and 5. minima closely appressed to the stem

Subgenus 1. Filago Gifola Cass. Capitula in axillary and terminal clusters. Involucral bracts acuminate or aristate, not spreading star-like in fruit. Outer achenes not enclosed in bracts. 1. F. vulgaris Lam. Common Cudweed F. germanica L., non Huds.; F. canescens Jord.; F. eriocephala auct.; Gifola germanica Dumort. nom. illegit.; Gnaphalium dichotomum Salisb. nom. illegit. Annual herb with fibrous roots. Stems 5–40 cm, greyishwhite, erect or ascending, densely woolly, simple or more or less regularly branched above the middle, sometimes branched from the base, leafy. Leaves 12–20(–30) × 1– 3(–4) mm, linear-lanceolate to lanceolate, obtuse or tapering to an acute apex, entire and usually undulate, rounded at base, covered with white woolly hairs. Capitula 1.4– 1.6 mm in diameter, more or less terete, in dense, globose, sessile, clusters of (15–)20–35(–40) terminating the main stems and branches, the clusters 10–12 mm wide and not overtopped by subtending leaves. Involucral bracts in a few rows, 4.0–4.5 × about 1.1 mm, yellowish and usually redtinged, linear-lanceolate, with a long, yellow arista at apex, erect and longitudinally folded, the outer densely woolly, the inner scarious. Flowers all tubular, shallowly 5-lobed, yellow, the inner female ones 20–25, the inner bisexual ones (1–)2–3(–4). Receptacle flat, pitted, the inner flowers without scales, the outer with scales doubtfully distinguish-

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able from inner involucral bracts. Achenes 0.5–0.8 × about 0.2 mm, brown, oblong to ovoid, somewhat compressed, papillose; pappus of inner achenes of scabrid hairs longer than the body of the achenes, in outer achenes absent. Flowers 7–8. 2n = 28. Native. Rather bare places on heaths, dry pastures, fields, pits and waysides, usually on sandy soils. Throughout most of the British Isles, but local in some areas and absent from most of west and north-west Scotland. Central and south Europe northwards to south Sweden and Poland; Siberia and western Asia; North Africa; Canary Islands; introduced in North America. A member of the European Southerntemperate element. 2. F. lutescens Jord. Red-tipped Cudweed F. apiculata G. E. Sm. ex Bab. nom. illegit.; F. germanica auct.; Gifola apiculata Chrtek & Holub nom. illegit. Annual herb with fibrous roots. Stems 10–25 cm, greyishwhite, erect or ascending, densely woolly, simple or rather irregularly branched, leafy. Leaves 15–20 × 3–6 mm, oblong-lanceolate to spathulate, obtuse or apiculate at apex, entire and hardly undulate, rounded at base, covered with greenish-yellow woolly hairs. Capitula 2.0–2.5 mm in diameter, conical-ovoid, weakly 5-angled, in clusters of (5–)10–20(–25) terminating the main stem and branches, the clusters (8–)10–14 mm wide, overtopped by 1–2 subtending leaves. Involucral bracts in a few rows, 4.0– 4.4 × 1.0–1.3 mm, reddish-purple before anthesis, then yellowish, oblong-ovate to lanceolate, with a straight, reddish arista up to 1.5 mm at apex, the outer densely, yellowishwoolly. Flowers all tubular, shallowly 5-lobed, yellow, the inner female ones 12–20 and about 3 mm, and the inner bisexual ones (2–)3–4(–5). Receptacle flat, pitted, the inner flowers without scales, the outer with scales doubtfully distinguishable from the inner involucral bracts. Achenes 0.5– 0.8 mm, brown, oblong-cylindrical, somewhat compressed, papillose; pappus of inner achenes of scabrid hairs, in the outer absent. Flowers 7–8. 2n = 28. Native. Sandy fields, waysides and pits. Rare and decreasing in south and east England from Hampshire to Yorkshire, formerly west to Worcestershire. Much of Europe from Denmark and southern Sweden southwards to Portugal, Sicily and Bulgaria; western Asia; North Africa. A member of the European Temperate element. Our plants are referable to subsp. lutescens which occurs throughout most of the range of the species. 3. F. pyramidata L. Broad-leaved Cudweed F. spathulata auct.; F. germanica Huds. Annual herb with fibrous roots. Stems (2–)5–30(–40) cm, ascending or erect, greyish-white, densely woolly, often with several decumbent or ascending branches from near the base of the rather short stem, all stems with divaricate branches above, leafy. Leaves (5–)10–15 × 3–6 mm, linear-oblong to spathulate, widest above the middle, obtuse or apiculate at apex, entire, hardly undulate, rounded at base, covered with greyish-white woolly hair. Capitula

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2.0–2.5 mm in diameter, pyramidal, sharply 5-angled, in clusters of 5–20 terminating the main stem and branches, the clusters 5–12 mm wide, overtopped by 2–4(–5) subtending leaves. Involucral bracts in a few rows, 2.5–4.5 × 1.0– 1.3 mm, pale yellow without reddish coloration, oblonglanceolate, with a recurved, yellowish arista at apex, keeled, distinctly divergent in fruit, the outer softly woolly on the back. Flowers all tubular, shallowly 5-lobed, yellow, the inner female ones (0–)5–7, the inner bisexual ones 4–7. Receptacle flat, pitted, the inner flowers without scales, the outer with scales doubtfully distinguishable from the inner involucral bracts. Achenes 0.5–0.8 mm, pale brown, cylindrical to oblong-ovoid, somewhat compressed; pappus of inner achenes of scabrid hairs longer than the body, in outer absent. Flowers 7–8. 2n = 28. Native. Sandy and chalky fields, waysides and pits. Very local and decreasing in southern England, formerly north Lincolnshire and in Jersey. South and west Europe northwards to the Netherlands and eastwards to Bulgaria and the Crimea; western Asia; North Africa; Canary Islands. A member of the Submediterranean-subatlantic element. Subgenus 2. Logfia (Cass.) P. D. Sell Logfia Cass.; Oglifa Cass. Capitula in small clusters or solitary. Involucral bracts obtuse to acute, never acuminate, spreading star-like in fruit. Outer achenes enclosed in involucral bracts and falling with them. 4. F. arvensis L. Field Cudweed Logfia arvensis (L.) Holub; Gnaphalium arvense L.; Gnaphalium lagopus Stephan ex Willd.; F. arvensis var. lagopus (Stephan ex Willd.) DC.; F. lagopus (Stephan ex Willd.) Parl.; F. arvensis subsp. lagopus (Stephan ex Willd.) Nyman Annual herb with fibrous roots. Stems 5–70 cm, erect or decumbent, simple or branched, leafy, white woolly. Leaves (6–)10–20 × 1–4 mm, oblong to linear-lanceolate, obtuse to abruptly acute at apex, entire, rounded at base, erect but not appressed to stem, generally overlapping, white woolly. Capitula 2.5–5.0 mm in diameter, broadly ovoid, terete, in clusters of (1–)3–12 in a racemose or paniculate inflorescence not overtopped by subtending leaves. Involucral bracts in a few rows, 4–5 × 2.0–2.5 mm, ovate-lanceolate, acute at apex, with woolly hairs except at the hyaline apex, slightly saccate at base, spreading star-like in fruit. Flowers all tubular, shallowly 5-lobed, yellow, the female inner ones 15–18, the inner bisexual ones about 3. Receptacle flat, pitted, the inner flowers without scales, the outer with scales doubtfully distinguishable from the inner involucral bracts. Achenes 0.6–1.1 × 0.2–0.3 mm, pale brown, the outer obliquely oblong, enclosed in involucral bracts and falling with them, the inner oblong-obovoid; pappus 2.5–3.5 mm, white, of scabridulous hairs. Flowers 7–9. 2n = 28. Introduced. Once a regular casual, but now only occasionally recorded. Native in most of Europe to about 65◦ N in Scandinavia; western Asia; Canary Islands.

5. F. minima (Sm.) Pers. Small Cudweed Gnaphalium minimum Sm.; Logfia minima (Sm.) Dumort.; Oglifa minima (Sm.) Rchb. fil. Annual herb with fibrous roots. Stems 5–25(–30) cm, prostrate, ascending or erect, slender, simple or irregularly branched, leafy, greyish hairy. Leaves 4–10 × 0.5–1.2 mm, linear to oblong-linear, acute at apex, entire, rounded at base, erect and appressed to the stem, greyish hairy. Capitula 1.5–2.0 mm in diameter, pyramidal-ovoid, 5-angled, in clusters of 3–7 at the ends of stems and branches and in the forks of branches, the clusters 2–5 mm wide and not overtopped by subtending leaves. Involucral bracts in a few rows, 1.5–3.5 × 0.8–1.0 mm, linear-lanceolate to lanceolate, narrowed to an obtuse apex, the middle ones keeled with a slightly saccate base, woolly on the back and glabrous and scarious near the apex, the inner similar to middle but short and narrower with a non-saccate base and rounded apex, the innermost narrowly oblong, scarious and quite glabrous, all spreading star-like in fruit. Flowers all tubular, shallowly 5-lobed, yellow, the outer female, the inner female many, 1.8–2.5 mm, the inner bisexual 3–5, 2.2– 2.5 mm. Receptacle flat, pitted, the inner flowers without scales, the outer with scales like the inner involucral bracts. Achenes of outer flowers 0.8–0.9 mm, oblong and smooth, enclosed in involucral bracts and falling with them, the inner flowers 0.5–0.6 mm and papillose; pappus absent on outer flowers, in inner flowers about 2 mm, white, of scabridulous hairs. Flowers 6–9. 2n = 28. Very variable in habit, branching and number of capitula, but all variants sometimes occur in one population. Native. On neutral to acid sands and gravels in open vegetation of heaths, grassland, dunes, dry fields, waysides, waste ground, walls, pits and quarries. Throughout much of the Great Britain and Ireland, but commoner in the east than the west and rare in Ireland. Through much of Europe northwards to about 60◦ N in Scandinavia with a few scattered localities further north; Siberia. A member of the European Temperate element. 6. F. gallica L. Narrow-leaved Cudweed Gnaphalium gallicum (L.) Hill; Logfia gallica (L.) Coss. & Germ.; F. filiformis Lam. nom. illegit. Annual herb with fibrous roots. Stems 2–25 cm, erect or ascending, slender, greyish hairy, often branched throughout, leafy. Leaves 8–25 × 0.8–1.5 mm, linear to filiform, acute at apex, entire, usually with a slightly involute margin, rounded at base, held loosely erect, but with the tip somewhat spreading, greyish-green with short, silky hairs. Capitula 2.0–2.5 mm in diameter, pyramidal-ovoid, 5angled, in clusters of 2–14, usually much overtopped by subtending linear-lanceolate leaves. Involucral bracts in a few rows, 3.0–4.5 × about 1.5 mm, with some small outer, the main outer broadly triangular-lanceolate, long acute at apex, with a saccate base and completely enclosing the marginal female flowers, woolly on the back except for the infolded margins, the inner (receptacular scales) becoming more narrowly lanceolate, less hairy on the back and less concave-based from the outside inwards, subtending but not

45. Antennaria enclosing some achenes of the central flowers, all spreading star-like in fruit. Flowers all tubular, shallowly 5-lobed, yellow, the outer female, the inner female 8–12 and 1.8–2.5 mm, the inner bisexual 2–3 and 2.2–2.5 mm. Receptacle flat, pitted, with involucral bracts running into scales. Achenes oblong, of outer flowers 0.8–0.9 × about 0.4 mm, without a pappus and falling with the enclosing bracts, of inner flowers about 0.6 × 0.2 mm, covered with transparent papillae and a pappus 2.0–2.7 mm, white and of scabridulous hairs. Flowers 7–9. 2n = 28. ?Native. Open sandy and gravelly ground. Occurring in Sark in the Channel Islands since 1902. Formerly in a few localities from Hampshire to Essex from 1696 to 1955; introduced to its last known Essex site in 1994. Native of west and south Europe northwards to Belgium and eastwards to Bulgaria and Turkey; eastern Asia; North Africa; Azores. The species has a Submediterranean-subatlantic distribution. Ifloga verticillata (L. fil.) Fenzl, Micropsis spathulata (Pers.) Cabrera, Chevreulia sarmentosa (Pers.) Blake, Facelis retusa (Lam.) Sch. Bip. and Stuartina hamata Philipson have been recorded as wool casuals. 45. Antennaria Gaertn. Dioecious perennial herbs with a woody stock and slender, branched stolons bearing numerous leaf rosettes. Stems erect, simple, leafy. Leaves alternate, densely white-woolly beneath. Capitula in a small terminal cluster. Involucral bracts in several rows, scarious, erect in fruit. Flowers all tubular, shallowly 5-lobed at apex, those of the female plants filiform, lacking anthers, the styles with truncate branches with are apically hairy, and with a pappus of a single row of slender hairs united below and falling together, while those of the usually smaller male plants have tailed anthers and an abortive ovary bearing clavate pappus hairs free to the base and falling separately. Corolla of male usually whiter, sometimes pink, of female usually rose-pink. Receptacle flat, without scales. Achenes cylindrical or somewhat compressed. About 70 to several hundred species depending on the concept of a species as many are apomictic. Mainly North America and Eurasia, but a few in South America. ¨ Braun-Blanquet, J. (1951). Pflanzensoziologische Uberlegungen als Hilfsmittel zur Erkennung systematischer einheiten am Beispiel von Antennaria hibernica dargelegt. Vegetatio 3: 298– 300. 1. Capitula on long peduncles up to 40 mm 1(iii). dioica var. pedicellata 2. 1. Capitula sessile or on very short peduncles 2. Leaves glabrous or nearly so on upper surface 1(i). dioica var. dioica 2. Leaves white woolly on upper surface. 1(ii). dioica var. hyperborea

1. A. dioica (L.) Gaertn. Mountain Everlasting Gnaphalium dioicum L.; A. montana Gray nom. illegit. Dioecious perennial herb with a creeping, woody stock and slender, branched stolons with numerous leaf rosettes

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which form mats. Stems 5–20(–30) cm, white-tomentose, erect, simple, leafy. Leaves medium green and glabrous or sparsely hairy or white-woolly on upper surface, whitewoolly beneath; basal 10–35(–40) × 3–10 mm, ovatespathulate, obtuse, apiculate or emarginate at apex, entire, shortly narrowed to a petiole-like base; cauline narrowly lanceolate to linear, acute with a short, green mucro at apex, entire, narrowed to a sessile base. Capitula about 12 mm wide in functionally female plants, about 6 mm on functionally male plants, 1–8(–12) in a terminal corymbose cluster, usually more or less sessile but sometimes on peduncles up to 40 mm. Involucral bracts in several rows, 6–8 × 0.5– 0.7 mm, the outer woolly below, scarious and glabrous above, the male capitula with them obovate-spathulate, obtuse at apex, white or sometimes pink and spreading above like ray flowers, the female capitula with them linear-lanceolate, more or less acute, usually rose-pink and erect. Flowers all tubular, shallowly 5-lobed, those of female plants usually rose-pink, filiform and lacking anthers, those of the usually smaller male plants often whiter, with tailed anthers and an abortive ovary. Receptacle flat, without scales. Achenes 0.9–1.0 mm, cylindrical or somewhat compressed; pappus of female flowers a single row of slender hairs united below and falling together, those of male plants with clavate hairs free to the base and falling separately. Flowers 6–7. Visited by various insects. 2n = 28. (i) Var. dioica Leaves glabrous or nearly so on upper surface. Capitula sessile or on short peduncles. (ii) Var. hyperborea (D. Don) DC. A. hyperborea D. Don Leaves white woolly on upper surface. Capitula sessile or with short peduncles. (iii) Var. pedicellata F. B. White A. hibernica Braun-Blanq. Leaves glabrous or nearly so on upper surface. Capitula on long peduncles up to 40 mm. Native. Heaths, dry pastures and dry mountain slopes, usually over limestone, basic igneous rock or base-rich boulder-clay, perhaps most commonly where the surfacesoil is somewhat leached, up to 914 m in Scotland. Locally common in northern and western Scotland and northern and western Ireland, occasional in southern Scotland and northern England, with scattered localities south to southern Ireland and Cornwall, much reduced in southern and eastern Great Britain, no longer in central-east and southeast England except for one locality in Northamptonshire. North and central Europe; Siberia; western Asia and North America, though other taxa may be involved. A member of the Eurasian Boreo-temperate element. The usual plant in Great Britain and Ireland is var. dioica. Var. hyperborea is recorded from the Scottish islands and mountains and the English Lake District and similar plants occur in Continental Europe especially in the Alps. Var. pedicellata is recorded from Scotland and Ireland. The ecological value

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of var. hyperborea and var. pedicellata is not understood, but they seem to be worth recording. 46. Anaphalis DC. Dioecious to variably sexed perennial herbs with rhizomes. Stems erect to ascending. Leaves alternate, not in a rosette, simple and entire, whitish. Capitula small, in a large, terminal, corymbose inflorescence. Involucral bracts in several rows, scarious, mainly pearly white. Flowers all tubular, shallowly 5-lobed at apex, male and female variously arranged, the males with wider corollas. Corolla yellowish. Anthers with short tails, apical appendage ovate. Style branches truncate with apical hairs. Receptacle flat or convex, without scales. Achenes spindle-shaped; pappus of 1 row of simple hairs. About 110 species, mainly in Asia, but a few species in North America; widely cultivated. 1. Leaves 8–15(–20) mm wide, with 1 prominent vein 1. margaritacea beneath 2. Leaves 5–30 mm wide, with 3 prominent, parallel veins 2. triplinervis beneath

1. A. margaritacea (L.) Benth. Pearly Everlasting Gnaphalium margaritaceum L.; Antennaria margaritacea (L.) Gray Perennial, sometimes slightly woody herb with rhizomes. Stems 30–100 cm, pale green, erect to ascending, robust, densely white-woolly, very leafy, branched only in the inflorescence. Leaves all cauline, numerous, 3–10 × 0.8– 1.5(–2) cm, medium green, linear or narrowly elliptical, rarely lanceolate, narrowed to an acute apex, more or less entire, revolute, gradually narrowed to a sessile base, whitewoolly, often becoming glabrous on the upper surface; vein 1, prominent beneath. Capitula 9–12 mm in diameter, in dense, terminal corymbs; peduncles short, densely whitewoolly. Involucral bracts in several rows, 2–4 × 1.5–2.0 mm, brown below, shining white above, elliptical, ovate or oblong, rounded-obtuse at apex, the outer white-woolly below, the inner glabrous. Flowers all tubular, shallowly 5lobed at apex, yellowish, male plants with abortive ovaries, female plants without anthers except for a few central bisexual flowers. Receptacle slightly convex, without scales. Achenes spindle-shaped, papillose, hairy; pappus 5–6 mm, whitish, of 1 row of simple hairs, slender in fertile flowers which are free to the base and fall separately, thickened above in male flowers. Flowers 8–9. 2n = 28. This species is extremely variable both in nature and in gardens, but it is said not to show distinct ecological or geographical variation. Plants with rather few, oblong leaves which are green on the upper surface are var. margaritacea (var. occidentalis Greene). They seem to be rare in our flora. Dwarf plants with a densely woolly upper surface of the leaves, var. subalpina A. Gray, seem also to occur. The common plants with us have linear or narrowly elliptical leaves, var. angustior (Miq.) Nakai (var. revoluta Suksd.) and have the upper surface of the leaves green and glabrous, while var. intercedens Hara is similar but has them whitewoolly.

Introduced. An established garden escape widely naturalised on railway banks, river sides, grassy and waste places and roadsides. Scattered throughout the British Isles and locally common in the west, especially in south Wales. In the Rhymney Valley in south Wales it has been known since 1698. Native of North America and eastern Asia. 2. A. triplinervis (Sims) C. B. Clarke Three-nerved Pearly Everlasting Antennaria triplinervis Sims Perennial, sometimes slightly woody herb with rhizomes. Stems 20–100 cm, pale green, erect, robust, densely whitewoolly, very leafy, branched only in the inflorescence. Leaves all cauline, numerous, 3–10 × 0.5–3.0 cm, medium green, lanceolate or elliptical, obtuse to more or less acute at apex, more or less entire, narrowed below, the lower subpetiolate, the upper sessile and semiamplexicaul, arachnoid-hairy on upper surface, grey-woolly beneath; with 3 longitudinal veins which are prominent beneath. Capitula 11–13 mm in diameter, in dense terminal corymbs; peduncles short, densely white-woolly. Involucral bracts in several rows, 4–7 × 1.5–2.0 mm, shining white with brown and green bases, lanceolate or ovate, obtuse to acute at apex, glabrous. Flowers all tubular, shallowly 5-lobed at apex, yellowish, male plants with abortive ovaries, female plants without anthers except for a few central bisexual ones. Receptacle slightly convex, without scales. Achenes spindle-shaped, papillose, hairy; pappus 5–7 mm, whitish, of simple hairs. Flowers 8–10. Introduced. Casual garden escape, but may have been overlooked as A. margaritacea. Recorded from near Diss in Suffolk, Glentarbuck in Dunbartonshire and Tholthorpe in Yorkshire. Native of the Himalayas. 47. Gnaphalium L. Annual to perennial herbs. Stems prostrate to erect, leafy. Leaves alternate, entire, usually tomentose. Capitula small, variously arranged from solitary to large clusters. Involucral bracts in several rows, whitish to yellowish or brown, subherbaceous to scarious. Flowers all tubular, shallowly 5-lobed, the inner bisexual, the outer female with narrower corollas. Corolla yellowish or purplish. Anthers with tails, apical appendage ovate, flat. Styles with oblong-linear branches. Receptacle flat, without scales. Achenes cylindrical or obovoid, not or little flattened; pappus of 1 row of simple hairs, sometimes ciliate below, free to the base or connate in a basal ring. About 200 species, mainly in temperate regions of both the Old and New Worlds and on tropical mountains, with a few in the Arctic. In Flora Europaea 4: 126–128 (1976) Gnaphalium, as regards the British and Irish species as defined here, has been divided into four genera. In this respect we have followed Stace (1997) and included them all in Gnaphalium as subgenera. If technical characters are not considered Filago and Gnaphalium can be recognised at a glance by their general appearance. G. viscosum Kunth has been recorded as a casual.

47. Gnaphalium Hilliard, O. M. & Burtt, B. L. (1981). Some generic concepts in Compositae – Gnaphaliinae. Bot. Jour. Linn. Soc. 82: 181– 232. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [G. sylvaticum.] Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [G. luteoalbum and G. norvegicum.] 1. Capitula in terminal, subglobose or subcorymbose 2. clusters 1. Capitula in elongated, racemose clusters, sometimes few 4. or rarely 1 2. Clusters of capitula overtopped by subtending leaves; 5. uliginosum involucral bracts brown 2. Clusters of capitula not overtopped by subtending leaves; 3. involucral bracts scarious, straw-coloured or whitish 3. Leaves white-woolly on both sides, not decurrent down 6. luteoalbum the stem 3. Leaves green on upper surface, white-woolly beneath, 7. undulatum decurrent down the stem 4. Annual to biennial without non-flowering shoots; involucral bracts acute to acuminate at apex; achenes less 4. purpureum than 1 mm 4. Perennials with leafy non-flowering shoots; involucral 5. bracts obtuse or emarginate; achenes more than 1 mm 5. Capitula less than 10 per stem; pappus hairs free, falling 3. supinum separately 5. Capitula normally more than 10 per stem; pappus hairs 6. united at base, falling as a unit 6. Cauline leaves 1- (or indistinctly 3)-veined, gradually 1. sylvaticum decreasing in size from the base up the stem 6. Cauline leaves 3(–5)-veined, scarcely decreasing in size 2. norvegicum up the stem until above half way

Subgenus 1. Omalotheca (Cass.) Endl. Omalotheca Cass. Perennial herbs with non-flowering shoots. Leaves alternate. Inflorescence terminal, spicate or solitary. Involucral bracts brownish; indurated tissue of the inner ones entire. Flowers all fertile, the outer female, the inner bisexual, the bisexual flowers reddish-purple at apex. Pollen grains with broad furrows, or bicular pores and slender spinules. Achenes 1.4–1.6 mm, with imbricate papillae and hairs 6– 7 times as long as wide, not emitting mucilage in water; pappus present. 1. G. sylvaticum L. Heath Cudweed Omalotheca sylvatica (L.) Sch. Bip. & F. W. Schultz Short-lived perennial herb with a woody stock, with short, ascending, non-flowering shoots as well as flowering ones. Stems (5–)20–60(–70) cm, erect, whitish-tomentose, simple, leafy. Leaves 2–8 × 0.2–0.8 cm, whitish-tomentose, becoming glabrous and greenish-grey on upper surface, the basal and lower cauline narrowly lanceolate, acute at apex, entire and narrowed into a petiole-like base, the middle and upper cauline gradually getting smaller upwards, linear-lanceolate to linear, acute at apex, entire, narrowed to a sessile base; 1- to 3-veined. Inflorescence spike occupying one-third to five-sixths the length of the stem, leafy,

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lax and often interrupted below. Capitula 6–7 mm in diameter, in clusters of 2–8 or sometimes solitary in leaf axils. Involucral bracts in several rows, 1.5–5.5 × 1.0–1.5 mm, the outer brown with the upper end and margins hyaline, broadly oblong, obtuse and more or less emarginate at apex and tomentose in the lower half, the inner longer, the innermost about equalling the flowers, usually with a pale green central stripe and pink or silvery upper end and margins. Flowers all tubular, 0.7–1.0 mm in diameter, yellowish, the outer about 70 and female, the inner 3–4 bisexual, and reddish-purple at apex. Corolla 3–4 mm, shallowly 5-lobed at apex. Receptacle flat, without scales. Achenes 1.4–1.6 mm, cylindrical, with imbricate papillae and stiff hairs 6–7 times as long as wide; pappus 3.5–3.8 mm, reddish, of 1 row of simple eglandular hairs united at the base and falling as a unit. Flowers 7–9. Little visited by insects and probably wind-pollinated. 2n = 56. (i) Var. sylvaticum G. rectum Sm., G. sylvaticum var. rectum (Sm.) Gaudin Plant up to 40 cm. Basal leaves up to 5 mm wide. Inflorescence long and not congested. (ii) Var. alpestre Br¨ugger G. sylvaticum var. nigrescens auct. Plant up to 14 cm. Basal leaves up to 8 mm wide, often more hairy on upper surface. Inflorescence fairly short and congested. Native. Calcifuge and found on moorland and forestry tracks in the north and dry, open woods, dunes, heaths and sandy field margins elsewhere. Essentially a lowland species but occurs up to 950 m in Scotland. Scattered throughout Great Britain and Ireland, but has much declined in recent years. Europe to 70◦ N in Scandinavia; western and central Asia; eastern North America; introduced in New Zealand. A member of the Eurosiberian Boreo-temperate element. The lowland plant is var. sylvaticum, the Scottish upland plant var. alpestre, but the exact distributions of the two variants are not known. 2. G. norvegicum Gunnerus Highland Cudweed Omalotheca norvegica (Gunnerus) Sch. Bip. & F. W. Schultz; G. sylvaticum var. norvegicum (Gunnerus) Hook. & Arn.; G. sylvaticum subsp. norvegicum (Gunnerus) Hook. fil. Perennial herb with a short woody stock and short, ascending, non-flowering shoots as well as flowering ones. Stems (5–)15–30(–40) cm, erect, simple, whitishtomentose, leafy. Leaves 5–12 × 0.4–1.8 cm, whitishtomentose, becoming greyish-green on upper surface, the basal and lower cauline oblanceolate, acute at apex, entire, gradually narrowed to a petiole up to as long as the lamina, 3–5-veined, the median and upper cauline deminishing abruptly in length only in the short inflorescence, similar to basal but narrower, linear-lanceolate, acuminate at apex and with shorter petioles to sessile. Inflorescence spike occupying one-tenth to one-quarter the length of the stem, compact, its subtending leaves as long as or longer than the inflorescence. Capitula 6–7 mm in diameter, solitary or

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2–3 together in the axils of the upper leaves. Involucral bracts in several rows, 1.5–5.0 × 1.0–1.5 mm, with an olive-green central stripe and dark brown scarious margins, unequal, more or less elliptical, obtuse at apex, the inner shorter than the flowers. Flowers all tubular, 0.5– 0.7 mm in diameter, yellowish, the outer 35–45, female, the inner 3–4 bisexual and reddish-purple at apex. Corolla 3.0–3.5 mm, shallowly 5-lobed at apex. Receptacle flat, without scales. Achenes 1.4–1.6 mm, pale brown, cylindrical, with imbricate papillae and stiff hairs 6–7 times as long as wide; pappus 3–4 mm, white, of 1 row of simple eglandular hairs, united at base and falling as a unit. Flowers 8. 2n = 56. Native. Mountain rocks and gravel in gullies and stream gorges up to 1,100 m. Very local in Forfarshire, Aberdeenshire, Inverness-shire and Ross-shire. Mountains of central and arctic and subarctic Europe, Balkan peninsula; Caucasus; western Asia; eastern North America; Greenland. A member of the European Arctic-montane element. 3. G. supinum L. Dwarf Cudweed Omalotheca supina (L.) DC. Dwarf, tufted perennial herb with a slender, creeping, branched stock, with numerous, short, leafy, non-flowering as well as flowering stems. Stems 2–12(–25) cm, whitishwoolly at least towards the apex, erect, simple, leafy. Leaves 0.5–2.0(–2.5) × 0.1–0.3 cm, whitish- or greywoolly on both surfaces, the basal linear-oblanceolate, subacute at apex, entire and narrowed below, the cauline linear, acute at apex, entire and slightly widened at base, 1veined. Inflorescence a short, more or less compact terminal spike somewhat lengthening in fruit. Capitula 7– 8 mm in diameter, 1–7 in a spike. Involucral bracts in 2–4 rows, 1.5–5.0 × 1.0–1.5 mm, with an olive central strip and broad, brown, scarious margins, broadly elliptical, obtuse at apex, woolly, the inner almost equalling the flowers, becoming stellate-patent. Flowers all tubular, 0.4– 0.5 mm in diameter, the outer few and female, the inner bisexual, and slightly shorter with shorter lobes. Corolla 3.0–3.4 mm, yellow, shallowly 5-lobed at apex. Receptacle flat, without scales. Achenes 1.0–1.5 mm, brown, obovoid, compressed, shortly hairy, the hairs 2-fid at apex; pappus 3.0–3.5 mm, whitish, of 1 row of stout simple eglandular hairs, free and falling separately. Flowers 7–8. 2n = 28. Native. Mountain rocks and gravel. Scotland from Stirlingshire to Sutherland, and in Skye. Mountains of central Europe and western Asia; arctic Europe; Greenland; North America. A member of the European Arctic-montane element. Subgenus 2. Gamochaeta (Weddell) Gren. Gamochaeta Weddell Annual to perennial herbs without non-flowering shoots. Basal leaves in a rosette. Inflorescence terminal, spicate, more or less leafy. Involucral bracts brownish-tipped; indurated middle portion of the inner ones not sulcate. Flowers all fertile, the outer female, the inner bisexual, all purple. Achenes 0.8–1.0 mm, with papillae, mucilaginous when

wet; pappus of barbed hairs, connate in a basal ring and falling as a unit. 4. G. purpureum L. American Cudweed Gamochaeta purpurea (L.) Cabrera: G. pensylvanicum Willd. nom. illegit. Annual or perennial herb with a tap-root and without nonflowering stems. Stems 10–40 cm, decumbent to erect, usually simple, appressed-tomentose, leafy. Leaves 1–6(–10) × 0.5–1.5(–2.0) cm, usually green and sparsely hairy above, white-tomentose beneath, the basal in a rosette, dead but persisting at anthesis, spathulate to oblanceolate, rounded at apex and often mucronate, entire and narrowed below, the cauline progressively smaller upwards, lanceolatespathulate, subacute at apex, narrowed below. Inflorescence a more or less elongated terminal spicate cluster. Capitula 4–5 mm in diameter, in clusters subtended by leaves. Involucral bracts in several rows, 3–5 × 1.0–1.2 mm, green with brown tips, sometimes tinged pink, ovate-lanceolate to lanceolate, acute to acuminate at apex, sparsely hairy to glabrous. Flowers all tubular, 0.6–0.8 in diameter, the outer few female, the inner bisexual. Corolla purple and 2–3 mm, shallowly 5-lobed at apex. Receptacle flat, without scales. Achenes 0.8–1.0 mm, olivaceous to yellowishbrown, often shiny, cylindrical, minutely papillose; pappus of barbed hairs, connate in a basal ring and falling as a unit. Flowers 7–8. 2n = 14, 18, 28. Introduced. Naturalised in a churchyard in Surrey since 1978. Native of North America; naturalised in Portugal and the Azores. Subgenus 3. Gnaphalium Filaginella Opiz Annual herbs without non-flowering shoots. Leaves alternate. Inflorescence terminal and axillary, clusters of capitula subtended by leaves. Involucral bracts herbaceous and scarious, mottled, indurated tissue of inner involucral bracts in irregular patches. Flowers all fertile, the outer female, the inner bisexual. Pollen grains with narrow, shallow furrows, ovate pores and short, broadly conical spinules. Achenes 0.4–0.7 mm, not papillose, glabrous or hairy, the hairs 2–4 times as long as wide, omitting mucilate in water; pappus of filiform, fragile free hairs not ciliate at the base and falling separately. 5. G. uliginosum L. Marsh Cudweed Filaginella uliginosa (L.) Opiz Annual herb with a slender tap-root and fibrous sideroots, without non-flowering shoots. Stems numerous, (1–)4–20 cm, whitish-tomentose, decumbent, ascending or erect, divaricately branched from base, leafy. Leaves 1–5 × 0.2–0.5 cm, whitish-tomentose, linear-oblong to oblanceolate, more or less acute at apex, entire but often undulate, narrowed to a sessile base. Inflorescence of terminal and axillary clusters. Capitula 4–5 mm in diameter, 3–10 in a cluster overtopped by subtending leaves. Involucral bracts 1–3 × 0.5–0.7 mm, longer inwards, the innermost at least equalling the flowers, the outermost less than half as long, brown and glabrous near the apex, paler and more or less hairy below, oblong to linear, subobtuse at

48. Helichrysum apex, spreading in fruit. Flowers all tubular, 0.2–0.3 mm in diameter, all female except for 5–8 bisexual ones in centre. Corolla 1.4–1.5 mm, yellowish, shallowly 5-lobed at apex. Receptacle flat, without scales. Achenes 0.4–0.7 mm, olive-brown, oblong-cylindrical, not papillose, more or less glabrous or with hairs 2–4 times as long as wide; pappus 1.4–1.5 mm, whitish, of 1 row of filiform, free, fragile hairs, falling separately. Flowers 7–9. Little visited by insects. 2n = 14. Native. Damp places in arable and waste land, banks of streams, ditches and ponds, woodland rides and waysides. Throughout Great Britain and Ireland on moderately acid soils. Most of Europe to about 68.5◦ N in Scandinavia; Asia; introduced in North America. A member of the Eurasian-temperate element. Our plant is subsp. uliginosum which occurs throughout much of the European range of the species. Subgenus 4. Laphangium (Hilliard & B. L. Burtt) P. D. Sell Pseudognaphalium Kirp.; Pseudognaphalium subgenus Laphangium Hilliard & B. L. Burtt Annual herbs without non-flowering shoots. Leaves alternate, semiamplexicaul. Inflorescence terminal, cymosecorymbose, clusters of capitula without subtending leaves. Involucral bracts scarious, entirely shining white or yellow, indurated middle portion of the inner ones sulcate. Flowers all fertile, the outer in 2–4 rows and female, the inner bisexual, all yellowish or reddish above. Pollen grains with narrow, shallow furrows, ovate pores and short, broadly conical spinules. Pappus of filiform, fragile, free hairs, ciliate at the base and falling in small groups. 6. G. luteoalbum L. Jersey Cudweed Pseudognaphalium luteoalbum (L.) Hilliard & B. L. Burtt Annual herb with a slender tap-root and fibrous sideroots, without non-flowering shoots. Stems 8–50 cm, whitewoolly, erect or decumbent at base then erect, the main stem and branches simple below but corymbosely branched above, leafy. Leaves white-woolly on both surfaces; basal 1.5–3.0(–7.0) × 0.5–0.7 cm, broadly oblanceolate, obtuse at apex, entire but more or less undulate, narrowed to a petiolelike base; cauline (1–)2–5(–7) cm, oblong to linear, the lower obtuse, the upper acute at apex, entire but undulate, narrowed to a semiamplexicaul base. Inflorescence terminal and subcorymbose. Capitula 4–5 mm in diameter, ovoid, in clusters of 4–12, without subtending leaves. Involucral bracts 1–4 × 0.7–1.2 mm, straw-coloured, mostly scarious, the outer broadly ovate and obtuse at apex, the inner oblong and obtuse at apex, glabrous except the outer which are slightly woolly below. Flowers all tubular, 0.2–0.3 mm in diameter, the outer female in 2–4 rows, the inner 4–7(–10) bisexual. Corolla 2.9–3.1 mm, yellowish, becoming reddish near the toothed apex. Receptacle flat, without scales. Achenes 0.4–0.5 mm, brown, cylindrical, tuberculate, glabrous or hairy; pappus 2.0–2.5 mm, whitish, filiform, fragile, free hairs, ciliate at base, falling in small groups. Flowers 6–8. Visited by flies and bees. 2n = 14. Native. Sandy fields, waste places and dunes. Very local in Guernsey, Jersey, Norfolk and Kent, formerly in Suf-

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folk and Cambridgeshire; a casual or established alien at scattered sites elsewhere. Europe, northwards to Holstein, southern Sweden and Latvia; temperate, western and central Asia. A member of the Eurosiberian Southerntemperate element; widely naturalised outside its native range. 7. G. undulatum L. Cape Cudweed Pseudognaphalium undulatum (L.) Hilliard & B. L. Burtt Annual bushy herb with a tap-root, without non-flowering shoots. Stems (20–)30–80(–90) cm, erect, sparingly whitewoolly, with spreading branches above, leafy. Leaves green on upper surface, white-woolly beneath, glandular on upper surface, the lower 2–8 × 0.5–1.2 cm, oblong-lanceolate or lanceolate, acute to acuminate at apex, entire and undulate, broad and decurrent at base the wings often reaching to the node below or sometimes lower; the upper linear-lanceolate, very acute at apex, entire and undulate and more or less cordate at base. Inflorescence a large, lax terminal corymb, sometimes a panicle. Capitula 4–5 mm in diameter, ovoid, in globose clusters not overtopped by subtending leaves. Involucral bracts in 4 rows, 1.5–3.0 × 0.7–1.2 mm, whitish, scarious, the outer ovate and acute at apex, the inner oblong and obtuse at apex, woolly at base. Flowers all tubular, 0.2– 0.3 mm in diameter, the outer 33–69 female, the inner 3–9 bisexual. Corolla 2–3 mm, yellowish, shallowly 5-lobed at apex. Receptacle flat, without scales. Achenes 0.5–0.6 mm, brown, cylindrical, minutely mamillate; pappus 2.5– 3.0 mm, of numerous, filiform, fragile, free hairs, ciliate at the base, falling in small groups. Flowers 6–8. Introduced. Rough and waste ground and cliffs. All main islands of the Channel Islands since 1888. Native of South Africa. Troglophyton parvulum (Harvey) Hilliard & B. L. Burtt and Vellereophyton dealbatum (Thunb.) Hilliard & B. L. Burtt have been recorded as wool casuals. 48. Helichrysum Mill. Mat-forming shrubs, woody perennials or annuals. Stems erect or spreading. Leaves alternate, simple and entire. Capitula terminal and solitary, or in a corymb. Involucral bracts in several rows, scarious, white, yellow, orange or red. Flowers all tubular, 5-lobed the inner bisexual, the outer female with the corolla narrower. Corolla yellow or cream. Anthers with tails, apical appendage ovate, flat. Style with oblong-linear, truncate branches. Receptacle slightly convex, without scales. Achenes more or less cylindrical; pappus of 1 row of hairs. About 600 species in Eurasia, Africa and Madagascar. H. capillaceum (Thunb.) Less., H. expansum (Thunb.) Less. and H. odoratissimum (L.) Less. have been recorded as wool casuals and H. italicum (Roth) Don as a casual garden escape. 1. Annual herb; leaves with sparse hairs; involucral bracts 1. bracteatum bright yellow, orange, red or white 1. Woody perennials or shrubs; leaves glabrous or grey- or white-woolly; involucral bracts usually milky white or 2. cream

432 2. Capitula solitary 2. Capitula in dense corymbs

1 4 8 . A S T E R AC E A E 2. bellidioides 3. petiolare

1. H. bracteatum (Vent.) Andrews Annual Everlasting-flower Xeranthemum bracteatum Vent. Annual herb with a tap-root and fibrous side-roots. Stem 20–60(–200) cm, pale green, erect, viscid-glandular-hairy and sparsely tomentose, becoming glabrous below and sometimes also above, branched above, leafy. Leaves 3.5– 10.0 × 0.5–1.5(–2.5) cm, medium green on upper surface, oblanceolate, narrowly elliptical or lanceolate, mucronate and acute at apex, entire, cuneate to the amplexicaul base, with sparse, short, rigid simple eglandular hairs and glandular hairs, spreading and widely spaced. Inflorescence corymbose. Capitula 1–5, 25–50 mm in diameter. Involucral bracts in 7–15 rows, membranous, glabrous, bright yellow, gold, orange or white to red, the middle ones 10– 15 mm, opaque, ovate and mucronate at apex, the outer much shorter and less brightly coloured, the inner shorter, narrower and often paler or partly transparent. Flowers tubular, yellow, 5-lobed. Receptacle convex, without scales. Achenes 2.0–2.5 mm, cylindrical, pale brown and darker flecked, glabrous; pappus of barbellate hairs in 1 row, slightly fused at base. Flowers 7–9. Introduced. Garden escape and wool casual. In a few scattered localities in Great Britain. Native of Australia. 2. H. bellidioides (G. Forst.) Willd. New Zealand Everlasting-flower Xeranthemum bellidioides G. Forst.; Gnaphalium bellidioides (G. Forst.) Hook. fil. Mat-forming, dwarf evergreen shrub. Stems (3–)5–15(–20) cm, rooting stems prostrate, flowering stems erect, slender, tomentose, much branched, leafy. Leaves 5–9(–15) × 2–4(–6) mm, medium green on upper surface, obovate to narrowly lanceolate, mucronate at apex, entire, cuneate to a sessile, amplexicaul base, usually glabrous but sometimes sparsely to densely white tomentose on the upper surface, tomentose on lower surface, loosely imbricate below, more open above. Capitula 14–22(–25) mm in diameter, solitary, pedunculate. Middle involucral bracts (5–)6–9(– 12) mm, milky white, rarely tinged rose or cream, membranous, elliptical to oblong, obtuse at apex, glabrous, spreading, tomentose. Flowers tubular, pale yellow, 5lobed. Receptacle convex, without scales. Achenes 0.8–1.4 mm, obovoid-cylindrical, slightly angled or compressed, glabrous; pappus a few, slender hairs in 1 row. Flowers 7–9. 2n = 28. This species varies considerably in stature and hairiness. Introduced. Naturalised in rocky turf by a stream at Tagon Voe in the Shetland Island. Native of New Zealand. 3. H. petiolare Hilliard & B. L. Burtt Silver-bush Everlasting-flower H. petiolatum auct.; Gnaphalium lanatum auct. Woody perennial herb. Stems 30–50 cm, ascending or erect, slender, pale green or suffused brownish-purple, whitewoolly. Leaves 15–2.5 × 1.5–2.5 cm, dark green, subrotund

to ovate, rounded-obtuse at apex, entire, rounded, subcordate or subtruncate at base, grey-woolly on both surfaces, but more dense below; petiole up to 1.5 mm, grey-woolly. Inflorescence of dense corymbs. Capitula numerous, 3–4 mm in diameter. Involucral bracts in about 5 rows, 3.0– 3.5 × 0.7–1.2 mm, cream, oblong or oblong-ovate, obtuse or bluntly mucronate at apex, glabrous. Flowers tubular, cream, 5-lobed. Achenes cylindrical; pappus of 1 row of hairs. Flowers 7–9. Introduced. Naturalised on dunes at Pentle Bay, Tresco and in a pit above Tremelethen of St Mary’s in the Isles of Scilly. Native of South Africa. (Rhodanthe charsleyae (F. Muell.) Paul G. Wilson, R. corymbiflora (Schltdl.) Paul G. Wilson, R. floribunda (DC.) Paul G. Wilson, R. manglesii Lindl. and R. moschata (Cunn. ex DC.) Paul G. Wilson have been recorded as wool casuals. Millotia perpusilla (Turcz.) P. S. Short is also a wool casual as are Myriocephalus rhizocephalus (DC.) Benth. and Calocephalus knappii (F. Muell.) Ewart & J. White. Ammobium alatum R. Br. is a casual garden escape.) 49. Cassinia R. Br. Evergreen shrubs. Stems erect. Leaves alternate small and entire, with a revolute margin. Capitula in small, dense, rounded corymbs. Involucral bracts in 3–5 rows. Flowers usually all tubular, shallowly 3-lobed and bisexual, rarely with a few outer flowers filiform and female. Corolla white or yellow. Anthers tailed, apical appendage ovate and flat. Style with oblong-linear, truncate branches. Receptacle convex, usually with scales. Achenes 4-angled; pappus in 1 row of denticulate hairs. About 20 species native to Australia and New Zealand. 1. C. leptophylla (G. Forst.) R. Br. Golden Bush Calea leptophylla G. Forst.; C. fulvida Hook. fil. Evergreen shrub up to 5 m, much branched. Stems erect, brown, more or less tomentose and sometimes glandularhairy; twigs yellowish. Leaves 1.5–10.0(–15.0) × 1–3(–4) mm, bright green on upper surface, oblong to narrowly obovate or elliptical, rarely linear or almost obtriangular, obtuse at apex, entire, with revolute margin, narrowed at base, from almost glabrous to moderately clothed with white tomentum on upper surface, with dense white tomentum on lower surface and sometimes with a sparse to dense covering of glandular hairs which give the leaves a yellowish appearance. Inflorescence of small, dense, rounded corymbs; peduncles short, tomentose. Capitula 3–4 mm in diameter, cylindrical. Involucral bracts in 3–5 rows, 1–4 × 0.3–0.4 mm, mostly translucent, sometimes tinted pink or red in bud, lanceolate or ovate-lanceolate, rounded-obtuse at apex, sparsely to moderately hairy. Flowers all tubular, 4–15(–25) per capitulum, 3.0–3.5 mm, white to yellow, 3-lobed at apex. Receptacle convex; scales few to many or absent, with a white lamina slightly exceeding the flowers. Achenes 1.0–1.7 mm, square in section and narrowed at the base, glabrous or with scattered antrorse hairs; pappus of denticulate, scarcely to distinctly thickened hairs in one row. Flowers 7–8.

50. Inula The broad concept for the species, with much geographical and ecological variation, accepted by C. J. Webb et al., Flora of New Zealand 4 (1988) is followed here. Introduced. Established by the side of the main road at Crianlarich in Perthshire and in a field on Raasay in the North Ebudes. Native of New Zealand. (Leontopodium alpinum Cass. has been introduced on chalk grassland at Pebblecombe in Surrey. Leysera capillifolia (Desf.) Spreng., L. tenella DC., Podolepis gracilis (Lehm.) Graham and P. longipedata A. Cunn. ex DC. have been recorded as wool casuals.) Tribe 4. Inuleae Cass. Tribe Buphthalmeae Lindl. Mostly perennial, sometimes annual herbs. Leaves alternate, simple. Capitula with tubular and ligulate flowers. Flowers yellow. Involucral bracts in several rows, herbaceous. Receptacular scales present or absent. Pappus of hairs, scales or a rim. Contains 38 genera and about 480 species mainly in the northern hemisphere particularly Europe, North Africa, Arabia, Middle East and central Asia with a few genera in tropical and southern Africa. 50. Inula L. Conyza L., non Less.; Eritheis Gray; Limbarda Adans. Perennial herbs, sometimes woody at base. Stems erect or decumbent, hairy. Leaves alternate, simple and entire. Capitula solitary or in a corymbose or paniculate inflorescence. Involucral bracts in many rows, herbaceous. Inner flowers tubular and bisexual with 5 short lobes; the outer ligulate and female with 3 lobes at apex, rarely all tubular. Anthers with branched tails, with firm apical appendages. Styles slender, hairs confined to the apical portion of the branches. Corolla yellow, rarely slightly reddish. Receptacle flat or slightly convex, without scales. Achenes cylindrical, oblong or 4-angled; pappus of 1 row of ciliate hairs. About 100 species in Europe, Asia and Africa. Grime, J. P. et al (1988). Comparative plant ecology. London. [I. conyza.] Macpherson, P. & Clement, E. J. (1999). Inula hookerii C. B. Clarke: established in Britain. B.S.B.I. News 81: 53. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [I. crithmoides.] 1. 1. 2. 2. 3. 3.

4. 4. 5.

8. crithmoides Stem and leaves fleshy, glabrous 2. Stem and leaves not fleshy, sparsely to densely hairy 7. conyzae Ligules absent or less than 10 mm 3. Ligules more than 10 mm Outer involucral bracts ovate; capitula 60–100 mm in 1. helenium diameter Outer involucral bracts oblong or linear-lanceolate or lanceolate or subulate-lanceolate; capitula 20–90 mm in 4. diameter 6. oculus-christi Outer involucral bracts lanate 5. Outer involucral bracts long hairy but not lanate Outer involucral bracts glabrous on the surface, but ciliate 2. salicina

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6. 5. Outer involucral bracts with hairs on surface and margin 6. Capitula 25–40 mm in diameter; achenes 1.0–1.5 mm 5. britannica 7. 6. Capitula 50–90 mm in diameter; achenes 2–3 7. Plant with glandular as well as simple eglandular hairs 3. orientalis 4. hookeri 7. Plant with simple eglandular hairs only

1. I. helenium L. Elecampane Corvisartia helenium (L.) M´erat; I. grandiflora Gray, non Willd. Robust perennial herb with a branching, aromatic, tuberous stock. Stem 60–250 cm, yellowish-green, sometimes tinted brownish or purple, stout, erect, furrowed, tomentose with dense unequal, pale simple eglandular and arachnoid hairs, leafy, simple or branched above. Leaves dull yellowishgreen on upper surface, with a very pale midvein, paler greyish-green beneath, with numerous, short simple eglandular hairs on upper surface, and dense, soft simple eglandular hairs beneath, especially on the veins, and dense minute glands; basal 17–70 × 8–25 cm, oblong-lanceolate, ovatelanceolate or ovate-elliptical, acute at apex, irregularly undulate-denticulate, cuneate or attenuate at base to a petiole up to 170 mm which is channelled above and rounded beneath; the cauline gradually decreasing in size, the lower similar to basal but more shortly stalked or subsessile, the middle and upper similar in shape to lower but becoming narrower, sessile and amplexicaul with broad, rounded auricles. Capitula few, 60–100 mm in diameter, in corymbs; peduncles robust, slightly swollen below the capitulum, yellowish-green, rigidly erect, with dense, unequal, pale, bulbous-based simple eglandular hairs. Involucral bracts in many rows, of 2 kinds; outer 10–13 × 4–5 mm, dull green, ovate, acute at apex, with spreading or recurved tips; inner 13–18 × 1.5–2.5 mm, green at base, brownish-purple above, lanceolate with a spathulate or acute apex; all with numerous to dense, short and medium, pale simple eglandular hairs. Flowers of 2 kinds, strong-smelling; outer up to 60 mm, ligulate, the ligules bright yellow, with 3 rather deep triangular lobes at apex; inner 15–18 mm, tubular, bright yellow and slightly deeper than outer, deeply and narrowly 5-lobed at apex. Receptacle slightly convex, pitted, with scarious margins to pits, without scales. Achenes 3–5 mm, rectangular, 4-angled, glabrous; pappus 5–7 mm, dirty white often tinged reddish, of about 30, subrigid simple eglandular hairs. Flowers 7–8. Visited by bees and hoverflies. 2n = 20. Introduced. Naturalised in fields, waysides and rough habitats. In scattered localities, especially old cottage gardens, throughout Great Britain and Ireland, but less common than in the past. Formerly much grown, especially for medicinal purposes, the rootstock being used as a remedy for coughs and colds and as a tonic, and candied with sugar as a popular sweetmeat. The chemical inulin, found in this species, has recently been accepted by mainstream medicine as a useful treatment for asthma. The Romans are said to have eaten the leaves as a potherb. Thought to be native in south-east Europe and western Asia, but widely naturalised elsewhere in Europe,

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and in North America and Japan. Named after Helen of Troy. 2. I. salicina L. Irish Fleabane I. cordata Boiss. Perennial herb with slender, hypogeal stolons. Stems 25– 50(–75) cm, pale yellowish-green, markedly ridged, stiffly erect, with few to numerous, short to medium, pale simple eglandular hairs especially in the upper part, simple or branched above, leafy. Leaves 2–6(–10) × 0.5–2.5 cm, dull medium green on upper surface, paler beneath, lanceolate, narrowly elliptical or oblong, acute at apex, entire to denticulate, narrowed to a cordate, amplexicaul base, markedly reticulate-veined, glabrous or nearly so on the upper surface, with sparse simple eglandular hairs on the veins beneath and short, rigid simple eglandular hairs on the margins. Capitula 20–40 mm in diameter, solitary or 2–5 in a corymb; peduncles ridged, with few to numerous simple eglandular hairs. Involucral bracts in many rows; outer 6–8 × 2.0–2.5 mm, pale green, oblong-lanceolate, acute at apex, glabrous on the surface, strongly ciliate on the margin; inner 10–11 × 1.0–1.5 mm, linear-lanceolate, acute at apex, with numerous, appressed simple eglandular hairs on the surface and ciliate on the margin. Flowers of 2 kinds, the outer 15–20 mm, ligulate, the ligules golden yellow and with 3 linear-lanceolate lobes at apex, the inner 10–15 mm, tubular, brownish-yellow and with 5 linear lobes at apex. Receptacle slightly convex, pitted, the pits with a scarious margin, without scales. Achenes 1.5–2.0 mm, oblong, glabrous; pappus 4–5 mm, whitish, of 30–35, rather rigid hairs. Flowers 7–8. 2n = 16. The Irish plant differs from the common Continental plant in being more hairy and the leaves denticulate, features noted by its discoverer, David Moore, but not described by most later authors. In these characters it is much more like subsp. aspera (Poir.) Hayek (I. aspera Poir.) which seems to be mainly in south-east Europe and south-west Asia. ?Native. On rough, grassy, stony ground with Schoenus nigricans, Molinia caerulea and Sesleria caerulea close to the winter flood-level. Confined to the limestone shores of Lough Derg in Tipperary and Galway where it is now almost extinct. The species is native in most of Europe and western Asia, reaching the Arctic Circle in Finland; it is a member of the Eurasian Temperate element. It is possible it was originally brought to Ireland by wildfowl and has spread vegetatively and only flowers irregularly. See also I. britannica. The above description is made entirely from Irish specimens. 3. I. orientalis Lam. Eastern Fleabane I. glandulosa Willd.; I. grandiflora Willd. Perennial herb with rhizomes. Stems 40–60 cm, pale yellowish-green, erect, with numerous spreading, yellowish or brown, stiff, medium to long simple eglandular hairs and short yellow glandular hairs, unbranched, more or less densely leafy. Leaves 5–12 × 1.5–4.0 cm, medium green on upper surface, paler beneath, ovate, ovate-elliptical or oblanceolate, acute at apex, entire, denticulate or minutely serrate, rounded or slightly attenuate at the semiamplexicaul base, with numerous to dense, medium to long, yellowish,

wavy simple eglandular hairs and minute glandular hairs on both surfaces and the margins. Capitula 50–90 mm in diameter, solitary. Involucral bracts in 4–5 rows, pale yellowishgreen; outer 7.5–12.0 × 1.0–1.5 mm, linear-lanceolate, long and narrowly acute at apex, with long, dense, brownish simple eglandular hairs and minute glandular hairs; inner 12–17 × 0.5–0.7 mm, linear, long-acute at apex, with long, brownish, simple eglandular hairs. Flowers of 2 kinds, the outer 17–25 mm, ligulate, the ligules golden yellow, very narrowly and minutely 3-lobed at apex and often with long, sparse simple eglandular hairs at base, the inner 6.5–7.0 mm, tubular, brownish-yellow, narrowly 5-lobed at apex. Receptacle slightly convex, pitted, the pits with a scarious margin, without scales. Achenes 2.0–2.5 mm, cylindrical, obscurely ribbed, glabrous or slightly hairy at apex; pappus 6.5–7.0 mm, brownish, of about 25 stiff hairs. Flowers 7–8. 2n = 16. Introduced. A persistent garden escape on Box Hill Common in Wiltshire and on a railway embankment at Tunbridge Wells in Kent. Native of the Caucasus. 4. I. hookeri C. B. Clarke Hooker’s Fleabane Perennial herb. Stem 30–45 cm, pale yellowish-green, slender, angled, with short, pale simple eglandular hairs below becoming more numerous and longer above, sparingly branched above, leafy. Leaves 2–13 × 0.5–4.0 cm, bright medium green on upper surface, paler and bluish-green beneath with a prominent midrib, elliptic-lanceolate, elliptical, lanceolate or oblong-lanceolate, obtuse to more or less acute at apex, minutely glandular-denticulate, cuneate at base, the lower shortly petioled, the upper sessile; all with short to medium, pale simple eglandular hairs on the upper surface and margin, usually tomentose beneath. Capitula 50–85 mm in diameter, solitary or 2–3 together at the ends of stems or leafy branches; peduncles rather short hairy. Involucral bracts in several rows, 10–25 mm, narrow linearsubulate, narrowed to a long, slender, acute apex, spreading and recurved, finely squarrose, with long, shaggy hair. Flowers of 2 kinds, the outer ligulate, the ligules long linear, pale yellow and 3-lobed at apex, the inner tubular and yellow. Receptacle convex, papillose. Achenes very small about 2–3 mm, deeply grooved; pappus about 4 mm, of 1 row of dirty white hairs. Flowers 7–9. 2n = 20. Introduced to gardens in 1849. A large colony naturalised in limestone woodland on the north-east side of Whitbarrow in Lancashire, another on a roadside at Kilnsey in Yorkshire and a third in an old sandy quarry west of Milncroft in Lanarkshire. Native of the Himalayas. 5. I. britannica L. Meadow Fleabane Perennial herb. Stems 15–75 cm, pale yellowish-green, erect, more or less striate, with numerous, short to long, pale simple eglandular hairs especially in the upper part, simple or branched above, leafy. Leaves 2–15 × 0.5–2.5(–4.0) cm, dull medium green on upper surface, paler and greyish beneath, narrowly elliptical, oblong-elliptical, lanceolate or ovate, acute at apex, entire to denticulate or serrulate, narrowed or rounded to a semiamplexicaul base, with few to numerous, short to medium, pale simple eglandular hairs on both surfaces and the margins. Capitula 25–40 mm in

50. Inula diameter, solitary or in a few-headed corymb; peduncles with numerous to dense, short to medium, stiff simple eglandular hairs. Involucral bracts in numerous rows, but more or less equal in length or the outer a little longer, 9–11 × 1– 2 mm, pale yellowish-green, linear-lanceolate, gradually narrowed to an acute apex, with dense, short to long, pale simple eglandular hairs over the surface and margin. Flowers of 2 kinds, the outer 15–25 mm, ligulate, the ligules golden yellow and with 3 triangular lobes at apex, the inner 10–15 mm, tubular, brownish-yellow, with 5 linear lobes at apex. Receptacle slightly convex, pitted, the pits with a scarious margin, without scales. Achenes 1.0–1.5 mm, oblong, hairy; pappus 4–5 mm, whitish, of 15–25 rather rigid hairs. Flowers 7–8. 2n = 32. Introduced. Naturalised at Cropstone Reservoir in Leicestershire from 1894 to at least 1932. There is a specimen in the Cambridge herbarium from a wet roadside near Lough Basagh in Galway in 1887 labelled I. salicina which is in fact this species. It is possible that both I. salicina and I. britannica have been introduced on the feet of wildfowl. I. britannica occurs throughout much of Europe and western Asia. It is a very variable species and a detailed study of the plant in Europe might tell us where our plants originated. 6. I. oculus-christi L. Hairy Fleabane I. suaveolens Pall. Perennial herb with a horizontal, creeping rhizome. Stem 15–60 cm, erect, furrowed, densely covered with long, white, appressed hairs, slightly branched above, leafy. Leaves medium green, sericeous-lanate; lower 12– 14 × 1.5–3.0 cm, oblong or oblong-elliptical, obtuse at apex, entire or denticulate, narrowed at base, petiolate; upper 2.0–6.5(–8.0) × 1.0–2.5 cm, lanceolate, sessile, more or less amplexicaul. Capitula 25–40 mm in diameter, solitary or 2–4 in dense corymbs; peduncles with bracts, sericeous-lanate. Involucral bracts in several rows, the outer 5–7 × about 1.1 mm, lanceolate, acute at apex and densely white lanate, the middle ones linear-lanceolate and 7– 8 × about 1.0 mm, the inner 10 × 0.3–0.5 mm and linear, all membranous, dorsally short-hairy in upper part, with golden sessile glands and densely long-ciliate. Flowers of 2 kinds, the outer 13–16 mm, ligulate, the ligules yellow and covered with sparse long hairs and golden glands with 3 lobes at apex, the inner 7 mm, tubular with 5 lobes at apex. Receptacle flat, pitted, without scales. Achenes 2–3 mm, brown, linear-oblong, longitudinally ribbed, covered with reddish, appressed hairs; pappus of numerous bristles. Flowers 6–8. 2n = 32. Introduced. Naturalised in derelict gardens and rough ground in two places in the Isle of Man since the 1960s. Native of eastern Europe and western Asia. 7. I. conyzae (Griess.) Meikle Ploughman’s Spikenard I. conyza DC.; Conyza squarrosa L.; I. squarrosa (L.) Bernh., non L.; Aster conyzae Griess.; Conyza vulgaris Lam. nom. illegit.; I. vulgaris Trevis. nom. illegit. Biennial to perennial herb with an oblique, irregularly thickened, aromatic stock. Stems 30–130 cm, erect, pale yellowish-green, sometimes flushed dark purple or red-

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dish, with numerous to dense, medium to long, pale, wavy simple eglandular hairs and numerous, minute glandular hairs, branched in the upper half, leafy. Leaves medium to rather dark, dull green on upper surface, paler beneath, with numerous, short, pale, stiff, simple eglandular hairs with white, swollen bases on the upper surface and numerous to dense, pale, soft simple eglandular hairs with white, swollen bases on the lower surface, long and dense on the veins and margins, shorter and fewer between the veins; lower 8–15 × 3–5 cm, elliptical, ovate, lanceolate or oblonglanceolate, obtuse-mucronate to more or less acute at apex, irregularly denticulate and gradually narrowed below to a hairy, winged, semiamplexicaul petiole; middle and upper gradually decreasing in size, elliptical, lanceolate or oblong, acute to obtuse at apex, petiolate like the lower or the uppermost sessile. Inflorescence paniculate-corymbose; peduncles with dense, short to medium, wavy, pale simple eglandular hairs with white swollen bases. Capitula numerous, 7–12 mm in diameter. Involucral bracts in many rows; outer 4–6 × 0.8–1.0 mm, green with pale bases, oblonglanceolate, acute at apex, with spreading or recurved tips and with medium, pale, appressed simple eglandular hairs; inner 9–11 × 0.4–0.7 mm, very pale green with narrow red borders in upper half, linear-lanceolate, long-acute at apex, with short, pale, appressed simple eglandular hairs on the surface and ciliate margins. Flowers usually of 2 kinds, the outer 7–9 mm, ligulate, yellowish and 3-lobed at apex, the inner 5–7 mm, tubular, with 5 triangular lobes at apex. Receptacle slightly convex, pitted, the pits with scarious margins. Achenes 2.0–2.5 mm, dark brown, oblong, strongly ribbed with sparse, appressed hairs; pappus 6– 8 mm, whitish, of stiff, denticulate hairs. Flowers 7–9. 2n = 32. Native. Calcicolous plant of dry or rocky slopes and cliffs and open scrub and woodland, and becoming more common as an introduction in rough and grassy places. Locally common through England, Wales and the Channel Islands. Central and south-east Europe northwards to Denmark; Near East; Algeria. A member of the European Temperate element. True Spikenard, or ‘nard’, was an expensive, spicy perfume made from the roots of the Himalayan plant Nardostachys jatamansii DC. Ploughman’s Spikenard was the rustic English equivalent, whose roots have a strong aromatic smell, and were sometimes dried and hung up in cottages as a room-freshener. 8. I. crithmoides L. Golden Samphire I. crithmifolia L. nom illegit. Senecio crithmifolius Scop.; Limbarda tricuspis Cass. nom. illegit.; Eritheis maritima Gray nom. illegit.; Limbarda crithmoides (L.) Dumort.; Limbarda crithmifolia Raf. Perennial herb with a branched, woody stock which is long-lived. Stems in groups 15–60(–100) cm, rather woody below, erect or ascending, pale yellowish-green, very fleshy, usually glabrous, rarely with a few glandular hairs, branched above, leafy. Leaves 2.0–4.5(–6.0) × 0.2–0.4(–0.9) cm, yellowish-green on upper surface, paler beneath, fleshy, linear to narrowly oblanceolate, obtuse at apex, those on the main stem often 3-toothed at apex, otherwise entire, gradually narrowed below to a sessile base, glabrous, the

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upper often with axillary leaf-clusters. Capitula 23–28 mm in diameter, few in a corymb; peduncles glabrous, with numerous leaf-like bracts. Involucral bracts in many rows, glabrous, yellowish-green, the inner with scarious margins, the outer 3–4 × 0.5–1.0 mm, linear and acute at apex, the inner 5–10 × 0.5–1.0 mm, linear-subulate and acute at apex. Flowers of 2 kinds; outer 14–25 mm, ligulate, the ligules golden yellow and 3-lobed at apex; inner 8–11 mm, orange-yellow, with 5 triangular lobes at apex. Receptacle slightly convex, pitted, the pits with scarious margins, without scales. Achenes 2–3 mm, greyish-brown, cylindrical, faintly ribbed, hairy; pappus somewhat reddish, of about 30, stiff hairs. Flowers 7–10. 2n = 18. Native. Salt-marshes, shingle banks, calcareous rocks and ditch-sides by the sea or on Festuca rubra–Armeria maritima turf a little inland. Coasts round the Thames estuary north to Suffolk and to the south coast of Kent, on the Hampshire and Dorset coast and round the Devonshire– Cornwall peninsula, south Wales, north-west Wales and Anglesey, and isolated localities in and near the Mull of Galloway; Channel Islands. In Ireland scattered round the southern coast north to Co. Louth. Atlantic and Mediterranean coasts of west and south Europe northwards to about 55◦ 30 N in Scotland and eastwards to Yugoslavia and Greece; inland in eastern Spain. A member of the Mediterranean-Atlantic element. 51. Dittrichia Greuter Cupularia Gren. & Godr., non Link Woody perennial or annual herbs. Stems ascending to erect, leafy, with viscid glandular hairs. Leaves alternate, entire or denticulate, cauline semiamplexicaul, with viscid glandular hairs. Capitula few to many, in a raciform inflorescence. Involucral bracts in several rows, herbaceous. Inner flowers tubular and bisexual with 5 lobes at apex, the outer ligulate, 3-lobed and female. Corolla yellow. Anthers with branched tails and firm apical appendages. Style slender, with hairs confined to the apical portion of the style branches. Receptacle honeycombed; without scales. Achenes cylindrical; pappus of 1 row of hairs, connate at base. Two species, Mediterranean region, southern Europe and western Asia; introduced in North and South America and Australia. Named after Manfred Dittrich (b. 1934). 1. Ligules 10–12 mm, distinctly exceeding the involucre; 1. viscosa pappus of 15 hairs 1. Ligules 4–7 mm, scarcely exceeding the involucre; 2. graveolens pappus of about 30 hairs

1. D. viscosa (L.) Greuter Woody Fleabane Erigeron viscosum L.; Inula viscosa (L.) Aiton; Pulicaria viscosa (L.) W. D. J. Koch; Cupularia viscosa (L.) Gren. & Godr.; Solidago viscosa (L.) Lam. Perennial resinous-scented herb with a stout root. Stems 40–130 cm, pale green, often suffused with brownishpurple, erect, usually robust and woody at base, rarely more slender, striate, densely viscid glandular-hairy and with longer simple eglandular hairs, branched in upper part, densely leafy. Leaves becoming gradually smaller upwards, sometimes bundles of small leaves in the axils of larger ones, 1–70 × 0.5–30.0 cm, medium green on upper sur-

face, paler beneath, oblong-lanceolate, linear-lanceolate or linear, obtuse to acute at apex, subentire to denticulate, narrowed at base, the lower sometimes shortly petiolate the upper sessile and semiamplexicaul, covered with dense, short, viscid glandular hairs. Inflorescence a long, raceform panicle; peduncles slender, with numerous, short, viscid glandular hairs and longer, simple eglandular ones. Capitula 15–25 mm in diameter. Involucral bracts in several rows, pale green in centre with broad, pale margins, linearlanceolate, acute at apex, with numerous, minute glandular hairs and very short, appressed simple eglandular hairs, the outer 1–2 × 0.5–0.7 mm, the inner 6–8 × 0.6–0.8 mm. Flowers of 2 kinds, the outer ligulate, the ligules 10–12 mm, distinctly exceeding the involucre, yellow, minutely 3-lobed at apex, the inner tubular, yellow with 5 lobes at apex. Receptacle honey-combed, without scales. Achenes 1.8–2.0 mm, pale brown, cylindrical, abruptly contracted at apex, hairy; pappus pale brown, of 15 stiff hairs, connate at their bases. Flowers 7–8. 2n = 18, 34. Introduced. Naturalised on waste ground at Languard Common, near Felixstowe in Suffolk and by a harbour in Sussex; elsewhere a rare casual in southern England. Native of southern Europe and the Mediterranean region. 2. D. graveolens (L.) Greuter Stinking Fleabane Erigeron graveolens L.; Inula graveolens (L.) Desf.; Pulicaria graveolens (L.) Nyman; Cupularia graveolens (L.) Gren. & Godr.; Solidago graveolens (L.) Lam. Annual herb smelling of camphor, with a slender tap-root and fibrous side-roots. Stems 20–50 cm, pale green, sometimes suffused brownish-purple, erect, rather slender but rigid, striate, with dense, very short, viscid glandular hairs and medium to long simple eglandular hairs, usually with many, slender, erecto-patent branches, very leafy. Leaves often of similar size throughout the length of the plant and with small ones in the axils of larger ones, 1.0–7.5 × 0.2–1.3 cm, dull medium green on upper surface, paler beneath, narrowly oblong-oblanceolate, oblong or linear, obtuse to acute at apex, entire or remotely denticulate, narrowed at base, the lower with a short, winged petiole the upper sessile and semiamplexicaul, clothed with very short glandular hairs. Inflorescence often taking up nearly the whole stem, with the longest branches from near the middle; peduncles slender, with dense, short glandular hairs and medium simple eglandular hairs. Capitula 8–14 mm in diameter. Involucral bracts in several rows, with minute glandular hairs, the outer 2.8–3.0 × 0.5–1.0 mm, green, linear-lanceolate and acute at apex, the inner 4–7 × 0.7–1.0 mm, green in the centre with a broad pale margin or entirely pale, linear-lanceolate, acute at apex. Flowers of 2 kinds, the outer ligulate, the ligules 4–7 mm, not or scarcely exceeding the involucre, yellow, minutely 3-lobed at apex, the inner tubular and yellow, 5lobed at apex. Receptacle honeycombed, without scales. Achenes 1.8–2.0 mm, pale brown, cylindrical, abruptly contracted at apex, hairy; pappus pale brown of about 30 stiff hairs, connate at their base. Flowers 7–8. 2n = 18, 20 + 1–2 B. Introduced. Frequent wool alien in fields and waste places. Scattered records in England. Native of south and west Europe, Mediterranean region and western Asia.

53. Buphthalmum 52. Pulicaria Gaertn. Annual or perennial herbs. Stems erect, leafy, branched. Leaves alternate, entire to denticulate, more or less amplexicaul. Capitula numerous, in a loose corymb. Involucral bracts in several rows, herbaceous. Inner flowers tubular, 5lobed and bisexual, the outer ligulate, 3-lobed and female. Corolla yellow. Anthers with branched tails and firm apical appendages. Styles with linear branches with hairs throughout their length. Receptacle flat or slightly convex, without scales. Achenes cylindrical; pappus in 2 rows the inner of hairs, the outer of free or fused scales. About 80 species in Europe, North Africa and Asia. Chatters, C. (1991). The status of Pulicaria vulgaris Gaertner in Britain in 1990. Watsonia 18: 405–406. Fitzgerald, R., Field, G. D. & Chatters, C. (1997). The status of Pulicaris vulgaris Gaertner in Britain in 1995. Watsonia 21: 279–280. Hare, A. D. R. (1990). Lesser Fleabane – a plant of seasonal hollows. British Wildlife 2: 77–79. Prince, S. D. & Hare, A. D. R. (1981). Lactuca saligna and Pulicaria vulgaris in Britain in Synge, H. (Edit.) The biological aspects of rare plant conservation. pp. 379–388. Chichester. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [P. vulgaris].

1. Perennial herb with stolons; cauline leaves cordate at base; capitula 15–30 mm in diameter; ligules usually 1. dysenterica much longer than involucral bracts 1. Annual herb with tap-root and fibrous side-roots; cauline leaves not cordate at base; capitula 6–12 mm in diameter; 2. vulgaris ligules about as long as involucral bracts

1. P. dysenterica (L.) Bernh. Common Fleabane Inula dysenterica L. Perennial herb with stolons. Stems 20–60 cm, yellowishgreen, sometimes suffused purplish, ridged, erect, more or less tomentose, branched above, leafy. Leaves 2–8 × 0.5– 2.5 cm, yellowish-green on upper surface, greyish-green beneath, oblong, oblong-lanceolate, lanceolate or elliptical, obtuse to more or less acute at apex, entire or undulatedenticulate, cordate at base and sessile and amplexicaul, with very short, stiff, tuberculate-based, appressed simple eglandular hairs on the upper surface and more or less tomentose beneath. Inflorescence a loose corymb; peduncles with dense woolly and shorter glandular hairs. Capitula 15–30 mm in diameter, hemispherical. Involucral bracts in several rows, 6.5–7.5 × 0.1–0.3 mm, pale green, narrowly linear, long drawn out to an acute apex, with long, pale simple eglandular and glandular hairs. Flowers of 2 kinds, the outer variable in length, ligulate, the ligules much longer than involucral bracts, 3-toothed at apex, yellow, the inner tubular, 5-toothed at apex and yellow, rarely all tubular. Receptacle flat or slightly convex, without scales. Achenes 1.0–1.5 mm, cylindrical, shortly hairy; pappus of an outer row of scales forming a small, denticulate cup, and of an inner row of long simple hairs. Flowers 8–9. Visited by many insects, chiefly flies. 2n = 18. Variable in the length of the ligules and in the amount of indumentum. Var. longiradiata Druce has very long ligules, var. discoidea Druce (subvar. flosculosa Corb.) none at all.

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There is every sort of intermediate between these two, sometimes in one colony. Native. Marshes, ditches, hedgebanks, wet and waste places. Common in lowland Great Britain north to northern England, rare in southern Scotland and northern Ireland. Europe northwards to Denmark and central Russia; North Africa, western Asia. A member of the Eurosiberian Southern-temperate element. 2. P. vulgaris Gaern. Small Fleabane Inula pulicaria L.; Inula uliginosa Sibth.; Inula cylindrica With. nom. illegit.; Inula pulicaris Stokes; P. uliginosa (Sibth.) Gray, non Hoffmans. & Link; P. prostrata Aschers. nom. illegit.; P. pulicaria (L.) Karst. nom. illegit. Annual herb with tap-root and fibrous side-roots. Stems 8–45 cm, pale yellowish-green, often flushed brownishpurple, erect, striate, with numerous, very short glandular hairs and longer curled simple eglandular ones, much branched, often to the base, leafy. Leaves 1–6 × 0.2–2.0 cm, dull medium green on upper surface, paler beneath, oblanceolate, lanceolate or elliptical, obtuse to acute at apex, entire or sinuate-denticulate, the upper rounded and semiamplexicaul at base, the lower narrowed into a winged petiole, more or less glabrous or shortly glandular-hairy, with some longer eglandular ones especially beneath. Inflorescence a loose corymb; peduncles with numerous long pale simple eglandular hairs and short glandular ones. Capitula 6– 12 mm in diameter. Involucral bracts in several rows, 4– 5 × 0.2–0.5 mm, pale green with purplish-red tips, linear, long drawn out to an acute apex, with numerous to dense, long, pale simple eglandular hairs and shorter, pale glandular ones. Flowers of 2 kinds, the outer in 1 row, shortly ligulate, the ligules about as long as the involucral bracts, 3-lobed at apex, yellow, the inner tubular, pale yellow, 5lobed at apex. Receptacle flat or slightly convex, without scales. Achenes 1.2–1.5 mm, cylindrical, shortly hairy; pappus of an outer row of scales forming a deeply laciniate cup, and an inner row of long simple hairs. Flowers 8–9. 2n = 18. Native. Moist sandy places, where water stands in the winter, but not during the growing season, such as pond margins, shallow ditches and muddy track ruts. Formerly widespread in the Channel Islands and England north to Leicestershire; now very local in Surrey, Hampshire and Wiltshire particularly in the New Forest. Europe north to Denmark and southern Sweden; Caucasus; western Asia; North and East Africa. A member of the Eurosiberian Temperate element. 53. Buphthalmum L. Perennial herbs. Stems simple or branched, erect. Leaves alternate, simple, entire or denticulate. Capitula solitary on long, leafy peduncles. Involucral bracts in several rows. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex, in 1 row and female. Corolla yellow. Anthers with short tails and firm apical appendages. Style with linear branches which have hairs throughout their length. Receptacle convex, with numerous scales which are folded round the achenes. Achenes of ligulate flowers 3-angled or more or less flattened, those of inner

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flowers with several angles; pappus a scarious, denticulate rim. Two species in southern and central Europe. 1. B. salicifolium L. Willow-leaved Yellow Ox-eye Perennial herb. Stem 15–70(–150) cm, medium green often suffused brownish-purple, erect, slender to robust, ridged, with numerous, short to medium, pale simple eglandular hairs, leafy, branched. Leaves 5–10 × 0.5–2.5 cm, dull, medium green, slightly paler beneath, lanceolate, oblanceolate, linear-lanceolate or linear, acute at apex, entire or undulate-denticulate, the lower shortly petiolate, the upper sessile and semiamplexicaul, with short to medium, pale, appressed simple eglandular hairs on both surfaces. Capitula solitary, 30–40 mm in diameter; peduncles long, hairy and leafy. Involucral bracts in several rows, 5–7 × 1.8– 2.0 mm, greyish-green, lanceolate, with a long-acute, sharp apex, with dense, short and medium, pale simple eglandular hairs. Flowers of 2 kinds; outer 7–20 mm, ligulate, the ligules yellow, with 3 short lobes at apex; inner 7–10 mm, tubular, brownish-yellow, with 5 lobes at apex. Receptacle convex, with numerous scales which are folded round the achenes. Achenes 3–4 mm, those of the ligulate flowers 3angled or more or less flattened, those of the inner flowers with several angles, glabrous; pappus a scarious, denticulate rim. Flowers 6–8. 2n = 20. Introduced. A persistent garden escape at the edge of Wandsworth Common in Surrey. Native of central Europe extending to south-east France, north Italy and central Yugoslavia. 54. Telekia Baumg. Molpadia (Cass.) Cass.; Buphthalmum subgen. Molpadia Cass. Perennial herbs. Stems erect, leafy. Leaves alternate, ovate, cordate, crenate-serrate, the lower petiolate. Capitula solitary to several, terminal and subcorymbose. Involucral bracts in several rows, herbaceous. Inner flowers tubular 5-lobed at apex and bisexual, the outer ligulate, 3lobed at apex and female. Corolla yellow. Anthers with branched tails and firm apical appendages. Style with linear branches which have hairs throughout their length. Receptacle convex, with scales. Achenes of both tubular and ligulate flowers similar, terete or slightly compressed, with several angles; pappus of fused scales. Two species in Europe and south-west Asia. Named after Samuel Teleki de Szek. 1. T. speciosa (Schreb.) Baumg. Yellow Ox-eye Buphthalmum speciosum Schreb.; Buphthalmum cordifolium Waldst. & Kit.; Inula caucasica Pers.; Molpadia suaveolens Cass.; Corvisartia caucasica (Pers.) G. Don ex Loudon.; T. cordifolia (Waldst. & Kit.) DC.; T. ovata K. Koch Perennial herb with a thick, oblique, knotty rhizome. Stem 50–200 cm, pale yellowish-green, often suffused brownish-purple, sulcate-striate, densely and finely, yellowish glandular-hairy, laxly branched above, leafy. Leaves yellowish-green on upper surface, paler beneath and sometimes with purplish colouring in the veins, nearly glabrous

or with short, pale simple eglandular hairs on the upper surface, with a mixture of short glandular and long simple eglandular hairs beneath, on the margin and on the petiole; basal 19–25 × 13–17 cm, usually broadly ovate, rarely subrotund, acute at apex, biserrate, cordate at base, on petioles 11–20 cm; middle cauline 15–20 × 8–15 cm, broadly ovate, acute at apex, biserrate, cordate at base on petioles 1.5–5.0 cm; upper cauline 10–20 × 6–12 cm, ovate, ovate-oblong or elliptic-lanceolate, acute at apex, biserrate, more or less sessile and semiamplexicaul at base. Capitula solitary or few in a corymb, 50–80 mm in diameter, hemispherical; peduncles 5–25 cm, leafy, with glandular and simple eglandular hairs. Involucral bracts in several rows, varying in size, 10–35 × 2–15 mm, the outer ones green, oblong, acute at apex and smooth-edged or with inconspicuous teeth, the median, lanceolate and less acute, brownish in lower half, green at apex, the innermost spathulate and subscarious, all glandular-punctate on both surfaces. Flowers of 2 kinds; outer 30–45 mm, ligulate, the ligules golden yellow, 3-lobed at apex, glandular on outer face; inner 5–7 mm, tubular, 5lobed at apex and glandular. Receptacle convex, with linear, denticulate, acute scales. Achenes 3.0–3.5 mm, terete or slightly compressed, ribbed; pappus of fused scales forming an undulate or lacerate corona. Flowers 7–8. 2n = 20. Introduced. Grown in gardens and naturalised in rough ground, by roadsides and by lakes and rivers. Scattered localities through most of Great Britain, but particularly in the north. Native of central Europe, the Balkans, central and south Russia and Caucasia. Tribe 5. Astereae Cass. Tribe Baccharideae Kostel.; Tribe Bellideae Cass. ex D. Don; Bellieae DC. ex Godr.; Tribe Erigeroneae Gren. & Godr. Usually perennial, rarely annual herbs, occasionally shrubs. Leaves alternate or all basal, simple. Capitula with flowers all tubular or tubular and ligulate. Flowers of various colours. Involucral bracts in 2–several rows, usually herbaceous. Receptacular scales absent. Pappus usually of hairs, sometimes absent or of strong bristles. Contains three subtribes, 174 genera and about 2,800 species, worldwide but especially in south-western North America, along the Andes, in southern Africa, Australia, New Zealand and central Asia. 55. Grindelia Willd. Perennial herbs. Stems erect, leafy. Leaves alternate, serrate, the cauline amplexicaul. Capitula solitary at the ends of branches forming an irregular corymb, very resinous in fruit. Involucral bracts in several rows, herbaceous, sticky, with recurved tips. Outer flowers ligulate, minutely 3-lobed, female, the inner tubular, 5-lobed at apex, bisexual or sterile. Corolla yellow. Anthers without tails, apical appendage ovate-lanceolate, flat. Style with short branches, sterile appendages ventrally glabrous, dorsally hairy. Receptacle flat or convex, without scales. Achenes compressed, subquadrangular, glabrous; pappus of 2–8 bristles. About 55 species occurring in North and South America. Named after David Hieronymus Grindel (1766–1836).

56. Calotis 1. G. stricta DC. Coastal Gum-plant G. oregana A. Gray; G. hendersonii Greene; G. lanata Greene; G. macrophylla Greene; G. andersonii Piper Perennial herb. Stems 20–75(–150) cm, pale green to deep purplish-red, ascending to erect, glabrous to more or less villous or tomentose, branched only at top, leafy, sometimes woody at base. Leaves more or less fleshy, medium green on upper surface, paler beneath, sometimes red-veined; basal 10–25 × 1–4 cm, oblanceolate, acute to rounded at apex, entire or serrulate near the apex, narrowed at base to a winged petiole; cauline 3–12 cm, broadly oblong to oblanceolate, obtuse to acute at apex, entire to slightly serrulate, amplexicaul at base; all resinous-punctulate. Capitula 1–numerous, 40–50 mm in diameter, in an irregular corymb. Involucral bracts in several rows, 8–14 × 2–3 mm, linear-lanceolate, the slender, tapering green tip erect or gradually recurving, erect or the outer ones spreading, resinous-punctulate. Flowers of 2 kinds, the outer 12–25 mm, ligulate, the ligules yellow, minutely 3-lobed at apex, the inner tubular and yellow, 5-lobed at apex. Receptacle flat or convex, pitted, without scales. Achenes 3.5– 7.0 mm, grey to reddish-brown, compressed, ridged and subquadrangular, with a knobbly apex; pappus of 2–6, minutely serrate, bristles. Flowers 9–10. 2n = 24. Introduced. Naturalised on sea-cliffs at Whitby in Yorkshire since 1977. Native of the west coast of North America. 56. Calotis R. Br. Cheiroloma F. Muell. Annual or perennial herbs. Stems erect or procumbent, leafy. Leaves alternate, oblanceolate or obovate, entire, dentate or pinnatifid. Capitula solitary on long peduncles or in dense cymes. Involucral bracts in more or less 2 or several rows, herbaceous. Flowers of 2 kinds, the outer ligulate, 3lobed at apex and female, the inner tubular, 5-lobed at apex, usually male, rarely bisexual. Anthers without tails, apical appendage ovate-lanceolate and flat. Style with oblong branches, sterile appendages ventrally glabrous, dorsally hairy. Receptacle conical, without scales. Achenes oblanceolate or obpyramidal; pappus of rigid awns, or of alternate awns and scales. Twenty-six species, mostly in Australia, but a few in south-east Asia. C. dentex R. Br. has been recorded as a wool casual. Davis, G. L. (1952). Revision of the genus Calotis R. Br. Proc. Linn. Soc. New South Wales 77: 146–188. 2. hispidula 1. Achenes densely hairy 2. 1. Achenes glabrous or nearly so 2. Pappus represented by rigid awns alternating with an 1. cuneifolia equal number of scales 2. Pappus represented by awns only, scales absent 3. lappulacea

1. C. cuneifolia R. Br. Purple Bur-daisy Perennial herb. Stems 10–60 cm, green often suffused brownish-purple, prostrate to erect, becoming woody at base, with few to numerous simple eglandular hairs, sparsely to much branched, leafy. Leaves 1.5–4.0 × 0.6–2.0

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cm, medium green on upper surface, paler beneath, obovate, 3- to 11-toothed at apex, cuneate at base, scabrous or very shortly hairy; petiole short, minutely hairy, abruptly expanded and amplexicaul at base. Capitula 10–20 mm in diameter, solitary on long peduncles. Involucral bracts in several rows, 2.0–3.5 × 1–2 mm, green, lanceolate, acute at apex, glandular-hairy. Flowers of 2 kinds, the outer 3– 6 mm, ligulate, the ligules white to mauve and 3-lobed at apex, the inner tubular, 5-lobed at apex and yellow. Receptacle pitted, conical, without scales. Achenes of ligulate flowers 0.8–1.8 mm, oblanceolate, flattened, glabrous, with narrow marginal ridges, but wingless, forming a bur-like head; pappus of 2–4 straight, rigid awns barbed distally and 2–4 smooth, inflexed scales broader than long; achenes of tubular flowers sterile. Flowers 7–9. 2n = 16, 20. Introduced. Wool alien in cultivated and waste ground. Scattered records in England. Native of Australia. 2. C. hispidula (F. Muell.) F. Muell. Hairy Bur-daisy Cheiroloma hispidulum F. Muell. Annual herb. Stems 10–30 cm, green, sometimes flushed brownish-purple, prostrate to ascending, rigid-hairy, branched, leafy. Leaves 0.5–2.5 × 0.2–0.8 mm, medium green on upper surface, paler beneath, oblanceolate, entire or 3- to 5-toothed at apex, cuneate at base, more or less rigid-hairy, sessile. Capitula 8–10 mm in diameter, in dense, leafy cymes. Involucral bracts in several rows, 2–4 × 1.0– 1.5 mm, greenish, lanceolate to spathulate, acute at apex, rigid-hairy. Flowers of 2 kinds, the outer ligulate, the ligules 0.8–1.2 mm, yellow and 3-lobed at apex, the inner tubular, 5-lobed at apex and yellow. Receptacle conical, with acute, ciliate scales. Achenes of both ligulate and tubular flowers 1.5–2.5 mm, oblanceolate, with narrow, marginal ridges but wingless and shortly but densely hairy; pappus of 5 or 6, straight, rigid awns hairy at the base and barbed distally, and 5–6, shorter, divided, hairy scales. Flowers 7–9. Introduced. Wool alien in cultivated and waste land. Scattered records in England. Native of Australia. 3. C. lappulacea Benth. Yellow Bur-daisy Glossogyne hennedyi R. Br.; C. microphylla Benth.; C. suffruticosa Domin; C. glabrescens C. T. White Perennial herb. Stems 10–40 cm, green to brownish-purple, ridged, woody towards the base, with few to numerous simple eglandular hairs becoming glabrous towards the base. Leaves 3–12(–20) × 0.5–2(–3) mm, medium green on upper surface, paler beneath; basal oblanceolate, usually pinnatifid, long-cuneate at base; cauline becoming gradually smaller upwards, linear-oblong to narrowly triangular or spathulate, acute with a cartilaginous apex, entire to shallowly pinnatifid with (1–)2–4 lobes, subtruncate and sessile; all with sparse simple eglandular hairs. Capitula 5–8 mm in diameter, solitary on long peduncles. Involucral bracts in 2–3(–4) rows, 2.0–2.5 × 0.5–1.0 mm, oblong to triangular, obtuse at apex, with sparse simple eglandular hairs. Flowers of 2 kinds, the outer 2.5–3.5 mm, ligulate, in 1–several rows, the ligules yellow and 3-lobed at apex the inner tubular, 5-lobed at apex and yellow. Receptacle hemispherical, without scales. Achenes 1.2–1.6 mm, obpyramidal, ribbed on 2 angles, glabrous, tuberculate on the faces, forming

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a globose, bur-like head 7–10 mm in diameter; pappus of 2(–3) long, more or less erect, barbed awns 5 with 2 groups of 1–6, short, horizontal, barbed awns between. Flowers 7–9. Introduced. Wool casual. A few scattered records in England. Native of Australia; introduced in New Zealand. Amellus microglossus DC. and A. strigosus (Thunb.) Less. have been recorded as wool casuals and Bellium bellidioides L. as a casual garden escape. 57. Solidago L. Perennial herbs with a rhizome or short stock. Stems simple, leafy; non-flowering leaf rosettes often present. Leaves alternate, serrate. Capitula usually numerous, small, fasciculate, thyrsoid or sometimes corymbose panicles. Involucral bracts in several rows, herbaceous. Inner flowers tubular, 5-lobed, bisexual, the outer ligulate, minutely 3lobed, female. Corolla yellow. Anthers without tails, apical appendage ovate-lanceolate, flat. Style with oblong-linear branches, sterile appendages, ventrally glabrous, dorsally hairy. Receptacle flat, without scales. Achenes oblong subterete or angled; pappus of 1–2 rows of more or less equal simple hairs. About 150 species, mainly in North America with a few species in Eurasia and South America. Hybrids occur in nature, often between dissimilar species, and have been artificially produced in cultivation. Burton, R. M. (1980). Solidago × niederederi Khek in Britain. Watsonia 13: 123–124. Druce, G. C. (1922). Solidago cambrica Huds. Rep. Bot. Soc. Exch. Cl. Brit. Isles 6: 283–290. Druce, G. C. (1928). Solidago virgaurea L. Rep. Bot. Soc. Exch. Cl. Brit. Isles 8: 307. Grime, J. P. et al. (1988). Comparative plant ecology. London. [S. virgaurea.] Morton, J. K. (1977). A cytological study of the Compositae (excluding Hieracium and Taraxacum) of the British Isles. Watsonia 11: 211–223. Turesson, G. (1925). The plant species in relation to habitat and climate. Hereditas 6: 147–236. Weber, E. (1997). Morphological variation of the introduced perennial Solidago canadensis L. sensu lato (Asteraceae) in Europe. Bot. Jour. Linn. Soc. 123: 197–210. 1. Leaves gland-spotted; capitula sessile, in small clusters forming a corymbose inflorescence; ligulate flowers more numerous than tubular flowers; receptacle pits fimbriate 6. graminifolia 1. Leaves not gland-spotted; at least most capitula pedunculate, forming a pyramidal to more or less cylindrical inflorescence; ligulate flowers fewer than 2. tubular flowers; receptacle pits not fimbriate 2. Leaves with 1(–2) pairs of main lateral veins from near the base, running parallel with the midrib for most of 3. their length 2. Leaves with many pairs of short often inconspicuous 6. lateral veins 3. Stems crisped-hairy at least in the top half; leaves usually 4. scabrid or shortly hairy on both surfaces 3. Stems more or less glabrous; leaves glabrous on both 5. surfaces or sparsely hairy beneath

4. Stem usually glabrous below; leaves sparsely scabrid; involucral bracts 2–3 mm; tubular flowers 2.4–2.8 mm 4(a). canadensis subsp. canadensis 4. Stem usually hairy throughout; leaves more scabrid; involucral bracts 2.5–4.0 mm; tubular flowers 3–4 mm 4(b). canadensis subsp. altissima 5. Leaves hairy on the veins beneath; involucral bracts 3.2–4.0 mm; achenes usually glabrous 5(a). gigantea subsp. gigantea 5. Leaves glabrous on lower surface; involucral bracts 5(b). gigantea subsp. serotina 3.5–5.0 mm; achenes hairy 6. Capitula in long, out-curving racemes at the end of erecto-patent or patent branches, the whole forming a 3. rugosa pyramidal inflorescence 6. Capitula in a simple racemose panicle or a dense 7. thyrsoid-panicle with loose branches 2. calcicola 7. Achenes 1–2 mm 8. 7. Achenes 2.8–3.2 mm 8. Stem 3–150 cm; inflorescence usually more or less open; involucral bracts 2.5–8.0 mm 1(a). virgaurea subsp. virgaurea 8. Stem 5–20 cm; inflorescence a compact, spike-like raceme; involucral bracts 6–10 mm 1(b). virgaurea subsp. minuta

Subgenus Solidago Capitula all pedunculate. Ligulate flowers fewer than tubular flowers. Receptacle pits not fimbriate. 1. S. virgaurea L. Goldenrod Perennial herb with a stout, obliquely ascending stock and rather fleshy, fibrous roots. Stems 5–150 cm, pale yellowishgreen, often suffused brownish-purple, erect, with numerous to dense, whitish, short, curly simple eglandular hairs and minute glandular hairs, simple or branched, leafy. Leaves medium yellowish-green on upper surface, paler beneath; basal 2–10 × 2–4 cm, ovate to lanceolate, elliptical or linear-lanceolate, obtuse to acute at apex, subentire to shallowly crenate-serrate, narrowed at base to a winged petiole; cauline gradually decreasing in size upwards, lower often similar to basal, the upper lanceolate or elliptical, more or less acute at apex, entire to shallowly crenate-serrate, attenuate or cuneate at base, sessile; all with many pairs of lateral veins, glabrous on upper surface and with numerous, short, curved hairs beneath and on the margins, particularly the veins and sometimes also on the upper surface. Inflorescence from a simple, racemose panicle to a broad, muchbranched, leafy one; peduncles short, with short, often curved simple eglandular hairs. Capitula numerous, 6–15 mm in diameter, narrowed at base. Involucral bracts in several rows, 2.5–10.0 × 0.8–1.2 mm, pale green, with paler margins, linear-lanceolate, lanceolate or narrowly ovate, obtuse to acute at apex, with very short, stiff simple eglandular hairs, especially down the centre line and on the margins. Flowers of 2 kinds, the outer 6–12, 4–9 mm, ligulate, the ligules yellow and spreading, minutely 3-lobed, the inner 10–30, tubular, yellow and 5-lobed. Receptacle flat, without scales. Achenes 2.8–3.2 mm, brown, oblong, shortly hairy; pappus 4–5 mm, whitish, of 1–2 rows of simple hairs.

57. Solidago Flowers 6–9. Much visited by various bees and flies, and automatically self-pollinated. 2n = 18. Very variable in stature, hairiness, size, shape and serration of leaves, branching of inflorescence and size of capitula. The two following subspecies are very variable in themselves and several genetically stable ecotypes in each need to be recognised, when more data becomes available (cf. Morton, 1977). (a) Subsp. virgaurea S. laltifolia Mill.; S. vulgaris Mill. nom. illegit.; S. angustifolia Mill.; S. minor Mill.; S. virgaurea var. latifolia (Mill.) Sm.; S. virgaurea var. vulgaris Mill. nom. illegit.; S. virgaurea var. angustifolia (Mill.) Sm.; S. virgaurea var. minor (Mill.) Sm.; S. virgaurea var. interrupta Druce; S. virgaurea var. dentatifolia Druce. Stems 3–150 cm, often much branched. Basal leaves more or less acute at apex. Inflorescence usually more or less open with numerous flowers. Involucral bracts 2.5–8.0 mm. Flowers 6–7. 2n = 18. (b) Subsp. minuta (L.) Arcangeli S. minuta L.; S. virgaurea var. arctica Hartm.; S. lapponica With.; S. virgaurea var. alpestris Rchb.; S. virgaurea var. minuta (L.) Simmons; S. virgaurea var. pumila (Willd.) Gaudin; S. minuta Mill., non L.; S. virgaurea var. plukenetiana Druce; S. virgaurea var. corymbosa Druce; S. virgaurea var. lanceolata Druce; S. cambrica Huds.; S. virgaurea var. cambrica (Huds.) Sm.; S. jailarum Juz. Stems 5–20 cm, not branched. Basal leaves rounded-obtuse at apex. Inflorescence a compact, few-flowered, spike-like raceme. Involucral bracts 6–10 mm. Flowers 8–9. 2n = 18. Native. Dry woods and grassland, rocks, cliffs, hedgebanks and dunes on acid and some calcareous strata. Frequent over most of Great Britain and Ireland, except in parts of central England, central Ireland and the Channel Islands. Europe; Asia. A member of the Eurasian Boreotemperate element. Subsp. virgaurea is the plant of woodlands and heaths of the lowlands throughout the range of the species. Subsp. minuta is the plant of open, exposed habitats of our mountains, in places coming down to coastal cliffs. It is widespread in the mountains of north, central and east Europe, and in the Arctic. 2. S. calcicola (Fernald) Fernald Lime Goldenrod S. virgaurea var. calcicola Fernald Perennial herb with a stout stock and fibrous roots. Stems often clustered, 60–130 cm, often purplish, erect, glabrous below, with simple eglandular and glandular hairs above, simple or with a few, erect branches, very leafy throughout. Leaves yellowish-green on upper surface, paler beneath; basal and lower cauline 10–15 × 3–5 cm, broadly oblanceolate, acuminate at apex, sharply and irregularly serrate above the middle, entire below, narrowed to a conspicuous, winged petiole; upper cauline 5–10 cm, narrowly ovate to elliptic-lanceolate, tapering about equally to the acuminate apex and base, coarsely and sharply serrate, sessile. Inflorescence large, dense and thyrsoid-paniculate with loose, raceform branches; peduncles short, with curved simple eglandular hairs. Capitula 8–15 mm in diameter, narrowed at

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base, forming a simple racemose panicle. Involucral bracts in several rows, 5–7 × 0.8–1.2 mm, green with paler margins, ovate-lanceolate to oblong or linear, obtuse to subacute at apex. Flowers of 2 kinds, the outer 12–15, ligulate, the ligules yellow, the inner tubular and yellow. Receptacle flat, without scales. Achenes 1–2 mm, oblong, shortly hairy; pappus 4–5 mm, whitish, of 1–2 rows of simple hairs. Flowers 7–9. 2n = 18. Introduced. Garden escape established in woodland at Kilmel Ford in Argyllshire. Native of eastern North America. This is the nearest North American species to our European S. virgaurea and may be better placed as a subspecies of it. 3. S. rugosa Mill. Rough-stemmed Goldenrod S. altissima Aiton, non L. Perennial herb with rhizomes. Stems solitary or numerous, 30–250 cm, pale yellowish-green, often suffused brownishpurple, erect, striate, with numerous, medium to long, pale simple eglandular hairs above, fewer or glabrous below, branched, leafy. Leaves numerous, 3.5–13.0 × 1.3–4.0 cm, medium green on upper surface, paler beneath, oblanceolate, oblong-oblanceolate, elliptical or lanceolate, acute or acuminate at apex, serrate, narrowed to a sessile base, rugose and roughly and shortly hairy especially on the veins, with many pairs of lateral veins. Capitula very numerous, 3–4 mm in diameter, in long, out-curving racemes at the end of erecto-patent or patent branches, the whole forming a pyramidal inflorescence; peduncles short, green, shortly hairy. Involucral bracts in 2–3 rows, 2–5 × 0.4–0.6 mm, yellowish-green in centre with broad, pale margins, oblongelliptical or oblong-lanceolate, obtuse at apex, glabrous or nearly so. Flowers of 2 kinds, the outer 6–10, 1.5–4.0 mm, ligulate, the ligules deep yellow, minutely 3-lobed, the inner 3–8, 2.5–3.5 mm, tubular, yellow, 5-lobed. Receptacle flat, pitted, without scales. Achenes 1.0–1.2 mm, pale brown, cylindrical, hairy; pappus 2–3 mm, dirty white, of very shortly denticulate hairs. Flowers 8–10. 2n = 18, 54. Very variable in leaf size and shape, hairiness and size and shape of inflorescence. Introduced. A garden escape naturalised near Bowling Harbour and in Glen Fruin, both in Dunbartonshire, and formerly in Argyllshire. Native of North America. 4. S. canadensis L. Canadian Goldenrod Aster canadensis (L.) Kunze; Doria canadensis (L.) Lunell Perennial herb with persistent rhizomes, clonal growth leading to dense stands. Stems solitary or numerous, 30– 250 cm, pale yellowish-green, prominently ridged and channelled, erect, with numerous, short, crisped simple eglandular hairs throughout or only in the upper part, densely leafy, branched in upper half and often with sterile shoots in the axils of the lower leaves. Leaves very numerous, crowded, all cauline the basal deciduous early, scarcely decreasing in size upwards, 4–15 × 0.5–3.7 cm, dull, medium yellowish-green on upper surface, paler beneath, lanceolate, oblong-lanceolate, or narrowly oblong-elliptical, gradually narrowed to an acute apex, obscurely to obviously serrate, with small sharp teeth, attenuate at base, sessile

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and semiamplexicaul, minutely scabridulous on the margins and veins beneath and sometimes throughout; with 2 pairs of main lateral veins running parallel to the midrib. Capitula very numerous, 3.0–3.5 mm in diameter, in long, outcurving racemes at the ends of branches, the whole forming a broadly pyramidal inflorescence; peduncles short, pale green, with short, crisp hairs. Involucral bracts in 2–3 rows, 2.0–4.5 × 0.2–0.6 mm, pale greenish-yellow, linearlanceolate, obtuse to acute at apex, glabrous or nearly so. Flowers strong-smelling, of 2 kinds, the outer 9–17, ligulate, the ligules deep yellow, 1.0–1.5 mm, and minutely 3-lobed at apex, the inner tubular, 2.4–4.0 mm, deep yellow, 5-lobed at apex. Receptacle flat, pitted, without scales. Achenes 0.9–1.2 mm, pale brown, subterete, many veined, very shortly hairy; pappus 2.0–2.5 mm, dirty white, of very shortly denticulate hairs. Flowers 8–10. Much visited by small insects.

numerous, 2–3 mm in diameter, in racemose panicles forming scorpioid cymes, the whole inflorescence broadly pyramidal; peduncles short, pale green, with dense simple eglandular hairs, the branches with longer, dense, curly hairs. Involucral bracts in 3–5 rows, 3.2–5.0 × 1.0–1.2 mm, pale yellowish-green, linear-lanceolate, acute at apex, glabrous. Flowers of 2 kinds, the outer 10–17 ligulate, the ligules yellow, clearly exceeding the involucre and tubular flowers and often curved, minutely 3-lobed, the inner (4–)5–10 (–13), tubular, yellow, 5-lobed. Receptacle flat, pitted, without scales. Achenes 0.9–1.2 mm, brown, cylindrical, ribbed, glabrous or hairy; pappus 2.0–2.5 mm, whitish, of minutely denticulate hairs. Flowers 7–10. Visited by flies. 2n = 18, 36, 54.

(a) Subsp. canadensis Stem up to 150 cm, usually glabrous below. Leaves sparsely scabrid. Involucral bracts 2.0–3.0 mm. Ligules 1.5–2.8 mm. Tubular flowers 2.4–2.8 mm. 2n = 18.

(b) Subsp. serotina (Kuntze) McNeill S. serotina Aiton, non Retz.; Aster latissimifolius var. serotinus Kuntze; S. gigantea var. serotina (Kuntze) Cronquist; S. gigantea var. leiophylla Fernald Leaves glabrous on lower surface. Involucral bracts 3.5–5.0 mm. Achenes hairy. 2n = 36.

(b) Subsp. altissima (L.) O. Bol`os & Vigo S. altissima L.; S. canadensis var. scabra Torr. & A. Gray Plant up to 250 cm. Stem usually hairy throughout. Leaves more scabrid. Involucral bracts 2.5–4.5 mm. Ligules 2.4– 4.1 mm Tubular flowers 3–4 mm. A very variable species and it is possible that some of our naturalised plants are hybrids with S. gigantea or S. rugosa. Introduced. Much grown in gardens and naturalised on waste land, banks, waysides and rough ground. Frequent throughout southern and central Great Britain and the Channel Islands and scattered localities in northern Britain and Ireland. Native of North America. Subsp. altisssima is the common race. Subsp. canadensis also occurs, but it is not known how frequent it is. × virgaurea = S. × niederederi Khek This hybrid has the inflorescence shape and capitulum size of S. canadensis and the leaf venation of S. virgaurea. It is sterile. Recorded from a railway bank in Kent in 1979. It is also recorded for Austria, Denmark and Sweden. 5. S. gigantea Aiton Early Goldenrod Perennial herb with rhizomes. Stems solitary or numerous and clustered, 50–250 cm, pale glaucous or yellowishgreen, often suffused purplish-red, rather stout, ridged and slightly angled, glabrous, often with a whitish bloom, very leafy, branched only in the inflorescence. Leaves all cauline, numerous, alternate, scarcely decreasing in size upwards, 4–9 × 1.0–1.5 cm, yellowish-green on upper surface, sometimes tinted reddish with age, paler beneath, very narrowly elliptical or narrowly lanceolate, acute or acuminate at the very slender apex, entire or with few, very small teeth towards the apex, gradually narrowed at base, glabrous on both surfaces or very shortly and sparsely hairy on the veins beneath, very shortly ciliate on the margin; with a prominent midrib beneath and 1–2 other veins running parallel with it; petiole short, pale green and glabrous. Capitula

(a) Subsp. gigantea Leaves hairy on the veins beneath. Involucral bracts 3.2–4.0 mm. Achenes usually glabrous. 2n = 18.

Introduced. Much grown in gardens and naturalised on waste land, banks, waysides, dunes and rough grassland. Frequent throughout south and central Great Britain and the Channel Islands and scattered localities in north Britain and Ireland. Only subsp. serotina has been recorded with certainty, but subsp. gigantea should be looked for. Both subspecies are native of North America. Subgenus Euthomia (Nutt.) Fernald Euthamia Nutt. Capitula all sessile. Ligulate flowers more numerous than tubular flowers. Receptacle pits minutely fimbriate. 6. S. graminifolia (L.) Salisb. Grass-leaved Goldenrod Chrysocoma graminifolia L.; Euthamia graminifolia (L.) Elliott; S. lanceolata L. Perennial herb with rhizomes. Stems 30–150 cm, pale brown, almost woody, erect, striate, glabrous to shortly hairy, leafy, much branched, the branches green. Leaves all cauline, gradually smaller upwards, 2–8(–15) × 0.2– 0.7(–1.2) cm, dark dull green on upper surface, only slightly paler beneath, linear to narrowly linear-lanceolate or lanceolate, narrowed to a more or less acute apex, entire, narrowed to a sessile base, with minute projections on the margins and midrib, otherwise glabrous but gland-spotted; midrib prominent beneath with 1–2 less prominent parallel veins either side of it. Capitula numerous, sessile, in erect, corymbose panicles; branches channelled, covered with minute projections. Involucral bracts in numerous rows, 1.5–5.0 × 0.6–1.0 mm, pale green, darker near the tips, linear-lanceolate or lanceolate, obtuse at apex, glabrous. Flowers of 2 kinds, the outer 15–25, 4.5–5.0 mm, ligulate, the ligules yellow, more numerous than and not exceeding the tubular flowers, minutely 3-lobed at apex, the inner tubular, 2.5–5.0 mm, yellow, 5-lobed at apex. Receptacle flat, with fimbriate pits, without scales. Achenes 0.5–1.0 mm,

58. Aster cylindrical, appressed-hairy; pappus 2.0–2.5 mm, white, of simple hairs. Flowers 5–10. 2n = 18. Very variable in width of leaf and hairiness. Introduced. Grown in gardens and naturalised on waste land, banks of streams and rivers and waysides. Very scattered records in England and central and south Scotland. Native of North America. 58. Aster L. Perennial herbs. Stems erect, leafy. Leaves alternate, simple and entire or shallowly toothed. Capitula numerous, in corymbs or elongated panicles. Involucral bracts in 2– several rows, herbaceous or partly membranous. Inner flowers tubular, 5-lobed and bisexual, the outer ligulate, in 1 row and female or neuter, sometimes absent. Corolla of inner flowers yellow, 5-lobed, of outer flowers blue, red or white, 3-lobed. Anthers without tails, apical appendage ovate-lanceolate and flat. Style with oblong-linear branches, prolonged into a sterile appendage ventrally glabrous, dorsally hairy and longer than the stigmatic area. Receptacle flat, without scales. Achenes oblong, ellipsoid, fusiform, obconical or oblanceolate, usually compressed; pappus of 1–2 rows of shortly denticulate hairs. About 250 species, mostly in North America, but extending into South America and widely distributed in the Old World. Various attempts have been made to divide the genus Aster into several smaller genera, but from a practical point of view this does not seem very sensible, as the characters used are not normally regarded as very important. The classification by Semple and Brouillet (1980) seems to be made more on basic chromosome number than important morphological features. The subgenera recognised in this account are the groups which some authors would regard as distinct genera. The Subgenus Aster, which includes most of the garden Michaelmas Daisies, contains species that are difficult to define owing to the presence of intermediates. This situation has been made worse by the bringing together of normally disjunct species into cultivation and the forming of hybrids. Asters that have escaped from gardens or even spread from gardens are frequent, and much more attention needs to be given to them before we know exactly what we have in Great Britain and Ireland. We have included here rather more species and varieties than we would normally, in an attempt to cover those taxa which may be found. We are especially indebted for help with this difficult group to Peter Yeo, who was preparing an account for The European Garden Flora at the same time as we were preparing ours. Cronquist, A. J. (1952). Aster L. in Gleason, H. A. The New Britton and Brown, Illustrated Flora of the North-eastern United States and adjacent Canada, Vol. 3, pp. 323–545. Lancaster, Pennsylvania. Cronquist, A. (1955). Compositae in Hitchcock, C. L. et al. Vascular plants of the Pacific Northwest. Vol. 5. Seattle, Washington. Fernald, M. L. (1950). Gray’s Manual of Botany. New York. Gray, A. J. (1987). The rayless genes in Aster tripolium: a longer term study in Gray, A. J. et al. Colonization, succession and stability. Oxford, London, Edinburgh.

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Hoffmann, M. H. (1996). Die in Zentraleuropa verwilderten und kultivierten nordamerikanischen Astern. Feddes Rept. 107: 163– 188. Huiskes, A. H. L., van Soelen, J. & Markusse, M. M. (1985). Field studies in the variability of populations of Aster tripolium L. in relation to saltmarsh zonation. Vegetatio 61: 163–169. Jones, A. G. (1978). Observations on reproduction and phenology of some perennial asters. Amer. Midl. Naturalist. 99: 184–197. Jones, A. G. (1980). A classification of the New World species of Aster (Asteraceae). Brittonia 32: 230–239. Labracque, J. & Brouillet, L. (1996). Biosystematique du complexe de l’Aster novi-belgii (Asteraceae: Asterae) au Qu´ebec. Canad. Jour. Bot. 74: 162–188. Semple, J. C. & Brouillet, L. (1980). A synopsis of North American asters: the Subgenera, Sections and Subsections of Aster and Lasallea. Amer. Jour. Bot. 67: 1010–1026. Semple, J. C. & Heard, S. B. (1987). The asters of Ontario: Aster L. and Virgulus Raf. (Compositae: Astereae). University of Waterloo Biology Series 30. (Very detailed illustrations.) Semple, J. C., Heard, S. B. & Xiang, C. (1996). The asters of Ontario (Compositae: Astereae): Diplactis Raf., Oclemena E. L. Greene, Doellingeria Nees and Aster L. (including Canadanthus Nesom, Symphyotrichum Nees, and Virgulus Raf.). University of Waterloo Biology Series 38. Stark, A. A. & Wijnands, D. O. (1970). On the variation of the flower heads of Aster tripolium L. in the Netherlands. Act. Bot. Neerl. 19: 436–444. 1. Basal and lower stem leaves with a cordate base and long 2. petiole 1. Basal leaves cuneate at base; cauline leaves cuneate to 3. attenuate at base and/or sessile 2. schreberi 2. Involucral bracts broadly rounded at apex 2. Involucral bracts narrowed to a more or less acute apex 3. cordifolius 3. Leaves all or upper only 1-veined, glandular-punctate and 4. not succulent, linear, lanceolate or narrowly elliptical 3. Leaves usually with well-developed lateral veins, if all 1-veined then succulent and not gland-dotted but 5. sometimes with glandular hairs 17. sedifolius 4. Involucral bracts more or les obtuse at apex 18. linosyris 4. Involucral bracts acute or acuminate at apex 5. Leaves succulent, without lateral veins or with few veins mostly running parallel with the midrib; mostly maritime 6. 5. Leaves not succulent, mostly with narrowly developed 8. later veins 6. Capitula without ligulate outer flowers 19(iii). tripolium var. flosculosus 7. 6. Capitula with ligulate outer flowers 7. Stems up to 20(–30) cm; leaves often forming a pseudorosette, very fleshy, lanceolate to elliptical; inflorescence condensed; cliffs or rocks 19(i). tripolium var. condensatus 7. Stems up to 100 cm; leaves slightly fleshy, lanceolate, to narrowly lanceolate-elliptical or linear; inflorescence a loose corymbose panicle; salt-marshes 19(ii). tripolium var. tripolium 8. Leaves and upper part of plant with numerous glandular 15. novae-angliae hairs 8. Leaves and upper part of plant without glandular hairs, 9. rarely with few glandular hairs on the involucral bracts 10. 9. Involucral bracts spine-tipped

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9. Involucral bracts not spine-tipped (but sometimes 11. hair-tipped) 10. Plant with elongated rhizomes forming clones 16(i). ericoides var. ericoides 10. Plant with corm-like rootstock 16(ii). ericoides var. pansus 12. 11. Involucral bracts obtuse at apex 14. 11. Involucral bracts acute at apex 12. Involucral bracts 8–12 mm, more or less equal, often with additional bracts; leaves more or less glabrous on 12. foliaceus the surfaces, but ciliate 12. Involucral bracts 3–9 mm, in a graduated series, without 13. additional large bracts; leaves more or less hairy 13. Outer involucral bracts obovate or oblanceolate, the 1. amellus inner oblong 13. Involucral bracts linear-lanceolate or narrowly 14. umbellatus lanceolate 15. 14. Upper cauline leaves not clasping the stem 23. 14. Upper cauline leaves more or less clasping the stem 16. 15. Involucral bracts with inrolled tips 17. 15. Involucral bracts with tips flat 16. Stem and leaves with numerous to dense hairs 4(i). pilosus var. pilosus 16. Stem and leaves glabrous or with a few hairs 4(ii). pilosus var. pringlei 17. Leaves of inflorescence much reduced and almost 18. bract-like 19. 17. Leaves of inflorescence not reduced 18. Inflorescence occupying one-third to two-thirds of the height of the plant, its branches spreading and forking 5(i). dumosus var. dumosus 18. Inflorescence occupying one-tenth to half the height of the stem, its branches stiffly ascending and only slightly 5(ii). dumosus var. strictior forking 19. Outer involucral bracts two-thirds or more the length of the inner, usually subtended by large leafy bracts. 11(a). lanceolatus subsp. hesperius 19. Outer involucral bracts one-third to two-thirds the length of the inner, not usually subtended by large leafy bracts 20. 20. Stems more or less densely hairy to the base 11(b,ii). lanceolatus subsp. lanceolatus var. hirsuticaulis 21. 20. Stems glabrous or hairs only in lines near the base 21. Involucral bracts less than 4 mm 11(b,i). lanceolatus subsp. lanceolatus var. interior 22. 21. Involucral bracts averaging greater than 4 mm 22. Leaves broadly oblanceolate; involucral bracts 4.0–5.5 mm; ligules white 11(b,iii). lanceolatus subsp. lanceolatus var. latifolius 22. Leaves linear to oblanceolate; involucral bracts 3.5–5.0(–6) mm; ligules white to bluish-violet 11(b,iv). lanceolatus subsp. lanceolatus var. lanceolatus 23. Stem and leaves more or less glaucous, especially the 24. leaves beneath 25. 23. Stem and leaves green 24. Involucral bracts clearly imbricated in several rows with the green tip short, broad and rhombic 7(i). laevis var. laevis 24. Involucral bracts subequal with long, narrow, green tips 7(ii). laevis var. geyeri 25. Involucral bracts 8–12 mm; upper leaves often strongly 26. auriculate-amplexicaul

27. 25. Involucral bracts 2–8 mm 26. Involucral bracts narrowly elliptical or broadly 12. foliaceus spathulate, shortly acute at apex 26. Involucral bracts linear to narrowly linear-lanceolate, 13. puniceus with a long, narrow acute apex 27. Involucral bracts whitish with a green centre in upper 28. half 27. Involucral bracts wholly or mainly greenish in upper 29. half 28. Capitula 15–20 mm in diameter; involucral bracts oblong or oblong-oblanceolate; ligulate flowers 8–11 6. concinnus mm 28. Capitula 30–50 mm in diameter; involucral bracts linear- or linear-lanceolate; ligulate flowers 17–25 mm 8. × versicolor 29. Middle cauline leaves mostly 2.5–5.0 times as long as wide, conspicuously amplexicaul; outer involucral bracts usually half to three-quarters as long as inner ones 8. × versicolor 29. Middle cauline leaves mostly 4–10 times as long as wide, usually only semiamplexicaul; outer involucral 30. bracts nearly as long as inner ones 30. Outer involucral bracts widest at or just above the middle with conspicuous green apical half, loosely or 9. novi-belgii unevenly appressed 30. Outer involucral bracts widest below the middle, the inner rather neatly appressed, the outer often lax 10. × salignus

Subgenus 1. Aster Perennial herbs with creeping rhizomes, or annuals. Stems erect. Involucral bracts with the green area in the upper third well defined and with a scarious margin, the green uniformly distributed or most dense apically, obovate or oblanceolate, oblong to narrowly lanceolate, sometimes leafy, ciliate. Section 1. Aster Perennial herbs. Leaves entire to serrate. Inflorescence a flat-topped, corymbose cyme. Involucral bracts with the green area truncate or cuneate and usually poorly delimited at the base, obovate, oblanceolate, oblong or broadly lanceolate, more or less obtuse at apex. Ligulate flowers usually in 1 row. Pappus of 1–2 rows of denticulate hairs. 1. A. amellus L. European Michaelmas Daisy A. amelloides Besser Perennial herb with creeping rhizomes and non-flowering rosettes. Stems 10–70 cm, yellowish-green becoming purple above, often flecked purplish, erect, but often decumbent at base, with a few to numerous, short, forwardly directed, curly, pale simple eglandular hairs, with erect branches above, leafy. Leaves 3–7 × 0.5–3.0 cm, faintly 3-veined, medium green on upper surface, paler beneath, with pale bulbous-based simple eglandular hairs on both surfaces and margin; basal and lower cauline obovate or oblanceolate, rarely elliptical, obtuse to acute at apex, entire or remotely crenate, long-attenuate at base to a winged petiole; median and upper oblong or lanceolate, obtuse at apex, entire, narrowed at base, sessile. Capitula few to numerous, rarely solitary, 25–40(–75) mm in diameter, usually in

58. Aster a flat-topped, cymose-corymbose inflorescence; peduncles rather long, often suffused brownish-purple, with numerous short, stiff, simple eglandular hairs. Involucral bracts in about 3 rows, 5–9 × 1.5–3.5 mm, with the green area pale, poorly delimited at base and with a reddish apex, the outer obovate or oblanceolate, the inner oblong, obtuse and often recurved at apex and glabrous on the surface with ciliate margins. Flowers of 2 kinds, the outer 10–40, 13–32 mm, ligulate, the ligules usually deep purplish-blue, rarely red or white and minutely 3-lobed at apex, the inner 4.5–8.0 mm, tubular, yellow and with 5 triangular lobes at apex. Receptacle flat, pitted, the pits with a dentate margin. Achenes 2–3 mm, pale brown, ellipsoid, compressed, hairy; pappus 4–8 mm, of a single whorl of minutely denticulate, whitish or brown-tinged hairs. Flowers 7–10. 2n = 18, 36, 54. A very variable species in need of further study. Introduced. A garden escape naturalised on waste ground at a disused railway station at South Wigston in Leicestershire. Native of central and south Europe and western Asia. 2. A. schreberi Nees Nettle-leaved Michaelmas Daisy A. macrophyllus auct. Perennial herb with tufts of leaves on short, lateral rhizomes, the rhizomes often elongated. Stems 50–110 cm, solitary, pale yellowish-green, often suffused purplish, erect, striate, usually glabrous below, becoming sparsely hairy above with short to long, pale simple eglandular hairs, branched above, leafy. Leaves 2–10 × 1–5 cm, medium green on upper surface, paler beneath, glabrous or with a few, short, stiff simple eglandular hairs on the upper surface and few to numerous ones beneath and on the margins and longer ones on the veins; lower ovate or lanceolate, acute at apex, coarsely serrate, cordate at base, the petioles up to 15 cm and with long simple eglandular hairs; median and upper similar to lower but rounded or cuneate at base and sessile. Capitula few to numerous, 25–30 mm in diameter, cylindrical, in a flat-topped cymose-corymbose or paniculate inflorescence; peduncles with few to numerous, short to medium, stiff, pale simple eglandular hairs, sometimes with a very few glandular hairs. Involucral bracts in 4–6 rows, 5.5–7.0 × 1.5–2.0 mm, pale green with a darker green area in the upper third which is darkest near the tip, lanceolate or oblong, broadly rounded at apex, with minute glandular hairs and a fimbriate margin. Flowers of 2 kinds, the outer 6–12, 6.5–10.0 mm, ligulate, the ligules white and with 3 minute lobes at apex, the inner 14–30, 5–6 mm, tubular, yellow becoming purple, with 5 narrow lobes at apex. Receptacle flat, pitted, the pits with a dentate margin. Achenes 3–4 mm, pale brown, fusiform, compressed, with 6–12 ribs, glabrous to sparsely strigose-hairy; pappus 5–6 mm, of a single whorl of minutely denticulate hairs. Flowers 8–10. 2n = 54. Introduced. Naturalised on a railway bank near Lochside Station in Renfrewshire since 1931. Native of eastern North America. Named after Johann Christian Daniel von Schreber (1730–1810). Originally called A. macrophyllus L., which differs in having densely glandular-hairy peduncles and involucral bracts and pale to dark bluish-violet (rarely

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faint pink or white) ligules. The true A. macrophyllus was persistent at Swanley in Kent for a few years. It occurs in gardens and may turn up elsewhere. Section 2. Dumosi Torr. & A. Gray Perennial herbs. Inflorescence a corymbose to paniculate cyme. Involucral bracts with the green area equally intense throughout, oblong, linear-lanceolate or linearoblanceolate. 3. A. cordifolius L. Blue Wood Aster A. paniculatus Willd. Perennial herb with a woody, branched rootstock, forming clones by elongated rhizomes. Stems few to numerous, 20– 120 cm, yellowish-green, sometimes tinted purplish, erect, glabrous or with fairly numerous simple eglandular hairs, branched in upper part, leafy. Leaves 2.5–12.0 × 1–6 cm, medium green, paler beneath, glabrous or with fairly numerous, stiff, pale simple eglandular hairs on both surfaces especially along the margins on the upper surface and on the veins beneath; lower ovate, acute or acuminate at apex, serrate, deeply cordate at base, long petiolate, usually deciduous by anthesis; middle cauline ovate, acuminate at apex, serrate, shallowly cordate at base, with narrowly winged petioles; upper and ones on branches ovate to lanceolate or linear-lanceolate, acute or acuminate at apex, serrate, sessile. Capitula few to numerous, 10–16 mm in diameter, campanulate-turbinate, in a paniculate-cymose inflorescence which is ovoid in outline; peduncles pale green, with few to dense, pale simple eglandular hairs and many small sterile bracts. Involucral bracts in 4–6 strongly graduated rows, appressed, 3.5–5.5 × 0.7–1.0 mm, uniform green with reddish-purple tips, oblong, more or less acute at apex, glabrous or with sparse simple eglandular hairs. Flowers of 2 kinds, the outer 10–16, 5–8 mm, ligulate, the ligules pale blue, white, pink, violet or lilac, and minutely 3-lobed at apex, the inner 14–20, 3–5 mm, tubular, yellow becoming purple, with 5 triangular lobes at apex. Receptacle flat, pitted, the pits with a dentate margin. Achenes 1.8–2.2 mm, obconical, compressed, with 1 rib per side; pappus 3.8–4.5 mm, of a single whorl of whitish to pale brown minutely denticulate hairs. Flowers 8–10. Visited by bees, wasps and hoverflies. 2n = 16, 32. Very variable in flower colour, hairiness and habit. Introduced. A persistent garden escape near Whittlesey in Cambridgeshire and a garden relic at Denver in Norfolk. Native of eastern North America. 4. A. pilosus Willd. Frost Aster A. ericoides auct. Perennial herb with a short branched stock or elongated rhizomes. Stems 20–120 cm, pale yellowish-green, sometimes suffused brownish-purple, sometimes striate, glabrous and smooth to rough with numerous to dense, stiff, pale simple eglandular hairs, much branched with spreading or ascending branches, leafy. Leaves up to 16 × 2 cm, medium yellowish-green, paler beneath, glabrous or with numerous to dense, short to medium, stiff, pale simple eglandular hairs on both surfaces and the margins; basal lanceolate, oblanceolate, spathulate or obovate, rounded at apex, entire

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or sparsely serrate at apex, gradually narrowed at base, petiolate or sessile and usually deciduous at anthesis, the median and upper linear to narrowly lanceolate or oblanceolate, acute to a hair-like point at apex, entire and narrowed to a sessile base. Capitula numerous, 10–20 mm in diameter, in a large cymose panicle, sometimes secund; peduncles rather short, glabrous or with sparse to dense, short to long, pale, rather stiff simple eglandular hairs and many bracts. Involucral bracts in 4–6 rows, 4.0–6.5 × 0.8–1.0 mm, outer green, the inner whitish, with an elongate-diamond-shaped uniform green area at apex, lanceolate, the acute, hair-like apex spreading with its margin inrolled, glabrous or with a few, pale simple eglandular hairs. Flowers of 2 kinds, the outer 20–35, 3–8 mm, ligulate, the ligules usually white, sometimes pink or pale blue and minutely 3-lobed at apex, the inner 25–45, 2.5–5.0 mm, tubular, yellow becoming purple, with 5 triangular lobes at apex. Receptacle flat, pitted, the pits with dentate margins. Achenes 0.8–1.0 mm, pale brown, obconical, compressed, with 1 rib on each side, with few to fairly numerous, strigose hairs; pappus 2.0– 3.5 mm, of a single whorl of minutely denticulate hairs. Flowers 9–10. Sometimes carries on flowering and producing new rosettes of small leaves throughout the winter. 2n = 32, 40, 48.

with 3-minute lobes at apex, the inner 15–30, 3.5–4.5 mm, tubular, white to pale yellow becoming purple, 5-lobed at apex. Receptacle flat, pitted, the pits with a dentate margin. Achenes 1.8–2.2 mm, obconical, compressed, with 1 rib on each side, sparsely strigose-hairy; pappus 3.5–4.5 mm, of a single whorl of minutely denticulate hairs. Flowers 8–10. 2n = 16, 32. A variable plant with several named varieties. The two most obvious are as follows.

(i) Var. pilosus Stems rough with stiff, numerous to dense, pale simple eglandular hairs. Leaves lanceolate, oblanceolate or linearlanceolate, hairy. 2n = 32, 48.

6. A. concinnus Willd. Narrow-leaved Smooth Aster A. laevis var. concinnus (Willd.) House Perennial herb with long, stout, woody, leafy rhizomes. Stems 20–100 cm, pale green, often flushed reddish-purple, erect, rather slender but rigid, striate, with obscure puberulent lines decurrent from the nodes, otherwise glabrous or nearly so. Leaves 3–8(–14) × 0.4–2.0 cm, rather pale green and even paler beneath, those of the rhizome often suffused purplish, glabrous, those of the rhizome lanceolate or narrowly elliptical, obtuse at apex, minutely serrulate on the margin and long attenuate at base to a winged, often purplish petiole, the cauline narrowly lanceolate, linearlanceolate or very narrowly elliptical, acute or acuminate at apex, minutely serrulate, gradually narrowed to a sessile, more or less semiamplexicaul base. Capitula numerous, 15– 20 mm in diameter, in a long, narrow panicle; peduncles pale green or brownish-purple, striate, glabrous. Involucral bracts in several rows, 2–5 × 0.3–0.7 mm, whitish, with a long, deltate, greenish patch above and tipped reddish, oblong or oblong-oblanceolate, acute at apex, glabrous. Flowers of 2 kinds, the outer 8–11 mm, ligulate, the ligules pale lilac and deeper in colour towards the minutely 3-lobed apex, the inner 5–9 mm, pale yellow becoming purplish, narrowly 5-lobed at apex. Receptacle flat, pitted, the pits with a dentate margin. Achenes 3.0–3.5 mm, pale brown, oblanceolate, compressed, hairy; pappus 5–6 mm, dirty white, of a single whirl of minutely denticulate hairs. Flowers 8–9. Introduced. Naturalised on waste ground by the railway at Hob Moor, near York and in Cardiganshire on the north bank of Afon Cletwr near Tre’r-ddol and near Aberystwyth railway station. Native of eastern North America.

(ii) Var. pringlei (A. Gray) Blake A. pringlei A. Gray; A. pilosus var. demotus Blake Stems smooth and almost hairless. Leaves mostly linear, almost glabrous. 2n = 48. Introduced. Var. pilosus is naturalised near Handley in Dorset and the species occurs as a casual garden escape in a few localities. Both varieties are grown in gardens. Native of eastern North America. 5. A. dumosus L. Bushy Aster A. foliosus Pers. Perennial herb with a branching stock. Stems few to numerous, 30–90 cm, pale green, erect, slender, glabrous or with few to dense, pale simple eglandular hairs, branched in upper half, leafy. Leaves 1.5–5.0(–10) × 0.1–0.3 cm, medium green, paler beneath, glabrous to sparsely scabrid near the margins on the upper surface, glabrous to sparsely and stiffly hairy beneath; lower cauline leaves linearoblanceolate, acute at apex, entire to slightly serrate, narrowed at base, sessile, often deciduous at anthesis; upper cauline leaves small linear to narrowly lanceolate, acute at apex, entire, sessile. Capitula numerous, 8–18 mm in diameter, campanulate-turbinate, in a much-branched cymose panicle, rather narrow in outline; peduncles long, with rather sparse simple eglandular hairs and many linear, sterile bracts. Involucral bracts in 4–6 graduated rows, 1.5– 4.5 × 0.7–1.0 mm, whitish below with a narrowly elliptical uniform green area sometimes slightly purplish in the upper part, oblong, acute at the flat apex, glabrous on the surface with ciliate margins. Flowers of 2 kinds, the outer 15–33, 5–7 mm, ligulate, the ligules white to pale purplish-pink,

(i) Var. dumosus Inflorescence occupying one-third to two-thirds the height of the plant, its branches spreading and forking. Ligulate flowers 20–26. (ii) Var. strictior Torr. & A. Gray Inflorescence occupying one-tenth to one-half the height of the stem, its branches stiffly ascending and only slightly forking. Introduced. Apparently only recorded previous to 1930, but some of the dwarf Michaelmas Daisies may belong here or they may be hybrids with other species. As it is likely to turn up the description is included. Native of North America.

7. A. laevis L. Glaucous Michaelmas Daisy Symphyotrichum laeve (L.) Å. L¨ove Perennial herb with a short, stout, woody, branching stock, rarely with a few, creeping rhizomes. Stems 30–120 cm,

58. Aster pale glaucous, often flushed reddish-purple, erect, rigid, with obscure puberulent lines decurrent from the nodes, otherwise glabrous or nearly so. Leaves (1–)2–18 × 0.5– 4.0 cm, dull, medium, glaucous green, paler and somewhat pruinose beneath, firm and rather thick, narrowly linear to broadly ovate, acute at apex, entire or obscurely dentate, the lower narrowed into winged petioles, and soon deciduous, the median and upper gradually smaller, auricled and more or less amplexicaul, glabrous. Capitula few to numerous, 15–30 mm in diameter, in a long, narrow panicle; peduncles pale green or glaucous, glabrous. Involucral bracts in several rows, 3–7 × 1.4–1.7 mm, very unequal or subequal, whitish, but with a green, deltate, narrow or rhombic patch above, linear to linear-lanceolate, with appressed, shortly tapered, acute apex, glabrous or ciliate. Flowers of 2 kinds, the outer 10–35, 8–15 mm, ligulate, the ligules pale to dark violet-blue or purple, rarely white and 3-lobed at apex, the inner 15–40, 4.5–8.5 mm, tubular, becoming reddish-yellow and with 5 triangular lobes at apex. Receptacle flat, pitted, the pits with a dentate margin. Achenes 3.3–3.6 mm, brown, oblanceolate, compressed, with 1–2 ribs on each side, glabrous or nearly so; pappus 5–6 mm, whitish to pale brown or reddish, of a single whorl of minutely denticulate hairs. Flowers 8–10. Visited by bees, wasps and hoverflies. 2n = 48, 54. (i) Var. laevis Involucral bracts clearly imbricated in several rows with the green tip short, broad and rhombic. (ii) Var. geyeri A. Gray A. brevibracteatus Rydb.; A. geyeri (A. Gray) Howell; A. pickettiana Suksd. Involucral bracts subequal with long, narrow, green tips. Introduced. Naturalised on waste and rough ground. Scattered localities in south and central England. Both varieties may occur. Native of North America. 8. A. × versicolor Willd. Late Michaelmas Daisy A. laevis × novi-belgii A. novi-belgii subsp. laevigatus (Lam.) Thell.; A. laevigatus Lam.; Symphyotrichum × versicolor (Willd.) G. L. Nesom Perennial herb with a short, stout stock. Stems 50–220 cm, pale yellowish-green, often flushed or saturated brownishpurple, striate, glabrous or nearly so. Leaves 2–15 × 0.5– 4.0 cm, thick, rather deep green, slightly paler, but not glaucous beneath, ovate, lanceolate, obovate, or oblonglanceolate, more or less obtuse at apex, entire, denticulate or shallowly crenate, the lower narrowed at base to a winged, semiamplexicaul petiole, the upper rounded at base, sessile and more or less amplexicaul, glabrous. Capitula 30–50 mm in diameter, mostly in a narrow, open panicle, sometimes corymbiform; peduncles pale green or purplish, ridged, glabrous or with short simple eglandular hairs and numerous sterile linear-lanceolate bracts. Involucral bracts in several rows, appressed or outer spreading at tips, 3.5– 7.0 × 0.7–1.0 mm, whitish, with a narrow, green, deltate or rhombic patch above, linear or linear-lanceolate, more or less acute at apex, glabrous. Flowers of 2 kinds, the outer 15–25 mm, ligulate, the ligules pale mauve to violet-blue

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and with 3 minute lobes at apex, the inner 10–14 mm, tubular, pale yellow at first finally turning dull purplish and narrowly 5-lobed above. Receptacle flat, pitted, the pits with toothed, membranous borders, without scales. Achenes 1.5– 2.5 mm, pale brown, oblanceolate, compressed, appressedhairy; pappus 4–7 mm, whitish, of several rows of minutely denticulate hairs. Flowers 10–11. The larger-flowered, taller, late-flowering cultivars of Michaelmas Daisies belong here. They can be distinguished from A. laevis by their regularly imbricated involucral bracts with more extensive green tips and from A. novi-belgii by their more regular and more appressed involucral bracts with shorter, less green tips. The hybrids often flower later than both parents. Some plants are very near A. novibelgii and differ mainly in their more appressed involucral bracts. Introduced. A fairly common garden plant which is naturalised in waste and rough places. Scattered records in Great Britain, but probably much under-recorded. It is not known in North America where the parents are native. 9. A. novi-belgii L. Confused Michaelmas Daisy A. longifolius auct.; A. brumalis Nees; A. floribundus auct.; Symphyotrichum novi-belgii (L.) G. L. Nesom Perennial herb. Stems 40–150 cm, pale yellowish-green, often suffused brownish-purple, erect, ridged and channelled, glabrous or with short, pale simple eglandular hairs above which are sometimes confined to the ridges, very leafy, much branched above. Leaves all cauline, much smaller on the branches, 1.0–12.0 × 0.3–2.0 cm, dull, rather dark green on upper surface, paler beneath, lanceolate, linear-lanceolate, linear or oblong, more or less acute at apex, entire or remotely toothed, firm, rounded at the semiamplexicaul, auricled base, glabrous or with some very short simple eglandular hairs beneath; midvein prominent beneath. Capitula 25–50 mm in diameter, campanulate, in a symmetrical, sometimes corymbiform panicle; peduncles green, ridged, with short woolly hair and numerous, recurved, linear bracts. Involucral bracts in several rows, 5– 8 × 0.7–1.0 mm, the outer often as long as the inner, loose at least at the tips and green, the inner linear-lanceolate, long-acute at apex, loose or appressed, white with a green spear-shaped area at tip, glabrous. Flowers of 2 kinds, the outer 30–50, 7–25 mm, ligulate, the ligules violet-blue or in cultivars white, violet or purple, rounded and 3-lobed at apex, the inner 5–10 mm, tubular, yellow, becoming purple or red, narrowly 5-lobed above. Receptacle flat, pitted, the pits with toothed membranous borders, without scales. Achenes 1.5–2.5 mm, oblong, appressed-hairy; pappus 4–7 mm, white, of several rows of minutely denticulate hairs. Flowers 9–11. Visited by bees, wasps and hoverflies. 2n = 18, 48, 49, 54. Very variable as a wild species and in gardens a complex of cultivars often make it difficult to distinguish from A. × versicolor and A. × salignus. J. Labracque and L. Brouillet (1996) have revised the aggregate in Quebec. Introduced. Grown in gardens and widely naturalised in waste and rough ground, waysides and railway banks. In scattered localities over Great Britain and recorded for Ireland. Native of North America.

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10. A. × salignus Willd. Common Michaelmas Daisy A. lanceolatus × novi-belgii A. longifolius auct.; Symphotrichum × salignum (Willd.) G. L. Nesom Perennial herb. Stems 5–130 cm, pale yellowish-green, usually suffused brownish-purple, erect, striate or ridged, glabrous below, usually shortly hairy above especially on the ridges, leafy, much branched above. Leaves all cauline, gradually decreasing in size upwards, very numerous, 20– 100 × 3–15(–20) mm, lanceolate, medium green on upper surface, paler beneath, lanceolate, oblong-lanceolate, acute at apex, entire to shortly serrate, narrowed to a usually shortly auricled, semiamplexicaul base, glabrous or the upper slightly hairy; midrib prominent beneath. Capitula 15–25 mm in diameter, usually in a much broader panicle than in A. lanceolatus; peduncles and branches yellowishgreen, often suffused brownish-purple, with numerous to dense, short, subrigid simple eglandular hairs. Involucral bracts in several rows, the outer nearly as long as the inner, 5–7 × 0.5–1.0 mm, widest below the middle, almost wholly medium green in the upper half and with reddish tips, oblong to linear-lanceolate, acute at apex, glabrous or with short hairs especially on the margins, the inner neatly appressed, the outer often lax. Flowers of 2 kinds, the outer 9–12 mm, ligulate, the ligules bluish or white turning bluish and rounded and 3-lobed at apex, the inner 4–5 mm, tubular, dirty yellowish-brown and narrowly 5-lobed above. Receptacle flat, pitted, the pits with toothed membranous borders, without scales. Achenes 1.5–2.0 mm, oblong, appressed-hairy; pappus 4.0– 5.0 mm, white, of several rows of minutely denticulate hairs. Flowers 9–11. Visited by bees, wasps and hoverflies. 2n = 18. Introduced. Grown in gardens and widely naturalised on waste and rough ground, waysides, railway banks and ditchsides. Probably the commonest Michaelmas Daisy throughout Great Britain and Ireland. A hybrid of garden origin reproducing by seed and often weedy in appearance.

branches; peduncles and branches ridged, clothed like the upper stem and leafy. Involucral bracts in several rows, 3.5–5.5(–6.0) × 0.4–0.6 mm, medium green with very pale green margins and red-tipped, very unequal, appressed, linear- or linear-lanceolate, flat and long-acute at apex, glabrous or with minute hairs. Flowers of 2 kinds, the outer 20–40, 5–12 mm, ligulate, the ligules usually white, sometimes bluish-violet 3-lobed and rounded at apex, the inner 3–6 mm, tubular, dirty yellowish-brown, narrowly 5lobed above. Receptacle flat, pitted, the pits with toothed, membranous borders, without scales. Achenes 1.5–2.0 mm, oblong, appressed-hairy; pappus 4.0–5.5 mm, white, of several rows of minutely denticulate hairs. Flowers 9–1. Visited by bees, wasps and hoverflies. 2n = 32, 64.

11. A. lanceolatus Willd. Narrow-leaved Michaelmas Daisy A. paniculatus Lam., non Willd.; A. tradescantii auct.; Symphyotrichum lanceolatum (Willd.) G. L. Nesom; A. salicifolius Aiton, non Lam. Perennial herb with creeping rhizomes. Stems 50–150 cm, pale yellowish-green, usually suffused brownish-purple, erect, striate or ridged, the channels sometimes brownishpurple with the ridges green, glabrous or with short, pale simple eglandular hairs above, which are sometimes confined to the ridges, leafy, branched in upper half. Leaves all cauline, gradually decreasing in size upwards, numerous, 2–15 × 0.3–1.5(–3.0) cm, pale yellowish-green above, paler beneath, narrowly to broadly oblanceolate, linear-lanceolate or the uppermost linear, acute at apex, entire to shortly serrate, gradually narrowed below to a sessile or subpetiolate base, glabrous or uppermost shortly hairy; midvein prominent beneath. Capitula 13–25 mm in diameter, sometimes subtended by large leafy bracts, in a narrow panicle, often arranged unilaterally on the

(iii) Var. latifolius Semple & Chmiel. Stems glabrous or only with lines of hairs at the base. Leaves broadly oblanceolate, not much reduced in size in the inflorescence. Involucral bracts 4.0–5.5 mm.

(a) Subsp. hesperius (A. Gray) Semple & Chmiel. A. hesperius A. Gray; A. hesperius var. wootonii Greene; A. wootonii (Greene) Greene; A. osterhoutii Rydb.; A. fluvialis Osterh.; A. durus Lunell; A. lautus Lunell; Symphyotrichum hesperius (A. Gray) Å. L¨ove Leaves linear or narrowly linear-lanceolate. Capitula usually subtended by large, leafy bracts. Involucral bracts 4–5 mm, the outer two-thirds the length of the inner or more. Ligulate flowers blue. (b) Subsp. lanceolatus Capitula not usually subtended by leafy bracts. Outer involucral bracts one-third to two-thirds as long as inner. (i) Var. interior (Wiegand) Semple & Chmiel. A. interior Wiegand; A. simplex var. interior (Wiegand) Cronquist; A. lanceolatus subsp. interior (Wiegand) A. G. Jones Stem glabrous or with only lines of hairs at the base. Leaves linear or narrowly linear-lanceolate. Involucral bracts less than 4 mm. (ii) Var. hirsuticaulis Semple & Chmiel. Stem moderately to densely hairy at base. Leaves linear or narrowly linear-lanceolate. Involucral bracts more than 4 mm.

(iv) Var. lanceolatus A. bellidiflorus Willd.; A. eminens Willd.; A. laxus Willd.; A. dracunculoides Willd.; A. recurvatus Willd; A. simples Willd.; A. stenophyllus Lindl.; Symphyotrichum simplex (Willd.) Å. L¨ove Stems glabrous or only with lines of hairs at the base. Leaves linear to oblanceolate, reduced in size in the inflorescence. Involucral bracts 3.5–5.0(–6.0) mm. Introduced. Grown in gardens and widely naturalised on waste and rough ground, waysides and ditch margins. Frequent throughout much of Great Britain and Ireland and probably under-recorded. Native of North America. Said to have grown in gardens from the early seventeenth century and to be the original Michaelmas Daisy. All our specimens need to be checked to see which variants occur.

58. Aster 12. A. foliaceus Lindl. Leafy Aster Perennial herb with fibrous roots and a creeping stock or rhizome. Stems 20–50 cm, pale yellowish-green often suffused brownish-purple, erect, rigid, striate, glabrous or with very few, short, pale simple eglandular hairs, much branched, leafy. Leaves fairly numerous, dull medium green above, paler beneath, glabrous on both surfaces, minutely ciliateappressed; lower 12–20 × 1.6–4.0 cm, oblanceolate, or narrowly oblong-elliptical or linear-elliptical, obtuse to acute at apex, entire, more or less attenuate at base, long-petiolate, the petioles winged and semiamplexicaul; middle and upper similar but gradually smaller and narrower, acute at apex, sessile and amplexicaul, sometimes auriculate. Capitula few to fairly numerous, 20–45 mm in diameter, cymosepaniculate; peduncles pale green, rather long, with minute simple eglandular hairs. Involucral bracts 8–12 × 2–4 mm, green for two-thirds of upper part or more with darker midrib and pale narrow margins, more or less equal but additional larger bracts sometimes present, oblong, narrowly elliptical or broadly spathulate, acute to rounded-obtuse at apex, glabrous on the surface with a ciliate margin. Flowers of 2 kinds, the outer 10–17 mm, ligulate, the ligules purple, blue or violet, and shallowly 3-lobed at apex, the inner 7– 11 mm, tubular, yellowish, narrowly 5-lobed at apex. Receptacle slightly convex, pitted, the pits with a dentate margin. Achenes 3.0–3.5 mm, pale brown, oblanceolate, glabrous or appressed-hairy; pappus 5–6 mm, whitish or pale brown. Flowers 8–9. Visited by bees and hoverflies. This species occurs in gardens and may well escape. It is similar in appearance to A. laevis, but lacks the pruinose leaves and has fewer large capitula. Many varieties have been named (cf. Cronquist, 1955; Fernald, 1950), some of which do not have the leafy involucral bracts after which the plant is named. Native of northern North America. 13. A. puniceus L. Red-stalked Michaelmas Daisy A. firmus Nees; A. hispidus Lam. Perennial herb. Stems numerous, 40–180 cm, pale green flushed purplish-red, erect, rigid, ridged, from almost glabrous to softly or rigid hairy with stiff, spreading, swollen-based simple eglandular hairs, without glandular hairs, leafy, much branched. Leaves 20–200 × 5–30 mm, dull medium green above, paler beneath with an even paler prominent midvein, lanceolate, oblong-lanceolate, long-acute to acuminate at apex, entire to coarsely serrate, the sessile base strongly auricled and more or less amplexicaul, minutely scabrid especially on the margins and midrib below. Capitula numerous, 15–25 mm in diameter, hemispherical, in a corymbose panicle; peduncles pale yellowish-green, with short, simple eglandular hairs. Involucral bracts 8–10 × 1.0–1.2 mm, lax with apices recurved, medium green with pale margins, linear to narrowly linear-lanceolate, with a long, narrow, acute apex, more or less shortly simple hairy. Flowers of 2 kinds, the outer 10–15 mm, ligulate, the ligules violet-blue, rarely lilac, pink or white and shallowly 3-lobed at apex, the inner 8–9 mm, tubular, straw-coloured with brownish-red tips and divided into 5-triangular lobes at apex. Receptacle slightly convex, pitted, the pits with a dentate margin. Achenes

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2–3 mm, brown, oblanceolate, more or less appressed-hairy; pappus 5–6 mm, white, of several rows of minutely denticulate hairs. Flowers 9–10. Visited by bees, wasps and hoverflies. 2n = 16. In its native North America it is wide-ranging and very variable, but it is questionable if any of its named varieties can be accepted. Introduced. An established garden escape naturalised over an extensive area about Clachan in Kintyre and by the Rivers Greeba and Neb in the Isle of Man and occasional as a casual elsewhere. Native of eastern North America. Subgenus 2. Doellingeria (Nees) A. Gray Doellingeria Nees; Aster section Doellingeria (Nees) Kitam. Perennial herbs with creeping rhizomes. Stem usually solitary and erect. Leaves entire. Involucral bracts with the green area forming a narrow band down the centre and most dense near the midrib, narrowly lanceolate or linearlanceolate, obtuse at apex. Ligulate flowers in 1 row. Pappus of 2 rows, the outer few and shorter than the numerous inner. 14. A. umbellatus Mill. Tall White Aster Doellingeria umbellata (Mill.) Nees Perennial herb with a thick, woody, creeping rhizome. Stems usually solitary, 30–200 cm, pale green or tinted purplish-brown, erect, rigid, glabrous or with numerous to dense, pale simple eglandular hairs. Leaves numerous, the lowest deciduous at anthesis, 4–16 × 1.0–2.5 cm, dull medium green, paler beneath, narrowly to broadly elliptical, lanceolate or linear-lanceolate, acute at apex, entire, gradually narrowed at base, scabrous to sparsely strigose-hairy on the upper surface, glabrous or with numerous to dense, pale simple eglandular hairs below or tomentose-canescent, the lower shortly petiolate, the remainder sessile. Capitula numerous, 10–18 mm in diameter, turbinate-cylindrical, in a flat-topped corymbose cyme, occasionally more open and paniculate or raciform; peduncles sparsely to densely rigid-hairy. Involucral bracts in 3–4 graduated series, 3– 7 × 0.8–1.2 mm, with a narrow green band down the centre and broad scarious margins, linear-lanceolate or narrowly lanceolate, obtuse at apex, with few, pale simple eglandular hairs and very few glandular hairs. Flowers of 2 kinds, the outer 4–15, 5–9 mm, ligulate, the ligules white and minutely 3-lobed at apex, the inner 10–40, 5.5–6.5 mm, tubular, yellow, 5-lobed at apex. Receptacle flat, pitted, the pits with dentate margins. Achenes 2.5–3.0 mm, obconical, compressed, 3- to 5-ribbed, sparsely strigose-hairy; pappus 5.0– 5.5 mm of 2 rows of pale brown bristles, the outer few and shorter than the numerous inner. Flowers 9–10. 2n = 18. (i) Var. umbellatus Leaves narrowly elliptical, lanceolate or linear-lanceolate and sparsely hairy. Inflorescence usually open. (ii) Var. pubens A. Gray Doellingeria pubens (A. Gray) Rydb.; A. pubentior Cronquist Leaves broadly elliptical, densely tomentose-canescent on under surface. Inflorescence usually more compact.

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Introduced. A persistent garden escape near Newton Stewart in Kirkcudbrightshire and recorded from Limpsfield in Surrey. Native of North America, Var. umbellatus occurs north-east of a broad zone of intermediates running though Ontario, Wisconsin and Minnesota and var. pubens south-west of this zone. It is not known to which variety our introduced plants belong. Subgenus 3. Virgulus (Raf.) A. G. Jones Virgulus Raf.; Lasallea Greene Perennial herbs usually with a woody rhizome. Stems erect or ascending. Involucral bracts green below, darker above, with paler margins and sometimes suffused brownishpurple, linear to narrowly linear-lanceolate or oblong, narrowed above to a long-acute apex. 15. A. novae-angliae L. Hairy Michaelmas Daisy Virgulus novae-angliae (L.) Reveal & Keener; Lasallea novae-angliae (L.) Semple & Brouillet Foetid, perennial herb, usually with a woody stock. Stems usually numerous, 30–220 cm, usually purplish-brown, erect or ascending, robust, with numerous medium to long, pale, spreading simple eglandular hairs and dense, minute, pale glandular hairs intermixed, very leafy, branched in upper half. Leaves all cauline, getting smaller upwards, those of the branches much smaller, 0.2–12.0 × 0.3–2.0 cm, dull yellowish-green, paler beneath, lanceolate or linearlanceolate, acute or acuminate at apex, entire, with rounded auricles at the semiamplexicaul base, densely covered with pale, minute glandular hairs and with some very short, very thick single hairs on the margin; midrib prominent beneath. Capitula numerous, 20–40 mm in diameter, hemispherical, in a large corymbose panicle; peduncles yellowishgreen, suffused brownish-purple, with dense, minute, pale or reddish glandular and a few, long simple eglandular hairs. Involucral bracts in several rows, lax or recurved, 7–9 × 0.5–0.8 mm, medium green, darker at the apex and with pale margins, sometimes suffused brownish-purple, linear to narrowly linear-lanceolate, narrowed to a longacute apex, the inner almost filamentous, with dense minute, often reddish glandular hairs. Flowers of 2 kinds, the outer 40 or more, 10–15 mm, ligulate, the ligules reddish-purple or mauve, more rarely pink or white and minutely 3-lobed at apex, the inner 7–9 mm, tubular, yellow and with 5triangular lobes at apex. Receptacle flat, pitted, the pits with dentate margins. Achenes about 2 mm, oblong, appressedhairy; pappus 4–5 mm, pale brown, of minutely denticulate hairs. Flowers 9–10. Visited by bees, wasps and hoverflies. 2n = 10. Introduced. Naturalised on waste and rough ground. In scattered localities in Great Britain, mainly in the centre and south. Native of North America. 16. A. ericoides L. Heath Aster Virgulus ericoides (L.) Reveal & Keener; A. multiflorus Aiton; Lasallea ericoides (L.) Semple & Brouillet Perennial herb with a woody, corm-like rootstock or elongated leafy rhizomes. Stems 40–120 cm, pale green, sometimes tinted purplish, ascending to erect, striate, with numerous to dense strigose hairs in upper part, branched above, leafy. Leaves up to 5 × 0.4(–0.7) cm, medium green on

upper surface, paler beneath, with numerous to dense strigose hairs on both surfaces and the margins; lower oblanceolate, obtuse but spine-tipped at apex, entire and sessile. Capitula numerous, 8–12 mm in diameter, campanulate, in a pyramidal inflorescence, the capitula distributed along the branches; peduncles short, with dense, pale simple eglandular hairs and numerous small, sterile bracts. Involucral bracts in 3 to 4 rows, 3.0–4.5 × 0.5–1.0 mm, green below with a darker green, diamond-shaped zone at apex, oblong, acute at apex and spine-tipped, with few to numerous, stiff simple eglandular hairs, the outer bracts spreading to reflexed. Flowers of 2 kinds, the outer 13–20, 3.0– 4.5 mm, ligulate, the ligules white and with 3 minute lobes at apex, the inner 7–15, 2.5–4.0 mm, tubular, yellow becoming brown and with narrow lobes at apex. Receptacle flat, pitted, the pits with a dentate margin. Achenes 1.2–1.5 mm, pale brown, fusiform-obconical, faintly ribbed, with dense strigose hairs; pappus 3.5–4.0 mm, white, of a single row of minutely denticulate hairs. Flowers 7–10. 2n = 10, 20. (i) Var. ericoides Plant with elongated rhizomes forming clones. (ii) Var. pansus (Blake) Boivin A. pansus Blake; Virgulus ericoides var. pansus (Blake) Reveal & Keener Plant with a woody, corm-like rootstock. Introduced. A casual garden escape recorded from a few places in Great Britain. Native of eastern North America. It is not known to which variety our records belong. Subgenus 4. Galatella (Cass.) P. D. Sell Galatella Cass. Perennial or rarely annual herb. Stems erect. Leaves entire, glandular-punctate. Involucral bracts with outer green and inner with broad scarious margins, lanceolate to oblonglanceolate, more or less obtuse at apex. Ligulate flowers in 1 row. Pappus of many, unequal minutely denticulate hairs. 17. A. sedifolius L. Stonecrop Aster A. punctatus Waldst. & Kit.; Galatella punctata (Waldst. & Kit.) Nees Perennial, rarely annual aromatic herb. Stems 25–120 cm, erect, pale yellowish-green, with dense, minute glandular hairs, leafy, unbranched or branched above. Leaves 1– 5 × 0.2–1.0 cm, yellowish-green, hardly paler beneath, linear to lanceolate, obtuse at apex, entire, slightly narrowed to a sessile base, glandular-punctate, sometimes grey-felted beneath with a very short, scabrid margin, upper 1-veined, the lower 3-veined. Capitula 30–35 mm in diameter, in dense corymbs; peduncles short, with dense minute glandular hairs and sometimes sterile bracts. Involucral bracts in several rows, 3–6 × 0.8–1.5 mm, medium green, the inner with broad scarious margins, lanceolate to oblonglanceolate, more or less obtuse at apex, at least the outer minutely glandular-hairy. Flowers of 2 kinds, the outer up to 10, 12–14 mm, ligulate, the ligules violet-blue and shortly 3-lobed at apex, the inner 5–6 mm, tubular, greenishyellow, 5-lobed at apex. Receptacle flat, pitted, the pits with a dentate margin. Achenes 2.5–4.0 mm, oblanceolate, appressed hairy; pappus 5–6 mm, white, of many, unequal,

58. Aster

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minutely denticulate hairs. Flowers 8–9. Visited by bees, wasps and hoverflies. 2n = 36. Introduced. A garden escape persistent for a few years by a track near the Blackwater River, at Ash Vale in Surrey. Native of south and east Europe.

tipped and ovate and inner mainly scarious and oblongovate, rounded-obtuse at apex. Ligulate flowers in 1 row. Pappus of numerous, minutely denticulate hairs, elongating after flowering.

Subgenus 5. Crinitaria (Cass.) P. D. Sell Crinitaria Cass.; Linosyris Cass., non Ludw.; Pseudolinosyris Novopokr.; Crinita Moench, non Houtt. Perennial herbs with a woody stock. Stems decumbent to erect. Leaves entire. Involucral bracts green, at least the inner with a broad scarious margin, the outer linear and acute at apex, the inner linear-lanceolate and acute at apex. Flowers all tubular. Pappus of numerous minutely denticulate hairs.

19. A. tripolium L. Sea Aster Aster maritimus Salisb. nom. illegit.; Tripolium vulgare Besler ex Nees; Tripolion maritimum Raf. nom. illegit.; Eurybia maritima Gray nom. illegit. Annual or short-lived perennial herb with a short, swollen, suberect rhizome and fleshy roots. Stems 15–100 cm, dull medium green, often suffused brownish-purple below, erect or ascending, stout and fleshy, channelled, glabrous, leafy, branched at least above, in tall plants sometimes with flowering branches almost to base. Leaves dull, dark green with paler midrib, barely paler beneath, more or less fleshy, glabrous; basal 50–120 × 12–30 mm, lanceolate, elliptical, lanceolate-elliptical or linear, acute at apex, entire, cuneate or attenuate at base, with a pale green, sometimes tinted purplish; petiole up to 50 mm, which is channelled on the upper side and rounded beneath; cauline numerous, gradually decreasing in size upwards, narrowly elliptical, lanceolate, oblong or linear, acute at apex, entire, narrowed to a sessile base or the lower sometimes petiolate. Capitula few to numerous, 20–30 mm in diameter when ligules present, 13–15 mm when absent, forming a loose or compact corymbose panicle; peduncles medium green, glabrous, with occasional linear bracts. Involucral bracts in several rows, 2–10 × 1.2–2.5 mm, outer pale green at tip with a broad, scarious margin and ovate, the inner mainly scarious, oblong-ovate, rounded-obtuse at apex, glabrous. Flowers of 2 kinds or all tubular, the outer when present 18–23 mm, ligulate, the ligules pale to deep mauve or whitish, minutely 3-lobed at apex, the inner or all flowers 10–12 mm, tubular, yellow, deeply divided at the apex into 5 narrow lobes. Receptacle flat, pitted, the pits with crenate, membranous borders. Achenes 2–4 mm, pale brownish, oblanceolate, compressed, ribbed, appressed-hairy; pappus 12–15 mm, white, of many, minutely denticulate hairs, elongating after flowering. Flowers 7–10. Visited by many flies and bees; self-pollination possible. 2n = 18. The presence or absence of ligulate flowers is genetically controlled and particularly in south-east England large populations of entirely ligulate or eligulate plants may be found. However, the number and size of the ligules in a capitulum are variable.

18. A. linosyris (L.) Bernh. Goldilocks Aster Chrysocoma linosyris L.; Crinitaria linosyris (L.) Less.; Chrysocoma nupera Gray nom. illegit.; Linosyris vulgaris Less.; Erigeron linosyris (L.) Clairv.; Galatella linosyris (L.) Rchb. fil. Perennial herb with a woody stock. Stems 10–70 cm, pale yellowish-green, decumbent to erect, slender, wiry, striate, minutely scabridulous, very leafy, branched only in the inflorescence at the very top. Leaves all cauline, decreasing in size upwards only a little, (10–)20–50 × about 1 mm, dull medium green, hardly paler beneath, narrowly linear, acute at apex, entire, sessile, minutely scabridulous on the margin, punctulate on the upper surface, 1-veined. Capitula 12–18 mm in diameter, in dense, terminal corymbs; peduncles medium green, markedly striate, minutely scabridulous, with several linear bracts just below the capitulum. Involucral bracts in several rows, the outer 10–12 × 0.5– 0.7 mm, green, linear, acute or acuminate at apex and lax, the inner shorter, green with broad, scarious margins, linearlanceolate, acute at apex, erect, scabridulous or shortly hairy on the margin. Flowers all tubular, 7–8 mm, bright yellow, divided deeply at apex into 5 linear lobes. Receptacle flat or slightly convex, pitted, the pits with dentate membranous borders, without scales or hairs. Achenes 2.5–4.0 mm, brown, oblanceolate, with few ribs, appressed-hairy; pappus 5–6 mm, whitish, of many minutely denticulate hairs. Flowers 8–10. 2n = 18, 36. Very variable in size and branching. Many plants from Berry Head, Devonshire, for example, are short with few capitula and are referable to var. minor (Wallr.) Rouy (Linosyris vulgaris var. minor Wallr.) while the Great Ormes Head, Caernarvonshire plants are mostly tall and much branched. It is questionable whether these variants are worth recognising as their distribution and ecology are not known. Native. A rare plant of limestone cliffs. Devonshire, Somerset, Glamorganshire, Caernarvonshire, Pembrokeshire and Humphrey Head, Cumberland. Most of Europe northwards to south-east Sweden. A member of the European Temperate element. Subgenus 6. Tripolium (Nees) P. D. Sell Tripolium Nees; Tripolion Raf. Annual or short-lived perennial herb with a short, swollen, suberect rhizome. Stems erect or ascending. Leaves more or less fleshy, entire. Involucral bracts with outer green

(i) Var. condensatus Druce Stems up to 20(–30) cm. Leaves numerous, often forming a pseudorosette, very fleshy, lanceolate to elliptical. Inflorescence condensed in a tight corymb. Ligulate flowers present. (ii) Var. tripolium Often annual. Stems up to 100 cm. Leaves fairly distant, slightly fleshy, lanceolate or narrowly lanceolate-elliptical to linear. Inflorescence a loose corymbose panicle. Ligulate flowers present. (iii) Var. flosculosus (Gray) P. D. Sell Eurybia maritima var. flosculosa Gray; A. tripolium var. discoideus Rchb.

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Usually perennial. Stems up to 100 cm. Leaves fairly distant, slightly fleshy, lanceolate, narrowly lanceolateelliptical to linear. Inflorescence a loose corymbose panicle. Ligulate flowers absent. Native. Salt-marshes, where it can be dominant over large areas, and maritime cliffs and rocks. All round the coasts of Great Britain and Ireland, but almost confined to estuaries in northern England and Scotland; rarely in saline areas inland. Most European coasts, but not in Iceland and the Faeroes; North Africa; Caspian Sea; Lake Baikal; inland saline areas in Europe and central Asia. A member of the Eurasian Temperate element. Our plant is subsp. tripolium which is the race of the coasts of north and west Europe. Var. condensatus is the plant of maritime cliffs and rocks. Var. tripolium occurs throughout the range of the subspecies, mainly in the upper parts of salt-marshes. Var. flosculosus is frequent from Lancashire and Yorkshire southwards with isolated localities elsewhere and is mainly in the lower part of salt-marshes. 59. Chrysocoma L. Small shrubs. Branches erect or ascending, thin, leafy. Leaves alternate, linear. Capitula solitary at the ends of branches. Involucral bracts in 2–3 rows, herbaceous with scarious margins. Flowers all tubular, 5-lobed at apex and bisexual. Corolla yellow. Anthers without tails and apical appendage ovate-lanceolate and flat. Style with oblonglinear branches with sterile appendages ventrally glabrous and dorsally hairy. Receptacle flat or convex, without scales. Achenes obovate in outline, compressed; pappus of 1 row of scabrid bristles. Twenty species in southern Africa. Bayer, E. (1981). Revision der Gattung Chrysocoma L. Mitt. Bot. Staats. M¨unchen 17: 259–292. 1. Leaves 0.6–2.2 mm wide, absent for 1–3 cm below the 1. coma-aurea capitula; capitula 10–15 mm in diameter 1. Leaves 0.2–0.6(–1.0) mm wide, present almost to the 2. tenuifolia capitula; capitula 4–10 mm in diameter

1. C. coma-aurea L. Shrub Goldilocks C. patula P. J. Bergius; C. coma-aurea var. patula (P. J. Bergius) DC.; C. aurea Salisb., non Bojer Small, ericoid shrub up to 50 cm. Bark brown, rough. Branches leafy but absent for 1–3 cm below the capitula, brown, erect or ascending, glabrous or slightly hairy when young. Leaves 3–17(–20) × 0.6–2.2 mm, alternate, medium green, linear, more or less obtuse at apex, entire, sessile, glabrous, not gland-dotted. Capitula 10–15 mm in diameter, campanulate, solitary at the ends of branches; peduncles long, slender, glabrous. Involucral bracts in 2–3 rows, 2–6 × 0.5–0.8 mm, narrowly green down the centre with broad, scarious margins, linear or linear-lanceolate, more or less acute at apex, glabrous, not gland-dotted. Flowers all tubular, bright yellow, 5-lobed at apex. Receptacle flat or convex, without scales. Achenes 2–3 mm, obovate in outline, compressed, sparsely hairy; pappus of 1 row of scabrid bristles. Flowers 7–9. 2n = 18.

Introduced. Grown in gardens and naturalised on walls, dunes and open ground. Porth Seal on St Martin’s and Pentle Bay on Tresco in the Isles of Scilly. Native of South Africa. 2. C. tenuifolia P. J. Bergius Fine-leaved Goldilocks C. microcephala DC.; Aster discoideus Sond. Small, twiggy, ericoid shrub up to 1 m. Bark pale brown, rough. Branches bare below, closely leafy towards the tips, pale brown, erect or ascending, very shortly appressedhairy. Leaves 1–15 × 0.2–0.6(–1.0) mm, alternate, medium yellowish-green, linear, obtuse at the recurved apex, entire, sessile, glabrous, but gland-dotted. Capitula 4–10 mm in diameter, campanulate, solitary at the ends of branches; peduncles long, slender, very shortly appressed-hairy. Involucral bracts in 2–3 rows, 2–4 × 0.8–1.0 mm, pale green, with broad scarious margins, ovate-lanceolate to lanceolate, obtuse to acute at apex, glabrous but glanddotted and with oil-sacs along the median line. Flowers all tubular, bright yellow, 5-lobed at apex. Receptacle flat or convex, without scales. Achenes 2.0–2.5 mm, obovate in outline, compressed, the margins thickened, the apical rim usually minutely toothed, sparsely hairy; pappus of 1 row of scabrid bristles. Flowers 7–9. Introduced. Fairly frequent wool alien. Scattered records in England. Native of South Africa. 60. Erigeron L. Annual to perennial herbs. Stems procumbent to erect, leafy. Leaves alternate, entire or dentate. Capitula in a lax corymb or elongate panicle. Involucral bracts in several rows, herbaceous with a more or less scarious margin. Inner flowers tubular, 5-lobed and bisexual, the peripheral flowers filiform and female, the outermost ligulate, 3-lobed and female. Corolla of tubular flowers yellow to whitish, of ligulate flowers whitish, pink. bluish, purple or mauve. Anthers without tails, apical appendage ovate-lanceolate and flat. Styles with oblong-linear branches, the sterile appendage ventrally glabrous and dorsally hairy and shorter than the stigmatic area. Receptacle flat, pitted, without scales. Achenes usually cylindrical and compressed; pappus of 1 row of hairs, sometimes with an outer row of very short hairs or scales, the ligulate flowers sometimes with only very short hairs or narrow scales. About 200 species in North and South America and Eurasia. Although they form natural groups, the separation of Erigeron and Conyza from Aster is not possible morphologically and many species are placed in their genus by the jizz of the plant. That there is a general agreement amongst authors as to which species belong to which genus is fascinating. Cronquist, A. (1947). Revision of the North American species of Erigeron, north of Mexico. Brittonia 6: 121–302. Frey, D. (1996). Genetic variability of the invasive Erigeron annuus in Europe. Bull. Geobot. Inst. ETH 1996: 79–81. Grime, J. P. et al. (1988). Comparative plant ecology. London. [E. acer.] Nesom, G. L. (1989a). The separation of Trimorpha (Compositae: Astereae) from Erigeron. Phytologia 67: 61–66.

60. Erigeron Nesom, G. L. (1989b). Infrageneric taxonomy of New World Erigeron (Compositae: Astereae). Phytologia 67: 67–93. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [E. borealis.] 1. 1. 2. 2. 3. 3. 4. 4. 5. 5. 6. 6. 7. 7. 8. 8.

8. acer Ligules not or scarcely exceeding the pappus Ligules more than 4 mm and clearly exceeding the pappus 2. 3. Capitula 30–50 mm in diameter 5. Capitula 10–25 mm in diameter 1. glaucus Succulent plant with sprawling stems 4. Non-succulent plant with erect stems Uppermost leaves lanceolate; involucral bracts more or 2(i). speciosus var. speciosus less hairy Uppermost leaves ovate; involucral bracts glabrous 2(ii). speciosus var. macranthus 5. karvinskianus Stems procumbent to ascending 6. Stems erect Stems rarely more than 20 cm; capitula usually solitary; 4. borealis ligules purple (rare alpine) Stems rarely less than 20 cm; capitula few to numerous; 7. ligules white to deep pink or bluish to rose-purple Leaves obviously clasping the stem; pappus of ligulate 3. philadelphicus and tubular flowers of long hairs only Leaves not or slightly clasping the stem; pappus of tubular 8. flowers of long hairs and an outer row of short scales Hairs on stem and peduncle spreading; stem leaves 6. annuus gradually decreasing in size upwards Hairs on stem and peduncle strigose; stem leaves abruptly 7. strigosus decreasing in size

Section Asteroideae Nutt. Erigeron section Pseudoerigeron Torr. & A. Gray; Erigeron section Stenactis Torr. & A. Gray; Asterigeron Rydb. Perennials with a thick root. Stems prostrate to erect. Capitula 30–50 mm in diameter. Flowers of 2 kinds. 1. E. glaucus Ker Gawl. Seaside Daisy Aster bonariensis Spreng.; Aster californicus Less.; Stenactis glauca (Ker Gawl.) Nees; Woodvillea calendulacea DC.; E. hispida Nutt.; E. maritimus Nutt. Caespitose perennial herb with a stout, thick stock and a tap-root. Stems 5–50 cm, pale green, stout, usually sprawling at the base and procumbent or ascending, very variable in hairiness from glabrous or nearly so to having dense, spreading simple eglandular hairs and sometimes glandular hairs. Leaves 2.5–15.0 × 1.0–5.0 cm, glaucous and rather succulent, glabrous to more or less hairy; basal large and conspicuous, obovate or broadly spathulate, usually rounded or obtuse or rarely acute at apex, usually rather distantly toothed above the middle, gradually narrowed at base to broad, winged petioles; upper smaller and narrower. Capitula 1–15, on long, leafy branches which sometimes arise near the base of the stem, 30–50 mm in diameter; peduncles clothed like stem. Involucral bracts in several rows, 10–12 × 0.7–1.5 mm, numerous, equal, yellowish-green, often with dark or thickened midrib, linear, acuminate at the sometimes lax apices, with dense, long, often viscid hairs. Flowers of 2 kinds; outer 9–15 mm, ligulate, the ligules bluish, mauve or white sometimes

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turning yellow when dry, 3-lobed at apex; inner mostly about 5 mm, tubular, yellow, with 5 short lobes at apex. Receptacle flat, pitted, without scales. Achenes 1.0–1.5 mm, pale brown, oblanceolate, from 2-veined and obscurely 4angled, to often 4-veined and 4-angled, or sometimes even 6-veined; pappus 8–9 mm, dingy or reddish, of 20–30 stiff hairs, sometimes with a short outer series. Flowers 5–8. 2n = 18. The leaf rosettes resemble those of Limonium vulgare. Introduced. Naturalised in rocky places and on cliffs and Cornish hedgebanks. Coastal localities especially in southern and western England; occasionally found as a casual inland. Native of the western coast of North America. 2. E. speciosus (Lindl.) DC. Garden Fleabane Stenactis speciosa Lindl. Perennial herb with long, thick rhizomes. Stems 20–40 cm, pale green, angled, channelled, erect, with numerous, short and medium, pale simple eglandular hairs, leafy, branched above. Leaves dull dark green, slightly paler beneath, with short, pale simple eglandular hairs on the margins; basal numerous, 2–5 × 1–2 cm, oblanceolate, oblong or elliptical, rounded at apex, entire, attenuate at base to a narrowly winged petiole 15–40 mm; cauline similar in size except for the uppermost, oblong, lanceolate or spathulate, obtuse to subacute at apex, entire, rounded at the semiamplexicaul base. Capitula 30–40 mm in diameter, scentless, solitary at the ends of branches; peduncles long, yellowish-green, channelled, with numerous to dense, very unequal simple eglandular hairs. Involucral bracts in several rows, 6–7 × 1.0–1.5 mm, medium green with reddish tips, linear-lanceolate, acute at apex, glabrous or with numerous, unequal simple eglandular hairs. Flowers of 2 kinds; outer 12–14 mm, ligulate, the ligules mauve, with 3 shallow lobes at apex; inner 5–6 mm, tubular, yellow, 5-lobed at apex. Receptacle flat, pitted, without scales. Achenes 1.0– 1.5 mm, pale brown, oblanceolate 2–4 veined, hairy; pappus 4–5 mm, white, of 2 rows of simple eglandular hairs. Flowers 6–8. (i) Var. speciosus E. salicinus Rydb. Uppermost leaves lanceolate. Involucral bracts more or less hairy. 2n = 18. (ii) Var. macranthus (Nutt.) Cronquist E. grandiflorus Nutt.; E. macranthus Nutt.; E. vreelandii Rydb.; E. eucephaloides Greene; E. leiophyllus Greene Uppermost leaves ovate. Involucral bracts glabrous. 2n = 18. Introduced. Both varieties are grown in gardens. The species is established on an old wall at Knole Park in Kent, on dunes between Ainsdale and Birkdale in Lancashire, and has been recorded as a casual in a few other places. Both varieties are native of western North America. Plants which might key out here and which have the leaves abruptly decreasing in size upwards are referable to E. formosissimus Greene which is also grown in gardens.

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Section Olygtrichium Nutt. Heterochaeta DC., non Besser ex Schult. & Schult. fil. Biennial or short-lived perennial herbs occasionally flowering in the first season. Stems erect. Capitula 15–25 mm in diameter, solitary or in a narrow, open panicle. Flowers of 2 kinds. 3. E. philadelphicus L. Robin’s Plantain E. purpureus Aiton; E. hyemalis Raf.; E. amplexicaulis Desf.; E. purpureus var. attenuatus Nutt.; Tessenia philadelphica (L.) Lunell; Tessenia philadelphica var. attenuata (Nutt.) Lunell; E. philadelphicus var. glaber Henry; E. pulchellus var. townsendii Peattie Biennial or short-lived perennial herb, occasionally annual or at least flowering in the first season. Stems (10–)20– 75 cm, yellowish-green, erect, nearly glabrous, with short, appressed hairs or with long, spreading simple eglandular hairs. Leaves 1.5–15(–30) × 1.5–3.0(–12) cm, yellowishgreen on upper surface, paler beneath, broadly oblanceolate or obovate, rounded at apex, coarsely crenate or lobed, narrowed at the base, the cauline more or less amplexicaul and less toothed, the basal petiolate, all with simple eglandular hairs. Inflorescence rather narrow and open; peduncles very slender, shortly appressed-hairy. Capitula 15–25 mm in diameter. Involucral bracts in several rows, 4–6 × 0.8– 1.0 mm, more or less equal, pale greenish or brownish with a brown midrib and broad, sometimes purplish hyaline margins, linear or linear-lanceolate, subobtuse at apex, nearly glabrous or with pale, appressed simple eglandular hairs on the midrib. Flowers of 2 kinds, the outer 5–10 mm, ligulate, the ligules white to deep pink or rose-purple and 3lobed at apex, the inner 2.5–3.2 mm, tubular and yellow with 5 narrow lobes at apex. Receptacle flat, pitted, without scales. Achenes 0.8–1.0 mm, pale brown, cylindrical, 2-veined, with a few simple eglandular hairs; pappus 2–3 mm, whitish, of 20–30, rather rigid simple eglandular hairs. Flowers 6–8. 2n = 18. Very variable in size and luxuriance. The largest plants have been called forma scatericola (Fern.) Cronquist (E. scatericola Fern.) and those that are small and slender forma angustatus Vict. & Rousseau (E. provancheri Vict. & J. Rousseau), but the variation is continuous. Introduced. Naturalised on walls and in rough ground. In a few scattered localities in Great Britain. Native of North America. Section Erigeron Perennial herbs with a short, creeping, woody stock. Stems erect. Capitula solitary, 15–22 mm in diameter. Flowers of 3 kinds. 4. E. borealis (Vierh.) Simmons Alpine Fleabane Trimorpha borealis Vierh.; E. alpinus auct.; Trimorpha alpina Gray; E. uniflorus auct.; Trimorpha uniflora auct. Perennial herb with a short, creeping, woody stock. Stems 7–20 cm, pale green, often suffused brownish-purple, markedly striate, erect, with short to long, numerous, pale simple eglandular hairs, unbranched, leafy except for uppermost part. Leaves medium yellowish-green on upper

surface, paler beneath, with numerous, short to long, pale, rather stiff simple eglandular hairs on both surfaces and the margins; basal and lower cauline 1–6 × 0.2–1.0 cm, narrowly elliptical, spathulate, oblanceolate, oblong or oblongoblanceolate, obtuse to acute at apex, entire, gradually narrowed at base to a winged petiole; upper cauline smaller, linear or oblong, obtuse or acute at apex, entire, narrowed, sessile, semiamplexicaul. Capitula solitary and terminal, 15–22 mm in diameter; peduncles with numerous to dense, medium to long, pale simple eglandular hairs. Involucral bracts in several rows, 6–10 × 0.5–0.8 mm, outer green with a pale margin, the inner reddish, outer narrowly linearlanceolate, acute at apex, the inner narrowly linear and acute at apex, with numerous, medium, white simple eglandular hairs. Flowers of 3 kinds, the outer 4–6 mm, ligulate and female, the ligules purple and spreading with 3 small lobes at apex, the median very slender, tubular, 5-lobed at apex and female, the inner tubular, broader, yellow and bisexual. Receptacle flat, pitted and without scales. Achenes 2.0–2.5 mm, yellowish, cylindrical, compressed, hairy; pappus 3.5– 4.0 mm, of slightly reddish, stiff simple eglandular hairs. Flowers 7–8. 2n = 18. Somewhat intermediate in characters between E. alpinus L. from the mountains of south and central Europe and E. neglectus A. Kern. (from which it is often scarcely distinguishable) from the eastern Alps. It would probably be better if these and several other allied taxa were treated as one species with several subspecies. Native. Mountain rock-ledges between 640 and 1100 m in Perthshire, Forfarshire and Aberdeenshire. Scandinavia, Finland, north-west Russia; Iceland; Greenland. A member of the Eurosiberian Arctic-montane element. Section Karvinskia G. L. Nesom Perennial herb with stems procumbent to ascending and woody below. Capitula in a lax leafy corymb at the ends of branches, 17–20 mm in diameter. Flowers of 2 kinds. 5. E. karvinskianus DC. Mexican Fleabane E. mucronatus DC.; Vittadinia triloba auct.; E. karvinskianus var. mucronatus (DC.) Aschers. Perennial herb. Stems 10–50 cm, woody below, procumbent to ascending, pale green, sometimes tinted purplish, glabrous or with a few simple eglandular hairs, striate. Leaves 10–35 × 1–8 mm, medium green, with more or less appressed hairs on both surfaces and ciliate on the margins, looking like a basal rosette at anthesis; the lower obovate or oblanceolate, obtuse at apex, usually 3-lobed, cuneate to a short petiole; the upper narrowly elliptical or linear-lanceolate, acute at apex, entire, cuneate at base, shortly petiolate or sessile; sometimes with very short axillary shoots, the leaves then appearing verticillate. Capitula 17–20 mm in diameter, towards the ends of the branches on sparsely hairy peduncles 3–10 cm, forming a lax, leafy corymb. Involucral bracts in several rows, 3–4 ×0.5–0.6 mm, green, with a narrow scarious margin and sometimes a brown centre, linear-lanceolate, narrowly acute at apex, with a few, appressed hairs. Flowers slightly scented, the inner tubular, yellow or greenish-yellow, 5-lobed at apex and bisexual, the outer in 2 rows and ligulate, the ligules

60. Erigeron 5–6 mm, white or lilac above and pink to pale purple beneath and 3-lobed at apex. Receptacle flat and pitted, without scales or hairs. Achenes about 1.5 mm, shining reddishbrown, cylindrical, compressed, with appressed hairs; pappus 2–3 mm, whitish, of 1 row of simple eglandular hairs intermixed with shorter ones. Flowers 6–8. 2n = 18, 32, 36, 54. Introduced. Garden escape well naturalised on walls, banks and stony ground. In scattered localities in southern England and Ireland north to Norfolk and Co. Armagh, especially in south-west England and in the Channel Islands where it has been known for over a hundred years. Native of Mexico. Named after Wilhelm Friedrich von Karwinsky (1780–1855). Section Phalacroloma (Cass.) Torr. & A. Gray Phalacroloma Cass.; Diplemium Raf. Annual to perennial herbs with thick, fibrous roots. Stems erect. Capitula 10–20 mm in diameter, in a large corymbose panicle. Flowers of 2 kinds. 6. E. annuus (L.) Pers. Tall Fleabane Aster annuus L.; E. heterophyllus Muhl. ex Willd.; Stenactis annuus (L.) Nees Annual to perennial herb with rather thick, fibrous roots. Stems 20–100(–150) cm, pale green, sometimes suffused purplish, slender to robust, erect, striate, with few to numerous, short to long, pale, spreading simple eglandular hairs, branched above, leafy. Leaves medium yellowish-green on upper surface, paler beneath and slightly glaucous, glabrous or with rather short, pale simple eglandular hairs on both surfaces; basal 2–10 × 1.7 cm, broadly ovate or subrotund, rounded-obtuse to subacute at apex, subentire to coarsely dentate, abruptly or sometimes gradually contracted to the long petiole up to 10 cm; cauline numerous, gradually decreasing in size upwards, broadly lanceolate to linear-lanceolate, more or less acute at apex, entire or dentate, narrowed to a semiamplexicaul base. Inflorescence a large corymbose panicle; peduncles short and very slender, with numerous, short, pale, spreading simple eglandular hairs. Capitula 10–20 mm in diameter. Involucral bracts in several rows, 3–5 × 0.8–1.0 mm, green, with a pale margin, narrowly linear-lanceolate, acute at apex, with short, pale simple eglandular hairs. Flowers of 2 kinds, the outer ligulate, the ligules 4–10 mm, 3-lobed at apex and white or occasionally bluish, the inner 2.0–2.8 mm, tubular, 5-lobed at apex and yellow. Receptacle flat, without scales. Achenes 1.4–1.5 mm, cylindrical, compressed, 2veined, hairy; pappus of 2 rows, the outer of slender scales, the inner of very fragile bristles which are absent in the achenes of the ligulate flowers. Flowers 5–7. Apomictic. 2n = 27. Introduced. A garden escape in waste places and a weed of newly sown grass. Scattered records in south-west England and a rare casual elsewhere. Native of the northern United States and adjacent Canada, introduced elsewhere in North America. Plants need to be certainly separated from the E. strigosus aggregate. An apomictic aggregate which needs dividing into species.

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7. E. strigosus Muhl. ex Willd. Strigose Fleabane E. annuus subsp. strigosus (Muhl. ex Willd.) Wagenitz Annual or biennial herb with fibrous roots. Stems 30–70 cm, pale green, sometimes suffused brownish-purple, rather slender, erect, striate, strigose-hairy to nearly glabrous, branched above, leafy. Leaves medium yellowish-green on upper surface, paler beneath, sometimes tinted reddish, with short, pale simple eglandular hairs to nearly glabrous; basal often deciduous, 2–16 × 0.5–2.5 cm, oblanceolate or elliptical, obtuse at apex, entire or dentate, attenuate to the petiole which is up to 15 cm; cauline rapidly decreasing in size upwards and often rather scanty, lanceolate to linear, acute at apex, entire or rarely dentate, sessile and sometimes semiamplexicaul. Inflorescence a very open panicle; peduncles long and slender, strigose-hairy. Capitula 12–15 mm in diameter. Involucral bracts in several rows, 2–5 × 0.8–1.0 mm, green with pale margins, narrowly linear-lanceolate, acute at apex, appressed-hairy. Flowers of 2 kinds, the outer ligulate, the ligules 4–6 mm, 3-lobed at apex and white or rarely pink or bluish, the inner 1.5–2.6 mm, tubular, 5-lobed at apex and yellow; sometimes all flowers tubular. Receptacle flat, pitted and without scales. Achenes 1.4–1.5 mm, cylindrical, compressed, 2-veined, hairy; pappus of 2 rows, the outer scale-like, the inner of fragile bristles. Flowers 5–7. Apomictic. 2n = 27. Introduced. Naturalised on rough ground on an airfield at Longmoor in Hampshire. Some of the records for E. annuus probably belong here. A weed over most of the United States and southern Canada. An apomictic aggregate which needs dividing into species. Section Trimorpha (Cass.) DC. Trimorpha Cass.; Erigeron subgenus Trimorpha (Cass.) Popov Annual or biennial herb with thick, fibrous roots. Stems erect or ascending. Capitula 12–18 mm in diameter, in a corymbose panicle. Flowers dimorphic. This section is sometimes separated as a distinct genus on account of its flowers (cf. Nesom, 1989a). 8. E. acer L. Blue Fleabane Trimorpha acris (L.) Gray; Trimorpha vulgaris Cass. nom. illegit. Annual or biennial herb with rather thick, fibrous roots. Stems 8–40 cm, pale green, usually suffused brownishpurple for much of its length, erect or ascending, with numerous to dense, short to long, white simple eglandular hairs, sometimes simple, usually branched above, sometimes with numerous branches from base, leafy. Leaves medium green, sometimes tinted reddish, with numerous to dense, short to long, pale simple eglandular hairs on both surfaces and the margins, those on the surface usually appressed; basal 2.0–7.5 × 0.5–2.0 cm, narrowly obovate to oblanceolate, rounded at apex, entire, narrowed below to short petioles; cauline linear-oblanceolate to linear, obtuse to acute at apex, entire. Inflorescence a corymbose panicle; peduncles long, very slender, bracteate, with numerous, short and medium, pale simple eglandular hairs. Capitula 12–18 mm in diameter. Involucral bracts in several subequal

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rows, 6–8 × 0.8–1.0 mm, green, the inner with pale margins, narrowly linear-lanceolate, finely acute at apex, with numerous to dense, short to medium, pale simple eglandular hairs. Flowers of 2 kinds, the outer ligulate and female, the ligules 2.5–4.5 mm, 3-lobed at apex and pink or purplishmauve, the median filiform, tubular, 5-lobed at apex, yellow and female, the inner broader, tubular, yellow and bisexual. Receptacle flat, pitted, without scales. Achenes 2–3 mm, yellowish, cylindrical, compressed, 2-veined, hairy; pappus 3–5 mm, reddish-white, of 1 row of simple hairs. Flowers 7–8. 2n = 18. Native. Dry, rather bare grassland, dunes, banks and walls. Locally frequent in England, Wales and the Channel Islands and rare in Ireland; scarce as an introduction in Scotland. Temperate regions of the northern hemisphere, but some infraspecific taxa may deserve specific rank. A member of the Circumpolar Boreo-temperate element. Our plant is subsp. acer which occurs throughout much of Europe. In 1905 E. S. Marshall collected a very distinct plant with small pale flowers and oblong leaves at 1,400 feet (425 m) on limestone cliffs above the Builg Burn in Glen Avon in Banffshire. 61. Conyza Less. nom. conserv. Conyzella Fabr.; Leptilon Raf.; Marsea Adans. Annual to rarely perennial herbs. Stems erect, leafy. Leaves alternate, entire to dentate or incise-dentate, usually narrow. Capitula small, numerous, in a many-branched panicle. Involucral bracts in several rows, herbaceous with a more or less scarious margin. Central flowers tubular, 4- or 5-lobed at apex and bisexual, peripheral flowers tubular, more numerous and female, the outermost flowers shortly ligulate, the ligules with 3 lobes and female or tubular and female. Corolla white, cream, lavender or pinkish. Anthers without tails, apical appendage ovatelanceolate and flat. Style with oblong-linear branches which have lanceolate to triangular sterile appendages ventrally glabrous and dorsally hairy. Receptacle flat, without scales. Achenes obovate-oblanceolate, compressed; pappus of 1 row of scabrid, often brown or purple-tinged hairs. About 60 species widely distributed in temperate and tropical regions, often as introduced weeds. C. blakei (Cabrera) Cabrera (Erigeron blakei Cabrera) and C. japonica (Thunb. ex Murray) Less. (Erigeron japonica Thunb. ex Murray) have been recorded as wool casuals. While I was working on the Asteraceae for the Flora it seemed to me that plants of the genus Conyza had become very common round Cambridge. I also became aware that they were extremely variable and that they did not fit into the existing taxa recorded for Great Britain and Ireland. This variation also appeared to have nothing to do with habitat, small almost unbranched plants growing side by side with huge robust plants, from deep soil in the Botanic Garden, uniform soil and uniform light in an open growth chamber and in cracks in pavements. In one uncultivated area in the Botanic Garden nine distinct taxa were all growing intermingled. None had been deliberately introduced into the Garden. They did not look like a hybrid swarm as exactly the same plant was found in widely different localities, but more like the apomictic Hieracia of the H.

murorum group which grow intermingled at Royston (but in Cambridgeshire). I thus took the seeds from eight different kinds and Pete Michna grew them for me in the Botanic Garden. They reproduced themselves exactly. I thus concluded they were either self-pollinated or apomictic. The following account is based on some 200 specimens in CGE mainly collected around Cambridge. More work needs to be done on their breeding mechanism. It is possible that C. canadensis, which is 2n = 18, may be different from C. sumatrensis, which is 2n = 54, as diploid flowering plants are not known to be apomictic. Most of the literature does not have detailed enough descriptions to allow me to understand what the author is talking about. It would be a long job to bring all the types together. Herbarium sheets are often inadequate, as many as six sheets being needed to show all the characters of a plant 1 m high. On the other hand I can walk round Cambridge naming most at a glance (P.D.S.). Burtt, B. L. (1948). On Erigeron bonariensis L. Kew Bull. 3: 369– 372. Crawley, M. (1995). The inexorable spread of Conyza sumatrensis. B.S.B.I. News 68: 37. Cronquist, A. (1943). The separation of Erigeron from Conyza. Bull. Torrey Bot. Club 70: 629–632. Gu´ed`es, M. & Jovet, P. (1975). Conyza albida Willd. ex Sprengel, the correct name for ‘C. altissima’ Naudin ex Debeaux (C. naudinii Bonnet). Taxon 24: 393–394. Healy, A. J. (1957). Contributions to a knowledge of the adventive flora of New Zealand, No. 5. Trans. Roy. Soc. New Zealand 84: 649–659. Jovet, P. & Vilmorin, R. de (1975). Conyza in Coste, H. Flore descriptive et illustr´ee de la France, 3´eme supplement 187–192, 308–311. Marshall, J. B. (1973). Conyza – taxa found in Britain. Watsonia 9: 372–373. Marshall, J. B. (1974). A note on Conyza sumatrensis (Retz.) E. Walker (C. floribunda Kunth). Watsonia 10: 166–167. McClintock, D. & Marshall, J. B. (1988). On Conyza sumatrensis (Retz.) E. Walker and certain hybrids in the genus. Watsonia 17: 172–173. Michael, P. W. (1977). Conyza. Proc. 6th Asian-Pacific Weed Science Soc. Conf. 1: 87–95. Mundell, T. (2001). Conyza bilbaoana is on its way to you. B.S.B.I. News 87: 62–65. Palmer, J. R. (1993). Conyza canadensis/C. sumatrensis intermediates. B.S.B.I. News 64: 22. Phillips, B. (1997). Conyza bilbaoana confirmed from Surrey. B.S.B.I. News 76: 60–61. R´emy, J. (1849). Conyza in Gay, C. Historia fisica y politica de Chile. Vol. 4. Botanica, pp. 69–77. Paris. Reynolds, S. (1997). Conyza bilbaoana also in Ireland. B.S.B.I. News 74: 44–46. Sennen, E. C. (1904). Note sur le Cirsium corbariense Sennan, sur le Conyza naudinii Bonnet et sur quelques hybrides. Bull. Soc. Bot. France 51: 425–427. Sennen, E. C. (1908). Plantes d’Espagne, ann´ees 1906 et 1907. Bull. Akad. Int. Geog. Bot. 18: 469–470. Sennen, E. C. (1912). Quelques formes nouvelles o`u peu connues. Bol. Soc. Aragon 11: 194. Sennen, E. C. (1929). Sobre des plantes nuevas en la flora Espa˜nola. Bol. Soc. Ib´er. 28: 66–67. Stanley, P. (1996). Conyza bilbaoana J. R´emy new to south Hampshire (v.c. 11) and to Britain. B.S.B.I. News 73: 47–49 and drawing.

61. Conyza Th´ebaud, C. & Abbott, R. J. (1995). Characterization of invasive Conyza species (Asteraceae) in Europe; quantitative trait and isozyme analysis. Amer. Jour. Bot. 82: 360–368. Webb, C. J. (1988). Conyza in Webb, C. J., Sykes, W. R. & Garnock-Jones, P. J. Flora of New Zealand, vol. 4, pp. 187–191. Christchurch, New Zealand. Wurzell, B. (1988). Conyza sumatrensis (Retz.) E. Walker established in England. Watsonia 17: 145–148. Wurzell, B. (1992). Consumer. B.S.B.I. News 62: 38–39. Wurzell, B. (1994). A history of Conyza in London. B.S.B.I. News 65: 34–38. 2. 1. Involucral bracts densely hairy 4. 1. Involucral bracts glabrous or with an occasional hair 2. Inflorescence of long, slender, rigidly erect, straight branches often from low down the stem; involucral 2. daveauiana bracts 3–4 × 0.4–0.5 mm 2. Inflorescence of shorter, stouter branches with a great tendency to be curved; involucral bracts 4–5 × 0.5–0.6 3. mm 3. Inflorescence starting about halfway up the stem, ellipsoid and fairly dense; involucral bracts suddenly 1. sumatrensis narrowed halfway to the top 3. Inflorescence of long, stout branches not forming any regular shape; involucral bracts gradually narrowed all 3. bonariensis the way to the tip 5. 4. Whole plant dull greyish-green in appearance 7. 4. Whole plant yellowish-green in appearance 5. Ultimate flowering branches short and wide; involucral 4. bilbaoana bracts obtuse at apex 5. Ultimate flowering branches long and narrow; involucral 6. bracts acute at apex 6. Bracts not protruding beyond the capitula 5(i). floribunda var. floribunda 6. Bracts long-linear protruding beyond the capitula 5(ii). floribunda var. linearifolia 7. Plant up to 40 cm, with a long narrow inflorescence not 6(i). canadensis var. simplex more than 3 cm wide 7. Plant up to 100 cm, with inflorescence more than 3 cm 8. wide, usually much more 8. Inflorescence broadest at the top (to 7 cm) and narrowed 6(ii). canadensis var. obovoidea below 8. Inflorescence very broad and often more or less 9. cylindrical 9. Lower leaves deeply incise-dentate 6(iv). canadensis var. incisa 10. 9. Leaves entire or shortly dentate 10. Inflorescence up to 10 cm wide and cylindrical 6(iii). canadensis var. canadensis 10. Inflorescence huge, up to 20 cm wide 6(v). canadensis var. robusta

1. C. sumatrensis (Retz.) E. Walker Guernsey Fleabane Erigeron sumatrensis Retz.; C. albida Willd. ex Spreng.; C. erigeroides DC. Annual herb with fibrous roots. Stem 20–200 cm, greyishgreen, erect, slender to robust, markedly ridged, with dense, short to long, pale, very unequal, more or less spreading simple eglandular hairs, branched from about halfway, the branches ascending, leafy. Leaves 2–9 × 0.3–2.0 cm,

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yellowish-green or greyish-green on both surfaces, linear to narrowly elliptical, more or less acute at apex, entire or with occasional teeth, the lower sometimes dentate, narrowed to the sessile or shortly petiolate base, with dense, short, ascending-subappressed stiff eglandular hairs and minute glandular hairs on both surfaces. Capitula numerous, 5–8 mm in diameter, usually in an ellipsoid, fairly dense panicle in the upper half of the stem; peduncles with dense, minute glandular and short eglandular, ascending hairs. Involucral bracts in several rows, 2–5 × 0.4–0.6 mm, pale yellowish-green, rarely minutely red- or purple-tipped, narrowly linear-lanceolate, long-acute at apex, with very short to medium, unequal, ascending-subappressed simple eglandular hairs and minute glandular hairs. Flowers of 2 kinds, the outer with very short ligules, which are 3-lobed at apex, the inner tubular, pale yellow or purplish-yellow and with 5 lobes at apex. Receptacle flat, pitted and without scales. Achenes about 1.0 mm, pale, obovate-lanceolate; pappus 3–4 mm, off-white, of 1 row of hairs. Flowers 7–9. 2n = 54. Introduced. Naturalised in waste and rough ground and in cracks in concrete, especially in protected sunny spots. Frequent in the London area, Channel Islands, and around Cambridge with scattered records elsewhere in southern England and in eastern Ireland. Native of South America. 2. C. daveauiana Sennen Small-headed Fleabane Annual herb with fibrous roots. Stems 50–300 cm, dull greyish-green, erect, much ridged, with numerous, medium to long, white simple eglandular hairs, much branched from the base upwards with long, straight ascending branches but not exceeding the main stem, leafy. Leaves 2.5–12.0 × 0.3– 2.5 cm, dull medium greyish-green on upper surface, paler beneath, the lower narrowly elliptical to oblong-elliptical, obtuse at apex, obtusely dentate and attenuate at base with winged petioles up to 2 cm, the median and upper linear, oblong or elliptical-oblong, gradually narrowed but obtuse at apex and attenuate at base; all with more or less appressed, short hairs on both surface and the margins. Capitula very numerous, 2–3 mm in diameter, often well over 100, in a long, wide, broad often obovoid, panicle; peduncles with dense, ascending-appressed simple eglandular hairs and some more lax ones and minute glandular hairs. Involucral bracts in several rows, 3.5–4.0 × 0.4–0.5 mm, brownish, with a pale margin, linear or narrowly linear-lanceolate, long-acute with a filamentous tip, with numerous to dense, pale, appressed and spreading simple eglandular hairs and minute glandular hairs. Flowers of 2 kinds, the several outer rows with cream, sometimes purple-tipped, shortly 3-lobed ligules slightly longer than the involucral bracts, the inner tubular and 5-lobed at apex. Receptacle flat, pitted and without scales. Achenes 1.0–1.5 mm, pale, obovateoblanceolate; pappus 3.0–3.5 mm, cream, of 1 row of hairs. Flowers 7–9. Introduced. Waste ground, gardens and cracks in concrete. Common around Cambridge and at Santon in Norfolk; probably mistaken elsewhere for C. sumatrensis from which it differs in its much larger inflorescence with branches from the base and much smaller capitula. Named after Jules Alexandre Daveau (1852–1929).

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3. C. bonariensis (L.) Cronquist Argentine Fleabane Erigeron bonariensis L.; Erigeron crispus Pourr.; Erigeron linifolius Willd.; C. ambigua DC.; Erigeron undulatus Moench; Erigeron ambiguus (DC.) Sch. Bip.; Conyzella linifolia (Willd.) Greene; Leptilon bonariense (L.) Small; Leptilon linifolium (Willd.) Small Annual herb with fibrous roots. Stem 20–60 cm, dull medium or greyish-green, erect, markedly ridged, with numerous, short to medium, pale, ascending-subappressed simple eglandular hairs and with some longer, more spreading hairs, branched from about halfway, leafy. Leaves 2– 7 × 0.1–0.4 cm, dull medium or greyish-green on upper surface, paler beneath, linear or linear-oblanceolate, more or less acute at apex, narrowed to a sessile base, with many, short, pale, ascending-subappressed eglandular hairs, longer on the midrib beneath, and minute glandular hairs. Capitula numerous, 7–11 mm in diameter, usually in a subcorymbose panicle; peduncles with dense, minute glandular and short eglandular, ascending hairs. Involucral bracts in several rows, 5–8 × 0.5–0.7 mm, medium green, often redtipped, narrowly linear-lanceolate, acute at apex, with dense, very short to medium, unequal, spreading simple eglandular hairs and minute glandular hairs. Flowers all tubular, with 5 lobes at apex. Receptacle flat, pitted, without scales. Achenes 1.0–1.5 mm, pale, obovate-oblanceolate; pappus 3–4 mm, cream, of one row of hairs. Flowers 7–9. 2n = 54. Introduced. Rather frequent casual in waste and cultivated ground, often as a wool alien. Naturalised in Middlesex since 1993 and scattered records in England, Scotland and the Channel Islands. Native of South America. 4. C. bilbaoana J. R´emy Bilbao’s Fleabane Annual to perennial herb with fibrous roots and sometimes developing a stout stock. Stems up to 150 cm, pale yellowish-green, sometimes tinted violet, stout, solid, markedly ridged, with numerous medium to long, pale, rather stiff, often bulbous-based, short and broad, simple eglandular hairs, with long, rigid branches almost to the base, the ultimate branches curved, leafy. Leaves in a basal rosette as well as numerous cauline, alternate; lamina 2– 8 × 0.2–3.0 cm, medium greyish-green on upper surface, paler beneath, oblanceolate or linear-lanceolate to linear, acute or obtuse at apex, entire to sparsely serrate, or the basal with pinnate lobes, narrowed to a sessile base, with short, appressed, pale simple eglandular hairs on the midrib beneath and on the margin, and sometimes an occasional one on the surface. Capitula very numerous, 3–4 mm in diameter, in a large, often ovoid panicle; branches and peduncles with short, ascending-appressed simple eglandular hairs and some minute glandular hairs. Involucral bracts in several rows 3–4 × 0.5–0.7 mm, pale yellowish-green with scarious margins, linear-lanceolate, obtuse at apex, glabrous or with an occasional simple eglandular hair. Flowers of 2 kinds, the outer with very short, pinkish-purple, oblique ligules with 3 lobes at apex, the inner tubular, yellow and usually with 5 lobes at apex. Receptacle flat, pitted, without scales. Achenes about 1.2–1.3 mm, pale, obovateoblanceolate, with few appressed hairs, pappus 2.0–2.5 mm, whitish, of 1 row of hairs. Flowers 8–10.

Introduced. Found in several places in waste ground in Southampton, Hampshire since 1992, on roadsides and commons in Surrey during the 1990s, on roadsides, car parks and port entrance in Kilkenny, waste ground at New Ross in Co. Wexford since 1992, a railway yard at Blennerville near Tralee in Co. Kerry since 1995 and several places in Dublin since 1996. It is native of South America and is naturalised in New Zealand. It was named after a young Chilean boy, Bilbao. 5. C. floribunda Humb., Bonpl. & Kunth Many-flowered Fleabane Annual or biennial herb with a strong tap-root and fibrous side-roots. Stems up to 150 cm, pale greyish-green, erect, robust, markedly ridged, with many, short and medium, spreading, often bulbous-based, pale simple eglandular hairs, often branched from the base, the branches long, straight and the ultimate branches long and narrow, very leafy. Leaves numerous, alternate; lamina 2–11 × 0.2– 2.0 cm, medium greyish-green on upper surface, paler beneath, narrowly elliptical-oblong, lanceolate or linear, acute at apex, entire or the lower shortly dentate, long attenuate at base, the lower petiolate the upper sessile, glabrous or with occasional hairs on the surfaces, with numerous short to medium simple eglandular hairs along the margins and petioles. Capitula very numerous 5–7 mm in diameter, forming a long panicle of many branches; branches and peduncles with a few to fairly numerous, ascendingappressed hairs. Involucral bracts in several rows, 3.5– 4.0 × 0.3–0.4 mm, pale green, with a scarious margin, linear-lanceolate, acute at apex, glabrous or with an occasional hair. Flowers of 2 kinds, the outer with very short, cream ligules with 3 lobes at apex, the inner tubular, yellow and with 5 lobes at apex. Receptacle flat, pitted, without scales. Achenes 1.3–1.4 mm, pale, narrowly obovate; pappus 2.5–3.0 mm, off-white, of 1 row of hairs. Flowers 7–11. (i) Var. floribunda Bracts not extending beyond the capitula so that the inflorescence seems to be a mass of flowers. (ii) Var. linearifolia P. D. Sell Bracts long and linear extending beyond the capitula so that the inflorescence appears to be a mixture of capitula and bracts. Introduced. Waste places, gardens and cracks in pavements. Both varieties are common in Cambridge and Histon. Specimens of the species have also been seen from Norfolk, Surrey, Somerset, Kent, Oxfordshire and Suffolk. Both varieties are probably widespread but have been recorded as other taxa. Native of South America and specimens seen from North America where it may be introduced. 6. C. canadensis (L.) Cronquist Canadian Fleabane Erigeron canadensis L.; Conysella canadensis (L.) Rupr.; Leptilon canadense (L.) Britton; Aster canadensis (L.) Krause; Marsea canadensis (L.) Badillo Annual herb with tap-root and fibrous side-roots. Stem 8– 100 cm, yellowish-green, erect, with few to dense, pale, unequal simple eglandular hairs, with only very short

62. Callistephus branches in a narrow inflorescence to long branches halfway to the base, very leafy. Leaves numerous, widely spaced to dense, alternate; lamina 3–12 × 0.1–2.0 cm, gradually decreasing in size upwards, yellowish-green on upper surface, paler beneath, the lower oblanceolate to oblongoblanceolate, the upper mostly linear, acute to obtuse at apex, entire, usually shallowly and remotely toothed, rarely with deep narrow teeth, narrowed at base, at least the lower with narrowly winged petioles, the upper sessile; all with short to medium, pale simple eglandular hairs on both surfaces and longer, forwardly directed ones on the midrib and margin, the midrib prominent beneath. Capitula few to very numerous, 3–5 mm in diameter, in narrow to very broad panicles on the upper half of the stem; peduncles with few to numerous, short, ascending, pale simple eglandular hairs. Involucral bracts in several series, 4–5 × 0.3–0.4 mm, pale yellowish-green, with a scarious margin, narrowly linear or oblong, acute at apex, glabrous or with a few, very short simple eglandular hairs. Flowers of 2 kinds; the outer in several rows, 3–4 mm, ligulate, the ligules whitish to pale lavender, narrow and erect with 3 lobes at apex, the inner tubular, pale yellow, and with 5 lobes at apex. Receptacle flat, pitted, without scales. Achenes about 1.5 mm, pale, obovateoblanceolate; pappus longer than achene, yellowish, of simple hairs. Flowers 7–11. Visited by small insects. 2n = 18. (i) Var. simplex P. D. Sell Stem up to 40 cm; the inflorescence long and narrow and not more than 3 cm wide. Leaves not more than 5 mm wide, linear and entire. (ii) Var. obovoidea P. D. Sell Stem up to 60 cm; the inflorescence obovoid, that is wide at the top to 7 cm and narrowed at the bottom. Leaves up to 7 mm wide, occasionally with 1–2 teeth. (iii) Var. canadensis Stem up to 90 cm; the inflorescence cylindrical often for half the length of the stem but not more than 9 cm wide. Leaves up to 1 cm wide, the lower often shallowly dentate. (iv) Var. incisa P. D. Sell Stem up to 100 cm; the inflorescence very large, up to 23 cm wide. Leaves up to 2 cm wide, the lower with long, narrow teeth, often cusped. (v) Var. robusta P. D. Sell Stem up to 100 cm; the inflorescence very large, up to 23 cm wide. Leaves up to 2 cm wide, subentire or shallowly dentate. Introduced. Naturalised in waste and rough ground, on walls, waysides, open sandy areas, dunes, heaths, gardens and cracks in pavements and brickwork. Common in southeast England and the Channel Islands, and rarer further north and west. Native of North America but widely naturalised elsewhere. All the varieties grow commonly in and around Cambridge, sometimes as solitary plants, sometimes as a small uniform colony and sometimes as a large mixed community. 61. × 60. × Conyzigeron Rauschert Conyza × Erigeron

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Marshall, E. S. (1907). A hybrid Erigeron. Jour. Bot. (London) 45: 164. Roper, I. M. (1911). Erigeron acer × canadense = E. × Hulsenii Kerner. Jour. Bot. (London) 49: 348.

1. × C. huelsenii (Vatke) Rauschert C. canadensis × E. acer Erigeron × huelsenii Vatke This hybrid is intermediate between the parents in capitulum size and hairiness, has pale mauve ligules and is sterile. It resembles Conyza bonariensis in habit but has wider leaves and longer hairs. Native. Occurs sporadically with the parents in disturbed sandy places in southern England. Also recorded from central and southern Europe. 62. Callistephus Cass. Annual herbs. Stems erect, leafy. Leaves alternate, ovate to spathulate, entire to deeply toothed or lobed. Capitula conspicuous, on long peduncles. Involucral bracts in several rows, the outer herbaceous and very leafy, the inner with a scarious margin. Inner flowers tubular, 5lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female, or all ligulate. Tubular corollas yellow, ligulate corollas white, blue, red or purple. Anthers without tails, apical appendage ovate-lanceolate and flat. Style with oblong-linear branches, the triangular-lanceolate, sterile appendages ventrally glabrous and dorsally hairy. Receptacle slightly convex, without scales. Achenes all similar, obovoid, compressed, with 2–4 ribs; pappus of an inner row of very long, scabrid, deciduous hairs and an outer short membranous crown. One species in China, but widely cultivated. 1. C. chinensis (L.) Nees Chinese Aster Aster chinensis L.; Callistemma chinensis (L.) Druce; C. hortensis Cass. nom. illegit. Annual herb with fibrous roots. Stems 10–40 cm, pale yellowish-green, sometimes tinted brownish-purple, erect, ridged, with few to fairly numerous, short to medium, white simple eglandular hairs, simple or branched, leafy. Leaves 1–10 × 0.6–3.5 mm; basal and lower mostly withered at flowering, medium yellowish-green on upper surface, paler beneath, ovate, obtuse at apex, coarsely serrate-dentate, shortly cuneate or rounded at base and petiolate; middle and upper ovate, obovate or narrow to broadly spathulate, toothed or entire, narrowed to a sessile base or short, winged petiole; all with scattered, short simple eglandular hairs on the lower surface and shortly ciliate. Capitula 25–100 mm in diameter, solitary on long peduncles. Involucral bracts dimorphic; the outer in 1–2 rows, 5–18 × 4–6 mm, medium yellowish-green, linear to spathulate, obtuse at apex and markedly ciliate, the inner equalling or shorter than outer, spathulate, and with a membranous margin or ciliate. Flowers of 2 kinds, the outer 10–30 mm, numerous, particularly in cultivars, ligulate, the ligules white to red, blue or purple with 3 lobes at apex; the inner tubular, with 5 lobes at apex and yellow, or ligulate in double-flowered variants. Receptacle slightly convex, without scales. Achenes all similar, 3.0–3.5 mm, obovoid, compressed, with 2–4 narrow ribs, with hairs and glands; pappus of 1 inner row of long,

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scabrid, deciduous hairs and an outer short, membranous crown. Flowers 7–9. 2n = 18, 36. Introduced. Garden throw-out on tips and waste ground. Scattered records in England. Native of China, and widely cultivated. 63. Olearia Moench nom. conserv. Shawia J. R. & G. Forst. nom. rejic.; Eurybia (Cass.) Cass. Evergreen shrubs or small trees. Leaves alternate or opposite, simple, entire or sharply toothed, whitish or brownish tomentose on lower side. Capitula solitary or numerous and more or less crowded in lateral or terminal panicles. Involucral bracts in several rows. Flowers all tubular with 5 lobes at apex or the outer ligulate with 3 lobes at apex. Corolla of tubular flowers yellow to reddish, of ligulate flowers white. Anthers with short tails and apical appendage ovatelanceolate and flat. Style with oblong-linear branches, with sterile appendages ventrally glabrous and dorsally hairy. Receptacle flat or convex, pitted, without scales. Achenes linear or oblong to narrowly oblanceolate, more or less compressed; pappus of 1–2 rows of hairs. About 130 species in Australia, Tasmania, New Zealand, New Guinea and Lord Howe Island. Allan, H. H. (1961). Flora of New Zealand. Wellington. Heads, M. (1998). Biodiversity in the New Zealand divavicating tree daisies: Olearea sect. nov. (Compositae). Bot. Jour. Linn. Soc. 127: 239–285. 1. 1. 2. 2. 3. 3. 4. 4. 5. 5. 6. 6. 7. 7.

2. Leaves alternate 7. Leaves opposite 2. nummulariifolia Capitula solitary 3. Capitula in corymbose panicles 1. paniculata Capitulum with a solitary flower 4. Capitulum with few to many flowers 5. Leaves entire 6. Leaves serrate-dentate Leaves 5–10 cm; capitula with 0–2 ligulate flowers 3. avicenniifolia Leaves 1.0–3.5 cm; capitula with 3–5 ligulate flowers 4. × hastii Leaves 2.5–4.0 cm wide, lateral veins at a narrow angle to 5. macrodonta the midrib Leaves 1–2 cm wide, lateral veins at a wide angle to the 6. ilicifolia midrib Leaves up to 6.5 cm, oblong, ovate-oblong or broadly 7. traversii ovate Leaves up to 1.5 cm, linear-obovate or spathulate 8. solandri

1. O. paniculata (J. R. & G. Forst.) Druce Akiraho Shawia paniculata J. R. & G. Forst.; Erybia forsteri Hook. fil.; O. forsteri (Hook. fil.) Hook fil. Evergreen shrub or tree forming an ovoid outline. Stem up to 3(–6) m, with pale greyish-brown deeply furrowed bark; branches angular, grooved; young twigs tomentose. Leaves alternate, 3–10 × 2–4 cm, coriaceous, shiny yellowish-green on upper surface, whitish or buffish beneath, elliptical-oblong to ovate-oblong, obtuse at apex, entire, flat to strongly undulate on margin, rounded or truncate at base, glabrous on upper surface, with thin, appressed, whitish or buffish tomentum beneath, with a short petiole.

Inflorescence terminal, corymbose, the capitula sessile and fascicled on slender branches of the corymb. Capitulum of only 1 flower. Involucral bracts in several rows, 2–3 × 1.5– 2.0 mm, lanceolate, obtuse at apex, the outer glabrous or nearly so, the inner glandular-hairy. Flowers solitary in each capitulum, tubular, 5-lobed at apex, whitish, bisexual. Receptacle flat, pitted, without scales. Achenes 2.0–3.5 mm, narrowly oblanceolate, hardly compressed, shortly hairy; pappus 3–6 mm, of subequal, scabrid hairs. Flowers 11– 12. 2n = c. 288. Introduced. Grown in Guernsey and Alderney as a hedging plant where it is a frequent relic and rarely self-sown. Native of New Zealand. 2. O. nummulariifolia (Hook. fil.) Hook. fil. Sticky Daisybush Eurybia nummulariifolia Hook. fil.; O. hillii Colenso Evergreen shrub. Stems up to 3 m; branches stiff; bark rough; twigs stout and scarred; young shoots often viscidglandular and clothed with whitish or yellowish stellate tomentum to almost glabrous. Leaves more or less alternate, 5–10 × 4–6 mm, bright green on upper surface, pale grey beneath, close-set, very coriaceous, obovate to subrotund, obtuse at apex, entire and revolute, cuneate or rounded at base, glabrous and reticulate on upper surface, with white, buff or yellowish tomentum beneath, with a very short petiole. Inflorescence of a solitary capitulum on short, bracteate peduncles. Capitulum 3–5 mm in diameter, with 5–12 flowers. Involucral bracts in several rows, 2–3 × 1.5–2.0 mm, ovate, obtuse at apex, tomentose to almost glabrous. Flowers of 2 kinds, the outer ligulate, the ligules 3-lobed at apex, rather broad and white, the inner tubular, 5-lobed at apex and yellow. Receptacle flat, pitted, without scales. Achenes 2.5–3.0 mm, linear, compressed, ribbed, glabrous or slightly hairy at tips; pappus 5–6 mm, of whitish hairs. Flowers 7–8. 2n = c. 216. Introduced. Garden escape high on the hills south of Carlingford Lough in Co. Louth, Ireland. Native of New Zealand. 3. O. avicenniifolia (Raoul) Hook. fil. Mangrove-leaved Daisybush Shawia avicenniifolia Raoul; Erybia avicenniifolia (Raoul) Hook. fil. Evergreen tree or shrub with a broad crown. Stems up to 6 m, erect; branches angular; twigs with dense white tomentum. Leaves alternate, 5–10 × 3–5 cm, dark green on upper surface, buff beneath, ovate, elliptical or oblong-lanceolate, obtuse at apex, entire, cuneate or rounded at base, glabrous on upper surface, with dense greyish to buff tomentum beneath; petiole up to 5 mm, tomentose. Inflorescence of numerous corymbs on long peduncles. Capitula 2–4 mm in diameter, with 2–4 flowers. Involucral bracts in several rows, 2–4 × 0.8–1.0 mm, pale brown with a darker tip, linear, oblong or linear-lanceolate, obtuse to acute at apex, hairy. Flowers 2–3, rarely 4, the ligulate flowers 1–2, sometimes wanting. Receptacle flat, pitted, without scales. Achenes 1.5–2.0 mm, linear, compressed, appressedhairy; pappus 4–5 mm, of whitish hairs. Flowers 7–8. 2n = 108.

63. Olearia Introduced. Grown in the Isles of Scilly where it is naturalised on the dunes. Native of New Zealand. 4. O. × haastii Hook. fil. Daisybush O. avicenniifolia (Raoul) Hook. fil. × O. moschata Hook. fil. Evergreen shrub. Stems up to 2(–3) m, pale brown; branches ridged, pale brown, with appressed indumentum; young twigs whitish-tomentose. Leaves alternate, 1.0–3.5 × 1–2 cm, medium green on upper surface, pale greenish-cream beneath, elliptical, ovate-oblong or ovate, rounded-obtuse at apex, entire, rounded or shortly cuneate at base, glabrous and reticulate-veined on upper surface, thickly tomentose beneath; petiole 2–7 mm, tomentose. Inflorescence corymbose, terminal; peduncles strongly ridged, tomentose. Capitula numerous, 10–12 mm in diameter. Involucral bracts in several rows, 2.5–3.0 × 1.5–2.0 mm, pale green, ovate to ovate-oblong, obtuse at apex, glabrous or minutely hairy. Flowers of 2 kinds; outer 3–5, 5–6 mm, ligulate, the ligules white and 3-lobed at apex; inner tubular, 5-lobed at apex and yellow. Receptacle flat, without scales. Achenes 2.5–3.0 mm, narrowly oblanceolate, hairy; pappus of unequal, scabrid hairs. Flowers 7–8. Introduced. Much grown in gardens and shrubberies and often well established, though rarely self-sown, on walls and in open ground. Scattered records in south-west and westcentral Great Britain. Native of New Zealand, where wild hybrids of this origin are rare, but it is commonly propagated vegetatively by gardeners and is the hardiest of the Olearias cultivated in Great Britain. Named after Sir Johann Franz Julius von Haast (1824–87). 5. O. macrodonta Baker New Zealand Holly Eurybia dentata var. oblongifolia Hook. fil.; O. dentata Hook. fil., non Moench Evergreen shrub or tree with a strong musky fragrance and a broad crown. Stems up to 6 m, erect; branches ascending; twigs slender, with closely appressed tomentum. Leaves alternate 5–10 × 2.5–4.0 cm, dark shining green on upper surface, whitish beneath, subobliquely ovate-oblong to oblong, acute to acuminate at apex, the margins undulate and closely, coarsely and sharply serrate-dentate, glabrous on upper surface when mature, appressed white tomentose beneath; lateral veins at a narrow angle to the midrib; petiole up to 8 mm. Inflorescence a large corymb at the ends of branches; peduncles slender. Capitula 7.5–8.5 mm in diameter, with 8–12 flowers. Involucral bracts few, 3–6 × 1.5– 2.0 mm, greenish with brown tips, linear-oblong, obtuse to subacute at apex, with short to medium simple eglandular hairs. Flowers of 2 kinds, the outer ligulate, the ligules short, white, 3-lobed at apex and narrow, the inner tubular, 5-lobed at apex and yellow. Receptacle flat, pitted, without scales. Achenes 2–3 mm, linear, compressed, grooved, hairy; pappus dull white to reddish, of unequal hairs. Flowers 7–8. Introduced. Grown in gardens and naturalised in hedges, scrub and on banks and rough ground mostly near the sea. Scattered records in western Great Britain north to Wigtownshire and in Ireland. Native of New Zealand.

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6. O. ilicifolia Hook. fil. Maori Holly Eurybia dentata var. linearifolia Hook. fil.; O. multibracteolata Colenso Evergreen shrub or small tree with a musky fragrance and a broad crown. Stems up to 5 m, erect; branches ascending; twigs tomentose only when young. Leaves alternate, 5–10 × 1–2 cm, rather dark, shining green on upper surface, paler or brownish beneath, stiff and coriaceous, linear-oblong to lanceolate, acute to acuminate at apex, the margins strongly undulate and sharply serrate-dentate, almost spinous, truncate at base, glabrous on upper surface when mature, thinly clothed with yellowish-white tomentum beneath, the main lateral veins at a very wide angle; petioles up to 20 mm. Inflorescence of large corymbs at the ends of branches; peduncles slender, white tomentose. Capitula 7.5–8.5 mm in diameter, with 10–15 flowers. Involucral bracts in several rows, 3.0–4.5 × 0.4–1.0 mm, rather lax and spreading, linear-oblong, obtuse to subacute at apex, the outer long-hairy on the backs, the inner ciliate at the tips. Flowers of 2 kinds, the outer ligulate, the ligules short, 3-lobed at apex and narrow, the inner tubular and 5-lobed at apex. Receptacle flat, pitted, without scales. Achenes 1.5–2.0 mm, linear-oblong, conspicuously ribbed, with slender, ascending hairs on the ribs; pappus up to 4 mm, of copious, very slender, white, unequal hairs. Flowers 4–8. 2n = 108. Introduced. Escaped from a garden and naturalised at Beachy Head in Sussex. Native of New Zealand. 7. O. traversii (F. Muell.) Hook. fil. Ake-ake Erybia traversii F. Muell. Evergreen shrub or small tree. Stems 5–10 m, erect, with pale brown, furrowed bark; branches ascending, tetragonous; twigs with appressed, silky tomentum. Leaves opposite, (1.5–)4.0–6.5 × (1.0–)2.5–3.0 cm, flat and coriaceous, green or greenish-grey on upper surface, whitish or brownish beneath, oblong, ovate-oblong or broadly ovate, acute and usually apiculate at apex, entire, rounded or truncate at base, slightly hairy becoming glabrous when mature on upper surface, appressed silky-hairy beneath, shortly petiolate. Inflorescence axillary, much-branched; peduncles silky-tomentose. Capitula about 6–7 mm in diameter, with 5–15 flowers. Involucral bracts few, 4–6 × 1.0–1.5 mm, linear-oblong, obtuse to subacute at apex. Flowers of 2 kinds, the outer female and with a minute tubular corolla with an oblique mouth, the inner bisexual, 5-lobed at apex, tubular and campanulate above. Receptacle flat and pitted, without scales. Achenes 2–3 mm, linear, compressed, striate, silky-hairy; pappus of unequal, brownish hairs in 1 row. Flowers 4–8. 2n =108. Introduced. Grown in hedges in Guernsey in the Channel Islands and the Isles of Scilly as a windbreak, but rarely flowers. Native of Chatham Island, New Zealand. Named after William Thomas Locke Travers (1819–1903). 8. O. solandri (Hook. fil.) Hook. fil. Coastal Daisybush Eurybia solandri Hook. fil.; Aster solandri (Hook. fil.) F. Muell. Evergreen shrub or small tree. Stems up to 4 m, erect; branches stout, spreading and angled; twigs often

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viscid-glandular and covered with pale yellowish hairs. Leaves of young plants opposite, up to 1.5 cm, linearobovate or spathulate, obtuse at apex, entire, narrowed at base, flat, white beneath and petiolate, of mature plants in opposite fascicles, 0.5–1.0 cm, linear-oblong, obtuse at apex, margins flat to slightly revolute, subcoriaceous, narrowed at base, glabrous on upper surface, with pale yellowish glandular tomentum beneath. Inflorescence of a solitary capitulum on short twigs. Capitulum 5–7 mm in diameter, with 8–20 flowers. Involucral bracts in 3–4 rows, 1.8–2.0 × 1.0–1.2 mm, bright reddish-brown, ovatelanceolate or ovate, obtuse or subacute at apex, hairy or viscid-glandular. Flowers strongly scented, of 2 kinds, the outer ligulate, the ligules white and 3-lobed at apex, the inner tubular, yellow and 5-lobed at apex. Receptacle flat, pitted, without scales. Achenes 1–2 mm, grey, oblong, compressed, deeply grooved, glabrous to hairy; pappus 5–6 mm, whitish with a pinkish tinge, of hairs. Flowers 8–10. Introduced. Garden escape persistent for at least ten years on Hayle Towans in Cornwall, and seedlings have been recorded from Penzance in Cornwall. Native of New Zealand. Named after Daniel Carlsson Solander (1736– 82). 64. Baccharis L. Dioecious, deciduous shrubs. Leaves alternate, simple, the larger toothed in distal half. Capitula more or less numerous, in loose, terminal, leafy panicles. Involucral bracts in several rows, herbaceous with scarious margins. Flowers all tubular, 5-toothed at apex, male or female. Corolla whitish, yellowish or greenish. Anthers not tailed, apical appendages ovate-lanceolate and flat. Style with oblonglinear branches, sterile appendages ventrally glabrous, dorsally hairy. Receptacle flat or convex, without scales. Achenes oblong, somewhat compressed; pappus of 1 row of plumose-tipped hairs, shorter in male plants. About 400 species in North and South America. Named after Bacchus the god of wine. 1. B. halimifolia L. Tree Groundsel Dioecious, deciduous shrub. Stems up to 4 m, numerous, erect, much branched, the branches more or less erect, striate-angled. Bark greyish-brown, rough, sometimes more or less sticky. Twigs pale brown, rough and scaling; young shoots pale green, glabrous. Leaves alternate, 2– 6 × 0.2–4.0 cm, medium yellowish-green on upper surface, paler beneath, thick, linear, narrowly elliptical, lanceolate, oblanceolate or ovate, rounded-obtuse to more or less acute at apex, dentate, especially in the upper half on the larger leaves, the leaves near the inflorescence much smaller and entire, all cuneate to attenuate at base to a short, thin petiole and glabrous and punctulate. Inflorescence a terminal, loose, leafy panicle. Male capitula of numerous, bisexual flowers, the ovary abortive, the style branches sometimes fused. Female capitula 2.0–20.5 mm in diameter, with numerous female flowers. Involucral bracts in several rows, 3.5–6.0 × 0.5–0.7 mm, green with a scarious margin and the upper part often suffused purplish, lanceolate to ovate, obtuse to acute at apex, glabrous. Flowers all tubular,

5-lobed at apex, whitish, yellowish or greenish. Receptacle flat or convex, without scales. Achenes 1.0–1.7 mm, oblong, somewhat compressed, 8- to 10-ribbed; pappus 3–4 mm, of 1 row of plumose-tipped, whitish hairs, shorter in male plants. Flowers 9–10. 2n = 18. Introduced. Grown by the sea in southern England and naturalised in Hampshire since at least 1942. Coastal states of the United States from Massachusetts to Texas and inland to Oklahoma. 65. Bellis L. Perennial herbs. Stems leafless, erect or prostrate. Leaves in a basal rosette, simple, toothed. Capitula solitary. Involucral bracts in 2 rows, subequal, herbaceous. Flowers of 2 kinds; inner tubular, bisexual, 5-lobed at apex; outer ligulate, 3lobed at apex and female, or all flowers tubular. Corollas of tubular flowers yellow, of ligulate flowers white, sometimes suffused red. Receptacle conical to nearly flat, without scales. Achenes obovate, compressed, with thickened margins; pappus absent. About 15 species mainly in Europe and the Mediterranean region; western Asia. Grime, J. P. et al. (1988). Comparative plant ecology. London. Hull, R. (1961). Variation of populations of Bellis perennis L. Proc. B.S.B.I. 4: 269–272. Lacaita, C. C. (1923). The caulescence of Bellis perennis. Jour. Bot. (London) 61: 99–104. McClintock, D. (1984). Discoid daisies. Watsonia 15: 119. Oliver, J. (1996). Daisy ray florets – number of rows and gene flow over quarter of a century. B.S.B.I. News 71: 28–31. 1. Leaves 3–18 × 0.5–2.5 cm; 3-veined; capitula 27–40 mm 2. sylvestris in diameter; involucral bracts 7–12 mm. 1. Leaves 1–6 × 0.4–1.5 cm; 1-veined; capitula 14–30 mm 2. in diameter; involucral bracts 3–7 mm 1(1). perennis forma perennis 2. Ligules present 1(2). perennis forma disciformis 2. Ligules absent

1. B. perennis L. Daisy Perennial herb with a short, erect stock and stout, fibrous roots, with a single stem or forming large patches with numerous stems. Stems 3–12(–20) cm, without leaves or sometimes lengthening and with leaves, pale yellowishgreen, erect or ascending, with numerous, pale, short, appressed simple eglandular hairs. Leaves few to numerous in a basal rosette, spreading or ascending; lamina 1.0– 6.0 × 0.4–1.5 mm, bright yellowish-green on upper surface, only slightly paler beneath, oblanceolate to broadly oblanceolate-spathulate, rounded and often slightly retuse at apex, subentire to crenate-serrate, usually rather abruptly cuneate at base, with numerous, short, pale, rather appressed simple eglandular hairs on both surfaces at least when young, 1-veined; petioles 10–30 mm, pale green, winged, with scattered, short simple eglandular hairs. Capitula solitary, hemispherical, 14–30 mm in diameter. Involucral bracts in 2 rows, 3–6(–7) × 1.5–2.5 mm, green, sometimes with dark tips, oblong, rounded-obtuse at apex, with numerous, short to medium simple eglandular hairs. Flowers of 2 kinds; outer ligulate, in 1 row, female, rarely absent, the ligules 4–8(–11) × 0.7–1.2 mm, white, sometimes tipped

66. Tanacetum purplish-red, linear, 3-lobed and rounded at apex, glabrous; inner flowers 2.5–3.0 mm, tubular, 4- to 5-lobed at apex, bisexual, yellow. Receptacle conical, pitted. Achenes 1.0– 2.0 mm, obovate, strongly compressed, with a distinct border, with very short simple eglandular hairs, not ribbed; pappus absent. Flowers 3–10. Visited by many small insects. 2n = 18. (1) Forma perennis Outer flowers with ligules. (2) Forma disciformis P. D. Sell B. perennis var. discoidea Hus; B. perennis forma discoidea D. C. McClint., non var. discoidea Hus. Outer flowers without ligules. Native. Often abundant in grazed grassland, mowed areas and lawns, on cliffs and in waste land where there is not much competition, up to 915 m in Scotland. On lawns it tends to spread vegetatively in patches. Throughout great Britain and Ireland. Europe and western Asia; introduced in North America, New Zealand and elsewhere. A member of the European Temperate element. Forma disciformis is a rare plant in populations of forma perennis. × sylvestris This hybrid usually has large leaves and red-tinted flowers, but sometimes the leaves are small with the flowers red or the leaves large with the flowers white. Introduced. At Bassingbourn in Cambridgeshire, B. sylvestris was planted in a border. Some plants spread into the lawn and crossed with the B. perennis already there, all of which were white-flowered. Although B. sylvestris quickly disappeared, hybrids could still be seen in the lawn after 17 years. They are recorded from elsewhere and the flowers are often pink (cf. J. Oliver, 1996). 2. B. sylvestris Cyr. Garden Daisy B. monstrosa auct. Perennial herb with a short stock and stout, fibrous roots, with 1 or 2 stems. Stems 10–45 cm, without leaves, pale yellowish-green, erect, with numerous to dense, short to medium, stiff, white, ascending-appressed simple eglandular hairs, unbranched. Leaves numerous, in a basal rosette, spreading or ascending, (1.5–)3–18 × 0.5–2.5 cm, dark or medium green on upper surface, paler beneath, oblanceolate to obovate, rounded at apex, subentire to shallowly sinuatedentate, long attenuate at base, with numerous, short, stiff, pale, rather appressed hairs on both surfaces; 3-veined; petioles 10–60 mm, pale green, winged, with few, short hairs. Capitula solitary, 27–40 mm in diameter, hemispherical. Involucral bracts in 2 rows, (5–)7–12 × 2.5–3.0 mm, dark green, oblong-lanceolate, subacute at apex, with numerous, short, pale simple eglandular ahirs. Flowers of 2 kinds, the outer ligulate, the ligules 8–14 mm and white, usually tinged with deep pink on both surfaces of the upper half, the inner tubular and deep orange-yellow, or all ligulate. Receptacle conical, pitted. Achenes 1.5–2.5 mm, obovate, strongly compressed, with numerous, very short simple eglandular hairs, not ribbed; pappus absent. Flowers 3–10. Visited by small insects. 2n = 36.

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Introduced. Planted in gardens and sometimes escaping into waste ground and lawns where it crosses with B. perennis. Native of southern Europe. Tribe 6. Anthemideae Cass. Annual to perennial herbs, rarely shrubs. Leaves usually alternate, simple to pinnate, often finely and deeply divided. Capitula with all tubular or tubular and ligulate flowers usually with yellow tubular flowers and white ligulate flowers, but exceptions are not rare. Involucral bracts in 2–several rows, herbaceous with scarious margins and apex. Receptacular scales present or absent. Pappus usually absent, sometimes a low rim. 66. Tanacetum L. Strongly aromatic perennial herbs. Stems erect, leafy. Leaves alternate, simple and toothed to deeply pinnately lobed or pinnate. Capitula numerous, in terminal corymbs. Involucral bracts in several rows, herbaceous or scarious. Inner flowers tubular, bisexual, 5-lobed, the outer ligulate and female, or sometimes all tubular. Corolla of tubular flowers yellow, of ligulate flowers white. Anthers without tails, apical appendage ovate to ovate-oblong and flat. Styles with linear-oblong branches, stigmatic areas in 2 separate lines. Receptacle flat or convex, without scales. Achenes cylindrical, oblong-cylindrical or fusiform; pappus a very short rim (corona). About 50 species in temperate regions of Europe, Asia, North Africa and North America; some species widely cultivated. T. coccineum (Willd.) Grierson, the Garden Pyrethrum, has been recorded as a garden escape and T. leucophyllum Regel as a casual. 1. 1. 2. 2. 3. 3. 4. 4. 5. 5. 6. 6. 7. 7.

2. Leaves toothed, divided less than halfway to midrib 3. Leaves divided much more than halfway to midrib 4(i). balsamita var. balsamita Flowers all tubular 4(ii). balsamita var. balsamitoides Outer flowers ligulate Plant rhizomatous; leaf-lobes and teeth mostly acute or 4. acuminate Plant without rhizomes; leaf-lobes and teeth mostly 5. obtuse Capitula with obvious ligulate outer flowers 2. macrophyllum Capitula with all flowers tubular or rarely with outer 3. vulgare obscurely ligulate 1(4). parthenium forma flosculosum Flowers all tubular 6. At least some flowers ligulate 1(3). parthenium forma hortense All flowers ligulate 7. Outer flowers ligulate, inner flower tubular Ligules longer than involucre 1(1). parthenium forma parthenium Ligules shorter than involucre 1(2). parthenium forma breviradiatum

Section 1. Pyrethrum (Zinn) Rchb. fil. Pyrethrum Zinn; Gymnocline Cass. Leaves divided more than halfway to midrib. Ligulate flowers nearly always present, the ligules white. Achenes without epicarpic mucilaginous cells.

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1. T. parthenium (L.) Sch. Bip. Feverfew Matricaria parthenium L.; Chrysanthemum parthenium (L.) Bernh.; Pyrethrum parthenium (L.) Sm.; Matricaria vulgaris Gray nom. illegit. Pungent-aromatic perennial herb with a more or less vertical rootstock, fibrous roots and sometimes with a persistent woody base. Stems often several, 20–80 cm, pale yellowishgreen, sometimes suffused brownish-purple, erect, angled, ridged, usually with rather sparse minute hairs, branched above, leafy. Leaves 2–10 × 2–7 cm, gradually decreasing in size upwards, yellowish-green, often almost yellow, ovate to ovate-oblong in outline, acute or obtuse at apex, bipinnatisect, the segments 3–7, ovate or ovate-oblong and with a short petiolule, the lobes 2–6 mm, oblong, obtuse and unequal, entire or crenate, puberulous and with dense sessile glands; petioles up to 8 cm, pale yellowish-green, channelled, puberulous. Inflorescence a terminal corymb; peduncles pale yellowish-green, ridged and angled, sparsely hairy. Capitula numerous to abundant, 15–25 mm in diameter. Involucral bracts in 3 rows, 1.5–4.0 × 0.8–1.0 mm, yellowish-green with a darker, keeled centre, oblong, obtuse at apex, the innermost with a lacerate-membranous apex, sparsely arachnoid-hairy. Flowers of 2 kinds or 1 kind, the outer row and sometimes the 2 outer rows ligulate 2.5–7.0 mm, the ligules white and bluntly 3-lobed; the inner or sometimes all tubular, about 3 mm, yellow, with 5 recurved lobes. Receptacle convex, pitted, without scales. Achenes 1.2–1.5 mm, pale greyish-brown, fusiform, somewhat curved, prominently 5-ribbed, without simple hairs, glandular; pappus a minute, crenate or lobed corona. Flowers 6–9. Visited by bees and flies. 2n = 18. (1) Forma parthenium Outer flowers longer than involucre and ligulate, the inner tubular. (2) Forma breviradiatum Sch. Bip. Outer flowers shorter than involucre and ligulate, the inner tubular. (3) Forma hortense (Schur) Beck Flowers all ligulate.

Fingerh.; Pyrethrum macrophyllum (Waldst. & Kit.) Willd; Gymnocline leucocephala Cass. nom. illegit. Perennial aromatic herb with a rhizome. Stems 40–100 (–150) cm, pale yellowish-green, often suffused brownishpurple, erect, robust, markedly ridged, with numerous, short to medium, pale simple eglandular hairs and minute glandular hairs, branched above, leafy. Leaves all cauline at time of flowering, 5–50 × 5–20 cm, medium yellowish-green on upper surface, paler beneath, broadly elliptical in outline, obtuse to acute at apex, pinnately divided almost to the midrib, the segments 8–16 pairs, oblong or oblong-elliptical in outline and lobulate up to halfway to the midrib, the lobules entire to serrate and the teeth sometimes spinulose, with short, pale simple eglandular hairs and minute glandular hairs, the lower long petiolate, sometimes with small segments at intervals, the upper sessile. Inflorescence a dense, terminal compound corymb; peduncles short, ridged, with very short and short, unequal, pale simple eglandular hairs and minute glandular hairs. Capitula abundant, 7–13 mm in diameter. Involucral bracts in several rows, 2–5 × 0.1–1.2 mm, pale yellowish-green with a narrow brown margin and apex, linear to oblong, the outermost very narrow, the inner gradually broader, obtuse at apex, with short, pale simple eglandular hairs and minute glandular hairs. Flowers of 2 kinds, the outer 2–4 mm, ligulate, the ligules white, shallowly 3-lobed at apex, the inner tubular, 5-lobed at apex and yellow. Receptacle convex, pitted, without scales. Achenes 1.6–2.0 mm, pale brown, cylindrical, 5-ribbed; pappus a denticulate corona 0.2–0.3 mm. Flowers 7–10. 2n = 18. Introduced. Naturalised in grassy places and waysides. Scattered localities in England and Scotland. Known at Jervaulx Abbey in Yorkshire since 1912. Native of south-east Europe and south-west Asia. Section 2. Tanacetum Leaves divided more than halfway to midrib. Ligulate flowers always absent. Achenes without epicarpic, mucilaginous cells.

Bachelor’s Buttons

(4) Forma flosculosum (DC.) Beck T. parthenium forma discoideum Sch. Bip. nom. nud. Flowers all tubular. Introduced. Formerly cultivated as a medicinal herb and used as a febrifuge, hence its common name Feverfew. Still found in many, particularly older gardens and escaping and becoming naturalised on walls, waste places and waysides. Common throughout Great Britain and Ireland. Probably native in south-east Europe, Turkey and Caucasus, and widely established throughout the rest of Europe, in North and South America and many other temperate regions of the world. The forms do not appear to form populations anywhere, but when cultivated together seem to flower at different times. Forma hortense appears to be fertile. 2. T. macrophyllum (Waldst. & Kit.) Sch. Bip. Rayed Tansy Chrysanthemum macrophyllum Waldst. & Kit.; Gymnocline macrophylla (Waldst. & Kit.) Bluff &

3. T. vulgare L. Tansy Chrysanthemum vulgare (L.) Bernh., non (Lam.) Gaterau; Chrysanthemum tanacetum Karsch., non Vis.; Pyrethrum tanacetum (Karsch) Grecescu Aromatic perennial herb with a creeping, stoloniferous stock and wiry roots. Stems 1–several, 30–150 cm, erect, pale yellowish-green, usually suffused brownish-purple, ribbed and striate, making it angled, with a scattering of minute hairs and some arachnoid hairs especially near the base of the leaves, corymbosely branched above. Leaves all cauline at time of flowering, gradually decreasing in size upwards, the lower petiolate, the upper sessile and semi-amplexicaul, 2–25 × 1–10 cm, dull medium green on upper surface, paler beneath, often tinted purplish-brown on the rhachis, broadly elliptical to ovate-oblong in outline, pinnatipartite to pinnatisect; the rhachis ribbed and with acute, incised teeth distally or throughout, the segments up to 12 pairs, 5–40 × 1–10 mm, linear-lanceolate to oblong-elliptical, pinnatisect to pinnatilobed, the lobes linear to linear-lanceolate and acute at apex, glabrous to sparsely hairy and glandular-punctate. Inflorescence a dense

67. Artemisia compound corymb; peduncles 3–8 mm, pale green, glabrous or with a few minute hairs. Capitula (5–)10– 70(–100), 7–12 mm in diameter, vertically flattened and more or less button-shaped, the outer flowers tubular, female and zygomorphic or rarely very shortly ligulate or actinomorphic, the inner tubular and bisexual. Involucre 5–8 mm in diameter, hemispherical; bracts in 3 rows, 2–3 × 0.8– 1.2 mm, pale yellowish-green with a slightly darker keeled midrib and brown-tipped, ovate or elliptical, rounded at apex, glabrous or with a few minute or arachnoid hairs. Flowers yellow, the outer obliquely truncate and 3-lobed, the inner 5-lobed at apex. Receptacle convex, punctatetuberculate, without receptacular scales. Achenes 1.2–1.8 mm, greenish-white, cylindrical-obovoid, truncate at apex, 5-ribbed, with scattered epicarpic, sessile, transparent, nonmucilaginous glands; pappus 0.2–0.4 mm, an unevenly lobed and toothed membranous corona. Flowering 7–9. Freely visited by a great variety of small insects. 2n = 18. Our plants are fairly uniform, but in other parts of its range it shows much variation in leaf dissection and hairiness. Native. River and sandy banks, grassy places, waysides, hedgerows and waste places. Formerly much cultivated as a medicinal or pot-herb, and many of its localities may originally be garden escapes. Throughout Great Britain and Ireland, but not common. Europe; Caucasus; Armenia; Siberia; naturalised in North America. A member of the Eurasian Boreo-temperate element. Section 3. Balsamita (Mill.) Sch. Bip. Balsamita Mill. Leaves divided less than halfway to midrib. Ligulate flowers usually present, the ligules white. Achenes with epicarpic, mucilaginous cells. 4. T. balsamita L. Costmary Pyrethrum balsamita (L.) Willd.; Balsamita major Desf.; Chrysanthemum balsamita (L.) Baill., non L.; Pyrethrum major (Desf.) Tzvelev Perennial herb with a rhizome. Stems 30–130 cm, pale green, erect, stout, obscurely and paler ridged, sparsely appressed-hairy or nearly glabrous, simple or branched above, densely leafy. Leaves dull yellowish green on upper surface, paler beneath, appressed-hairy on both surfaces, closely gland-dotted, pleasantly aromatic; basal and lower cauline up to 150 × 5.5 cm, broadly oblong, obtuse or rounded at apex, crenate-dentate, cuneate at base, with slender, flattened channelled petioles up to 120 mm with a pair of disjunct, dentate pinnae near the base and often miniature leaves in the axils; cauline becoming very gradually smaller and more shortly petiolate upwards, the uppermost 3–4 × 1.0–1.5 cm and subsessile. Inflorescence a rather dense corymb; peduncles pale green, ridged, appressedhairy. Capitula 6–16 mm in diameter, campanulate, numerous. Involucral bracts in 4 rows, 1.5–3.5 × 0.5–1.0 mm, oblong, the outer herbaceous and obtuse or subacute at apex, the innermost with conspicuous, rounded laceratemembranous apices, glandular and thinly hairy. Flowers all tubular or rarely the outermost ligulate, the ligulate flowers 3.5–7.0 mm, the ligules white and 3-lobed at apex, the tubular flowers about 2.5 mm, 5-lobed, yellow, and thinly glandular. Receptacle flat, pitted, without scales. Achenes

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all similar, about 1.5 mm, pale brown, oblong-cylindrical, 5to 8-ribbed, with epicarpic, non-mucilaginous glands; pappus a short, irregularly toothed rim. Flowers 9–10 though it often fails to flower and rarely seeds. 2n = 18, 54. (i) Var. balsamita T. balsamita var. tanacetoides Boiss. Flowers all tubular. (ii) Var. balsamitoides (Sch. Bip.) P. D. Sell T. balsamitoides Sch. Bip.; T. balsamita subsp. balsamitoides (Sch. Bip.) Grierson Outer flowers ligulate, the ligules white, inner tubular and yellow. Introduced. Grown for ornament and for its mint-scented foliage for cooking. Rarely escapes and becomes naturalised. Scattered records in south and central Great Britain. Native of the Caucasus. The usual plant is var. balsamita. Var. balsamitoides is very rare. Both varieties grow in the same area in Anatolia. The spicy, aromatic leaves have been used in salads and in pot-pourri. A herb tea was made by infusing them in boiling water, and they were formerly used for flavouring ales. 67. Artemisia L. Seriphidium (Besser ex Hook.) Fourr.; Absinthium Hill Annual to perennial herbs or small shrubs, often aromatic. Stems more or less erect, leafy. Leaves alternate, entire to finely divided. Capitula small, usually pendulous, in racemose, paniculate or capitate inflorescences, rarely solitary. Involucral bracts in several rows. Flowers all tubular, 4–5lobed at apex, with the outer female and the inner bisexual, or all bisexual. Corolla usually yellowish, rarely reddish. Anthers without tails, apical appendage linear-lanceolate. Style with oblong-linear branches, stigmatic areas in 2 lines. Receptacle flat to hemispherical, hairy, without scales. Achenes obovate, oblong, ellipsoid or cylindrical, sometimes slightly compressed; pappus absent. About 520 species, predominantly in the northern hemisphere, but with a few species in South America, Africa south of the Sahara and in the Hawaiian Islands. To all intents and purposes species of the genus Seriphidium look and smell like species of Artemisia and differ only in having lost their outer female flowers; moreover they appear to be allied to different parts of the genus Artemisia. A. afra Jacq., A. anethifolia Weber, A. pontica L. and A. scoparia Waldst. & Kit. have been recorded as wool casuals and A. herba-alba Asso as an esparto casual. Blakelock, R. A. (1954). Artemisia norvegica Fries in Scotland. Kew Bull. 1954: 173–184. Brenan, J. P. M. (1950). Artemisia verlotorum Lamotte and its occurrence in Britain. Watsonia 1: 209– 223. (Gives large bibliography.) Cronk, Q. C. B. (1989). Some early notices of anthocyanin-rich variants of common British plants. Watsonia 17: 352–355. Grime, J. P. et al. (1988). Comparative plant ecology. London. [A. absinthium and A. vulgaris.] James, C. M., Wurzell, B. S. & Stace, C. A. (2000). A new hybrid between a European and Chinese species of Artemisia (Asteraceae). Watsonia 23: 139–147. Keck, D. D. (1946). A revision of the Artemisia vulgaris complex. Proc. Calif. Acad. Sci. 25: 421–468.

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Leach, S. J. (1984). Notes on the distribution of Artemisia maritima L. in eastern Scotland. Watsonia 15: 36–38. Ling Yeou-ruenn (1992). The old world Artemisia Linn. (Compositae). Bull. Bot. Res. Harbin 12: 1–108. Øvstedal, D. O. & Mjaavatten, O. (1992). A multivariate comparison between three NW European populations of Artemisia norvegica (Asteraceae) by means of chemometric and morphometric data. Pl. Syst. Ecol. 181: 21–32. Persson, K. (1974). Biosystematic studies in the Artemisia maritima complex in Europe. Opera Bot. 35: 1–188. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [A. campestris and A. norvegia.] 1. 1. 2. 2. 3. 3. 4. 4. 5. 5. 6. 6. 7. 7. 8. 8. 9.

9. 10. 10. 11.

11. 12.

12.

13. 13.

2. Most or all leaves entire 3. Most or all leaves deeply divided 4. ludoviciana Leaves more or less tomentose 10. dracunculus Leaves glabrous or nearly so 7. abrotanum Stems more or less woody to top 4. Stems not woody or only at base Stems less than 10 cm; capitula usually solitary, 6. norvegica subsp. scotica sometimes 2–3(–5) Stems more than 10 cm; capitula normally more than 10 5. Annual or biennial with a simple root system and 6. without non-flowering shoots Perennial with strong underground portion and 7. non-flowering shoots Leaves in inflorescence projecting laterally well beyond 8. biennis capitula Capitula projecting beyond leaves of inflorescence 9. annua 8. Mature leaves densely hairy on upper surface 11. Mature leaves glabrous or nearly so on upper surface Leaves grey appressed-hairy on both surfaces; 3. absinthium receptacle hairy Leaves whitish or greyish tomentose on both surfaces; 9. receptacle glabrous Ultimate leaf-lobes about 1 mm wide; all flowers bisexual; plants of salt-marsh and sea walls 12. maritima subsp. maritima Ultimate leaf-lobes wider; outer flowers filiform and 10. female; introduced plants of waste places and dunes 4. ludoviciana Involucre 2.5–4.0 mm 5. stelleriana Involucre 4–6 mm Plant scarcely aromatic; all leaf-lobes less than 2 mm wide; achenes usually only produced by marginal 11. campestris flowers Plant aromatic when fresh; all or most leaf-lobes more 12. than 2 mm wide; achenes produced by all flowers Plant strongly rhizomatous; pith occupying about one-third of total width of stem; flowers October to 2. verlotiorum December Plant not or scarcely rhizomatous; pith occupying four-fifths of total width of stem; flowers July to 13. September 1(i). vulgaris var. vulgaris Stem and branches pale green Stem and branches suffused brownish-purple 1(ii). vulgaris var. coarctata

1. A. vulgaris L. Mugwort Aromatic, more or less caespitose perennial herb with a branching rootstock, without overwintering rosettes. Stems 1–numerous, 60–210 cm, pale yellowish-green to com-

pletely suffused brownish-purple, erect, markedly longitudinally ridged, with large, white central pith occupying four-fifths of total width of stem and narrow, green peripheral tissues, arachnoid-hairy, usually much-branched, in large plants to the base, leafy. Leaves 2.5–8.0 × 2.0–5.0 cm, pale to dark green on upper surface, more or less white or rigid beneath, glabrous or with a few simple hairs on upper surface, densely, appressed, arachnoid-hairy beneath, ovate or elliptical in outline, acute at apex, pinnately divided, the terminal segments obovate and pinnatisect with the lobes 3–6 mm wide, lanceolate or oblong and sharply acute at apex, the lateral lobes similar but narrower and ascending, the area between the segments winged, only the main veins translucent, the basal petiolate and auricled, the cauline gradually decreasing in size with the upper sessile and amplexicaul. Capitula numerous, 2.0–2.5(–3.5) mm in diameter, narrowly campanulate to ovoid, in dense, sparsely leafy, racemose panicles with striate, nearly straight, erecto-patent branches; rhachis and peduncles densely arachnoid-hairy; bracts linear, more or less acute at apex, entire, arachnoid-hairy. Involucral bracts 3.5–4.0 × about 1.0 mm, pale green, the outer with broad scarious margins, lanceolate and acute at apex, the inner oblong and obtuse at apex. Flowers all tubular, reddishbrown, the outer filiform and female, the inner bisexual. Receptacle more or less flat, glabrous, without scales. Achenes produced by all flowers, about 1 mm, cylindrical, glabrous; pappus absent. Flowers 7–9. Wind-pollinated. 2n = 16. (i) Var. vulgaris Stem and branches pale green. (ii) Var. coarctata Forselles Stem and branches suffused brownish-purple. Native. Grassy and waste places, waysides and hedgerows. Common throughout Great Britain and Ireland except in the mountains. Most of the temperate regions of the northern hemisphere to 70◦ N in Norway and 74◦ N in Siberia, but this is an aggregate species. A. vulgaris sensu stricto is a member of the Eurosiberian Temperate element and is naturalised elsewhere. Closely allied taxa occur elsewhere. Var. coarctata is the variant most common with us (cf. Cronk, 1989). 2. A. verlotiorum Lamotte Chinese Mugwort Pleasantly and strongly aromatic, more or less caespitose perennial herb with long rhizomes, overwintering rosettes and sterile as well as fertile stems. Stems 1–numerous, 60–150 cm, pale green but often suffused brownishpurple, erect, markedly longitudinally ridged, with small, white central pith occupying one-third of total stem and narrow, green peripheral tissues, densely and persistently arachnoid-hairy and intermixed with minute glands, branched, very leafy. Leaves 2–18 × 2–18 cm, dark green on upper surface, pale greyish-green beneath, glabrous or nearly so on upper surface, appressed arachnoid-hairy and with a conspicuous network of small veins beneath, ovate or obovate in outline, acute at apex, pinnately divided, the terminal segment obovate and pinnatisect with the lobes 3–8 mm wide, linear or linear-lanceolate and sharply acute

67. Artemisia at apex, the lateral lobes similar and ascending, the area between the segments winged, the basal petiolate and auricled, the cauline gradually decreasing in size with the upper sessile and more or less amplexicaul. Capitula numerous, 2–5 mm in diameter, narrowly campanulate to ovoid, in very leafy panicles with arcuate-divaricate branches; rhachis and peduncles arachnoid-hairy; bracts linear, acute at apex, arachnoid-hairy, the capitulum sessile in the uppermost one. Involucral bracts in several rows, 1.2–4.0 × 0.5–2.0 mm, green, tinged purplish, ovate, acute at apex, arachnoidhairy. Flowers all tubular, the outer filiform, female and greenish, the inner bisexual, the male parts purplish or reddish. Receptacle flat, without scales. Achenes 1.0–1.5 mm, brown, obovoid, slightly compressed, more or less smooth; pappus absent or a small scarious ring. Achenes can be produced by all flowers, but not produced in Britain. Flowers 9–12. Wind-pollinated. 2n = 50, 52. Introduced. Rough ground, waste places and waysides. Frequent in the London area especially near the River Thames, scattered records elsewhere in south and central England and in Caernarvonshire and Guernsey. First recorded in 1908, part of a general spread in Europe, but spreading only slowly in England. Native of southwest China. Introduced in New Zealand. Named after Jean Baptiste Verlot (1815–91) and his brother. × vulgaris = A. × wurzellii C. M. James & Stace This hybrid is intermediate in pith characters and is sterile, the small capitula appearing from October to December but with abortive stamens. 2n = 34. Recorded from Essex, Surrey, Middlesex and Kent and should be looked for elsewhere. Endemic. 3. A. absinthium L. Wormwood Absinthium vulgare Lam. Strongly aromatic perennial herb. Stems 30–130 cm, pale green sometimes tinted purplish at the more or less woody base, markedly ridged and channelled, erect, grey appressed-hairy, leafy, branched in upper two-thirds. Leaves 2.5–6.0(–10.0) × 2.5–6.0 cm, pale greenish-grey on both surfaces, broadly ovate in outline, rounded at apex, the lower tripinnate, the median bipinnate and the uppermost pinnate or sometimes undivided, the lobes and segments punctate, linear or narrowly elliptical-linear, obtuse at apex and entire, densely or matted grey appressed-hairy on both surfaces. Capitula very numerous, 3–5 mm in diameter, campanulate to globose, slightly broader than long, forming a racemose inflorescence along each branch, the whole inflorescence narrowly lanceolate in outline; peduncles short, grey appressed-hairy, drooping. Involucral bracts in several rows, 2–3 × 0.8–1.0 mm, the outer green, oblong and obtuse at apex, the inner green with a wide scarious margin, ovate, obtuse at apex, grey appressed-hairy. Flowers 3–4 mm, all tubular, yellow, the outer filiform and female, the inner bisexual, all fertile. Receptacle flat, hairy, without scales. Achenes 1.4–1.6 mm, cylindrical, glabrous; pappus absent. Flowers 7–8. Wind-pollinated. 2n = 18. Native. Waste places, rough ground, and waysides. Frequent, but not common in England, Wales and the Channel Islands; scattered records in Scotland and Ireland. Temper-

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ate Europe and Asia northwards to Lapland, Karelia and southern Siberia; introduced in North and South America and New Zealand. A member of the Eurosiberian Temperate element. It is used for making a liqueur, absinthe, and for making vermouth wine. 4. A. ludoviciana Nutt. Western Mugwort A. brittonii Rydb.; A. diversifolia Rydb.; A. falcata Rydb.; A. gnaphalodes Nutt.; A. herriotii Rydb.; A. lindheimeriana Scheele; A. papularis (A. Nelson) Rydb.; A. purshiana Besser; A. serrata Nutt.; A. vulgaris subsp. ludoviciana (Nutt.) H. M. Hall & Clem.; A. candicans Rydb.; A. ludoviciana var. candicans (Rydb.) D. D. Keck.; A. platyphylla Rydb.; A. incompta Nutt. Aromatic perennial herb with a rhizome. Stems 30– 70(–100) cm, pale green, erect, striate, persistently whitetomentose to glabrescent, sometimes branched above, leafy, rather woody at base. Leaves 3–11 × 0.3–1.5 cm, greyish to whitish, linear, narrowly lanceolate or narrowly elliptical, more or less acute at apex, entire or the lower irregularly toothed or sometimes distinctly lobed, the lobes more than 2 mm wide, more or less tomentose, sessile, usually only gradually decreasing in size upwards. Capitula numerous, 2–4 mm in diameter, campanulate to globose, forming a compact but elongated inflorescence along the stem and branches; or much more open and diffuse with longer branches; peduncles short and white- or grey-tomentose. Involucral bracts in several rows, 2.5– 4.0 × 1.2–1.5 mm, ovate, rounded-obtuse at apex, greyishor whitish-tomentose. Flowers all tubular, the outer filiform and female, the inner bisexual. Receptacle flat, glabrous, without scales. Achenes 1.2–1.5 mm, ellipsoid-cylindrical, glabrous; pappus absent. Flowers 7–8. Wind-pollinated. 2n = 18. This is an aggregate of which many segregate species and infraspecific taxa have been named (cf. Keck, 1946). Divided leaved variants have been confused with A. stelleriana. Introduced. A casual garden escape or oil-seed alien in a few scattered localities in Great Britain. Native of North America. 5. A. stelleriana Besser Hoary Mugwort Non-aromatic perennial herb with thin, creeping rhizomes and sterile, leafy shoots. Stems usually solitary, 20–60 cm, erect or ascending, ridged, rather woody at base, densely white felted, leafy. Leaves 3–7 × 1.5–3.0 cm, densely whitefelted on both surfaces; lower and median ovate, obovate or broadly elliptical in outline, obtuse at apex, pinnate or pinnatifid, the segments more than 2 mm wide, linear, oblong or oblanceolate and entire at apex, sometimes lobed or bluntly toothed, more or less petiolate; upper pinnate, ternate or entire and sessile. Capitula numerous, 4–10 mm in diameter, cup-shaped, in a dense, spike-like panicle; peduncles erect or spreading, white-felted. Involucral bracts in several rows, 4–6 × 2.0–2.5 mm, brownish-membranous, lanceolate, obtuse at apex, white-felted. Flowers all tubular, the outer thread-like and female, the inner bisexual. Receptacle flat, glabrous, without scales. Achenes 3–4 mm, dark brown,

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narrowly linear-oblong, smooth; pappus absent. Flowers 7–9. Wind-pollinated. 2n = 18. Introduced. Naturalised on maritime dunes in Kirkcudbrightshire since 1979, Clyde Islands since 1976 and Ayrshire since 1994, formerly naturalised in Cornwall and Co. Dublin. Native of north-east Asia. Named after Georg Wilhelm Steller (1709–46). 6. A. norvegica Fr. Norwegian Mugwort Dwarf, tufted, aromatic perennial herb with a short rootstock. Stems 3–10 cm, usually flushed brownish-purple, with numerous to dense, medium to long, pale simple eglandular hairs intermixed with very short glandular hairs, unbranched, leafy. Leaves 1–2 × 0.5–1.0 cm, medium yellowish-green on upper surface, paler beneath, with few to numerous, long, pale simple eglandular hairs on both surfaces and the margins; basal ovate in outline, more or less acute at apex, bipinnately divided, the lobes linear and obtuse to acute at apex, petiolate; cauline smaller, less divided and sessile. Capitula 8–13 mm in diameter, usually solitary, sometimes 3(–5), nodding; peduncles clothed like stem. Involucral bracts 3.5–4.5 × 1.0–1.2 mm, pale green with a brownish-purple scarious margin and apex, oblong, with a rounded-obtuse apex, glabrous. Flowers all tubular, the outer thread-like and female, the inner bisexual, the corolla yellow and long-hairy. Receptacle flat, glabrous, without scales. Achenes oblong, compressed; pappus absent. Flowers 7–9. 2n = 18. Native. On rather bare mountain tops at about 700–870 m. In three sites in Ross-shire. Our plant is subsp. scotica (Hult´en) Å. & D. L¨ove (A. norvegica var. scotica Hult´en), which is endemic. Subsp. norvegica occurs in Norway and there is another race in the Urals. This species is a member of the European Arctic-montane element. The closely allied A. arctica Less. with several disjunct races occurs in Asia and North America (cf. Hult´en, 1954). 7. A. abrotanum L. Southernwood A. paniculatum Lam., non Besser; A. procera Willd.; A. herbacea Willd. Strongly aromatic shrub with an apple- or lemon-like scent. Stems up to 150 cm, pale green suffused brownish-purple, erect, striate, woody almost to top, glabrous, leafy, with numerous erect branches in the upper half. Leaves 3–7 × 1.5–4.0 cm, dull greyish-green on upper surface, paler beneath, lanceolate or ovate in outline, obtuse at apex, 1to 3-pinnatifid, the segments narrowly linear and obtuse, glabrous on upper surface, minutely greyish-hairy beneath; petiole slender and about one-third as long as the lamina. Capitula numerous, 3–4 mm in diameter, in the angles of simple, leaf-like bracts, forming narrow racemose inflorescences along the stem and branches; peduncles short, slender, down-curved, glabrous. Involucral bracts in several rows, 1.0–1.2 × 0.5–1.0 mm, green, the inner with scarious margins, ovate, obtuse at apex, glabrous. Flowers all tubular, yellowish, the outer filiform and female, the inner bisexual. Receptacle flat, glabrous, without scales. Achenes 1.5– 2.0 mm, brown, obovoid, glabrous; pappus absent. Flowers 9–10. 2n = 18. Introduced. A persistent garden escape on tips and waste ground. Scattered localities in south and central Great

Britain. Origin unknown, but widely cultivated for ornament and flavouring. A herb tea is made by infusing its leaves in boiling water. 8. A. biennis Willd. Slender Mugwort Non-aromatic annual or biennial herb with fibrous roots. Stems 30–150 cm, pale green, sometimes tinted brownishpurple, erect, ridged, glabrous or with a few hairs, with long, suberect branches, leafy. Leaves 5–15 × 2–4 cm, medium yellowish-green on upper surface, paler beneath, glabrous or slightly hairy, gland-dotted; lowest ovate in outline, acute at apex, 1- to 2-pinnatifid, the lobes linear and toothed, petiolate; median and upper gradually decreasing in size and sessile, but projecting laterally well beyond the capitula. Capitula 1.5–4.0 mm in diameter, spherical, in a series of spike-like clusters up the stem and branches; peduncles very short, glabrous or hairy. Involucral bracts 2–3 × 0.7– 1.0 mm, green with a broad, scarious margin, broadly ovate or subrotund, rounded at apex, glabrous. Flowers all tubular, the outer thread-like and female, the inner bisexual. Receptacle convex, glabrous, without scales. Achenes 1.0– 1.2 mm, ellipsoid, 4- to 5-ribbed, glabrous; pappus absent. Flowers 7–9. Wind-pollinated. 2n = 18. Introduced. Grain and wool casual on waste ground and mud by a reservoir, more or less naturalised in a few sites in southern England. Native of Asia and North America. 9. A. annua L. Annual Mugwort Aromatic annual herb with fibrous roots. Stems 30–200 cm, erect, pale green or flushed brownish-purple, ridged, glabrous or with sparse hairs, much-branched, leafy. Leaves 3–5 × 2–4 cm, yellowish-green on upper surface, paler beneath, glabrous or sparsely hairy, gland-dotted; lower ovate in outline, obtuse at apex, 2- to 3-pinnatisect, segments oblong, lobes oblong, teeth blunt; petiolate; median smaller and less divided; upper smaller and sessile; uppermost simple or with small lateral lobes less than 10 × 0.5–1.0 mm and not extending laterally as far as the capitula. Capitula 1.5–4.0 mm in diameter, spherical, in a wide, spreading panicle; peduncles short, glabrous. Involucral bracts 1.0– 1.5 × 0.3–0.5 mm, green with a membranous margin, ovate to oblong, obtuse at apex, glabrous. Flowers all tubular, the outer thread-like and female, the inner bisexual. Receptacle convex, glabrous, without scales. Achenes 0.6–0.8 mm, oblong-ovoid, compressed, obscurely ribbed, slightly bordered along margin, glabrous; pappus absent. Flowers 7–9. Wind-pollinated. 2n = 18. Introduced. Grain and wool casual on waste ground. A few, scattered records in southern England. Native of Russia. 10. A. dracunculus L. Tarragon A. glauca Pall. ex Willd.; A. inodora Willd.; A. redowskyi Ledeb.; Oligosporus condimentarius Cass. Aromatic perennial herb. Stems 60–200 cm, pale greenishyellow, faintly striate, erect, glabrous, leafy, much branched. Leaves dull, dark green, only slightly paler beneath, glabrous, 2–10 × 0.2–1.0 cm, linear, narrowly elliptical or lanceolate, the basal 3-fid at apex, the rest more or less acute, entire or weakly toothed, narrowed to a sessile base. Capitula numerous, 2–3 mm in diameter, globose, forming

68. Santolina a narrow, racemose inflorescence along the terminal and ascending branches; peduncles short, pale green, glabrous, curved downwards. Involucral bracts in several rows, 1.5– 3.0 × 1.3–1.5 mm, the outer greenish or brownish, oblongelliptical and obtuse at apex, the inner with a wide scarious margin, ovate and with a rounded apex, all glabrous. Flowers all tubular, yellow, the outer filiform and female, the inner functionally male. Receptacle flat, glabrous, without scales. Achenes about 2 mm, obovoid, glabrous; pappus absent. Flowers 9–10. 2n = 18, 36, 54, 72. Introduced. A persistent garden escape on tips and waste ground. Scattered localities in southern England. Native of south-east Europe through central Asia to North America. This is the French Tarragon whose green or dried leaves are used for seasoning fish sauces, pickles and salads and for making tarragon vinegar, which is an ingredient of sauce tartare and French mustard. Russian Tarragon is another species, A. dracunculoides Pursh.

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branched above, leafy. Leaves 2–5 × 1–2 cm, whitish or greyish, oblong to ovate in outline, obtuse at apex, mostly bipinnate, the ultimate segments linear, 0.5–1.0 mm wide and obtuse at apex, tomentose but not punctate, the lower petiolate and auricled at base, the upper sessile and sometimes entire. Capitula numerous, 1–2 mm in diameter, ovoid, erect or drooping, in a narrow, elongate panicle on stems and branches; peduncles short and woolly. Involucral bracts 1.2–4.0 × 0.4–2.0 mm, the outer greenish, the inner with a scarious margin, ovate or oblong, rounded at apex, woolly. Flowers all tubular, yellowish or reddish, all similar and bisexual, but the central ones sometimes sterile. Receptacle flat and glabrous, without scales. Achenes apparently very rarely produced, 1.8–2.5 mm, cylindrical; pappus absent. Flowers 8–9. Wind-pollinated. 2n = 54. All our plants are referable to subsp. maritima which usually has rather few, long, non-flowering rosettes and a broad panicle. Variants with the panicle branches short, erect and crowded have been called var. pseudogallica (Rouy) Ridd., Hedley & Price (A. maritima forma pseudogallica Rouy; A. gallica auct.; A. maritima var. gallica auct.) but numerous intermediates with var. maritima occur and it seems to have no ecological significance. Native. Dry parts of salt-marshes, sea walls and rough ground by the sea. Local on the coasts of Great Britain north to central Scotland, especially in eastern England and in east and west Ireland. The species occurs on the coasts of the Atlantic, Baltic and Mediterranean, calcareous substrate in western Switzerland and inland saline areas of Germany, Hungary, Caucasus, Armenia and across central Asia to Lake Baikal. Several subspecies are involved, some of which are sometimes regarded as species. Subsp. maritima is confined to the coasts of north-west Europe from western France to Denmark and southern Sweden, and is a member of the Suboceanic Temperate element.

11. A. campestris L. Field Wormwood Dracunculus marschallianus Ledeb. nom. illegit.; A. glabescens Stokes nom. illegit. Scarcely aromatic perennial herb with a branched, creeping, woody stock producing tufts of short, non-flowering shoots. Stems (10–)20–80 cm, pale green, often suffused brownishpurple, decumbent then ascending to erect, striate, more or less woody below, glabrous or nearly so, branched, the branches often long and suberect, leafy. Leaves 1–8 × 1–4 cm, greyish-green, oblong to broadly ovate in outline, acute at apex, the lower 2- to 3-pinnatisect, the middle 1- to 2-pinnatisect, the upper simple, the ultimate segments less than 2 mm wide, long-linear, obtuse to acute at apex and keeled beneath, sericeous when young becoming glabrous, the lower petiolate and auricled at base, the upper sessile. Capitula numerous, 3–4 mm in diameter, broadly ovoid, in a narrow elongated panicle along stem and branches; peduncles short, striate, glabrous. Involucral bracts 1.5–2.5 × 1.0–1.5 mm, greyish-green, sometimes tinged brownish-purple with broad scarious margins, ovate, rounded-obtuse at apex, glabrous. Flowers all tubular, yellow or reddish, the outer female and fertile, the inner bisexual, but mostly sterile. Receptacle flat, glabrous, without scales. Achenes usually only produced by marginal flowers, 1.5–1.8 mm, ellipsoid-cylindrical, glabrous; pappus absent. Flowers 8–9. Wind-pollinated. 2n = 16, 18, 36. Native. Grassy places by roads and on heathlands. Very rare in the Breckland of Norfolk and Suffolk, formerly in Cambridgeshire; naturalised at Crymlyn Burrows in Glamorganshire. Scattered through the northern hemisphere to 75◦ N in Novaya Zemyla, in a number of subspecies. A member of the Eurosiberian Temperate element. Our plant is subsp. campestris which occurs through much of Europe.

68. Santolina L. Strongly aromatic evergreen shrubs. Stems decumbent to more or less erect, leafy. Leaves alternate, closely pinnately lobed, whitish- to grey-tomentose. Capitula solitary on erect stems. Involucral bracts in several rows, gradually increasing in length inwards. Flowers all tubular, 5lobed at apex, usually all bisexual, the tube often compressed and more or less winged, spurred at base or saccate and enclosing the apex of the achene. Corolla yellow to whitish. Anthers without tails, apical appendage lanceolateoblong, flat. Style with linear-oblong branches, stigmatic area in 2 separate lines. Receptacle slightly hemispherical; with scales half surrounding the achenes. Achenes oblong, weakly 3- to 5-angled, glabrous; pappus absent. Eight species in southern Europe (mainly Spain) and North Africa (Morocco and Algeria).

12. A. maritima L. Sea Wormwood Seriphidium maritimum (L.) Poljakov Strongly aromatic perennial herb with a short, usually branching, vertical woody stock producing non-flowering rosettes and a fairly long, horizontal rhizome. Stems 5–60 cm, pale brown to greenish, decumbent to erect, slightly woody at base, striate, often grey- to whitish-tomentose,

1. Leaf-lobes not more than 2 mm, more or less tomentose; flowers deep yellow 1(a). chamaecyparissus subsp. chamaecyparissus 1. Leaf-lobes 2.5–7.0 mm, glabrous or nearly so; flowers whitish to pale yellow 1(b). chamaecyparissus subsp. tomentosa

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1. S. chamaecyparissus L. Lavender-cotton Strongly aromatic evergreen shrub. Stems 20–60(–100) cm, pale brown, woody, decumbent to more or less erect; young shoots numerous, slender and wiry, pale green, with dense woolly hairs, leafy. Leaves alternate, 2–6 × 0.3–0.6 cm, greyish-green or whitish, linear in outline, obtuse at apex, neatly and closely pinnately lobed, the lobes 1–7 mm, fleshy, linear and obtuse at apex, with dense woolly hairs on both surfaces or glabrous; petioles up to 6 mm, with dense woolly hairs. Capitula 8–15 mm in diameter, solitary at the ends of stems or branches; peduncles long, woolly-hairy. Involucral bracts in several rows, 5–6 × 1.2–1.5 mm, pale yellowishgreen with brown, scarious tips, linear-lanceolate, obtuse at apex, glabrous or with rather sparse to dense, woolly hairs. Flowers 4–5 mm, all tubular, 5-lobed at apex, usually all bisexual, whitish or yellow, the tubes compressed, winged, and with a basal, one-sided appendage enclosing the apex of the achene. Receptacle slightly hemispherical, hollow and pitted, with curved, oblong, obtuse scales half enclosing the achenes. Achenes about 2 mm, oblong, 3- to 5-angled, glabrous; pappus absent. Flowers 7–8. 2n = 18, 36. (a) Subsp. chamaecyparissus Leaves usually grey- to white-tomentose; lobes not more than 2 mm. Involucral bracts usually woolly hairy. Flowers deep yellow. (b) Subsp. tomentosa (Pers.) Arcang. S. tomentosa Pers. Leaves usually glabrous or slightly hairy; lobes 2.5–7.0 mm. Involucral bracts usually glabrous. Flowers whitish or pale yellow. Introduced. Much grown in gardens and persistent on tips, in rough ground and old gardens and rockeries. Scattered records in south and central England. Native of the Mediterranean region; introduced in North America. Subsp. tomentosa has been recorded from Leckwith in Glamorganshire and at least some of the other records belong to subsp. chamaecyparissus. 69. Otanthus Hoffmanns. & Link Diotis Desf., non Schreb. Shrubby perennial herbs. Stems erect to ascending, stout, woody. Leaves alternate, oblong to oblong-obovate, simple, entire to crenulate, white-tomentose. Capitula in 3- to 10-flowered corymbs. Involucral bracts in few rows, the outer densely hairy. Flowers all tubular, 5-lobed at apex and bisexual, the tube prolonged downwards into 2 spurs which almost enclose the ovary. Corolla yellow. Anthers without tails, apical appendage lanceolate-oblong. Style with linear-oblong branches, stigmatic areas in 2 separate lines. Receptacle convex, with scales. Achenes obovate, strongly compressed, partly enclosed by the thick, corky, persistent corolla; pappus absent. One species in south-west Europe and the Mediterranean region. 1. O. maritimus (L.) Hoffmanns. & Link Cottonweed Filago maritima L.; Athanasia maritima (L.) L.; Diotis maritima (L.) Desf. ex Cass.; Santolina maritima (L.)

Crantz; Gnaphalium maritimum (L.) Hill; Diotis candidissima Desf. nom. illegit. Shrubby perennial herb. Stems 15–50 cm, usually much branched, erect to ascending, stout, woody, densely whitetomentose. Leaves 5–20 × 4–5 mm, oblong to oblongobovate, subacute at apex, more or less entire or crenulate, sessile, fleshy, tomentose. Inflorescence of 3–10 capitula in terminal corymbs. Capitula 6–9 mm in diameter, globose; peduncles up to 15 mm, tomentose. Involucral bracts in few rows, 4–5 × 1.5–2.0 mm, ovate, obtuse at apex, the outer tomentose, the inner glabrous with a tomentose apex. Flowers about 3 mm, all tubular and bisexual, yellow, the tube strongly compressed and prolonged downwards into 2 spurs which almost enclose the ovary, sparsely glandular; the lobes about 0.5 × 0.4 mm, and ovate-deltoid. Receptacle convex; scales about 4 × 2 mm, oblong, subacute at apex, carinate, tomentose dorsally towards the apex. Achenes about 1 × 1 mm, obovate, strongly compressed, pale brown with a narrow cartilaginous margin, the apex of the achene partly enclosed in the accrescent, compressed, winged, spongy, brownish base of the corolla. Flowers 8– 10. 2n = 18. Native. Sandy seashores and stable shingle. Now known only in the south of Co. Wexford in Ireland. Formerly in a few localities in southern England and the Channel Islands. South and west Europe, including the Mediterranean islands, northwards to Ireland and eastwards to Greece and Turkey; North Africa; south-west Asia. A member of the Mediterranean-Atlantic element. 70. Achillea L. Aromatic perennial herbs. Stems erect, leafy. Leaves alternate, shallowly toothed to deeply pinnatifid. Capitula usually small, usually in corymbs, rarely solitary. Involucral bracts in many rows, with scarious margins. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3lobed at apex and female, rarely all flowers ligulate. Corolla white to deep pink or yellow. Anthers without tails, apical appendage lanceolate-oblong and flat. Style with oblonglinear branches, stigmatic areas in 2 separate lines. Receptacle flat or slightly convex, with narrow, scarious scales. Achenes oblanceolate, obovate or oblong, strongly compressed, truncate above; pappus absent. Contains 115 species mainly in the temperate regions of the Old World; a few species naturalised in North America and the Southern Hemisphere. 2. 1. Ligules yellow 4. 1. Ligules white or pink 2. Middle cauline leaves simple, incise-serrate to 6. ageratum serrate-dentate 3. 2. Middle cauline leaves deeply divided 3. Plant usually less than 1 m; leaves 0.4–0.7 cm wide, linear or oblong in outline; all involucral bracts rounded7. tomentosa obtuse at apex 3. Plant often over 1 m; leaves 3–7 cm wide, lanceolate to elliptical in outline, outer involucral bracts acute at apex 8. filipendulina 5. 4. Leaves simple

70. Achillea 4. 5. 5. 6. 6. 7. 7.

8.

8.

9. 9. 10. 10. 11. 11. 12. 12.

7. Leaves compound or deeply divided 1. × decolorans Leaves obviously serrate 6. Leaves entire or minutely serrulate Outer flowers ligulate, inner tubular 1(1). ptarmica forma ptarmica 1(2). ptarmica forma ligulosa Flowers all ligulate Middle cauline leaves elliptical to ovate in outline, usually flat and with not more than 10 pairs of segments 8. Middle cauline leaves lanceolate to oblong or linear in outline, the lobes and segments turning in various directions so they are 3-dimensional or flat and with 9. more than 15 pairs of segments Primary segments of middle cauline leaves elliptical in outline, regularly pinnatifid to pinnatisect; at least the 4. nobilis outer involucral bracts acute at apex Primary segments of middle cauline leaves lanceolate in outline, irregularly pinnatifid; involucral bracts all 5. ligustica obtuse at apex Leaves flat; involucral bracts 3–5 mm 2. distans subsp. tanacetifolia Leaves with segments curled and thus not flat; 10. involucral bracts 2–4 mm Stems up to 15 cm; leaves up to 6 cm; inflorescence 11. with few compact corymbs Stems up to 60(–80) cm; leaves up to 15 cm; 12. inflorescence with few to numerous corymbs 3(i). millefolium var. compacta Leaves sparingly hairy Leaves densely shaggy-hairy 3(ii). millefolium var. villosa Leaves with dense, imbricate segments, densely 3(iii). millefolium var. densiloba shaggy-hairy Leaves with rather open segments, sparingly or fairly hairy but never densely hairy 3(iv). millefolium var. millefolium

Section 1. Ptarmica (Mill.) W. D. J. Koch Ptarmica Mill. Leaves undivided, but sometimes serrate. Ligules white. 1. A. ptarmica L. Sneezewort Ptarmica vulgaris DC.; A. sylvestris Gray nom. illegit.; Ptarmica acuminata Ledeb. Perennial herb with a creeping, woody stock and fibrous roots. Stems 20–60 cm, pale yellowish-green, erect, angled, with short to medium, pale, wavy simple eglandular hairs and minute, pale glandular hairs in upper part, branched in the upper quarter, leafy. Leaves all cauline, numerous, 1.5–8.0 × 0.2–0.6 cm, dull medium yellowish-green on upper surface, paler beneath, linear, acute at apex, sharply serrulate, the teeth pale-tipped, the midrib prominent beneath, without obvious hairs, but minutely gland-dotted, sessile; bunches of smaller leaves often present in their axils. Capitula 12–15(–20) mm in diameter, in rather lax corymbs; peduncles rather short at the ends of long branches, more or less straight, erect, with dense short to medium, pale, wavy simple eglandular hairs and minute, pale glandular hairs. Involucral bracts in many rows, 3–5 × 1.0–1.5 mm, pale yellowish-green with a brown, scarious margin, linear-lanceolate to triangular-lanceolate, rounded-obtuse at apex, woolly with medium to long, pale, arachnoid hair and minute, pale glandular hairs. Flowers of 2 kinds, the

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outer 8–13, 7–9 mm, ligulate, the ligules white, broadly ovate to subrotund and with 3 broad, rounded lobes at the apex, the inner numerous, 5–6 mm, tubular, pale green below, yellow ridged above and whitish at the apex which has 5, triangular lobes at apex; in garden escapes sometimes all ligulate. Receptacle flat or slightly convex, not much elongating in fruit, with pale green, linear-lanceolate, acute scales. Achenes 1.5–2.0 mm, pale grey, oblanceolate, strongly compressed, truncate at apex, not ribbed; pappus absent. Flowers 7–8. Freely visited by bees and flies. 2n = 18. (1) Forma ptarmica Flowers outer ligulate, inner tubular. (2) Forma ligulosa Vilm. Flowers all ligulate.

Bachelor’s Buttons

Native. In damp meadows and marshy places and by streams up to 730 m in the Lake District. Frequent in Great Britain and most of Ireland; casual in the Channel Islands. Europe except the Mediterranean regions; south-western Asia and Siberia; introduced in North America. A member of the Eurasian Boreo-temperate element. The native plant is forma ptarmica. Forma ligulosa is grown in gardens and churchyards and and sometimes escapes into grassy and waste places. Infusions from the leaves and flowers have been used medicinally, and it was formerly used as a salad plant. × macrophylla L. = A. × decolorans Schrad. A. serrata Sm. Differs from A. ptarmica in having markedly serrate leaves. A hybrid of garden origin which occasionally escapes from cultivation. Section Achillea Section Millefolium (Adans.) W. D. J. Koch Millefolium Adans. Leaves much divided. Ligules white or yellow, sometimes pink. 2. A. distans Waldst. & Kit. ex Willd. Tall Yarrow Perennial herb with a deep root. Stems 30–130 cm, pale yellowish-green, erect, furrowed, with dense, long, pale, shaggy simple eglandular hairs and numerous, pale, minute glandular hairs, simple or branched above, leafy. Leaves 4–25 × 1–12 cm, medium green on upper surface, paler beneath, ovate, lanceolate or oblong in outline, more or less flat, deeply pinnatifid, the primary segments usually more than 15 pairs, oblong or lanceolate, acute at apex and serrate or pinnatifid, the rhachis dentate, the lobes lanceolate and serrate, with numerous to dense long, pale, shaggy simple eglandular hairs, the lower petiolate, the remainder sessile. Capitula 4–6 mm in diameter, in dense, terminal corymbs; peduncles short with dense, long simple eglandular hairs. Involucral bracts in many rows, 3–5 × 1.0–1.2 mm, pale with a narrow, brown margin and a prominent midrib, the outer lanceolate and acute, the inner ovate and obtuse, with long, pale simple eglandular hairs on surface and margin. Flowers of 2 kinds, the outer 4–5 mm, ligulate, the ligules white, and 3-lobed at apex, the inner 3.0–3.5 mm, tubular,

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cream and with 5 triangular teeth at apex. Receptacle more or less flat, with scarious, lanceolate scales which are jagged at apex and with some hairs. Achenes small, obovate, compressed, with a narrow wing; pappus absent. Flowers 7–9. 2n = 54. Introduced. Naturalised in grassy places. Derbyshire and Yorkshire. Native of central Europe. Our plant is subsp. tanacetifolia Janch. (A. tanacetifolia All., non Mill.). Tanacetum macrophyllum has been mistaken for it. 3. A. millefolium L. Yarrow Millefolium officinale Ledeb.; Millefolium vulgare Ledeb.; A. tenuifolia Salisb., non. Lam. Strongly scented perennial herb with a deep root and farcreeping rhizome. Stems 3–60(–80) cm, pale yellowishgreen, erect, furrowed, more or less woolly with long, pale, soft hairs, branched only in the inflorescence, leafy. Leaves 3–15 × 0.3–3.0 cm, medium green, oblong or oblonglanceolate in outline, obtuse in outline at apex, 2–3 times pinnate, the main segments broadly ovate in outline, the ultimate segments linear or linear-lanceolate, with a pale, apiculate apex, more or less softly hairy, the basal petiolate, the upper sessile. Capitula 4–7 mm in diameter, in dense, terminal corymbs; peduncles 2–5 mm, pale yellowish-green, more or less softly hairy. Involucral bracts in many rows, 2–4 × 1.2–1.4 mm, pale yellowish-green with brown or blackish scarious margins, the outer ovate, the inner oblong, obtuse at apex, sparingly softly hairy. Flowers of 2 kinds, the outer 4.5–6.5 mm, the tube pale green, the ligule white or sometimes pink or reddish, subrotund and 3-lobed at apex, the inner 2.5–3.0 mm, tubular, pale green below, white or cream above and with 5 triangular lobes at apex. Receptacle flat, elongating in fruit, with scarious, lanceolate, acute scales. Achenes 1.5–2.0 mm, shining greyish, obovate, strongly compressed, with a narrow wing; pappus absent. Flowers 6–9. Much visited by a great variety of insects. 2n = 54. (i) Var. compacta Br´eb. A. millefolium var. alpestris auct. Stems up to 15 cm, fairly densely hairy. Leaves up to 6 cm, with dense segments, sparingly hairy. Inflorescence with few corymbs. (ii) Var. villosa Hartm. A. millefolium var. lanata auct. Stems up to 15 cm, densely shaggy-hairy. Leaves up to 6 cm, with rather dense segments, densely shaggy-hairy. Inflorescence with few corymbs. (iii) Var. densiloba P. D. Sell Stems up to 60 cm, more or less hairy. Leaves up to 7 cm, densely shaggy-hairy, the segments dense and imbricate. Inflorescence with fairly numerous corymbs. (iv) Var. millefolium A. millefolium var. conspicua Druce Stems up to 60(–80) cm, sparingly hairy so that the stem looks green. Leaves up to 15 cm, with wide rather open segments, sparingly hairy. Inflorescence with numerous corymbs.

Native. Meadows, pastures, grassy banks, hedgerows, waysides, cliff-tops and sandy places by the sea and rocky places in mountains. Common throughout Great Britain and Ireland. Europe to 71◦ 10 N and in western Asia; introduced in North America, Australia and New Zealand. A member of the Eurasian Boreo-temperate element. All our plants are referable to subsp. millefolium. The common plant is var. millefolium. Var. compacta occurs in coastal areas in western Great Britain and in the mountains. It also occurs in France. Var. villosa is on the north coast of Scotland south to Argyll, the Hebrides, Shetland and the west coast of Ireland. It also occurs in the Baltic, and in the Faeroes and Iceland. Var. densiloba occurs in sandy areas on the East Anglian coast. A. millefolium has been used medicinally for a great variety of purposes from early times and contains two alkaloids, achillein and moschatinm, and an ethereal oil which is responsible for its aromatic scent. 4. A. nobilis L. Noble Yarrow Millefolium nobile (L.) Fourr. Perennial herb with a woody stock and fibrous roots, without stolons. Stems (10–)15–60 cm, pale yellowish-green, sometimes suffused reddish below, striate, with more or less numerous, short to medium, pale simple eglandular and numerous minute glandular hairs, unbranched, leafy. Leaves numerous, gradually decreasing in size upwards, 1.5–3.0 × 1.0–1.5 cm, medium green on upper surface, paler beneath, ovate in outline, more or less flat, pinnatisect, the rhachis toothed, the primary segments less than 10 pairs, (4–)6–8 mm, elliptical in outline and usually regularly pinnatifid to pinnatisect, the lobes linear or narrowly oblanceolate, and apiculate at apex, with more or less numerous, medium to long, pale, wavy simple eglandular hairs on both surfaces, all more or less sessile or the lower shortly petiolate. Capitula 4–6 mm in diameter, in dense, convex corymbs; peduncles short, with numerous, short to medium, pale simple eglandular hairs. Involucral bracts in many rows, 2–3 × 0.4–0.7 mm, pale, with a narrow brown margin and midrib, the outer narrowly lanceolate and acute at apex, the inner oblong and obtuse and jagged at apex, with numerous short to medium, pale simple eglandular hairs and pale minute glandular hairs. Flowers of 2 kinds; the outer 2.0–2.5 mm, ligulate, the ligules white, and with 3 lobes at apex, the inner 1.8–2.0 mm, tubular, cream and with 5, triangular lobes at apex. Receptacle more or less flat, with scarious, lanceolate, acute scales. Achenes very small, obovate, compressed; pappus absent. Flowers 7–9. Introduced. Grassy or waste places. Casual or persistent wool and grain alien, possibly also a garden escape. Native of south and central Europe and western Asia. As far as is known our plant is subsp. nobilis with white ligules and none of the subspecies with pale yellow ligules has been recorded. 5. A. ligustica All. Southern Yarrow Perennial herb with a woody stock and wiry roots. Stems (30–)50–100 cm, green, sometimes with reddish striations, erect, terete, with numerous, long, pale, wavy simple eglandular hairs and minute glandular hairs, unbranched, leafy. Leaves numerous, medium green on upper surface, paler

71. Chamaemelum beneath, with few to numerous, short to medium, pale simple eglandular hairs on both surfaces and densely glanddotted; lower and middle cauline 2–3(–4) × 1.5–2.0(–3.0) cm, ovate in outline, flat, 2-pinnatifid to 2-pinnatisect, with 3–5 pairs of segments, the primary segments lanceolate in outline and irregularly pinnatifid, the lobes linear and acute at apex; upper cauline gradually decreasing in size, more or less sessile. Capitula numerous, 5–7 mm in diameter, in dense, convex corymbs; peduncles short, with minute glandular hairs and occasional longer simple eglandular hairs. Involucral bracts in many rows, 2.5–3.5(–4.0) × 0.5– 0.7 mm, reddish, with a pale margin and midrib, oblong or oblong-lanceolate, obtuse at apex, ascending-appressedhairy. Flowers of 2 kinds, the outer 3.0–3.5 mm, ligulate, the ligules white, and 3-lobed at apex, the inner 2.0–2.5 mm, tubular, white and with 5 triangular lobes at apex. Receptacle more or less flat; scales lanceolate, cuspidate at apex. Achenes very small, obovate, compressed, winged; pappus absent. Flowers 6–8. Introduced. Naturalised at Newport Docks, Monmouthshire since 1953. Native of the Mediterranean region. 6. A. ageratum L. Sweet Yarrow Perennial herb, often woody at base. Stems 10–80 cm, pale yellowish-green, erect, striate, covered with very small, crisped and glandular hairs, simple or branched in upper part, leafy. Leaves 0.5–5.0 × 0.2–1.2 cm, oblanceolate to oblong or linear, rounded-obtuse at apex, the basal pinnatifid, the cauline incise-serrate to serrate-dentate or crenatedentate, the basal narrowed to a petiole, the upper sessile, glandular-punctate on both surfaces, sometimes also with pale simple eglandular hairs especially on the lower leaves. Capitula up to 15, 3.0–4.5 mm in diameter, in dense corymbs; peduncles and branches short, with dense, short glandular and crisped hairs. Involucral bracts in many rows, 1.5–3.0 × 0.7–1.0 mm, pale, with a scarious margin, ovate, more or less obtuse at apex, glandular. Flowers of 2 kinds, the outer 1.8–2.0 mm, ligulate, the ligules yellow and with 3 lobes at apex, the inner about 1.5 mm, tubular and with 5 triangular lobes at apex. Receptacle flat, with lanceolate, acute scales. Achenes 1.0–1.2 mm, obovate, compressed, winged; pappus absent. Flowers 7–8. 2n = 18. Introduced. Persistent garden escape or casual in a few places. Native of the west Mediterranean region. 7. A. tomentosa L. Yellow Milfoil Millefolium tomentosum (L.) Fourr. Perennial herb with a woody stock and wiry roots. Stems 10–50 cm, medium green, erect, terete, with long, white, shaggy simple eglandular hairs, unbranched, leafy. Leaves numerous, medium green on upper surface, paler beneath, with dense, long, pale, shaggy simple eglandular hairs on both surfaces; basal and lower cauline 2–8 × 0.4–0.7 cm, linear or oblong in outline, 2-pinnatisect, with 15–numerous pairs of segments, the lobes linear, apiculate at apex and entire; upper cauline similar, but gradually decreasing in size; lower shortly petiolate, upper sessile. Capitula (12–)15–many, 5–7 mm in diameter, in dense, convex corymbs; peduncles short, shaggy-hairy. Involucral bracts in many rows, 2.5–4.0 × 1.0–1.5 mm, very pale with a

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brown margin, oblong or oblanceolate, rounded-obtuse at apex, with numerous to dense, long, pale, shaggy simple eglandular hairs. Flowers of 2 kinds, the outer 3.0–3.5 mm, ligulate, the ligules bright yellow, with 3-lobes at the apex, the inner 2.5–3.0 mm, tubular, 5-lobed at apex and yellow. Receptacle more or less flat, with linear, acute scales. Achenes very small, brown, with a white wing. Flowers 7–9. 2n = 18. Introduced. Grown in gardens and a casual or persistent escape in a few places. Native of south-west Europe, extending eastwards to central Italy. 8. A. filipendulina Lam. Fern-leaf Yarrow Tanacetum angulatum Willd.; A. eupatorium M. Bieb.; A. filicifolia M. Bieb. Perennial, strongly aromatic herb, with a woody stock and long, thin rhizomes. Stems (80–)100–150 cm, medium green with brown striations, erect, terete but obtusely angled, solid, with numerous to dense, long, wispy, pale simple eglandular hairs and pale, minute glandular hairs, unbranched, leafy, but upper part leafless. Leaves numerous, medium green, paler beneath, with medium to long, appressed, pale, wispy simple eglandular hairs and pale, minute glandular hairs on both surfaces; basal and lower cauline 10–20 × 3–7 cm, lanceolate or elliptical in outline, pinnatipartite with 10–15 pairs of segments, the segments oblong to lanceolate, more or less obtuse at apex and dentate-serrate or lobed; upper cauline gradually decreasing in size and less divided; the basal with petioles up to 50 mm or sessile and the upper sessile. Capitula up to 300 or more, 2.0–3.5 mm in diameter, in dense, flat-topped corymbs 40– 100 mm in diameter; peduncles very short, pale green, with numerous pale simple eglandular and minute, pale glandular hairs. Involucral bracts in many rows, 2–5 × 0.5–0.7 mm, pale green, the outer with darker tips, linear-setaceous to linear-lanceolate and acute at apex, the inner lanceolate and subacute to obtuse at apex, all with sparse pale simple eglandular hairs and minute glandular hairs. Flowers of 2 kinds, the outer 2–4 mm, ligulate, the ligules golden yellow, and 3-lobed at apex, the inner 2.0–3.5 mm, tubular, 5-lobed at apex and golden yellow. Receptacle more or less flat; scales scarious with a green midrib, oblanceolate with an acute apex. Achenes 0.5–0.7 mm, oblong, compressed; pappus absent. Flowers 7–9. 2n = 18. Introduced. A common border plant in gardens and a persistent escape in a few places. Native of the Caucasus, Iran, Afghanistan and central Asia. 71. Chamaemelum Mill. Aromatic perennial herbs. Stems procumbent to ascending. Leaves alternate, deeply and finely dissected. Capitula solitary at the ends of branches, long-stalked. Involucral bracts in several rows, largely scarious or with a broad scarious margin. Inner flowers tubular, 5-lobed at apex and bisexual, the tube saccate at the base and enclosing the apex of the achene, the outer flowers ligulate and 3-lobed at apex and rarely all tubular or all ligulate. Corolla of inner flowers yellow, of outer white. Anthers without tails, apical appendages lanceolate-oblong, flat. Styles with linear-oblong branches, stigmatic areas in 2 separate lines. Receptacle hemispherical

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or conical, with scales. Achenes narrowly obovoid, covered with myxogenic cells in longitudinal rows; pappus absent or an inconspicuous membranous rim. Six species in the Mediterranean region, Canary Islands, North Africa, Middle East and west and south Europe. Stewart, A., Pearman, D. A. & Preston, C. D. (1994), Scarce plants in Britain. Peterborough. [C. nobile.] 2. mixtum 1. Cauline leaves 1-pinnatisect or serrate 2. 1. Caulines leaves 2- to 3-pinnatisect 2. Outer flowers ligulate, inner tubular 1(1). nobile forma nobile 3. 2. Flowers all ligulate or all tubular 1(2). nobile forma duplicatum 3. Flowers all ligulate 1(3). nobile forma nudum 3. Flowers all tubular

1. C. nobile (L.) All. Chamomile Anthemis nobilis L.; Ormenis nobilis (L.) J. Gay; Chamomilla nobilis (L.) Gren. & Godr. Perennial, sweet-scented herb with a short, much-branched stock. Stems 8–30(–40) cm, pale yellowish-green, often suffused brownish-purple, striate, prostrate and rooting, decumbent or ascending, more or less arachnoidhairy, branched, leafy. Leaves 1.0–5.0 × 0.5–1.5(–2.0) cm, greyish-green, oblong or oblanceolate in outline, acute at apex, 2- to 3-pinnatisect, the ultimate segments short, acute and linear-subulate, more or less arachnoid-hairy, sessile. Capitula 18–25 mm in diameter, solitary at the ends of branches; peduncles long, densely short-hairy. Involucral bracts in several rows, 5–6 × 1.2–1.5 mm, more or less scarious, broadly oblong, rounded-obtuse at apex, arachnoidhairy down the centre. Flowers usually of 2 kinds, the outer 7–12 mm, ligulate, the ligules white and 3-lobed at apex, the inner tubular 5-lobed at apex and yellow, rarely all tubular or all ligulate, the tubular flowers saccate at the base and enclosing the achene. Receptacle hemispherical to conical, solid with oblong, concave, obtuse scales often laciniate at apex. Achenes 1.0–1.5 mm, brown, narrowly obovoid, narrowed and rounded above, smooth except for 3 faint striae on the inner face; pappus absent or an inconspicuous membranous rim. Flowers 6–7. 2n = 18. Spreads vegetatively and sometimes by seed, but often non-flowering when heavily grazed. There is also a nonflowering form, cv. Treneague, which is used in lawns, walks, seats and banks, where cutting, rolling and treading encourage the plant to spread. (1) Forma nobile Flowers outer ligulate, inner tubular. (2) Forma duplicatum (Gray) P. D. Sell C. nobile var. duplicatum Gray Flowers all ligulate. (3) Forma nudum (Gray) P. D. Sell C. nobile var. nudum Gray Flowers all tubular. Native. Sandy commons, pastures and grassy waysides and saline gravel by the sea. Channel Islands, southern England, coastal Wales and south-west Ireland; formerly

in a few scattered localities elsewhere and probably mostly introduced. Has much declined in recent years, principally because of the cessation of grazing, it being unable to compete with taller herbs and scrub. It survives in the New Forest and Dartmoor because of grazing by ponies and cattle and on cliffs of the south-west because of climatic conditions. Western Europe from Belgium southwards; North Africa; Azores. A member of the Suboceanic Southerntemperate element. Forma duplicatum and forma nudum are rare. This species yields an essential oil used to flavour ice-cream, confectionary and alcoholic and non-alcoholic beverages. The flowering capitula used to be infused to make chamomile tea, but the species seems to have been replaced by Matricaria recutita, which has similar properties. 2. C. mixtum (L.) All. Mediterranean Chamomile Anthemis mixta L.; Ormenis mixta (L.) Dumort. Sweet-scented annual herb with a tap-root and fleshy, fibrous side-roots. Stems usually several, 8–30(–60) cm, pale green, often suffused brownish-purple especially towards the base, decumbent to erect, striate, arachnoidhairy, branched, leafy. Leaves 0.5–4.0 × 0.2–1.5 cm, greyish-green, oblong in outline, acute at apex, the lower 1to 2-pinnatisect, the lobes linear-lanceolate, entire or serrate and mucronate, the cauline 1-pinnatisect or serrate, arachnoid-hairy, sessile. Capitula 15–30 mm in diameter, solitary at the end of branches; peduncles long, more or less densely hairy. Involucral bracts in several rows, 3.8– 4.2 × 0.4–0.5 mm, pale green, with wide paler, scarious margins, oblong, obtuse at apex, arachnoid-hairy. Flowers of 2 kinds, the outer 8–10 mm, ligulate, the ligules white and shortly 3-lobed at apex, the inner tubular, 5-lobed at apex, yellow and saccate at the base and enclosing the achene. Receptacle hemispherical or conical, with oblong, concave, obtuse scales often laciniate at apex. Achenes 1.0– 1.5 mm, brown, narrowly obovoid, narrowed and rounded above, smooth except for 3 faint striae on the inner face; pappus an inconspicuous, membranous rim. Flowers 6–8. 2n = 18. Introduced. Bird-seed, grain, wool and granite casual. Native of the Mediterranean region and south-west Europe, north to west-central France. 72. Anthemis L. Aromatic annual to perennial herbs. Stems ascending, decumbent or erect, leafy. Leaves alternate, 1- to 3-pinnately divided, the ultimate segments usually linear. Capitula solitary, short- to long-stalked. Involucral bracts in several rows, usually some with a scarious margin, and often dark-margined. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female or neuter, sometimes all tubular. Corolla of inner flowers yellow, of outer flowers yellow or white. Anthers without tails, apical appendages lanceolate-oblong, flat. Style with linear-oblong branches; stigmatic areas in 2 separate lines. Receptacle hemispherical to conical, with scales subtending some or all of the flowers. Achenes not or somewhat compressed, rarely with myxogenic cells; pappus of a small, often oblique membranous rim or absent.

72. Anthemis About 210 species in Europe, Asia, North Africa and one species in east Africa. Fernandes, R. B. (1975). Taxonomic notes on the genus Anthemis L. Bot. Jour. Linn. Soc. 70: 6–17. Kay, Q. O. N. (1971). Anthemis arvensis L. in Biological flora of the British Isles. Jour. Ecol. 59: 637–648. Kay, Q. O. N. (1971). Anthemis cotula L. in Biological flora of the British Isles. Jour. Ecol. 59: 623–636. 1. Flowers all tubular 5(a,ii). tinctoria subsp. tinctoria var. discoidea 2. 1. Outer flowers ligulate 3. 2. Ligules yellow 6. 2. Ligules white 3. All or most involucral bracts with a distinct dark brown 5(d). tinctoria subsp. fussii or blackish margin and apex 3. All involucral bracts without a distinct dark brown 4. margin or the inner diffusely brownish at apex 4. Leaves green and sparsely hairy 5(a,i). tinctoria subsp. tinctoria var. tinctoria 5. 4. Leaves greyish-green and densely hairy 5. Leaves often more than 2.5 cm; ligules pale yellow 5(b). tinctoria subsp. subtinctoria 5. Leaves 1.5–2.5(–3.5) cm; ligules golden yellow 5(c). tinctoria subsp. australis 7. 6. At least the outer receptacular scales toothed at apex 8. 6. Receptacular scales not toothed at apex 7. Capitula 40–63 mm in diameter; achenes not or slightly 1. punctata subsp. cupaniana ribbed 7. Capitula 15–35 mm in diameter; achenes with rather 3. ruthenica thick, obtuse ribs 8. Receptacle with scales only in upper part, the scales 9. linear; achenes with ribs tuberculate 8. Receptacle with scales all over, the scales oblong or oblanceolate; achenes only striate or if ribbed the ribs 10. not tuberculate 9. Stems ascending or erect; leaves not fleshy 4(i). cotula var. cotula 4(ii). cotula var. vectensis 9. Stems prostrate; leaves fleshy 10. Achenes not compressed, 9- to 10-ribbed; receptacular 11. scales not becoming rigid 10. Achenes more or less compressed, 2–11 striate; 12. receptacular scales becoming rigid in fruit 11. Leaves not fleshy; ultimate divisions oblong lobes 2(i). arvensis var. arvensis 11. Leaves fleshy, ultimate divisions short teeth 2(ii). arvensis var. anglica 12. Peduncles clavate in fruit; achenes 7- to 11-striate on 6. altissima each side 12. Peduncles not clavate in fruit; achenes 2- to 3-striate on 7. austriaca each side

Section 1. Hiorthia (DC.) R. Fern. Anacyclus section Hiorthia DC.; Anthemis section Lyonnettia (Cass.) Griseb.; Anthemis section Rumata Fed.; Lyonnettia Cass. Perennial herbs. Involucral bracts with brown margins. Ligules white. Receptacle hemispherical-conical; scales persistent, dark brown at apex. Achenes obpyramidal, not or slightly ribbed, not tuberculate.

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1. A. punctata Vahl Sicilian Chamomile Perennial herb with a woody stock. Stems (20–)30–60 cm, pale yellowish-green, often suffused brownish-purple, erect, with numerous to dense, short to medium, appressed, pale simple eglandular hairs and minute glandular hairs, branched often nearly to base, the branches ascending to suberect, leafy. Leaves 1–12 × 0.5–5.5 cm, medium yellowish-green on upper surface, paler beneath, oblong to ovate in outline, obtuse or acute at apex, 1- to 2-pinnate, the segments linear to oblong and obtuse or acute at apex, sessile or shortly petiolate, with numerous, short to medium, pale simple eglandular hairs and glandular-punctate beneath. Capitula 40–63 mm in diameter, solitary at the end of stems and branches; peduncles long, without bracts, striate, with numerous to dense, short to medium, pale simple eglandular hairs and minute glandular hairs. Involucral bracts in several rows, 4–7 × 1–2 mm, green in centre with a narrow, brown margin, triangular-lanceolate to triangular-ovate, the outer obtuse the inner acute at apex, with numerous to dense, pale simple eglandular hairs. Flowers of 2 kinds, the outer ligulate, the ligules up to 24 mm, 3-lobed at apex and white, the inner tubular, 5-lobed at apex and yellow. Receptacle hemispherical-conical, with oblong scales that are 3-dentate and dark brown towards the apex. Achenes 2– 3 mm, obpyramidal, not or slightly ribbed, not tuberculate; pappus a corona up to 1.0 mm high. Flowers 6–7. 2n = 18. Introduced. Grown in gardens and naturalised in rough and marginal ground and on cliffs, mostly near the sea. Very scattered localities in southern Great Britain. Our plant is subsp. cupaniana (Tod. ex Nyman) R. Fern. (A. cupaniana Tod. ex Nyman; A. montana subsp. cupaniana (Tod. ex Nyman) Arcang.; A. punctata var. incana Guss.; A. punctata var. sicula Guss.) which is native of Sicily. Subsp. punctata occurs in north-west Africa. Section 2. Anthemis Anthemis section Erianthemis Griseb. Annual herbs. Involucral bracts with pale brown or scarious margins. Ligules white. Receptacle conical; scales subpersistent, not brown at apex. Achenes obovoid or subcylindrical, ribbed, not tuberculate. 2. A. arvensis L. Corn Chamomile Chamaemelum arvense (L.) Schreb. Aromatic annual herb with fibrous roots. Stems 10– 50(–80) cm, pale yellowish-green, sometimes brownishpurple towards the base, ascending or erect, striate, with numerous to dense, pale, curly simple eglandular hairs, branched, sometimes nearly to the base, leafy. Leaves 1.5– 5.0 × 0.5–2.0 cm, medium yellowish-green, oblong to obovate in outline, acute at apex, 1- to 3-pinnatipartite, the ultimate segments short, oblong and acute at apex, sessile or shortly petiolate, upper surface glabrous or nearly so, with numerous to dense, medium to long, often appressed simple eglandular hairs beneath. Capitula 20–40 mm in diameter, solitary at the ends of stems and branches; peduncles long and without bracts, striate, with numerous, short and medium, pale, subappressed simple eglandular hairs. Involucral bracts in several rows, 3.5–4.0 × 1.5–2.0 mm,

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with a green line down the midrib and a broad, pale brown scarious margin, oblong, rounded-obtuse at apex, with numerous to dense, pale simple eglandular hairs. Flowers of 2 kinds, the outer ligulate and female, the ligules white, 3-lobed at apex and spreading, the inner tubular, 5-lobed at apex and yellow. Receptacle conical, with oblanceolate, cuspidate scales. Achenes 1.2–3.0 mm, whitish, obovoid, about 9- to 10-ribbed, glabrous, the ribs not tuberculate, rugose on top; pappus a crenate border. Flowers 5–8(–10). Fragrant and much visited by bees and flies. 2n = 18. (i) Var. arvensis Leaves not fleshy, ultimate divisions oblong lobes. (ii) Var. anglica (Spreng.) Syme A. anglica Spreng. Leaves fleshy, ultimate divisions short teeth. Native. Often gregarious in arable land, waste places and marginal ground, usually on calcareous or sandy soils up to 300 m, and also a grass-seed alien. Locally frequent in south and central Great Britain; rare and mainly casual in northern Great Britain and the Channel Islands; extinct in Ireland where it was formerly an introduction in the east. Europe northwards to south Norway, central Sweden and Lake Onega; Turkey; North Africa; introduced elsewhere. Var. arvensis is the common plant. Var. anglica formerly occurred on the Co. Durham coast near Sunderland and may have occurred at Crookston near Edinburgh. It should be looked for on the coasts. Our plants are all subsp. arvensis. 3. A. ruthenica M. Bieb. Eastern Chamomile A. neilreichii Ortmann; A. pontica auct. Annual herb with fibrous roots. Stems 10–50 cm, pale yellowish-green, sometimes tinted reddish below, erect, striate, with medium to dense, long, pale, curly simple eglandular hairs, branched, sometimes almost to base, very leafy. Leaves 0.5–4.0 × 0.3–2.0 cm, greyish-green, oblong to ovate in outline, obtuse at apex, 1- to 2-pinnatisect, the lobes triangular and acute at apex, sessile or shortly petiolate, with dense long, pale simple eglandular hairs. Capitula 15–35 mm in diameter, solitary at the end of stems and branches; peduncles fairly long, without bracts, with dense, long, subappressed, pale simple eglandular hairs. Involucral bracts in several rows, 5–6 × 1.0–1.4 mm, with dense, long, pale simple eglandular hairs, the outer lanceolate and acute at apex, the inner oblong, with a wide scarious area at the margin and obtuse apex. Flowers of 2 kinds, the corolla tube swollen and spongy below the middle of the fruit, the outer ligulate, the ligules 8–17 mm, 3-lobed at apex and white, the inner tubular, 5-lobed at apex and yellow. Receptacle very acutely conical, with oblanceolate to obovate-cuneate scales, lacerate or dentate at the acuminate apex. Achenes 1.5–2.0 mm, those of the outer tubular flowers subcylindrical with rather thick, obtuse ribs; pappus an obtuse, but not thickened rim, or with a short auricle on the adaxial side. Flowers 6–7. 2n = 18. Introduced. A grain and bird-seed casual. In a few scattered localities in Great Britain. Native of eastern Europe and Caucasus.

Section 3. Maruta (Cass.) Griseb. Maruta Cass.; Cotula Rupr., non L. Annual herbs. Involucral bracts with pale brown scarious margins. Ligules white. Receptacle conical; scales more or less caducous, not brown at apex. Achenes turbinate, ribbed, the ribs tuberculate. 4. A. cotula L. Stinking Chamomile A. foetida Lam. nom. illegit.; Maruta foetida Gray nom. illegit.; Maruta cotula (L.) DC.; Chamaemelum foetidum Baumg. nom. illegit.; Maruta vulgaris Bluff & Fingerh.; Chamaemelum cotula (L.) All.; Cotula foetida S. G. Gmel. Foetid annual herb with fibrous roots. Stems (10–)20– 60(–70) cm, pale yellowish-green, sometimes flushed brownish-purple below, erect or ascending, sometimes prostrate, striate, with rather sparse, short simple eglandular hairs, branched, sometimes nearly to base, leafy. Leaves 1.5–6.0 × 0.5–2.0 cm, medium yellowish-green, oblong, oblong-obovate or obovate in outline, more or less acute at apex, 1- to 3-pinnate, the ultimate segments linear and more or less acute at apex, glabrous or nearly so, sessile or shortly petiolate. Capitula 12–30 mm in diameter, solitary at the ends of stems and branches; peduncles long, without bracts or leaves for a considerable distance, striate, with sparse, short simple eglandular hairs. Involucral bracts in several rows, 2.0–3.5 × 0.6–0.8 mm, with a green central stripe and broad, scarious brownish margin, oblong, obtuse at apex, slightly arachnoid-hairy. Flowers of 2 kinds, the outer ligulate, usually neuter without styles, the ligules 5– 14 mm, 3-lobed at apex, white and at first spreading but later reflexed, the inner tubular, 5-lobed and yellow. Receptacle long-conical, with linear, acute scales near the apex. Achenes 1.5–1.8 mm, yellowish-white, turbinate, about 8to 11-ribbed, the ribs tuberculate; pappus an inconspicuous, crenate, membranous border. Flowers 7–9. Visited mainly by flies. 2n = 18. (i) Var. cotula Stems ascending or erect. Leaves not fleshy. (ii) Var. vectensis (F. N. Williams) P. D. Sell A. foetida var. vectensis F. N. Williams; A. cotula var. maritima auct. Stems prostrate. Leaves fleshy. Native. Arable land, waste places and rough ground up to 300 m, often on heavier soils than A. arvensis. Locally common in south and central England, rarer in the north. Southern Europe northwards to south Norway; north and west Asia; introduced elsewhere. A member of the European Southern-temperate element. The common weed is var. cotula. Var. vectensis occurs on sandy beaches in the Isle of Wight and Somerset and should be looked for elsewhere on the south coast. Section 4. Anthemaria Dumort. Biennial to perennial herbs. Involucral bracts with or without brown margins. Ligules yellow. Receptacle flat to more or less hemispherical; scales persistent, not brown at apex. Achenes triangular-oblong, more or less compressed, striate, not tuberculate.

72. Anthemis 5. A. tinctoria L. Yellow Chamomile Cota tinctoria (L.) J. Gay ex Guss. Slightly foetid biennial to perennial herb with a stout stock. Stems 20–60 cm, pale yellowish-green, sometimes suffused brownish-purple below, erect, ridged, with rather sparse to dense, appressed arachnoid hair, usually with long branches, sometimes nearly to base, leafy. Leaves 2–7 × 1–3 cm, pale yellowish-green on upper surface, hardly paler beneath, oblong or lanceolate in outline, acute at apex, deeply pinnatisect or pinnately lobed, the lobes linear and dentate, the teeth apiculate at apex, glabrous or nearly so on upper surface, sparsely appressed, arachnoid-hairy beneath, or to densely hairy on both surfaces, sessile or shortly petiole. Capitula 20–40 mm in diameter, solitary at the ends of branches; peduncles long, without leaves for a considerable distance, ridged, sparsely to densely appressed arachnoidhairy. Involucral bracts in several rows, 3–7 × 1.0–1.5 mm, pale yellowish-green throughout or the outer tipped blackish-brown, and the inner with a blackish-brown margin in the upper half or throughout, lanceolate to narrowly ovate, the outer acute, the inner rounded-obtuse at apex, sparsely to densely, appressed arachnoid-hairy. Flowers of 2 kinds, the outer ligulate and female, the ligules golden yellow and shortly 3-lobed at apex, the inner tubular, golden yellow and with 5, short, rounded lobes at apex, or rarely all tubular. Receptacle flat to more or less hemispherical in fruit, covered with lanceolate, acute-cuspidate scales hardly exceeding the tubular flowers. Achenes about 2 mm, triangularoblong, more or less compressed, 4-angled and striate on each face, truncate at apex, smooth and glabrous; pappus a very short border. Flowers 6–8. Much visited by a great variety of insects. 2n = 18. (a) Subsp. tinctoria Stems frequently branched, sparingly arachnoid-hairy. Leaves sparingly arachnoid-hairy. Involucral bracts sometimes slightly brownish at apex, slightly arachnoid-hairy. Ligules usually golden yellow. (i) Var. tinctoria Ligulate flowers present. (ii) Var. discoidea (All.) DC. Chamaemelum discoidea All.; A. discoidea (All.) Willd. Flowers all tubular. (b) Subsp. subtinctoria (Dobrocz.) So´o A. subtinctoria Dobrocz. Stems frequently branched, more or less densely arachnoidhairy. Involucral bracts sometimes slightly brownish at apex; densely arachnoid-hairy. Ligules pale yellow. (c) Subsp. australis R. Fern. Stems frequently unbranched, more or less densely arachnoid-hairy. Leaves more or less densely arachnoidhairy. Involucral bracts sometimes slightly brownish at apex. Ligules golden yellow. (d) Subsp. fussii (Griseb.) Beldie A. tinctoria var. fussii Griseb. Stems frequently branched, sparingly arachnoid-hairy. Leaves sparingly arachnoid-hairy. Involucral bracts with

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a distinct brown or blackish margin and apex, sparingly arachnoid-hairy. Ligules golden yellow. Introduced. Waste places and rough and marginal land. Casual or naturalised rather frequently in south and central Great Britain, rarer in the north. South and central Europe northwards to Scandinavia and Finland; western Asia; introduced in North America. The species has a Eurosiberian Temperate distribution. The flowers yield a yellow dye. Specimens of the first three subspecies given above have been seen from Great Britain and the fourth is grown in gardens. Var. discoidea is occasional. Section 5. Cota (J. Gay ex Guss.) Rchb. fil. Cota J. Gay ex Guss. Annual or biennial herbs. Inner involucral bracts with more or less brown margins. Ligules white. Receptacle more or less hemispherical; scales persistent, not brown at apex. Achenes more or less compressed, striate, not tuberculate. 6. A. altissima L. Southern Chamomile Cota altissima (L.) J. Gay ex Guss.; A. cota L. Annual herb with fibrous roots. Stems 20–120 cm, pale yellowish-green, sometimes suffused brownish-purple, erect, robust, striate, with few to numerous, short simple eglandular hairs or glabrous, branched, sometimes nearly to base, leafy. Leaves 1.5–7.0 × 1–4 cm, medium yellowishgreen, broadly ovate in outline, obtuse at apex, 2- to 3-pinnatipartite, the segments patent, the lobes linear and mucronate-spinulose, sessile or shortly petiolate, glabrous or slightly hairy. Capitula (20–)25–40(–50) mm in diameter, solitary at the ends of stems and peduncles; peduncles fairly long, striate, sparsely hairy, becoming clavate in fruit. Involucral bracts in several rows, 5–6 × 0.6–1.0 mm, rather long-hairy, the outer without a scarious margin, ovate-lanceolate and acute at apex, the inner with a hyaline or brownish-scarious margin and apex, oblong-lanceolate to elliptic-oblong and obtuse at apex. Flowers of 2 kinds, the outer up to 20 mm, ligulate, the ligules 3-lobed and white, the inner tubular, 5-lobed and yellow. Receptacle hemispherical, with oblong-spathulate scales, contracted into a rigid point at the truncate or emarginate apex and longer than the flowers, becoming rigid in fruit. Achenes 2.0–2.5 mm, brownish, obpyramidal-subcompressed, very narrowly winged, (7–)8- to 10(–11)-striate on each side, those of the ligulate flowers triangular, those of the tubular flowers quadrangular; pappus an acute rim or short corona. Flowers 6–7. 2n = 18. Introduced. Grain and carrot-seed casual; formerly a wool casual. A few scattered records in Great Britain. Native of southern Europe, Caucasia and central Asia; frequent casual further north. 7. A. austriaca Jacq. Austrian Chamomile A. cotiformis Velen.; A. hyrcana Sosn. ex Grossh.; A. zangelana Sosn. ex Grossh.; Cota austriaca (Jacq.) Sch. Bip. Annual to biennial herb with fibrous roots. Stems (7.5–)10–60 cm, pale yellowish-green, often suffused brownish-purple, erect, striate, with few to numerous, pale

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arachnoid hairs, branched, leafy. Leaves 1.5–5.0 × 0.5–2.0 cm, medium yellowish-green, oblong to obovate in outline, acute at apex, regularly pinnatipartite, the segments linear and pectinate, the lobes linear-oblong to lanceolate and acute with a mucronate-cuspidate tip, sessile or shortly petiolate, with sparse to dense, pale simple eglandular hairs. Capitula (10–)20–40 mm in diameter, solitary at the ends of stems or branches; peduncles long and without bracts, striate, with few to numerous, short and medium, pale, appressed simple eglandular hairs. Involucral bracts in several rows, 4–9 × 1.5–3.0 mm, the outer pale green, lanceolate or ovate-lanceolate, obtuse and slightly hairy, the inner with wide hyaline, sometimes brownish-scarious margins, oblong to elliptic-oblong and acute at apex. Flowers of 2 kinds, the outer ligulate, the ligules (9–)10–15 mm, white and sometimes more or less 2- to 3-lobed, the inner tubular, 5-lobed and yellow. Receptacle subhemispherical, the oblong scales with a long-acute apex, becoming rigid in fruit and subequalling the flowers. Achenes 1.7–2.5 mm, brownish, obovate, compressed, with acute lateral angles and obscurely (2–)3-striate on each side; pappus an acute rim or very short corona. Flowers 6–7. 2n = 18. Introduced. Wool, grass-seed and wild flower seed casual, possibly also a bird-seed casual. Scattered localities in southern Great Britain. Native of east-central and southeast Europe; Turkey and Caucasia; casual further north. 72 × 80 × Tripleurothemis Stace Anthemis × Tripleurospermum Hybrids between these two genera have receptacular scales only in some capitula and only show traces of oil glands on the achenes. 72(4) × 80 × T. maleolens (P. Fourn.) Stace A. cotula × T. maritimum subsp. inodorum × Anthemimatricaria celakovskyi Geisenh. ex Domin nom. inval.; × Anthemimatricaria maleolens P. Fourn. This hybrid is intermediate in the leaves and indumentum, its mature achenes have 5–10 smooth, whitish ribs of varying thickness with traces of oil glands below a well-developed, spreading membranous pappus, receptacular bracts are present only in some capitula on a plant and are intermediate between the involucral bracts and receptacular scales of A. cotula. They are less vigorous than the parents and completely sterile. 2n = 18. Native. Known only from a single plant growing with both parents in a ley meadow near Moulsford in Berkshire in 1966 and two plants growing with both parents in barley stubble near Bridgnorth in Shropshire in 1969. Also recorded from France, Czechoslovakia and Germany. (× T. gruetterana (Asch.) P. D. Sell, A. arvensis × T. maritimum subsp. inodorum (Anthemimatricaria gruetterana Asch.) has been grown from seed collected from a plant of A. arvensis near Oxford in 1964. It has also been recorded from Germany.) 73. Xanthophthalmum Sch. Bip. Pinardia Cass.; Chrysanthemum auct. Annual herbs. Stems decumbent to erect. Leaves alternate, simple, shallowly to deeply lobed or pinnatisect. Capitula

solitary at the ends of stems or branches. Involucral bracts in several rows, with scarious margins. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female. Corolla yellow or yellow and cream. Anthers without tails, apical appendages lanceolate-oblong, flat. Styles with linear-oblong branches; stigmatic areas in 2 separate lines. Receptacle convex, without scales. Achenes of 2 kinds, those of the tubular flowers not or hardly angled and unwinged or with a single adaxial wing, those of the ligulate flowers more or less 3-angled with the ribs often winged. Two species in the Mediterranean region and western Asia; widely introduced elsewhere. 1. Upper leaves subentire, incise-dentate or lobed; achenes 1. segetum of ligulate flowers without an adaxial wing 1. Upper leaves bipinnatisect; achenes of ligulate flowers 2. with an adaxial wing 2(i). coronarium var. coronarium 2. Ligules yellow 2. Ligules white with a yellow base 2(ii). coronarium var. discolor

1. X. segetum (L.) Sch. Bip. Corn Marigold Chrysanthemum segetum L. Annual herb. Stems 20–60(–80) cm, simple or branched, glaucous, glabrous, decumbent to erect, obscurely sulcate. Leaves 1.5–10.0 × 0.6–5.0 cm, glaucous, oblong to obcuneate in outline, obtuse or acute at apex, glabrous; basal deeply lobed or coarsely pinnatisect with oblong, irregularly serrate lobes and tapering at the base to an indistinct, flattened petiole; cauline sessile or subsessile, becoming smaller upwards, the uppermost often oblong, subentire and amplexicaul. Capitula 30–70 mm in diameter, solitary at the ends of stems and branches; peduncles 2–10 cm, distinctly swollen below the capitulum, becoming thickened in fruit. Involucral bracts in several rows, 3–8 × 2–4 mm, oblong, rounded with a wide, brownish, membranous margin at apex. Flowers of 2 kinds, the inner 3–4 mm, tubular, yellow, 2-winged, dilated above the middle, expanded at the base and partly covering the apex of the immature achene, with 5, recurved, ovate, acute lobes at the apex, the outer 10–15 × 6–7 mm, ligulate, the ligules yellow, and rounded or obscurely 3-lobed at apex. Receptacle shallowly convex, without scales, accrescent after anthesis. Achenes of tubular flowers 2.4–3.0 × 1.2–1.4 mm, shortly subcylindrical, 10-ribbed; of ligulate flowers 2.0–2.5 × 1.5–1.6 mm, pale brown, oblong, trigonous, with 2 lateral wings, truncate at apex and base, adaxial face prominently 3-ribbed; pappus and oil glands absent. Flowers 6–8. Visited by various insects especially flies. 2n = 18. Introduced. Weed of acid arable fields, waste places and roadsides. Locally frequent throughout Great Britain and Ireland and sometimes a troublesome weed of loamy and sandy soils. Probably native of the eastern Mediterranean region and western Asia, but well established throughout Europe to 70◦ N in Norway; introduced in North Africa and North and South America. The species as a native plant and ancient introduction has a European Southerntemperate distribution.

75. Coleostephus 2. X. coronarium (L.) P. D. Sell Crown Daisy Pinardia coronaria (L.) Less.; Chrysanthemum coronarium L.; Glebionis roxburghii Cass. nom. illegit. Annual herb. Stems 20–70(–100) cm, simple or branched, slightly glaucous, erect, glabrous. Leaves 1.5–9.0 × 0.6–5.0 cm, slightly glaucous, oblong to obovate in outline, obtuse at apex, deeply bipinnatisect, the ultimate divisions narrowly oblong and irregularly serrately lobed with the apices apiculate, sessile and semiamplexicaul or shortly petiolate. Capitula (13–)15–18(–20) mm in diameter, solitary at the ends of stems or branches; peduncles 2–10 cm. Involucral bracts in several rows, 2–10 × 2–5 mm; outer ovate, with a brownish marginal band and a narrow, whitish scarious margin; inner without marginal band but with a wider scarious margin and rounded, scarious apical appendage. Flowers of 2 kinds, the inner 3–4 mm, tubular, yellow, gradually dilated above the middle, slightly expanded at base but not covering the top of the immature achene, with 5, recurved, ovate, acute lobes at apex, the outer 10–15 mm, ligulate, the ligules yellow or creamy-white with a yellow base, and rounded, emarginate or obscurely 3-lobed at apex. Receptacle strongly convex, not much accrescent after anthesis; without scales. Achenes covered with sessile, non-mucilaginous glands; those of the tubular flowers 2.8–3.2 × 2.4–2.6 mm, laterally compressed, with an adaxial wing, prominent ribs on the abaxial face and rounded, superficial ribs on the lateral faces, sometimes the central ones cylindrical and unwinged; those of ligulate flowers 2.8–3.2 × 2.4–2.6 mm, 3-angled, with the angles winged, the abaxial face with 3 slender ribs and lateral faces without ribs; pappus absent. Flowers 6–8. 2n = 18. (i) Var. coronarium Flowers uniformly bright yellow. (ii) Var. discolor (d’Urv.) P. D. Sell Chrysanthemum coronarium subsp. discolor (d’Urv.) Rech. fil.; Chrysanthemum coronarium var. discolor d’Urv. Ligulate flowers creamy-white with a yellow base. Introduced. Wool, grain and bird-seed casual and garden escape in waste and cultivated places. Both varieties occur. Native of the Mediterranean region eastwards to Iran; widely cultivated for ornament and sometimes naturalised elsewhere. 74. Leucanthemella Tzvelev Perennial herbs. Stems erect, leafy. Leaves alternate, simple, sharply serrate-dentate. Capitula solitary or 2–8 on long peduncles in a lax corymb. Involucral bracts in 2–3 rows. Inner flowers tubular, entire or 5-lobed at apex and bisexual, the outer in 1 row, ligulate, entire or 3-lobed at apex and sterile, with the tube strongly compressed but unwinged. Corolla of tubular flowers yellow, those of ligulate flowers white or reddish. Anthers without tails; apical appendages lanceolate-oblong, flat. Styles with linear-oblong branches; stigmatic areas in 2 separate lines. Receptacle strongly convex, without scales. Achenes all similar, cylindrical, with thick, white, obtuse ribs, without mucilaginous cells or vallecular secretory canals; pappus minute or absent.

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Two species in eastern Europe, eastern Russia, China, Korea and Japan. 1. L. serotina (L.) Tzvelev Autumn Oxeye Daisy Chrysanthemum serotinum L.; Chrysanthemum uliginosum (Waldst. & Kit. ex Willd.) Pers.; Pyrethrum uliginosum Waldst. & Kit. ex Willd.; Tanacetum serotinum (L.) Sch. Bip. Perennial herb. Stems 30–150 cm, pale yellowish-green, ridged, erect, very shortly appressed-hairy, very leafy, branched in upper part. Leaves all cauline, numerous, hardly decreasing in size except on the branches, 2–8 × 0.7– 2.5 cm, dull medium green on upper surface, slightly paler beneath, lanceolate, oblanceolate, oblong and oblonglanceolate, acute at apex, the lowest 2- to 4-lobed, the median serrate-dentate, the upper subentire, denticulate or serrate-dentate with a few teeth, the lower narrowed to a sessile base, the upper cuneate or rounded to a sessile base, with minute hairs and gland-dotted on both surfaces; veins prominent beneath. Capitula 55–80 mm in diameter, solitary at the ends of branches or 2–8 in a lax corymb; peduncles pale green, ridged, minutely hairy. Involucral bracts in 2–3 rows, 10–14 × 3.3–3.5 mm, yellowish-green with brownish, wrinkled scarious margins, broadly ovate, rounded-obtuse at apex. Flowers of 2 kinds, the outer in 1 row, 25–35 mm, ligulate, sterile, the ligules white or reddish and entire or shallowly 3-lobed at apex, the inner 8–10 mm, tubular, bisexual, yellow, entire or shallowly 5-lobed at apex. Receptacle convex, pitted, without scales or hairs. Achenes 2–3 mm, cylindrical, with thick, white, obtuse ribs; pappus minute or absent. Flowers 9–11. 2n = 18. Introduced. Garden escape or throw-out naturalised on rough ground and by ditches and ponds. Scattered records in southern England. Native of south-east Europe. 75. Coleostephus Cass. Annual herb. Stems erect or ascending. Leaves alternate, serrate-dentate. Capitula solitary at the ends of stems or branches. Involucral bracts in 2–3 rows, with a brownish, scarious margin. Inner flowers tubular, bisexual and basally much swollen and spongy in fruit, the outer ligulate, 3-lobed at apex and female and fertile. Corolla of both ligulate and tubular flowers yellow. Anthers without tails; apical appendages lanceolate-oblong, flat. Style with linear-oblong branches; stigmatic areas in 2 separate lines. Receptacle plano-convex, without scales. Achenes of ligulate flowers compressed, with an elongate, membranous tubular pappus enveloping the corolla tube; achenes of tubular flowers cylindrical, with an auricle enveloping the lower half of the corolla tube. Three species in southern Europe and North Africa; sometimes cultivated. 1. C. myconis (L.) Rchb. fil. Mediterranean Marigold Chrysanthemum myconis L.; Pyrethrum myconis (L.) Moench; Myconia chrysanthemoides Sch. Bip.; Chrysanthemum hybridum Guss. Annual herb with fibrous roots. Stems 10–45 cm, pale green, striate, erect or ascending, slightly angled, glabrous or with occasional hairs, leafy, branched. Leaves 4–10 ×

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0.5–3.0 mm, medium green, slightly paler beneath, glabrous or with a few hairs; lower obovate to obovate-spathulate, rounded-obtuse at apex, regularly serrate-dentate, the teeth triangular, petiolate; middle leaves ovate-oblong and semiamplexicaul; upper oblong or linear, obtuse to acute at apex, toothed, sessile and semiamplexicaul. Capitula numerous, solitary at the end of stems and branches, 25–30 mm in diameter; peduncles ridged, glabrous. Involucral bracts in 2–3 rows, 7–8 × 3–4 mm, pale green, with a pale brownish scarious margin in upper part, oblong but slightly narrowed in the middle, broadly rounded at apex, glabrous. Flowers of 2 kinds, the outer 10–11 mm, ligulate and female, the ligules yellow, broadly elliptic and shallowly 3-lobed at apex; the inner 2.5–4.0 mm, tubular and bisexual, golden yellow, 5-lobed and papillose at apex. Receptacle planoconvex, pitted, without scales. Achenes of ligulate flowers about 3 mm, compressed, sterile, with an elongate, membranous tubular pappus enveloping the corolla tube; achenes of tubular flowers about 2 mm, brown, cylindrical, curved, strongly ribbed and with an auricle 1.2–1.8 mm, enveloping the lower half of the corolla tube. Flowers 7–10. 2n = 18. Introduced. Wool and granite casual. In a few scattered localities in Great Britain. Native of southern Europe and western Asia. 76. Leucanthemum Mill. Perennial or rarely annual herbs with red-tipped roots. Stems erect or ascending. Leaves alternate, entire to serrate or deeply lobed. Capitula solitary or few at the ends of stems and branches. Involucral bracts in several rows, usually with dark, scarious margins. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female. Corolla of tubular flowers yellow, those of ligulate flowers white. Receptacle usually convex, without scales. Anthers without tails; apical appendages lanceolate-oblong, flat. Style with linear-oblong branches; stigmatic areas in 2 separate lines. Achenes all similar, obconical-cylindrical, usually with 10 prominent ribs and epicarpic mucilaginous cells on most ribs, vascular strands between the ribs and vallecular secretory canals present; pappus absent or a corona or auricle. Thirty-three species in Europe and North Africa, some species widely cultivated. L. vulgare subsp. ircutanum, L. vulgare subsp. vulgare and L. × superbum sometimes appear to occur mixed in wild flower seed and form a continuous flowering period from May to August in the order given. Bocher, T. W. & Larsen, K. (1957). Cytotaxonomical studies in the Chrysanthemum leucanthemum complex. Watsonia 4: 11–16. Howarth, S. E. & Williams, J. T. (1968). Chrysanthemum leucanthemum L. in Biological flora of the British Isles. Jour. Ecol. 50: 565–595. Vogl, R. (1991). Die Gattung Leucanthemum Mill. (Compositae– Anthemideae) auf der iberischen Halbinsel. Ruizia 10: 1–261. 1. Leaves up to 15 cm; capitula 60–100 mm in diameter; 2. × superbum involucral bracts 11–15 mm 1. Leaves 1–8 cm; capitula 25–60(–75) mm in diameter; 2. involucral bracts 6–8 mm

2. Stems woody at base; lower cauline and basal leaves large, fleshy and subrotund; involucral bracts with a dark 1(c). vulgare subsp. crassifolium brown border 2. Stem not woody; lower and basal leaves smaller and less 3. fleshy; involucral bracts with a pale brown border 3. Stems usually nearly glabrous; lower leaves regularly dentate; capitula 45–60(−75) mm in diameter; flowering 1(b). vulgare subsp. ircutianum in May or June 3. Stems 1 usually obviously hairy; lower leaves deeply divided with alternating lobes or teeth; capitula 25–48 mm 4. in diameter; flowering late June to August 4. All leaves very narrow and deeply divided, many teeth or 1(d). vulgare subsp. cantabricum lobes almost spinulose 4. Leaves broader and not so deeply divided, the teeth 5. broader 5. Stems 1 or more, up to 75(−100) cm 1(a,i). vulgare subsp. vulgare var. vulgare 5. Stems up to 12 cm 1(a,ii). vulgare subsp. vulgare var. nanum

1. L. vulgare Lam. Oxeye Daisy Chrysanthemum leucanthemum L.; Tanacetum leucanthemum (L.) Sch. Bip. Perennial herb with a slender, branched, oblique stock, sometimes woody, producing non-flowering rosettes. Stems 1–several, 6–75(–100) cm, pale green, sometimes suffused brownish-purple, erect or ascending, faintly to markedly striate, glabrous or with few to numerous, pale simple eglandular hairs, simple or branched, leafy. Leaves alternate, dull medium to dark green, paler beneath, glabrous or with few simple eglandular hairs; basal and lower cauline 1–8 × 0.5– 4.0 cm, obovate-spathulate to oblong-spathulate, rounded at apex, dentate to incise-dentate or crenate, abruptly contracted to a long petiole; upper cauline oblong or linear, obtuse at apex, dentate to pinnatifid, with very narrow to wide lobes, with a sessile, amplexicaul base. Capitula 25– 60(–75) mm in diameter, solitary on long peduncles; peduncles medium green, glabrous. Involucral bracts in several rows, 6–8 × 1.2–1.8 mm, green, with pale to dark purplish or brown scarious margins and tips, narrowly lanceolate to oblong, obtuse at apex, glabrous. Flowers of 2 kinds, the outer 12–30 mm, ligulate and female, the ligules white and 3-lobed at apex, rarely absent; inner 3–5 mm, tubular, yellow and shortly 5-lobed at apex, very rarely all tubular. Receptacle convex, faintly pitted, without scales. Achenes 2–3 mm, pale grey, obconical-cylindrical, strongly 10ribbed; pappus absent or a membranous border in ligulate flowers, absent in tubular flowers. Flowers 5–8. There seems to be no certain way of morphologically dividing diploid and tetraploid plants into two species, although it has often been attempted. In fact two of the tetraploids are easier to distinguish from each other, than the third tetraploid is from the diploid. All have therefore been placed as infraspecific taxa under one species. (a) Subsp. vulgare Chrysanthemum leucanthemum subsp. praecox Horvatic; L. praecox (Horvatic) Horvatic Stem not usually woody, with few to numerous hairs. Lower leaves deeply divided with rather broad lobes or teeth. Capitula 25–40 mm in diameter. Involucral bracts with

77. Chrysanthemum pale brown margins. Flowering from late June to August. 2n = 18. (i) Var. vulgare Stems up to 75(–100) cm. (ii) Var. nanum P´erard Stems up to 12 cm. (b) Subsp. ircutianum (DC.) P. D. Sell L. ircutianum DC.; Chrysanthemum ircutianum (DC.) Turcz. Stem not usually woody, usually nearly glabrous, often tinted brownish-purple. Lower leaves regularly dentate, not deeply divided. Capitula 45–60(–75) mm in diameter. Involucral bracts with pale brown margins. Flowering in May and June. 2n = 36. (c) Subsp. crassifolium (Lange) Rouy L. pallens var. crassifolium Lange; L. crassifolium (Lange) Lange; Chrysanthemum crassifolium (Lange) Guinea; L. vulgare var. crassifolium (Lange) Merino; L. ircutianum subsp. crassifolium (Lange) Vogt Stems with long branches from a very woody base, markedly striate, often brownish-purple, glabrous or slightly hairy. Lower leaves large, fleshy, subrotund and crenate-dentate. Capitula 30–40 mm in diameter. Involucral bracts with very dark brown margins. Flowering in May and June. 2n = 36. (d) Subsp. cantabricum (Sennen) P. D. Sell L. cantabricum Sennen; Chrysanthemum leucanthemum var. cantabricum (Sennen) Guinea; L. ircutianum subsp. cantabricum (Sennen) Vogt Stem slightly hairy, not woody, often tinted brownishpurple. Lower and upper leaves deeply incise-dentate, many of the teeth long and narrow, even filamentous. Capitula 30–40 mm in diameter. Involucral bracts with pale brown margins. Flowering in June and July. Native. Grassy places especially on rich soils, introduced on railway banks, roadsides and waste places. Throughout Great Britain and Ireland. A morphologically and cytologically very variable species or species-complex which occurs throughout Europe to Lapland and Siberia, and is introduced in North America and New Zealand. A member of the Eurosiberian Boreo-temperate element. Subsp. vulgare is the native plant of grassland. Var. vulgare is the common plant, while var. nanum is found on cliff-tops in Cornwall and should be looked for elsewhere in similar situations. It also occurs in France. Subsp. ircutianum is found on railway banks, roadsides and in waste places and is probably introduced. Subsp. crassifolium is known only from cliff-tops on serpentine at Mullion and probably Cadgwith in Cornwall. It is found also in Portugal, Spain and France. Subsp. cantabricum is known only from dry limestone near Cong in Co. Mayo, Ireland and Dinas Powis in Glamorganshire. It is a very distinctive plant and should be looked for elsewhere on the limestone. It also occurs in Spain and France. 2. L. × superbum (Bergmans ex J. Ingram) Kent Shasta Daisy L. lacustre (Brot.) Samp. × maximum (Ramond) DC. Chrysanthemum × superbum Bergmans ex J. Ingram; L. maximum auct.; Chrysanthemum maximum auct.

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Perennial herb with slender, branched, oblique, woody stock producing non-flowering leaf rosettes. Stems 1– numerous, up to 120(–150) cm, pale green, often suffused brownish-purple, erect, faintly striate, simple or branched, glabrous, leafy. Leaves alternate, dull dark green on upper surface, with a pale midrib, paler beneath, glabrous; basal and lower cauline 3–15 × 1–4 cm, elliptical-oblong or ovate, obtuse to more or less acute at apex, serrate, with ascending, rather blunt teeth, attenuate to a short or medium petiole; cauline gradually decreasing in size upwards, the upper lanceolate, more or less acute at apex, shallowly serrate to subentire, cuneate-based, sessile and semiamplexicaul. Capitula 60–100 mm in diameter, solitary at the ends of stems or branches; peduncles yellowish-green, ridged, glabrous. Involucral bracts in several rows, 11–15 × 4–6 mm, green, with a scarious margin and a brown band in between, ovate, subacute to obtuse at apex, glabrous. Flowers of 2 kinds or flore pleno; outer 32–40 mm, ligulate, 3-lobed at apex and female, the inner 6–8 mm, tubular, bisexual and yellow, with 5 triangular-ovate lobes at apex. Receptacle convex, faintly pitted, without scales. Achenes all similar, 2.5–3.5 mm, obconical-cylindrical, prominently 10-ribbed, unwinged; pappus a membranous border, sometimes incomplete or absent. Flowers 6–8. Visited by bees, flies, butterflies and moths. Introduced. Abundant in gardens and fully fertile; well naturalised in waste and rough ground, railway banks, disused quarries and grassy waysides. In scattered localities throughout Great Britain and the Channel Islands. Of garden origin. L. maximum (Ramond) DC. (Chrysanthemum maximum Ramond) occurs very rarely as an escape and is smaller with crenate-dentate rather than serrate toothing. 77. Chrysanthemum L. nom. conserv. Dendranthemum (DC.) Des Moul. Perennial herbs. Stems erect or ascending, sometimes almost woody, leafy. Leaves alternate, pinnatisect, usually with 2 pairs of segments which are entire or toothed. Capitula solitary at the ends of stems or branches or in loose corymbs. Involucral bracts in several rows, scarious or with a scarious margin. Flowers of 2 kinds, the inner tubular, 5lobed at apex and bisexual, the corolla tube with sessile glands, the outer 3-lobed at apex, ligulate and female, or with all flowers ligulate. Corolla of tubular flowers yellow, of ligulate flowers yellow, pink, white, red or orange or a mixture of colours. Anthers without tails, apical appendages subtriangular. Style with linear-oblong branches; stigmatic areas in 2 separate lines. Receptacle markedly conical, pitted, without scales. Achenes narrowly obovoid, faintly 5- to 8-ribbed, thin-walled; pappus absent. About 37 species in Asia, mainly China and Japan, with 1 species extending into eastern Europe; widely cultivated as ornamentals. Brummitt, R. (1997). Chrysanthemum once again. The Garden 122: 662–663. Committee for Spermatophyta (1998). Report 46, Conserve Chrysanthemum L. (Compositae) with a conserved type. Taxon 47: 443–444. Sealy, J. R. (1938). Chrysanthemum rubellum and its allies. Jour. Roy. Hort. Soc. 63: 265–273.

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Trehane, P. (1995). Proposal to conserve Chrysanthemum L. with a conserved type (Compositae). Taxon 44: 439–441. Tzvelev, N. N. (1995). English translation of Dendranthema (DC.) Des Moul. from Schischkin, B. K. and Bobrov, E. G. (Edits.) Flora URSS, vol. 26, pp. 423–449. Moscow and Leningrad. 1. Leaves sparsely hairy to glabrous beneath; not normally 4. zawadskii double-flowered 1. Leaves with numerous hairs beneath; often 2. double-flowered 2. Capitula 25–40(–50) mm in diameter, aggregated in loose 1. indicum corymbs 2. Capitula 50–150 mm in diameter, solitary at the ends of 3. stems or branches 3. Capitula 50–150 mm in diameter; flowers usually all 2. × morifolium ligulate, colours variable 3. Capitula 50–80 mm in diameter; outer flowers ligulate 3. × rubellum and pink, inner flowers tubular and yellow

1. C. indicum L. Indian Chrysanthemum Dendranthemum indicum (L.) Tzvelev; Matricaria indica (L.) Desr.; Pyrethrum indicum (L.) Cass.; Tanacetum indicum (L.) Sch. Bip. Perennial herb with a thick, branched rootstock. Stems 25– 100 cm, pale yellowish-green, erect, usually more or less branched, with more or less numerous bifid and simple eglandular hairs, very leafy. Leaves alternate, 2–8 × 1–4 cm, dingy green or greyish-green on upper surface, paler beneath, with sparse, short hairs on upper surface, profusely hairy and often tomentose beneath, and with dotted glands; basal leaves rapidly dying, the lower cauline oblong-ovate, obtuse at apex, deeply pinnately divided, with a large terminal segment and usually 2 pairs of oblong lateral ones which are more or less lobed or serrate, truncate or slightly cordate at base, with a petiole 1–2 cm, almost wingless and without or with false auricles; upper cauline, smaller, partly entire and with shorter petioles. Capitula (1–)5–30(–50), 25–40(–50) mm in diameter, aggregated in loose compound corymbs; peduncles thin and more or less hairy. Involucral bracts in several rows, 5–6 × 1–2 mm, pale green with a rather wide pale or brownish membranous margin, the inner with a large rounded appendage, more or less hairy to nearly glabrous. Flowers of 2 kinds, the outer 13–18 mm with the corolla yellow, pink or white etc., the inner about 3 mm and tubular, or often all ligulate (double). Receptacle markedly conical, pitted, glabrous, without scales. Achenes 1.5–2.5 mm, narrowly obovoid, with 5 longitudinal ribs; pappus absent. Flowers 8–10. Introduced. Cultivated in gardens and probably occasionally occurring on tips. The wild form with one row of yellow ligulate flowers is rarely cultivated. Most garden varieties are double-flowered with pink, white, purple or variable coloured ligules. The wild plant is possibly native of China, Korea and Japan. 2. C. × morifolium Ramat. Florist’s Chrysanthemum Dendranthemum × grandiflorum (Ramat.) Kitam.; Dendranthemum morifolium (Ramat.) Tzvelev; Anthemis grandiflora Ramat.; C. sinense Sabine; Pyrethrum sinense (Sabine) DC.; Tanacetum sinense (Sabine) Sch. Bip.;

Tanacetum morifolium (Ramat.) Kitam.; Dendranthemum sinense (Sabine) Des Moul. Perennial herb with a thickened, sometimes almost woody stock, forming rhizomes. Stems 25–100 cm, dingy green or greyish-green, with more or less dense bifid or simple eglandular hairs, more or less branched, with numerous leaves. Leaves alternate, 2–15 × 1–8 cm, greyish-green, with numerous hairs to nearly glabrous on upper surface, with dense hairs and sometimes almost tomentose beneath, with numerous, dotted glands; basal and lower cauline rapidly dying; cauline varying greatly, oblong-ovate to almost subrotund, pinnately divided into segments or lobes, the petioles short to long. Capitula 1–10, 50–150 mm in diameter, solitary at the ends of stems and branches; peduncles more or less hairy. Involucral bracts in several rows, 8–15 × 2–4 mm, the outer pale green with a rather narrow membranous margin, the inner with a wide pale or brownish membranous margin and a widened appendage at apex. Flowers usually all ligulate, the ligules (17–)22– 54(–64) mm, white, pink or red, of various hues or mixtures. Achenes develop rarely, reproduction by grafting. Flowers 8–10. Introduced. The main ancestors of the Florist’s Chrysanthemums are thought to be C. indicum, C. vestitum Hemsl., C. erubescens Stapf, C. ornatum Hemsl., C. japonense Nakai and C. makinoi Matsum. There is some evidence that Chrysanthemums were grown in China as early as 500 BC. Unsuccessful attempts were made to import them into Europe from Asia in the late seventeenth century, but it was not until 1789 that three cultivars were successfully imported into France from China. Only one of these survived and it arrived in England in 1790. New cultivars followed and about 60 were known by 1825. From then on many new cultivars were developed by hybridisation and selection and many thousands are now available. Commonly grown in gardens and often surviving for a time when thrown out on waste ground and tips. 3. C. × rubellum Sealy Korean Chrysanthemum Dendranthemum rubellum (Sealy) Philp; Chrysanthemum erubescens auct. Perennial herb with a branched rootstock. Stems 80–90 cm, pale yellowish-green, erect, branched in the upper half, with numerous bifid and simple eglandular hairs, very leafy. Leaves alternate, decreasing in size upwards, 3.5–5.0 × 3.5–5.0 cm, bright green on upper surface, paler beneath, broadly triangular-ovate to ovate, more or less obtuse at apex, pinnatisect, with 2 pairs of segments, with the segments coarsely toothed or lobed, the teeth and lobes subacute and mucronate, shortly hairy near the base of the midrib on the upper surface to almost glabrous, whitish-hairy beneath; petioles 20–70 mm, hairy. Capitula 50–80 mm in diameter, solitary at the ends of stems or branches; peduncles hairy. Involucral bracts in several rows, 5.0–10 × 1.0–4.0 mm, the outer green, narrowly lanceolate, with a rounded, scarious apex and ciliate, the inner almost completely scarious, oblong and truncate at apex. Flowers of 2 kinds, the outer 24–30 mm, ligulate, the ligules pink, the inner about 5 mm, tubular. Receptacle

79. Matricaria markedly conical, pitted, without scales. Achenes 1.8– 2.5 mm, narrowly obovoid; without a pappus. Flowers 8–10. Introduced. Grown in gardens and recorded as an escape in Kent; should be looked for elsewhere. Its origin is unknown. 4. C. zawadskii Herbich Zawadski’s Chrysanthemum Dendranthemum zawadskii (Herbich) Tzvelev; Chrysanthemum sibiricum Turcz.; Tanacetum zawadskii (Herbich) Pawl.; Pyrethrum zawadskii (Herbich) Nyman Perennial herb with a branched rootstock. Stems 15–50 cm, erect, pale yellowish-green, with numerous, bifid and simple eglandular hairs, with a few branches in the upper half, densely leafy. Leaves alternate, 2–8 × 1.5–4.0 cm, dull rather dark green on upper surface, paler beneath, sparsely and appressed-hairy to nearly glabrous but with numerous dotted glands; basal and lower cauline ovate to nearly subrotund, obtuse at apex, pinnatisect, with 1–3 pairs of segments, the segments 2–3 mm wide, linear and abruptly acute at apex, cuneate at base, with rather long, narrowly winged petioles; median leaves smaller, less divided and with shorter, broadly winged petioles; upper leaves entire or pinnately lobed. Capitula 1–5, 40–60 mm in diameter, solitary at the ends of the stems or branches, usually not forming a corymb; peduncles sparsely hairy. Involucral bracts in several rows, 4.5–6.0 × 1–3 mm, pale green, with rather wide, pale or brownish membranous margins, the outer broadly linear to oblong, the inner linear to linear-oblong, glabrous to more or less hairy. Flowers of 2 kinds, the outer 13–30 mm, ligulate, the ligules white, pink, pinkishviolet or various tones and mixtures, the inner 2.5–3.0 mm and tubular. Receptacle markedly conical, pitted, glabrous, without scales. Achenes 1.8–2.5 mm, narrowly obovoid; without a pappus. Flowers 9–10. 2n = 54. Introduced. Grown in gardens and possibly occurring as a throw-out on tips. Native of the Carpathians, Urals and north and central Russia. Greatly variable, forming many isolated populations distinguished chiefly by the colour of the ligulate flowers and form of the leaves. Named after Aleksander Zawadski (1798–1868). Anacyclus valentinus L. has been recorded as a grain casual, Cladanthus arabicus (L.) Cass. as a casual and Pentzia calcria Kies, P. cooperi Harv. and P. incana (Thunb.) Kuntze as wool casuals. 78. Oncosiphon K¨allersj¨o Annual herbs. Stems erect or ascending, leafy. Leaves alternate, bipinnatisect with linear segments. Capitula solitary at the ends of long branches. Involucral bracts in several rows, with a scarious apex. Flowers all tubular, 4-lobed at apex. Anthers without tails, apical appendage lanceolateoblong, flat. Style with linear-oblong branches; stigmatic area in 2 separate lines. Receptacle flat to slightly convex, without scales. Achenes more or less terete, 4-ribbed, glandular between the ribs; pappus a small, entire crown. Eight species in South Africa and Namibia. O. piluliferum (L. fil.) K¨allersj¨o and O. suffruticosum (L.) K¨allersj¨o have been recorded as wool casuals.

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K¨allersj¨o, M. (1988). A generic reclassification of Pentzia Thunb. (Compositae–Anthemideae) from South Africa. Bot. Jour. Linn. Soc. 96: 299–322.

1. O. grandiflorum (Thunb.) K¨allersj¨o Swollen-flowered Maywood Tanacetum grandiflorum Thunb.; Pentzia grandiflora (Thunb.) Hutch.; Matricaria grandiflora (Thunb.) Fenzl ex Harv., non Poir. Annual herbs with a tap-root and fibrous side shoots. Stems 20–30 cm, pale yellowish-green, erect or ascending, flexuous, with short curly hairs, furrowed, branched, leafy. Leaves alternate, 3–4 × 2–3 cm, pale yellowishgreen, broadly ovate or ovate-oblong in outline, acute at apex, bipinnatisect, the segments all narrowly linear, with a short sharp point at apex, more or less hairy. Capitula 10–15 mm in diameter, solitary at the ends of long branches; peduncles long, hairy. Involucral bracts in several rows, the outer 3.0–3.5 × 0.6–0.7 mm, pale green, linear, with a membranous area at apex and with dense, short, curly hairs, the inner 3–4 × 1.2–1.4 mm, oblong, with a broad, membranous area at apex, densely hairy. Flowers all tubular, yellow, with 4, short corolla lobes with dorsal appendages and a very much swollen and brittle tube. Receptacle flat to slightly convex, without scales. Achenes yellowish-brown, more or less terete but tapering at base, 4-ribbed, glandular between the ribs; pappus a small, entire crown. Flowers 6–7. 2n = 16. Introduced. A persistent wool alien. Native of South Africa. 79. Matricaria L. Chamomilla Gray Annual herbs. Stems erect or ascending, leafy. Leaves alternate, 2- to 3-pinnatisect, with numerous linear segments. Capitula solitary at the ends of stems and branches. Involucral bracts in 2 or 3 rows, with a scarious margin. Inner flowers tubular, bisexual, 4- to 5-lobed at apex, tube swollen in fruit, the outer when present ligulate, 3-lobed at apex and female, sometimes all tubular. Corolla of tubular flowers yellow, those of ligulate flowers white. Anthers without tails, apical appendage lanceolate-oblong, flat. Style with linear-oblong branches; stigmatic area in 2 separate lines. Receptacle conical, hollow, without scales. Achenes mucilaginous when wet, slightly compressed, obliquely truncate above, abaxial face convex, adaxial face with 3–5 narrow, whitish, longitudinal ribs; pappus absent or a very short rim. Seven species in Eurasia, North America and North Africa with some species widespread elsewhere as weeds. The genus, as typified and accepted here is as defined by C. Jeffrey (1979), not S. Rauschert (1974). M. aurea (Loefl.) Sch. Bip. has been recorded as a casual and M. decipiens (Fisch. & C. A. Mey.) K. Koch as a grain and wool alien. Grime, J. P. et. al. (1988). Comparative plant ecology. London. [M. discoidea.] Jeffrey, C. (1979). Note on the lectotypification of the names Cacalia L., Matricaria L. and Gnaphalium L. Taxon 28(4): 349– 351.

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Rauschert, S. (1974). Nomenklatorische Probleme in der Gattung Matricaria L. Folia Geobot. Phytotax. 9: 249–260. Salisbury, E. (1961). Weeds and aliens. London. 1. White ligulate flowers present, soon deflexed; 1. recutita sweet-smelling 1. Flowers all tubular and yellowish-green, smelling 2. strongly of pineapple 2. Capitula 5–8 mm in diameter; pappus minute and entire 2(a). discoidea subsp. discoidea or absent 2. Capitula 8–12 mm in diameter; pappus with 2 dark 2(b). discoidea subsp. occidentalis brown, lateral lobes

1. M. recutita L. Scented Mayweed M. chamomilla L. (1763), non L. (1753); Chamomilla recutita (L.) Rauschert; M. suaveolens L. nom illegit.; Chamomilla vulgaris Gray nom. illegit. Annual herb with fibrous roots, sweetly scented when fresh. Stems (2–)10–60 cm, pale yellowish-green, erect, ridged, rather slender, flexuous, glabrous, leafy, branched above, sometimes nearly to base. Leaves alternate, 1–7 × 1.5–2.5 cm, medium green, oblong or oblong-oblanceolate in outline, rounded at apex, finely bipinnatisect, all the lobes narrowly linear or almost filiform and subacute or mucronate at apex, glabrous, sessile or with short petioles, swollen at base and semiamplexicaul. Capitula (1–)8–120(–300), solitary and terminal on stems and branches and forming a loose corymb, 10–25 mm in diameter; peduncles 30– 100 mm, with pale ridges and dark green channels, glabrous. Involucral bracts in 2(–3) rows, 1.5–3.0 × 1.0–1.5 mm, green, with pale brown scarious margins, oblong, obtuse at apex. Flowers of 2 kinds, the outer 6–8 mm, ligulate and female, the ligules white, shortly 3-lobed and often markedly reflexed soon after the flowers open, the inner about 1.5 mm, tubular and bisexual, the tube expanded rather suddenly in the upper half, yellow, thinly glandular externally, with 5, deltoid-acute lobes at apex. Receptacle conical, regularly pitted, hollow, without scales. Achenes about 1.0 × 0.4 mm, brownish, oblong-cylindrical, slightly curved, thinly glandular, with 4–5, pale, longitudinal ribs on the adaxial face, truncate at apex; pappus absent or represented by a membranous rim. Flowers 6–8. Visited by flies and small bees. 2n = 18. The reflexed ligules and protruding receptacle, imitating a circumcised penis (i.e. recutita), and the sweet scent help to recognise this species, but are not diagnostic. To be certain of the identification the mature achenes should be examined. Native. Cultivated and waste ground, particularly on light soils. Frequently grows with Tripleurospermum maritimum subsp. inodorum but not so common as that plant. Locally common in England, Wales and the Channel Islands, very scattered records in Scotland and a rare casual in Ireland. Europe, where it may be introduced in the north, and western Asia to India. Introduced in North America and Australia. A member of the European Southern-temperate element. The flowering capitula of this plant are infused to give chamomile tea. The word Chamomilla,
, is a name in Dioscorides of a plant that smelt of apples.

This species grows in Greece, but Chamaemelum nobile, which is called chamomile, is a plant of western Europe. For an explanation of Linnaeus’ muddle over the use of the epithet chamomilla see under Tripleurospermum maritimum. The chief healing substance of this species is the deep blue oil extracted from the fresh flowers which has great anti-inflammatory powers. 2. M. discoidea DC. Pineapple Weed Strongly aromatic annual herb with a pineapple-like scent and fibrous roots. Stems simple or numerous, (2–)5–45 cm, pale yellowish-green, sometimes purplish at base, erect, striate or ridged, glabrous, leafy, branched, often from the base. Leaves alternate, 1–6 × 0.7–2.0 cm, rather crowded, oblong to oblong-oblanceolate in outline, pinnatisect, the segments finely linear, flat, acute and shortly aristate at apex, glabrous. Capitula numerous, 5–10(–12)mm in diameter, enlarging as they mature, in irregular corymbs at the ends of branches; peduncles 2–30 mm, pale green, ridged, glabrous. Involucral bracts in several rows, 4–12 × 1–2 mm, yellowish-green with wide scarious margins, oblong, obtuse at apex, glabrous. Flowers 0.7–0.9 mm, all tubular, yellowish-green, 4-lobed at apex. Receptacle conical, hollow, without scales. Achenes 1.2–1.5 mm, pale brown, slightly compressed, with 3–4 ribs on the adaxial face, abaxial face convex, truncate at apex; pappus an obscure rim or a unilateral tooth or lobed crown. Flowers 6–8. Little visited by insects. 2n = 18. (a) Subsp. discoidea M. matricarioides auct.; M. suaveolens (Pursh) Buchenau, non L.; Chamomilla suaveolens (Pursh) Rydb.; Santolina suaveolens Pursh; Tanacetum suaveolens (Pursh) Hook.; Lepidanthus suaveolens (Pursh) Nutt.; Akylopsis suaveolens (Pursh) Lehm.; Chamomilla discoidea (DC.) J. Gay Capitula 5–8 mm in diameter. Pappus minute and entire or absent. (b) Subsp. occidentalis (Greene) P. D. Sell M. occidentalis Greene; Chamomilla occidentalis (Greene) Rydb. Capitula 8–12 mm in diameter. Pappus with 2 dark brown, lateral lobes. Introduced. First recorded in 1871, it is common and increasing in waysides and waste places, especially where the soil is compacted on tracks, paths, farmyards and gateways. Throughout Great Britain and Ireland. The common plant is subsp. discoidea. It is probably native of north-east Asia, but occurs in North America from Alaska to Lower California and eastwards to Newfoundland and Missouri. It is a common weed throughout Europe. Subsp. occidentalis has been recorded for western Ireland. It occurs in North America from Middle California to southern Oregon. 80. Tripleurospermum Sch. Bip Matricaria sensu Flora Europaea Annual to perennial herbs. Stems erect, prostrate or ascending, leafy. Leaves alternate, 2- to 3-pinnatisect, the ultimate

80. Tripleurospermum segments linear. Capitula solitary at the ends of stems and branches. Involucral bracts in several rows, with a scarious margin. Inner flowers tubular, bisexual and 5-lobed at apex, the outer usually ligulate, 3-lobed at apex and female. Corolla of inner flowers yellow, of outer flowers white. Anthers without tails, apical appendage lanceolate-oblong, flat. Styles with linear-oblong branches; stigmatic area in 2 separate lines. Receptacle hemispherical to conical, more or less solid, without scales. Achenes more or less compressed laterally, with 3 conspicuous, smooth ribs on the adaxial face and 2 resin glands at the apex of the abaxial face; pappus a small corona, sometimes absent. Thirty-eight species mainly in Europe and Asia, and a few species in North Africa and North America. One species a widespread weed. In 1753 Linnaeus named Tripleurospermum maritimum subsp. inodorum as Matricaria chamomilla. In 1755, realising he had used the epithet chamomilla for the species which was not the wild chamomile, he changed it to the species he had earlier called M. recutita and called the M. chamomilla of 1753 M. inodora. M. chamomilla of 1755 is thus a later homonym of M. chamomilla 1753 and a synonym of M. recutita. M. inodora is an illegitimate superfluous name of M. chamomilla 1753. When transferred to the genus Chamomilla by K. Koch in 1843, it becomes legitimate, since M. chamomilla cannot be transferred as it would create an illegitimate tautonym. At subspecific rank it can be transferred to Tripleurospermum but at the rank of species in that genus it is illegitimate unless conserved. This does not affect the typification of Matricaria. Grime, J. P. et al. (1988). Comparative plant ecology. London. [T. inodorum.] Kay, Q. O. N. (1969). The origin and distribution of diploid and tetraploid Tripleurospermum inodorum (L.) Schultz Bip. Watsonia 7: 130–141. Kay, Q. O. N. (1972). Variation in sea mayweed (Tripleurospermum maritimum (L.) Koch) in the British Isles. Watsonia 9: 81–107. Kay, Q. O. N. (1991). Tripleurospermum inodoratum (L.) Sch.-Bip. in Biological flora of the British Isles. Jour. Ecol. 82: 681–697. Lester-Garland, L. V. (1921). The maritime forms of Matricaria inodora. Jour. Bot. (London) 59: 170–174. 1. Annual; achenes with ribs on adaxial face of mature achene clearly separated by at least one-third of their width, oil glands at upper end of abaxial face less than 1.5 times as long as broad and almost circular in outline. 1(d). maritimum subsp. inodorum 1. Biennial or perennial; achenes with ribs on adaxial face of mature achenes contiguous or slightly separated, oil glands at upper end of abaxial face usually more than 2. twice as long as wide 2. Involucral bracts broadly triangular, with a conspicuous, blackish-brown, scarious margin at least 0.4 mm wide 1(a). maritimum subsp. nigriceps 2. Involucral bracts oblong or narrowly triangular, with a pale brown or brown, scarious margin less than 0.3 mm 3. wide 3. Stems without purplish coloration or only near the base; leaves more open with segments 0.6–1.0 mm wide 1(b). maritimum subsp. maritimum

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3. Stem purplish-red to at least halfway up; leaves with dense segments 0.5–0.6 mm wide and appearing 1(c). maritimum subsp. vinicaule brush-like

1. T. maritimum (L.) W. D. J. Koch Scentless Mayweed Annual to short-lived perennial herb with fibrous roots. Stems often several, 10–80 cm, pale yellowish-green, flushed purple or the whole deep brownish-purple, procumbent, ascending or erect, faintly channelled, glabrous or sparsely hairy, leafy, branched often from base, sometimes woody at base. Leaves alternate, 1.5–6.0 × 1.0–4.0 cm, gradually decreasing in size upwards, soft or fleshy, yellowish-green, oblong, elliptical or ovate in outline, 2to 3-pinnatisect, the segments narrowly linear, obtuse to acute or bristle-tipped at apex, glabrous, with a few hairs or sparsely scaberulous, the lower petiolate, the upper sessile. Capitula numerous, 30–60 mm in diameter, solitary at the ends of branches and forming a loose corymb; peduncles channelled, glabrous or with an occasional hair. Involucral bracts in several rows, 5–8 × 1.5–2.0 mm, green with pale brown to blackish-brown scarious margins, oblong to triangular-ovate, irregularly rounded at apex, glabrous. Flowers of 2 kinds, scentless or slightly to quite strongly scented, the outer 10–20 mm, ligulate, white, entire or undulate at apex and female, the inner 4–5 mm, tubular and bisexual, yellow, with triangular lobes at apex. Receptacle hemispherical, pitted, without scales. Achenes 1.4–3.5 × 0.5–2.0 mm, oblanceolate, compressed, ribbed on adaxial face, with 2 resin glands at the apex of the abaxial face; pappus a short, truncate corona. Flowers 6–9, with occasional flowering throughout the year. Visited mainly by flies. (a) Subsp. nigriceps P. D. Sell Matricaria inodora var. phaeocephala auct.; Matricaria maritima var. phaeocephala auct.; Tripleurospermum maritimum subsp. phaeocephalum auct.; Matricaria maritima subsp. phaeocephala auct. Biennial to perennial herb. Stems yellowish-green, sometimes flushed reddish-purple, ascending or erect. Rosette leaves not toothbrush-like, the segments 0.6–1.0 mm wide, fleshy. Capitula 45–60 mm in diameter. Involucral bracts broadly triangular, with conspicuous blackish-brown, scarious margins at least 0.4 mm wide. Achenes 2.5–3.5 mm; ribs on adaxial face contiguous or slightly separated; oil glands more than twice as long as wide. 2n = 18. (b) Subsp. maritimum Matricaria inodora var. maritima (L.) Wahlenb.; Matricaria maritima L.; Pyrethrum maritimum (L.) Sm. Biennial to perennial herb. Stems yellowish-green, or reddish only at the base, ascending, prostrate or erect. Rosette leaves not toothbrush-like, the segments 0.6–1.0 mm wide, fleshy. Capitula 30–45(–60) mm in diameter. Involucral bracts oblong or narrowly triangular, with pale brown or brown, scarious margins not more than 0.3 mm wide. Achenes 2–3 mm; ribs on adaxial face contiguous or slightly separated; oil glands more than twice as long as wide. 2n = 18.

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(c) Subsp. vinicaule P. D. Sell Matricaria inodora var. salina auct.; T. maritimum var. salinum auct. Biennial to perennial herb. Stems purplish-red to at least halfway up, ascending or prostrate. Rosette leaves with close-packed segments 0.5–0.6 mm wide so that they appear toothbrush-like, fleshy. Capitula 30–45 mm in diameter. Involucral bracts oblong or narrowly triangular with pale brown or brown scarious margins not more than 0.3 mm wide. Achenes 1.8–2.5 mm; ribs on adaxial face contiguous or slightly separated; oil glands more than twice as long as wide. 2n = 18. (d) Subsp. inodorum (K. Koch) Hyl. ex Vaar. Matricaria chamomilla L. nom. rejic.; Matricaria inodora L. nom. illegit.; Chamomilla inodora K. Koch; Chrysanthemum inodorum L. nom. illegit.; Pyrethrum inodorum Moench nom. illegit.; T. inodorum Sch. Bip. nom. illegit.; Matricaria maritima subsp. inodora (K. Koch) A. R. Clapham; Matricaria perforata M´erat; T. perforatum (M´erat) Lainz; Dibothrospermum agreste Knaf Annual herb. Stems yellowish-green, sometimes reddish at the very base and rarely higher up, usually erect or ascending, rarely prostrate. Rosette leaves not toothbush-like with segments 0.6–1.0 mm wide, not usually fleshy. Capitula 30– 45 mm in diameter. Involucral bracts oblong or narrowly triangular, with pale brown scarious margins not more than 0.3 mm wide. Achenes 1.4–2.2 mm; ribs on adaxial face separated by at least one-third of their width; oil glands less than 1.5 times as long as wide and more or less circular. 2n= 18, 36. Native. Shingle, sand, cliffs, dunes, walls and waste land by the sea and a weed of cultivated and waste ground inland. Throughout Great Britain and Ireland but scarce in the mountains and central Ireland. Circumpolar; widely introduced as a weed elsewhere. A member of the Circumpolar Wide-boreal element. Subsp. nigriceps occurs on the coasts of northern Scotland, the Orkney Islands, Fair Isle and the Shetland Islands. It is also found in Faeroes and Iceland. Subsp. phaeocephalum (Rupr.) Hamet-Ahti, with which it has been confused, is more dwarf with oblong involucral bracts and is confined to the Arctic. Subsp. vinicaule occurs along the coast of southern England north to the Wash and north Cornwall and in the Channel Islands. It also occurs on the west coast of France, northern Portugal and Spain. The type of T. maritimum var. salinum (Wallr.) Kay is subsp. inodorum. Subsp. maritimum is found in the remaining central coastal areas of Great Britain and Ireland. It also occurs on the Continent in coastal areas of the North Sea and Baltic. Subsp. inodorum is the inland weed of cultivated and waste places. Formerly commonly, now rarely forming sheets of white on the stubble-lands in the autumn as do the coastal races on shingle. Hybrids or intermediates between subsp. inodorum and the coastal races occur not infrequently near the sea where there are suitable habitats. They are vigorous and more than 80 per cent fertile and populations showing intragression occur. All the races which occur in Great Britain and Ireland as far as is known are diploid. One example, which may have been introduced, of tetraploid subsp. inodorum, a plant of central

and eastern Europe not morphologically recognisable, has been recorded. 81. Cotula L. Leptinella Cass. Annual to perennial herbs. Stems procumbent to suberect. Leaves alternate or mostly clustered at nodes, entire to deeply pinnately divided. Capitula terminal and axillary, pedunculate. Involucral bracts in 2–3 rows. Flowers pedicillate, the pedicels persistent after the achenes have fallen, the inner tubular, 4-lobed at apex and bisexual or functionally male, the outer tubular and female, the corolla small or absent. Corolla white or yellow. Anthers with tails, apical appendage lanceolate-oblong, flat. Style with branches linear-oblong; stigmatic area in 2 separate lines. Receptacle flat, without scales. Achenes of female flowers compressed, those of bisexual flowers plano-convex; pappus absent. Fifty-five species mainly in the southern hemisphere, especially South Africa, but also in Australia, South America and East Africa, extending north to North Africa and Mexico. A few are widespread, weedy species. The genus Leptinella has been separated on the grounds that the female flowers possess a corolla and male and female are distinct, but some species attributed to it have bisexual inner flowers. C. bipinnata Thunb., C. pusilla Thunb., C. sororia DC., C. turbinata L. and C. zeyheri Fenzl ex Harvey have been recorded as wool casuals. 1. Leaves entire to irregularly pinnately divided with usually fewer than 6 pairs of segments and fleshy; capitula 6–12 1. coronopifolia mm in diameter; flowers bright yellow 1. Leaves pinnately or bipinnately divided with usually more than 6 pairs of segments and not succulent; capitula 2. 3–10 mm in diameter; flowers white or pale yellow. 2. Annual; capitula bisexual, the female outer flowers 2. australis without a corolla, the inner bisexual and white 2. Perennial, occurring in large patches; capitula unisexual, 3. all flowers with a dull yellow corolla 3. Stem, leaves and involucral bracts, glabrous or sparsely hairy; leaves dentate or shallowly lobed at least in upper part, sometimes deeply divided in lower part, but lobes 3. dioica entire or with short, blunt teeth 3. Stem, leaves and involucral bracts conspicuously hairy; leaves lobed nearly to midrib throughout, the upper margins of the lobes incise-dentate with acute teeth 4. squalida

1. C. coronopifolia L. Buttonweed C. integrifolia Hook. fil.; C. coronopifolia var. integrifolia (Hook. fil.) T. Kirk Annual or short-lived perennial herb with fibrous roots. Stems 8–30 cm, pale green often suffused reddish-brown, prostrate or ascending, striate, glabrous, branched, leafy, often rooting from the lower nodes. Leaves alternate, (1.0–)1.5–8.0 × 0.2–1.5 cm, bright green on upper surface, sometimes flushed brownish, paler beneath, fleshy, linear, lanceolate or spathulate or oblong-oblanceolate in outline, obtuse at apex, entire, dentate or irregularly pinnately lobed, the lobes usually fewer than 6 pairs, narrow and sometimes with narrow teeth, with a whitish, sheathing base, glabrous, but with scattered sunken glands. Flowering stem

81. Cotula slender, axillary and terminal. Capitula solitary, bisexual, 6–12 mm in diameter, hemispherical; peduncles exceeding the leaves, glabrous. Involucral bracts in 2–3 rows, 4–6 × 0.8–1.2 mm, greenish, edged reddish, oblong-elliptical, broadly obtuse at apex, glabrous. Flowers of 2 kinds, the outer (in 1 row) female, pedicillate, tubular, bright yellow, and with a corolla, the inner bisexual, sessile, with a yellow, flattened, tubular corolla with 4 lobes at apex. Receptacle flat, without scales. Achenes all flattened, hairy on inner surface and glabrous on outer, those of female flowers 1.5– 2.0 × 1.0–1.2 mm and with thick, corky, entire wings on the lateral angles, those of the bisexual flowers 1.1–1.2 × 0.5– 0.6 mm, and thinly ribbed on the lateral angles; pappus absent. Flowers 7–8. 2n = 20, 40. Introduced. Wet, usually saline habitats in coastal marshes. Naturalised in a marsh near Birkenhead, Cheshire since about 1880, and in Co. Cork in Ireland; a rare casual elsewhere. Possibly native in South Africa, Australia and South America, but now a cosmopolitan weed. 2. C. australis (Sieber ex Spreng.) Hook. fil. Annual Buttonweed Anacylus australis Sieber ex Spreng.; Strongylosperma australe (Sieber ex Spreng.) Less.; C. venosa Colenso Annual herb with fibrous roots. Stems 5–15(–40) cm, pale green, slender, decumbent to suberect, striate, with few to numerous, lax, long, spreading simple eglandular hairs, weakly diffusely branched, leafy. Leaves alternate, (0.5–)1.0–3.5(–5) × 0.2–2.0 cm, greenish, oblong or spathulate in outline, obtuse at apex, deeply 1- to 2pinnatisect, the segments usually more than 6 pairs, linear, entire and acute, sessile and amplexicaul, with few to numerous simple eglandular hairs and sunken glands. Flowering stem very slender, axillary or terminal with ascending-appressed simple eglandular hairs and shining glands. Capitula solitary, bisexual, 3–7 mm in diameter; peduncles with sparse to dense, antrorse simple eglandular hairs. Involucral bracts in 2 rows, 2–3 × 0.5–1.0 mm, oblong, obtuse at apex, glabrous or with a few, scattered simple eglandular hairs. Flowers of 2 kinds, the outer female in 2–3 rows, pedicellate and lacking a corolla, the inner bisexual, subsessile and with a pale yellow, tubular corolla. Receptacle flat, without scales. Achenes of female flowers 1.2–1.5 × 0.7–1.0 mm, flattened, hairy on both surfaces and the lateral angles with thin, glabrous, entire marginal wings, those of the bisexual flowers flattened on inner surface, 0.9– 1.1 × 0.4–0.5 mm, glabrous and thinly ribbed on the lateral angles; pappus absent. Flowers 7–8. 2n = 18, 20, 36, 40. Introduced. Frequent wool alien in arable or waste land. Scattered records in England and naturalised on a wall at Newton Abbot in Devonshire since 1946. Native of Australia and New Zealand. 3. C. dioica (Hook. fil.) Hook. fil. Hairless Leptinella Leptinella dioica Hook. fil. Perennial herb forming open to dense patches up to 30 cm in diameter. Stems up to 30 cm, yellowish-green, sometimes suffused brownish-purple, creeping and rooting at the nodes, slender to stout, glabrous or slightly simple hairy and with minute glandular hairs, branched, leafy. Leaves mostly

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clustered at the nodes, 2.0–3.0 × 0.5–0.9 cm, dull green on upper surface, paler beneath, more or less flaccid, spathulate or narrowly to broadly obovate in outline, obtuse at apex, crenate towards the apex, often becoming pinnatifid to pinnatisect, below, the 6–8 pairs of segments oblong, subacute and entire or with a few teeth on the lower pairs, glabrous or nearly so, but with sunken glands; petioles up to 15 mm, glabrous. Flowering stems axillary, longer or shorter than the leaves, more or less hairy. Capitula unisexual; male 4–7 mm in diameter, the involucral bracts in few to 2 rows, with broad purple scarious margins, subrotund-oblong and hairy; female 8–10 mm in diameter, the involucral bracts several in 3–4 rows and incurved and concealing the flowers. Flowers male funnel-shaped; female subconical, inflated at base, pale yellow and minutely 4-lobed at apex. Receptacle flat or conical without scales. Achenes 1.8–2.0 mm, obovoid, slightly curved, rounded on back, glabrous; pappus absent. Flowers 7–8. 2n = 260. Very variable in size and division of leaves and size of capitula. Introduced. Grown in gardens and becoming naturalised in lawns, but not often flowering. In a few, scattered localities. Native of New Zealand. 4. C. squalida (Hook. fil.) Hook. fil. Leptinella Leptinella squalida Hook. fil. Perennial herb forming patches up to 40 cm in diameter. Stems yellowish-green, often suffused brownishpurple, slender, branched, wiry, creeping, rooting, usually more or less hairy, leafy. Leaves mostly clustered at nodes, 3–5 × 0.5–1.0 cm, dull yellowish-green, narrowly obovate, oblanceolate or oblong in outline, acute at apex, pinnate in the lower part, pinnatisect to pinnatifid in the upper part, the pinnae 8–15 pairs, with the upper margins deeply incised into triangular, acute, apiculate teeth and the lower margins curved and usually entire, the lowest pinnae minute and narrow, with dense, long, pale, appressed simple eglandular hairs and sunken glands; petioles 10–20 mm, slender, with dense, long, pale simple eglandular hairs, with sheathing scarious base about 5 mm. Flowering stem 10–75 mm, slender, with rather long, pale, subappressed hairs. Capitula unisexual; male 4–5 mm in diameter, involucral bracts few and in 2 series, more or less purple-tipped and marginal, broadly obovate-oblong, obtuse at apex and silky hairy; female 6–9 mm in diameter, involucral bracts in 3– 4 rows, purple-margined, broadly obovate, rounded at apex and incurved over the flowers. Flowers male funnel-shaped, pale yellow and 4-lobed at apex; female ovoid, pale yellow and 4-lobed. Achenes obconical and almost trigonous, curved, grooved, glabrous; pappus absent. Flowers 7–9. 2n = 260. Introduced. A garden plant becoming naturalised in mown lawns and often not flowering. Scattered records in Great Britain and Ireland. Native of New Zealand. Tribe 7. Senecioneae Cass. Tribe Jacobaeae Dumort.; Tribe Tussilagineae Cass. Annual to perennial herbs, sometimes shrubs, rarely weak climbers. Leaves alternate or all basal. Capitula with all tubular or ligulate and tubular flowers. Flowers both ligulate

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and tubular usually yellow. Involucral bracts usually in 1 or 2 rows, often in 1 main row and 1 much shorter row, herbaceous. Receptacular scales absent. Pappus of 1–many rows of simple hairs. Contains 3 subtribes, 120 genera and about 3,200 species worldwide, but especially abundant in Central and South America and in Tropical and South Africa. 82. Senecio L. Annual to perennial herbs, rarely woody. Stems ascending to erect, leafy. Leaves alternate, subentire to dentate or deeply pinnately divided. Capitula solitary at the ends of branches or in corymbs. Involucral bracts in 1 main row with short supplementary ones. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female, or all flowers tubular. Corolla yellow, ligulate ones rarely white or purple. Anthers without tails, apical appendage ovate-lanceolate to oblong, flat; filament collar dilated. Style with oblong-linear, truncate branches; stigmatic areas in 2 separate lines. Receptacle flat, without scales. Achenes more or less cylindrical; pappus of simple or denticulate hairs. About 1,250 species worldwide but most species, about 500, in South America, and 350 in Africa; some species weedy and widespread. S. brasiliensis Less., S. glossanthus (Sond.) Belcher (Erechtites glossanthus Sond.), S. juniperinus L. fil., S. nudiusculus DC. and S. oederiifolius DC. have been recorded as wool casuals. Abbott, R. J., Ashton, P. & Forbes, D. G. (1992). Introgressive origin of the radiate groundsel, Senecio vulgaris var. hibernicus Syme; Aut-3 evidence. Heredity 68: 425–435. Abbott, R. J. & Forbes, D. G. (1993). Outcrossing rate and selfincompatibility in the colonizing species Senecio squalidus. Heredity 71: 155–159. Abbott, R. J., Ingram, R. & Noltie, H. J. (1983). Discovery of Senecio cambrensis Rosser in Edinburgh. Watsonia 14: 407– 408. Abbott, R. J., Irwin, J. A. & Ashton, P. A. (1992). Genetic diversity for esterases in the recently evolved stabilized introgressant, Senecio vulgaris var. hibernicus Syme, and its parental taxa S. vulgaris L. var. vulgaris and S. squalidus L. Heredity 68: 546– 556. Abbott, R. J. & Lowe, A. J. (1996). A review of hybridization and evolution in British Senecio in Hind, D. J. N. (Edit.) Compositae systematics: Proc. Int. Compositae Conf. Kew 1: 679–689. Abbott, R. J., James, J. K., Irwin, J. A. & Comes, H. P. (2000). Hybrid origin of the Oxford Ragwort, Senecio squalidus L. Watsonia 23: 123–138. Alexander, J. C. M. (1979). The Mediterranean species of Senecio sections Senecio and Delphinifolius. Notes Roy. Bot. Gard. Edinb. 37: 387–428. Allen, D. E. (1967). The taxonomy and nomenclature of the radiate variants of Senecio vulgaris L. Watsonia 6: 280–282. Ashton, P. A. & Abbott, R. J. (1992). Multiple origins and genetic diversity in the newly arisen allopolyploid species, Senecio cambrensis Rosser (Compositae). Heredity 68: 25–32. Ashton, P. A. & Abbott, R. J. (1992). Isozyme evidence and the origin of Senecio vulgaris (Compositae). Pl. Syst. Evol. 179: 167–174. Belcher, R. O. (1956). A revision of the Genus Erechtites (Compositae), with enquiries into Senecio and Arrhenechthites. Ann. Missouri Bot. Gard. 43: 1–85.

Brenan, J. P. M. (1948). Senecio squalidus L. × vulgaris L. Rep. Bot. Soc. Exch. Cl. Brit. Isles 13: 364. Brettell, R. I. S. & Leslie, A. C. (1978). Senecio squalidus L. × S. vulgaris L. in Cambridgeshire. Watsonia 12: 155–156. Burbidge, F. W. & Colgan, N. (1902). A new Senecio hybrid (× S. albescens). Jour. Bot. (London) 40: 401–406. Burton, R. M. (1979). Senecio cineraria DC. × S. erucifolius L. in Kent. Watsonia 12: 333–334. Campbell, J. M. & Abbott, R. J. (1976). Variability of outcrossing frequency in Senecio vulgaris. Heredity 36: 267–274. Chater, A. O. (1974). Taxonomic and nomenclatural notes on Senecio L. Bot. Jour. Linn. Soc. 68: 272–276. Comes, H. P. (1995). Senecio vulgaris L. subsp. denticulatus (O. F. Muell.) P. D. Sell and S. vulgaris L. subsp. vulgaris var. vulgaris on Jersey (Channel Islands). Watsonia 20: 185–194. Comes, H. P. & Abbott, R. J. (1998). The relative importance of historical events and gene flow on the population structure of a Mediterranean ragwort, Senecio gallicus (Asteraceae). Evolution 52: 355–367. Comes, H. P. & Kadereit, J. W. (1990). Aspects of hybridization between the closely related Senecio vulgaris L. and Senecio vernalis Waldst. & Kit. Flora 184: 381–388. Comes, H. P., Kadereit, J. W., Pohl, A. & Abbott, R. J. (1997). Chloroplast DNA and isozyme evidence on the evolution of Senecio vulgaris (Asteraceae). Pl. Syst. Evol. 206: 375–392. Crisp, P. & Jones, B. M. G. (1970). Senecio squalidus L., S. vulgaris L. and S. cambrensis Rosser. Watsonia 8: 47–48. Gibbs, P. E. (1971). Studies on synthetic hybrids of British species of Senecio. I. Senecio viscosus L. × S. vulgaris L. Trans. Bot. Soc. Edinb. 41: 213–218. Harper, J. L. & Wood, W. A. (1957). Senecio jacobaea L. in Biological flora of the British Isles. Jour. Ecol. 45: 617–637. Harris, S. A., (2002). Introduction of Oxford ragwort S. squalidus L. (Asteraceae), to the United Kingdom. Watsonia 24: 31–43. Harris, S. A. & Ingram, R. (1992). Molecular systematics of the genus Senecio L. II. The origin of S. vulgaris. Heredity 69: 112– 121. Haskell, G. (1953). Adaptation and the breeding system in groundsel. Genetica 26: 468–484. Hora, C. J. (1997). A handsome and vigorous hybrid Ragwort. B.S.B.I. News 77: 34. Hull, P. (1974). Self-fertilization and the distribution of the radiate form of Senecio vulgaris L. in central Scotland. Watsonia 10: 69–75. Hull, P. (1976). The influences of different degrees of interspecific hybridisation with Senecio squalidus on the frequency of two morphs of Senecio vulgaris. Heredity 36: 67–72. Ingram, R. & Noltie, H. J. (1984). Ray morphology and the origin of variability in Senecio cambrensis Rosser, a recently established polyploid species. New Phytol. 96: 601–607. Ingram, R. & Noltie, H. J. (1995). Senecio cambrensis in Biological flora of the British Isles. Jour. Ecol. 83: 537–546. Ingram, R., Weir, J. & Abbott, R. J. (1980). New evidence concerning the origin of inland radiate groundsel, S. vulgaris var. hibernicus Syme. New Phytol. 84: 543–546. Irwin, J. A. & Abbott, R. J. (1992). Morphometric and isozyme evidence for the hybrid origin of a new tetraploid radiate groundsel in York, England. Heredity 69: 431–439. Kadereit, J. W. (1983). Senecio vernalis Waldst. & Kit. in Britain. B.S.B.I. News 35: 8. Kadereit, J. W. (1984). The origin of Senecio vulgaris (Asteraceae). Pl. Syst. Evol. 145: 135–153. Kadereit, J. W. (1984). Studies on the biology of Senecio vulgaris L. ssp. denticulatus (O. F. Muell.) P. D. Sell. New Phytol. 97: 681–689.

82. Senecio Kadereit, J. W. (1984). Senecio × subnebrodensis Simk., an earlier name for Senecio squalidus L. × viscosus L. Watsonia 15: 36. Kadereit, J. W. & Sell, P. D. (1986). Variation in Senecio jacobaea L. (Asteraceae) in the British Isles. Watsonia 16: 21–23. Kent, D. H. (1956–1966). Senecio squalidus L. in the British Isles. 1. Early records (to 1877) in Proc. B.S.B.I. 2: 115–118 (1956); 2. The spread from Oxford (1879–1939) in Proc. B.S.B.I. 3: 375–379 (1960); 3. East Anglia in Trans. Norf. Norw. Nat. Soc. 18(5): 30–31 (1957); 4. Southern England (1940→) in Proc. B.S.B.I. 5: 210–213; 5. The Midlands (1940→) in Proc. B.S.B.I. 5: 214–216; 6. Northern England (1940→) in Proc. B.S.B.I. 5: 217–219; 7. Wales in Nature Wales 8: 175–178; 8. The recent spread in Scotland in Glasgow Nat. 18: 407–408; 9. Ireland in Irish Nat. Jour. 14: 203–204. Lowe, A. J. & Abbott, R. J. (1996). Origins of the new allopolyploid species Senecio cambrensis and its relationship to the Canary Islands endemic Senecio teneriffae (Asteraceae). Amer. Jour. Bot. 83: 1365–1372. Lowe, A. J. & Abbott, R. J. (2003). A new British species, Senecio eboracensis (Asteraceae), another hybrid derivative of S. vulgaris L. × S. squalidus L. Watsonia 24: 375–388. Mabberley, D. J. (1983). The Dusty Miller’s tale, or Senecio cineraria DC. restored. Watsonia 14: 279–280. Murphy, J. P. (1981). Senecio × albescens Burbidge & Colgan at Killiney, Co. Dublin: a seventy-eight-year-old population. Watsonia 13: 303–311. Richards, A. J. (1975). The inheritance and behaviour of the rayed gene complex in Senecio vulgaris. Heredity 34: 95–104. Rosser, E. M. (1955). A new British species of Senecio. Watsonia 3: 228–232. Sell, P. D. (1973). The rediscovery of the Fen Ragwort in Cambridgeshire. Nat. Cambridgeshire 16: 37. Stace, C. (1977). The origin of radiate Senecio vulgaris L. Heredity 99: 383–388. Trow, A. H. (1909). Some segregates of S. vulgaris, Linn. Rep. Bot. Soc. Exch. Cl. Brit. Isles 1908: 379–382. Walters, S. M. (1963). Senecio rupestris Waldst. & Kit. and Senecio squalidus L. Proc. B.S.B.I. 5: 382. Walters, S. M. (1974). The rediscovery of Senecio paludosus L. in Britain. Watsonia 10: 49–54. Wigginton, M. J. (Edit.) (1999). British red data book. Vol. 1. Vascular plants. Peterborough. [S. cambrensis and S. paludosus.] 2. 1. All flowers of capitulum tubular 1. Outer flowers of capitulum ligulate, although the ligules 6. sometimes small 3. 2. Lower cauline leaves entire to dentate but not lobed 4. 2. Lower cauline leaves lobed 3. Leaves glabrous or nearly so; inner involucral bracts 24. minimus (6–)8(–10) 3. Leaves arachnoid-tomentose beneath; inner involucral 25. quadridentatus bracts 11–13 4. Inner involucral bracts without blackish-brown tips 13(a). jacobaea subsp. dunensis 5. 4. Inner involucral bracts with blackish-brown tips 5. Stems up to 15 cm, often branched nearly to base, arachnoid hairy in upper part; leaves very fleshy, often arachnoid-hairy on under surface 19(a,i). vulgaris subsp. vulgaris var. crassifolius 5. Stems up to 45(–50) cm, with few to numerous branches, glabrous to sparsely arachnoid-hairy; leaves not or slightly fleshy, glabrous to sparsely arachnoid-hairy. 19(a,ii). vulgaris subsp. vulgaris var. vulgaris 7. 6. Ligules purple

6. 7. 7. 8. 8. 9. 9. 10. 10. 11. 11. 12. 12. 13. 13. 14. 14. 15. 15. 16. 16. 17. 17. 18. 18. 19.

19.

20. 20. 21. 21. 22. 22. 23. 23. 24. 24.

25. 25. 26.

26.

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9. Ligules yellow or white 3. glastifolius Leaves coarsely dentate 8. Leaves deeply pinnately divided 5. grandiflorus Perennial; ligules 10–15 mm 23. elegans Annual; ligules 6–8 mm 12. smithii Ligules white 10. Ligules yellow 11. Leaves entire to deeply toothed 21. Leaves pinnately divided into lobes 12. Leaves tomentose or arachnoid-hairy beneath 14. Leaves glabrous or nearly so beneath 11. doronicum Leaves subentire to sinuate-denticulate 13. Leaves entire to sharply serrate-dentate 2. pterophorus Capitula 10–15 mm in diameter 10. paludosus Capitula 30–40 mm in diameter Leaves linear and entire, usually less than 5 mm wide 6. inaequidens 15. Leaves more than 5 mm wide Peduncles and involucral bracts without glandular hairs 16. 20. Peduncles and involucral bracts with glandular hairs 17. Leaves entire 18. Leaves dentate Leaves 8–30 cm; peduncles arachnoid-hairy at first 8. doria 18. Leaves 2–12(–15) cm; peduncles glabrous 16(d). squalidus subsp. aethnensis Leaves entire 19. Leaves dentate Leaves green; upper cauline leaves markedly auriculate-semiamplexicaul 16(a,vi). squalidus subsp. rupestris var. subinteger Leaves glaucous-green; upper cauline leaves more narrowed at base and not obviously auricled 16(c). squalidus subsp. squalidus Leaves lanceolate, ovate, oblong or oblong-lanceolate; 7. fluviatilis ligulate flowers 6–8, 8–12 mm Leaves linear to oblong-lanceolate; ligulate flowers 5–6, 9. ovatus subsp. alpestris 12–15 mm 22. Involucral bracts with glandular hairs 25. Involucral bracts without glandular hairs 22. viscosus Plant viscid; achenes glabrous 23. Plant not or scarcely viscid; achenes hairy Leaf segments narrowly linear; peduncles glabrous 4. pinnulatus Leaf segments much broader; peduncles glandular-hairy 24. Stems up to 20 cm; capitula solitary or in a small 20(i). sylvaticus var. nanus corymb at the top of the stem Stems up to 70 cm; capitula numerous in a large, flat-topped terminal corymb 20(ii). sylvaticus var. sylvaticus 1. ambiguus Plant tomentose throughout 26. Plant not tomentose throughout Annual to perennial, usually easily uprooted, without a thick stock or short rhizome; at least outer involucral 27. bracts with black tips Biennial or perennial, firmly rooted, usually with a very short, thick stock and sometimes also a short rhizome; 36. involucral bracts without black tips

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27. Capitula bell-shaped in flower; ligules usually more 28. than 8 mm 33. 27. Capitula cylindrical in flower; ligules less than 8 mm 29. 28. Leaves usually not more than 2.5 cm wide 30. 28. At least some leaves more than 2.5 cm wide 29. Leaves with narrow, acute lobes, glabrous or with an occasional hair beneath 16(a,i). squalidus subsp. rupestris var. calabricus 29. Leaves undulate with obtuse, spreading lobes, usually 21. vernalis arachnoid-hairy beneath 30. At least some leaves bipinnatisect 16(a,iii). squalidus subsp. rupestris var. bipinnatifidus 31. 30. Leaves simply pinnatisect and or deeply laciniate 31. Lower leaves not pinnatisect more than halfway to the midrib 16(a,iv). squalidus subsp. rupestris var. rupestris 32. 31. Lower leaves pinnatisect or laciniate almost to midrib 32. Leaves regularly pinnatisect 16(a,ii). squalidus subsp. rupestris var. pinnatifidus 32. Leaves regularly pinnatisect and laciniate with long narrow teeth 16(a,v). squalidus subsp. rupestris var. laciniatus 34. 33. Achenes 2.5–3.5 mm 35. 33. Achenes less than 2.5 mm 17. cambrensis 34. Ligules (8–)13(–15), 2.0–2.5 mm wide 18. eboracensis 34. Ligules 8(–10), 1.2–1.7 mm wide 35. Ligules 3.5–5.5 mm 19(a,iii). vulgaris subsp. vulgaris var. hibernicus 35. Ligules 2.3–3.0 mm, becoming revolute after flowering 19(b). vulgaris subsp. denticulatus 37. 36. Achenes glabrous 40. 36. At least the inner achenes hairy 37. Stems up to 50 cm; inflorescence fairly congested; capitula 25–45 mm in diameter 14(a). aquaticus subsp. ornatus 37. Stems up to 130 cm; inflorescence open; capitula 15–25 38. mm in diameter 38. Capitula 15–20 mm in diameter 14(c). aquaticus subsp. erraticus 39. 38. Capitula 20–25 mm in diameter 39. Lower leaves dentate, but not lobed 14(b,i). aquaticus subsp. aquaticus var. barbareifolius 39. Lower leaves with a large terminal segment and spreading lateral segments 14(b,ii). aquaticus subsp. aquaticus var. aquaticus 41. 40. All achenes shortly hairy 44. 40. Achenes of ligulate flowers glabrous 41. Leaves green and glabrous or nearly so beneath 15(i). erucifolius var. viridulus 42. 41. Leaves greyish or whitish hairy beneath 42. Leaves with a large terminal lobe much bigger than the 15(iv). erucifolius var. latilobus lateral lobes 42. Leaves with terminal lobe as big as or not much larger 43. than lateral lobes 43. Slender plant; leaves lanceolate, oblong-lanceolate or ovate in outline, both terminal and lateral lobes linear, greyish or whitish hairy beneath, often densely so; 15(ii). erucifolius var. erucifolius capitula small 43. Tall, robust plants; leaves mostly broadly ovate in outline, the segments often widely oblong, greyish hairy 15(iii). erucifolius var. communis beneath; capitula large

44. Stems markedly swollen below the basal leaves, up to 40(–45) cm; internodes rather short; inflorescence dense; ligules 5–7(–9) × 1–2 mm 13(b,i). jacobaea subsp. jacobaea var. condensatus 44. Stems not swollen below, basal leaves, up to 150 cm; leaves widely spaced; inflorescence open; ligules 43. 7–10 × 1.7–3.0 mm 45. Stem usually below 100 cm; leaves with narrow segments; ligules 7–9 × 1.7–3.0 mm, early flowering 13(b,ii). jacobaea subsp. jacobaea var. jacobaea 45. Stem often over 100 cm and deep purple; leaves with broad segments; ligules 8–10 × 2–3 mm; late-flowering 13(b,iii). jacobaea subsp. jacobaea var. nemorosus

Section 1. Incanae (DC.) Hoffm. Woolly perennial herbs often with a woody stock. Leaves more or less pinnately divided. Ligules yellow. Achenes hairy. 1. S. ambiguus (Biv.) DC. Silver Ragwort Cineraria ambigua Biv.; S. cineraria DC.; Cineraria bicolor Willd.; S. bicolor (Willd.) Tod., non Viv.; Cineraria nebrodensis Guss.; S. bicolor subsp. cineraria (DC.) Chater; S. bicolor subsp. nebrodensis (Guss.) Chater; Othonna maritima L., non S. maritima L. fil.; Jacobaea tomentosa Moench nom. illegit. Perennial herb with a stout rootstock. Stems 25–100 cm, erect, robust and woody at least at base, striate, greyishwhite woolly, branched above, leafy. Leaves alternate, 3– 15 × 1.5–7.0 cm, either equally leafy throughout the stem or mostly crowded towards the base of the flowering stems and forming rosettes on the non-flowering shoots, yellowishgreen to greyish-green on upper surface, greyish-white to white beneath, ovate to lanceolate in outline, obtuse at apex, pinnatifid, the segments irregular in shape with blunt, rounded teeth, arachnoid-hairy to nearly glabrous on upper surface, whitish tomentose beneath. Capitula numerous, 10–15 mm in diameter, rounded at base, in lax to dense, compound corymbs; peduncles whitish-woolly. Involucral bracts in 2 rows, the inner 5–8 × 0.8–1.0 mm, linear and obtuse at apex, the outer up to 5, 1–2 mm, linear and acute at apex, all usually whitish woolly, sometimes becoming nearly glabrous. Flowers of 2 kinds; outer 3–6 mm, ligulate and the ligules yellow and 3-lobed at apex, the inner tubular, 5-lobed at apex and brownish-yellow. Achenes 1.8–2.0 mm, subcylindrical, brown, ribbed, minutely hairy; pappus of numerous, brownish simple eglandular hairs. Flowers 6–8. 2n = 40. On native material we have found it difficult to accept two species, S. ambiguus and S. cineraria. It seems better to have one species occurring throughout the Mediterranean with numerous subspecies, probably more than have been described. Plants naturalised in Great Britain and Ireland are of more than one kind. The common kind naturalised on cliffs and commonly grown in gardens, with large stem leaves and woolly heads, does not seem to be a described variant, although some plants from the Mediterranean match it. Others with a cluster of leaves below and rather dwarf match S. cineraria DC. sens. strict.

82. Senecio Introduced. Naturalised on cliffs, walls, shingle and waste ground, mainly near the sea. South and south-west England, Wales, Channel Islands and Co. Dublin in Ireland. Native of the Mediterranean region. × erucifolius = S. × thuretii Briq. & Cavill. S. × patersonianus R. M. Burton Sometimes with stolons, woolly-hairy but less so than in S. ambiguus, leaves thinner than in S. ambiguus and the outer involucral bracts about one-third as long as the inner ones, not less than one-quarter as in S. × albescens. In two localities in Kent, one south of Deal, the other south of Walmer. × jacobaea = S. × albescens Burb. & Colgan This hybrid is intermediate in hairiness, leaf shape and habit, but has hairy achenes of the tubular flowers as in S. jacobaea. It is fertile and backcrosses. 2n = 40. Throughout much of the range of S. ambiguus in Great Britain and in Co. Dublin in Ireland. Section 2. Rigidi Harv. Perennial herbs sometimes with a woody stock, glabrous or slightly arachnoid-hairy. Leaves subentire or dentate. Ligules yellow or purple. Achenes hairy. 2. S. pterophorus DC. Shoddy Ragwort S. polyanthemus var. subserratus DC.; S. pterophorus var. apterus Harv. Perennial herb with a woody stock. Stems 30–150 cm, erect, pale yellowish-green, sometimes tinted brownish-purple, striate, woody at base, thinly arachnoid-hairy to nearly glabrous, branched in upper part, very leafy. Leaves alternate, numerous, becoming smaller upwards, 1–15 × 0.2– 1.5 cm, medium green on upper surface, greyish beneath, linear to linear-lanceolate or elongate-lanceolate, acute at apex, subentire to deeply dentate-serrate, narrowed to a sessile base, glabrous to sparsely arachnoid-hairy on upper surface, greyish tomentose beneath. Capitula 10–15 mm in diameter, rounded at base, in a spreading, corymbose panicle; peduncles pale yellowish-green, slightly angled, woolly in places. Involucral bracts in 2 rows; inner 4–6 × 0.4– 0.5 mm, pale yellowish-green, linear, acute at apex; outer several, 2–4 × 0.4–0.5 mm, pale yellowish-green, linearlanceolate, acute at apex; all glabrous or woolly at tips and base. Flowers of 2 kinds, the outer 11–13, 7–9 mm, ligulate, the ligules yellow and minutely 3-lobed at apex, the inner 4–5 mm, tubular, 5-lobed apex and yellow. Receptacle flat, without scales. Achenes 1.4–1.5 mm, pale brown, cylindrical, striate; pappus of numerous, whitish simple eglandular hairs. Flowers 7–9. 2n = 20. Introduced. An occasional wool alien in fields and waste places. A few scattered records in Great Britain. Native of South Africa and introduced as a weed in southern Australia. 3. S. glastifolius L. fil. Woad-leaved Ragwort Perennial herb with a tap-root. Stem up to 100 cm, pale yellowish-green, tinted brownish-purple, erect, woody towards the base, ridged, glabrous, branched above, leafy. Leaves alternate, 1.5–7.5 × 0.4–2.5 cm, pale

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yellowish-green on upper surface, paler beneath, rigid, oblong to oblong-lanceolate, more or less acute at apex, coarsely and unequally dentate, semiamplexicaul at base, glabrous. Capitula 15–20 mm in diameter, subtruncate at base, in lax corymbs; peduncles slender, striate, glabrous, with several linear bracts. Involucral bracts in 2 rows; inner 7–9 × 0.5–0.7 mm, green with a pale margin, narrowly linear, subacute at apex; outer 4–5 × 0.5–0.6 mm, green with a broad pale margin, narrowly linear-lanceolate, subacute at apex; all glabrous. Flowers of 2 kinds, the outer ligulate, the ligules 12–18 mm, 3-lobed at apex, purple and spreading; the inner tubular, 5-lobed at apex and yellow. Receptacle flat; without scales. Achenes 3.5–4.0 mm, cylindrical, striate, shortly hairy; pappus of numerous, whitish simple eglandular hairs. Flowers 7–8. Introduced. Garden escape, naturalised in open woodland and on a rubbish tip in the Isles of Scilly. Native of South Africa. Section 3. Leptolobi Harv. Glabrous perennial herbs. Leaves deeply pinnately divided. Ligules yellow or purple. Achenes hairy. 4. S. pinnulatus Thunb. Narrow-lobed Ragwort Perennial herb with a tap-root. Stem 25–100 cm, yellowishgreen sometimes suffused brownish-purple, erect, striate, woody at base, glabrous, much-branched, leafy. Leaves alternate, 4–15 × 1–8 cm, yellowish-green, broadly ovate in outline, acute at apex, distinctly pinnatipartite, the segments 3–5 pairs, narrowly linear, acute at apex and revolute and entire or with a remote small tooth on margin, glabrous or nearly so, often auricled at base, sessile. Capitula 12– 16 mm in diameter, narrowed to base, in loose paniculate corymbs; peduncles slender, glabrous, with linear bracts. Involucral bracts in 2 rows, the inner 4.5–5.0 × 0.8–1.0 mm, pale yellowish-green with a membranous margin and black tips, linear, obtuse at apex, glabrous on surface and with a tuft of glandular hairs at apex, the outer 0.5–0.8 × 0.3– 0.5 mm, lanceolate, acute at apex, glandular-hairy, especially on margins. Flowers of 2 kinds, the outer 8–10, 7–8 mm, ligulate, the ligules 3-lobed at apex and yellow, the inner tubular, 5-lobed at apex, and yellow. Receptacle flat, without scales. Achenes 2.0–2.5 mm, pale brown, cylindrical, angled, densely appressed-hairy; pappus 4–5 mm, of numerous, whitish simple eglandular hairs. Flowers 7–9. Introduced. A wool casual in fields and waste places. A few, scattered records in Great Britain. Native of South Africa. 5. S. grandiflorus P. J. Bergius Purple Ragwort S. venustus Aiton Perennial herb with a tap-root. Stem 50–150 cm, pale yellowish-green, often suffused with brownish-purple, erect, striate, glabrous, leafy in the lower part, branched above. Leaves alternate, 3–15 × 1–8 cm, pale green, very broadly ovate in outline, acute at apex, deeply pinnately lobed, the lobes narrowly linear, entire or denticulate and very acute at apex, sessile and semiamplexicaul at base, glabrous. Capitula 20–30 mm in diameter, rounded or subtruncate at base, in loose spreading panicles of partial

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corymbs; peduncles pale green, striate and glabrous. Involucral bracts in 2 rows; inner 7–8 × 0.7–1.0 mm, green with a pale margin, linear, narrowed to an obtuse apex; outer several, 3.5–4.0 × 0.7–0.8 mm, green with a pale margin, linear, obtuse at apex; all glabrous. Flowers of 2 kinds, the outer 10–15 mm, ligulate, the ligules purple and 3-lobed at apex, the inner tubular, 5-lobed at apex and yellow. Receptacle flat, without scales. Achenes 3.5–4.0 mm, black, cylindrical, furrowed, with minute hairs along the furrows; pappus of numerous, whitish simple eglandular hairs. Flowers 7–8. 2n = 40. Introduced. Garden escape naturalised in grassland at Fermain Bay and Hauteville in Guernsey. Native of South Africa. Section 4. Fruticulosi DC. Suffruticose perennial herbs, glabrous or nearly so. Leaves usually narrow and entire. Ligules yellow. Achenes hairy. 6. S. inaequidens DC. Narrow-leaved Ragwort S. lautus auct.; S. burchellii auct. Suffruticose perennial herb with stout main roots and fibrous side-roots. Stems 20–80 cm, pale yellowish-green, erect, markedly striate, glabrous or nearly so, leafy, widely branched. Leaves alternate, 2–6(–10) cm × 1–3(–5) mm, dull medium green on upper surface, paler beneath, longlinear, subobtuse to acute at apex, entire or remotely denticulate, with incurved margins and prominent midrib, sessile and often more or less amplexicaul, glabrous. Capitulum 10–25 mm in diameter, campanulate, rounded at base, in lax corymbs; peduncles pale yellowish-green, glabrous. Involucral bracts in 2 rows; inner 6–8 × 0.5–0.6 mm, pale yellowish-green, tipped blackish, linear, obtuse with a tuft of hairs at apex, otherwise glabrous; outer 2–3 × 0.8–1.0 mm, green, tipped blackish-brown or all blackish-brown, glabrous. Flowers of 2 kinds; outer ligulate, 7–15, the ligules 5–10 mm, 3-lobed at apex and yellow; inner tubular, 6–10 mm, 5-lobed at apex and dirty brownish-yellow. Receptacle flat, without scales. Achenes 2.0–2.5 mm, subcylindrical, hispid between the ribs; pappus whitish, of numerous simple eglandular hairs. Flowers 6–10. 2n = 40. Introduced. Frequent wool alien and naturalised on a sandy beach in Kent. Scattered records in England and Scotland. Native of South Africa. Naturalised in Belgium, Holland, France and Italy. Section 5. Doria (Fabr.) Rchb. Doria Fabr. Perennial, nearly glabrous herbs, sometimes with stolons. Leaves undivided. Ligules 8–12, yellow. Achenes glabrous or minutely hairy. 7. S. fluviatilis Wallr. Broad-leaved Ragwort S. salicetorum Godr.; S. transsilvanicus Schur; S. grandidissimus Schur; S. sarracenicus auct.; S. repens Stokes nom. illegit. Perennial herb with a short, creeping stock and long, fleshy stolons up to 60 cm. Stems 80–150(–200) cm, pale

green, sometimes flushed with brownish-purple, markedly ridged, erect, sometimes glabrous below, usually slightly arachnoid-hairy and with short, pale glandular hairs above, much branched above, leafy. Leaves alternate, 4–20 × 1–6 cm, rather dark green on upper surface, much paler beneath, gradually decreasing in size upwards, lanceolate, ovate, oblong or oblong-lanceolate, acute at apex, shortly serrulate, the teeth acute, incurved and cartilaginous, narrowed to the sessile, semiamplexicaul base, nearly glabrous or with minute glandular hairs. Capitula numerous, 15–30 mm in diameter, ovoid-campanulate, rounded at base, in compound corymbs; peduncles slender, ridged, with numerous, short, pale glandular hairs. Involucral bracts in 2 rows, the inner 5–8 × 1.2–1.5 mm, brownish-green with scarious margins and brown-tipped at apex, oblong to oblong-lanceolate and narrowed to an obtuse apex, the outer about 5, up to half as long as inner, linear, acute at apex and lax; all with numerous, short, pale glandular hairs. Flowers of 2 kinds, the outer 6–8, 8–12 mm, ligulate, the ligules bright yellow, minutely 3-lobed at apex and spreading, the inner 5–6 mm, tubular, brownish-yellow, with 5 narrow lobes at apex. Receptacle flat, with dentate pits, without scales. Achenes 3–4 mm, cylindrical, ribbed, glabrous; pappus 6–7 mm, of numerous, whitish, simple eglandular hairs. Flowers 7–9. Visited by bees and flies. 2n = 40. Introduced. Naturalised by streams and in fens and swampy ground. Scattered localities in Great Britain and Ireland north to central Scotland; formerly more common. Native of central and south Europe from Spain, northern Italy and the north Balkan peninsula and southern Russia north to Holland and Estonia, and in Siberia. It has a Eurosiberian Temperate distribution. 8. S. doria L. Golden Ragwort Jacobaea doria (L.) P. Gaertn., B. Mey. & Scherb. Perennial slightly foetid herb with an oblique, praemorse stock. Stems 40–150 cm, pale to medium yellowish-green, markedly striate, erect, hollow, terete, glabrous, branched above, leafy. Leaves alternate, numerous, the basal dying before anthesis, the cauline 6–30 × 2.5–9.0 cm, gradually decreasing in size upwards, dull, metallic, medium green with a pale midrib on upper surface, bluish-green beneath with a prominent, pale midrib, narrowly elliptical-oblong, oblong-lanceolate or panduriform and the upper ovate, rounded-obtuse to acute or acuminate at apex, denticulate, the teeth spreading and obtuse, the lower gradually attenuate and more or less decurrent and semiamplexicaul, the upper rounded-cordate and amplexicaul, glabrous. Capitula 12–25 mm in diameter, narrow, rounded at base, in large, terminal compound corymbs with some short, lower branches; peduncles short, at first arachnoid-hairy. Involucral bracts in 2 rows and descending down the peduncles, the inner 5–8 × about 1.0 mm, pale yellowish-green, linear, obtuse with a dark spot at apex and a tiny tuft of hairs, the outer 2.0–2.5 × about 1.0 mm, linear-lanceolate, acute with a dark spot at apex, sometimes with a little arachnoid hair. Flowers of 2 kinds the outer 4–6, 7–10 mm, ligulate, the ligules minutely 3-lobed at apex and bright golden yellow, the inner 5–6 mm, tubular, golden yellow and narrowly

82. Senecio 5-lobed at apex. Receptacle flat, without scales. Achenes 3–4 mm, cylindrical, ribbed, glabrous; pappus 9–12 mm, of numerous, whitish simple eglandular hairs. Flowers 6–8. Visited by bees and flies. 2n = 40. Introduced. Naturalised by streams and in wet meadows. Kirkcudbrightshire, Isle of Man and Co. Offaly in Ireland; perhaps elsewhere as it has been in the past. Native of south and east Europe and North Africa. 9. S. ovatus (P. Gaertn., B. Mey. & Scherb.) Willd. Wood Ragwort Jacobaea ovata P. Gaertn., B.Mey. & Scherb. Perennial herb with a shortly creeping stock. Stems up to 150 cm, erect, pale yellowish-green, ridged, glabrous or minutely and sparsely hairy, branched above, leafy. Leaves alternate, 3–15 × 0.2–3.0 cm, yellowish-green on upper surface, slightly paler beneath, oblong-lanceolate to linear, long-acute at apex, entire or sparsely and minutely denticulate, gradually narrowed and semiamplexicaul at base, the lower shortly petiolate, glabrous or nearly so, the midrib prominent beneath. Capitula 10–20 mm in diameter, narrowed to a rounded base, in compact corymbs with longer branches from upper leaf-axils; peduncles slender, striate, with unequal glandular and simple eglandular hairs. Involucral bracts in 2 rows; inner 5–7 × 0.5–0.7 mm, green with a broad scarious margin, linear-lanceolate, obtuse at apex, glandular-hairy with a tuft of hairs at apex; outer 2–3 × 0.2– 0.4 mm, green with a pale margin, linear, acute at apex, glandular-hairy. Flowers of 2 kinds, the outer 5–6 ligulate, the ligules 12–15 mm, 3-lobed at apex and yellow, the inner tubular, 5-lobed at apex and yellow. Receptacle flat, without scales. Achenes 2.5–3.0 mm, brown, cylindrical, ribbed, minutely hairy; pappus 5–7 mm, whitish, of simple hairs. Flowers 6–7. 2n = 40. Introduced. Naturalised in damp, shady places. Recent records are from Lancashire and Yorkshire and there are scattered old records from elsewhere. All our plants are subsp. alpestris (Gaudin) Herborg (S. alpestris Gaudin; S. sarracenicus auct.; S. fuchsii auct.) which is native of the Alps. Section 6. Crociseris Rchb. Perennial, nearly glabrous herbs. Leaves undivided, but often denticulate. Ligules (10–)12–25, yellow. Achenes glabrous. 10. S. paludosus L. Fen Ragwort Cineraria serratifolia Gray nom. illegit. Perennial herb with a shortly creeping stock. Stems 80–200 cm, erect, robust, markedly striate, slightly arachnoid-hairy, branched in the upper part, leafy. Leaves alternate, gradually becoming smaller upwards, 7–18 × 0.5–2.0 cm, dull medium green on upper surface, paler and greyish-green beneath, linear, linear-lanceolate or linear-elliptical, longacute with a very narrow apex, sharply serrate or serrulate, the lower narrowed to a short, broad stalk-like base, the upper sessile and shortly amplexicaul, all more or less glabrous on the upper surface, arachnoid-hairy beneath. Capitula numerous, 30–40 mm in diameter, campanulate, in a large, simple terminal corymb; peduncles arachnoid-hairy

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and with numerous linear bracts. Involucral bracts in 2 rows, the outer numerous and about half as long as the inner, narrowly linear, acute at apex and glabrous or arachnoid-hairy, the inner 6–8 × 0.8–1.1 mm, linear-lanceolate, subobtuse at apex, glabrous or arachnoid-hairy with a little tuft of hairs at the apex. Flowers of 2 kinds, the outer 10–16, 15–20 mm, ligulate, the ligules bright yellow, spreading and with 3 small lobes at apex, the inner 8–10 mm, tubular, 5-lobed at apex and deep treacly yellow. Receptacle flat, without scales. Achenes 2.5–3.0 mm, cylindrical, ribbed, glabrous; pappus 7–9 mm, whitish, of simple hairs. Flowers 5–7. Visited by various insects. 2n = 40. Native. Formerly in fen-ditches in Lincolnshire, Cambridgeshire, Norfolk and Suffolk until 1857. A small population was discovered in a fen-ditch near Ely in Cambridgeshire in 1972 (cf. P. D. Sell, 1973; S. M. Walters, 1974). Most of the seeds produced were sterile, but 21 seeds were germinated in the Cambridge Botanic Garden. Much of central Europe from Spain, northern Italy and the Balkan peninsula northwards to Holland, southern Sweden and northern Russia; northern Asia. A member of the Eurosiberian Temperate element. 11. S. doronicum (L.) L. Chamois Ragwort Solidago doronicum L. Perennial herb with a short stock. Stem 20–60 cm, pale yellowish-green, erect, rigid, solid, striate, with few to numerous, short to long, pale, wavy simple eglandular hairs, branched in the inflorescence, leafy. Leaves alternate, 3– 10 × 2–4 cm, thick and coriaceous, dull medium yellowishgreen on upper surface, paler beneath, with numerous very short glandular and simple eglandular hairs or arachnoidhairy; basal and lower cauline narrowly elliptical, obtuse to subacute at apex, subentire to sinuate-denticulate, longattenuate at base into a winged petiole; upper cauline lanceolate to oblong-lanceolate, acute at apex, sinuatedenticulate, sessile and semiamplexicaul. Capitula 40–50 mm in diameter, with a broad, rounded base, solitary or several at the end of stems or branches; peduncles long, with arachnoid and very short glandular hairs. Involucral bracts in 2 rows; inner 10–15 × 1.0–1.2 mm, green with a narrow, pale margin, narrowly linear-lanceolate, more or less acute at apex; outer as long as and similar to inner; all with varying amounts of arachnoid and minute glandular hairs. Flowers of 2 kinds, the outer ligulate, the ligules 12–25 mm, 3-lobed at apex and golden yellow, the inner tubular, 5-lobed at apex and golden yellow. Receptacle flat, without scales. Achenes 6–8 mm, pale brown, cylindrical, glabrous; pappus 7–8 mm, whitish, of simple hairs. Flowers 6–7. 2n = 40, 80. This is an aggregate species showing much variation in size, leaf-toothing and indumentum. Introduced. Naturalised on river banks in Perthshire since 1985. No specimens of this plant have been traced. An old naturalised record for Yorkshire is referred to subsp. transylvanicus (Boiss.) Nyman which is not accepted by Flora Europaea. The description given should enable the whole aggregate to be run down, but the segregates need revision. The group is native to the mountains of Europe.

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Section 7. Hualtata Cabrera Perennial herbs with arachnoid hairs. Leaves undivided, but dentate. Ligules white. Achenes glabrous. 12. S. smithii DC. Magellan Ragwort S. verbascifolius Hombr. & Jacq.; Cineraria gigantea Sm., non S. giganteus Desf. Robust perennial herb with a short, robust rhizome. Stem 60–120 cm, erect, stout, striate, arachnoid-hairy, branched above, leafy. Leaves alternate, pale glossy green, thick; basal 10–25(–35) × 9–14(–20) cm, ovate, ovate-oblong or oblong, obtuse at apex, dentate with acute, triangular teeth, cuneate to cordate at base, the petiole about equalling the lamina, arachnoid-hairy, sometimes broadly winged in the lower half, sometimes the wing forming a broadly ovate, fibrillous-dentate flange; upper smaller, ovate to lanceolate, acute to acuminate at apex, fimbriate-dentate at least in the lower half, sessile and amplexicaul; all more or less lanatearachnoid, but often becoming nearly glabrous on upper surface. Capitula 30–50 mm in diameter, subtruncate at base, in a lax to dense corymb; peduncles arachnoid-hairy, mixed with glandular hairs. Involucral bracts in 2 rows, the inner 8–14 × 0.8–1.0 mm, green in the centre with pale margins, linear, gradually narrowed to an acute apex and arachnoidhairy particularly in the upper half, the outer 8–20, over half as long as inner, similar but even narrower. Flowers of 2 kinds, the outer 15–20, ligulate, the ligules 15–25 mm, white and 3-lobed at apex, the inner tubular, yellow and 5-lobed at apex. Receptacle flat, without scales. Achenes 4.5–6.0 mm, cylindrical, deeply grooved, glabrous; pappus 7–10 mm, of whitish simple eglandular hairs. Flowers 7–8. 2n = 40. Introduced. Naturalised in grassy places and by streams. The extreme north of Caithness, the Orkney Islands and particularly the Shetland Islands. Views have been put forward that it was brought back by local men returning from Antarctic whaling, and others that it is connected with Shetlanders who tended sheep farms in the Falkland Islands, but it is likely most material arrived through normal gardening and found the climate to its liking. Native of Tierra del Fuego and southern Chile and Argentina. Section 8. Jacobaea (Mill.) Dumort. Jacobaea Mill. Perennial or biennial herbs nearly glabrous or with arachnoid hair. Leaves more or less pinnately divided. Ligules yellow. Achenes glabrous or hairy. 13. S. jacobaea L. Common Ragwort S. laciniatus Gray, non Bertol.; Jacobaea nemorosa Fourr.; Jacobaea vulgaris Gaertn. Biennial to perennial herb with a short, erect stock, a tap-root and fleshy side-roots. Stems 20–150 cm, pale yellowish-green, often suffused purple or brownishpurple especially below, erect, ridged, glabrous or slightly arachnoid-hairy, branched above, sometimes lower down, leafy. Leaves alternate, dark green on upper surface, paler beneath, glabrous or arachnoid-hairy beneath; basal 7– 30 × 2–6 cm, in a rosette, usually dying before anthe-

sis, obovate or oblong in outline, obtuse at apex, lyratepinnatifid, with a large, ovate, obtuse, dentate terminal lobe and 0–6 pairs of much smaller, oblong lateral lobes which are sinuate toothed or pinnatifid and petiolate; lower cauline similar and petiolate, the upper pinnatifid to bipinnatifid with a narrower terminal lobe and sessile and semiamplexicaul. Capitula 15–25 mm in diameter, rounded below, in a large, flat-topped, compound corymb; peduncles with numerous to dense arachnoid hairs. Involucral bracts in 2 rows, the inner 5–6 × 1.0–1.5 mm, green, with lines in centre and pale margins, linear-lanceolate, obtuse at apex and with a few minute hairs, the outer 2–5, a quarter to half as long as inner, linear, obtuse or acute at apex. Flowers of 2 kinds, the outer 5–9 mm, ligulate, the ligules 3-lobed at apex and bright golden yellow, the inner tubular 5-lobed at apex and golden yellow, or all tubular. Receptacle flat, without scales. Achenes 1.8–2.2 mm, pale brown, darker ribbed, those of the ligulate flowers glabrous, those of the tubular flowers hairy; pappus 5–6 mm, of whitish, simple eglandular hairs. Flowers 6–10. Visited by various bees and flies. 2n = 40. (a) Subsp. dunensis (Dumort.) Kadereit & P. D. Sell S. dunensis Dumort.; S. jacobaea var. flosculosus Lam. & DC.; S. laciniatus var. maritimus Gray Stems solitary 20–30(–60) cm. Peduncles often with dense arachnoid hairs. Ligulate flowers absent or rudimentary. Outer achenes hairy. (b) Subsp. jacobaea Stems 1–3, up to 150 cm. Leaves with little arachnoid hair. Peduncles with more or less numerous arachnoid hairs. Ligulate flowers usually present. Achenes outer row glabrous. (i) Var. condensatus Druce S. jacobaea var. abrotanoides Druce Stems markedly swollen below the basal leaves, 1–3, up to 40(–45) cm; internodes rather short. Leaves and inflorescence dense. Leaves rather small and with narrow segments. Ligules 5–7(–9) × 1–2 mm. Flowering early in June and continuing through the summer. (ii) Var. jacobaea S. flosculosus Jord.; S. jacobaea var. stenoglossus Brenan & N. D. Simpson; S. jacobaea var. discoideus Wimm. & Grab.; ?S. jacobaea var. nudus Weston Stems not markedly swollen below the basal leaves, usually solitary, up to 100 cm. Leaves rather widely spaced, with narrow segments. Inflorescence often large and open. Ligules 7–9 × 1.7–3.0 mm, rarely absent and then usually only 1–2 plants in a colony. Flowering mainly in July and August and going over by the time var. nemorosus comes into flower. (iii) Var. nemorosus (Jord.) Loret & Barrandon S. nemorosus Jord. Stems not markedly swollen below the basal leaves, several, up to 150 cm, often over 100 cm, often deep purple. Leaves large, up to 30 cm, rather widely spaced, with broad segments. Inflorescence large and open. Ligules 8–10 × 2–3 mm. Flowering in August and September.

82. Senecio Native. Sand-dunes, shingle, grasslands on light, disturbed calcareous soils, beech woodland, fens, streamsides and limestone pavements and introduced into other grasslands, waysides and waste places. Common throughout Great Britain and Ireland, reaching an altitude of 670 m in Scotland. Europe to about 66◦ N in Sweden and Finland; Caucasus; western Asia; introduced in Australia, New Zealand, South Africa and North and South America. A member of the Eurosiberian Temperate element. Named after St James (Jacobus) whose saint’s day is 25 July. S. jacobaea is toxic to cattle and horses whether living or dead, and it is avoided by rabbits. It is, however, readily eaten by sheep. Whole populations of the plant are defoliated by the orange-and-black ringed caterpillars of Hypochrita jacobaea (L.). Var. jacobaea is the most widespread variant but is absent from shingle and most dunes. It occurs throughout the range of the species. Var. condensatus is the plant of shingle and most dunes. It has not been seen outside our area. Var. jacobaea is sometimes brought into these coastal habitats with the building up of the sea defences. Some of the plants of chalk grassland approach var. condensatus but are more slender with open inflorescence and are usually mixed with var. jacobaea. Var. nemorosus occurs in fens, on ditch-sides and other wet places where it is a large and handsome plant. It is also recorded for France. Subsp. dunensis is a plant of coastal sand and pasture in Sutherland, the Orkney Islands, the Shetland Islands, Co. Kerry, Co. Wexford and the west coast of Ireland. It occurs also on the coasts of Belgium, Holland, France and Scandinavia. 14. S. aquaticus Hill Marsh Ragwort Jacobaea aquatica P. Gaertn., B. Mey. & Scherb.; S. jacobaea subsp. aquaticus (Hill) Bonnier & Layens Biennial herb with a short, more or less erect, praemorse stock and rather fleshy roots. Stem up to 130 cm, pale green, often suffused brownish-purple, erect, striate, more or less arachnoid-hairy, branched in the upper part, sometimes nearly to the base, leafy. Leaves alternate, medium yellowish-green or bluish on upper surface, paler beneath, glabrous or nearly so; basal and lower cauline 7–15 × 2–4 cm, oblong-obovate, oblong-ovate or ovate in outline, rounded-obtuse at apex, undivided or lyrate-pinnatifid, terminal lobe large, ovate or ovate-oblong and dentate or serrate-dentate, lateral lobes much smaller, oblong, obtuse at apex and sometimes dentate, with long, slender petioles; middle and upper cauline ovate or ovate-oblong in outline, obtuse at apex, pinnatifid, with the lateral lobes pointing forwards or rarely spreading, the lobes crenate to coarsely serrate, sessile and semiamplexicaul. Capitula 15–45 mm in diameter, in irregular, lax corymbs; peduncles long, sometimes slender, more or less arachnoid-hairy. Involucral bracts in 2 rows, the inner 3.5–5.5 × 1.2–1.4 mm, green, with a broad, whitish margin, lanceolate, acuminate at apex, but with an obtuse tip, the outer few and much narrower, all rather sparsely arachnoid-hairy. Flowers of 2 kinds, the outer 12–15 mm, ligulate, the ligules golden yellow to slightly orange and 3-lobed at apex, the inner tubular, yellow and narrowly 5-lobed at apex. Receptacle flat, without scales. Achenes 2.5–3.0 mm, pale brown, cylindrical, all more or less glabrous; pappus 5–6 mm, whitish, of

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simple hairs, readily falling. Flowers 7–8. Visited by flies. 2n = 40. (a) Subsp. ornatus (Druce) P. D. Sell S. aquaticus var. ornatus Druce Stems up to 50 cm. Lower leaves with large, dentate terminal segment. Inflorescence fairly congested, with rather stout, erect branches up to 11 cm. Capitula 25–45 mm in diameter. (b) Subsp. aquaticus Stems up to 80 cm. Lower leaves undivided or with a large terminal lobe. Inflorescence open with fairly stout branches up to 20 cm. Capitula 20–25 mm in diameter. (i) Var. barbareifolius Wimm. & Grab. S. aquaticus var. aquaticus auct. mult.; S. erraticus subsp. barbareifolius (Wimm. & Grab.) Beger; S. aquaticus subsp. barbareifolius (Wimm. & Grab.) Walters Lower leaves dentate, but undivided. (ii) Var. aquaticus S. aquaticus var. pennatifidus Gren. & Godr.; S. aquaticus var. intermedius Druce; S. aquaticus var. major Bab.; S. aquaticus var. flosculosus Gray; S. aquaticus var. lanatus Gray; S. aquaticus var. dubius Druce; S. aquaticus forma discoideus Druce; S. aquaticus forma aurantiacus Druce Lower leaves with a large terminal segment and spreading lateral segments. The very rare plants without ligulate flowers seem always to belong to this variety. (c) Subsp. erraticus (Bertol.) V. A. Matthews S. aquaticus var. barbareifolius auct.; S. erraticus subsp. barbareifolius auct.; S. erraticus Bertol.; S. jacobaea subsp. erraticus (Bertol.) Sudre; S. aquaticus var. erucoides (Wimm. & Grab.) V. A. Matthews Stems up to 130 cm. Lower leaves with a large terminal segment and spreading lateral segments; upper leaves with patent segments. Inflorescence open, with widely spreading slender branches up to 45 cm. Capitula 15–20 mm in diameter. Native. Marshes, damp meadows and streamsides. Occurs throughout Great Britain and Ireland, but much more common in the west and reaches 400 m in altitude. West and central Europe from northern Italy northwards to 62◦ 47 N in Scandinavia and eastwards to Posen and Lower Silesia. A member of the European Temperate element. Subsp. ornatus is the plant of northern Scotland, Hebrides and the Orkney and Shetland Islands. It appears to be endemic. Most of the remaining plants are subsp. aquaticus which occurs throughout the range of the species. Both varieties are widespread. Unfortunately var. barbareifolius has always been regarded as the type, as well as the name usually being considered to be synonymous with S. erraticus, but Hill’s plant is clearly what is usually known as var. pennatifidus. Subsp. erraticus is a much misunderstood plant. The plants that C. C. Babington first identified as ?S. erraticus and later described as S. aquaticus var. major are var. aquaticus. His plants from Guernsey which he called S. erraticus do seem to be the right plant and are probably native. Plants from a few other localities in southern England may also be referable to it. It occurs throughout the range of the species except in north-west and west Europe.

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× jacobaea = S. × ostenfeldii Druce This hybrid is very variable. It differs from S. aquaticus in its small fruiting involucres and from S. jacobaea in its more numerous flowers per capitulum (90–120 against 50–75). Fertility is variable but the most obvious hybrids have a high proportion of sterile pollen and less than 15 per cent developed achenes, the ones of the tubular flowers being distinctly pubescent but more minutely so than in S. jacobaea. It backcrosses however and sometimes forms hybrid swarms. 2n = 40. Native. Occurs throughout much of Great Britain and Ireland, but is more common in the west where intermediate habitats are more frequent. It occurs in Continental Europe, but its distribution is not known. 15. S. erucifolius L. Hoary Ragwort Jacobaea erucifolia (L.) P. Gaertn., B. Mey. & Scherb. Perennial herb with a short, creeping stock and short rhizomes. Stems 30–130 cm, yellowish-green, often deeply suffused brownish-purple, erect, ridged and channelled, with minute hairs and sometimes sparsely arachnoid-hairy, leafy, with ascending branches above. Leaves alternate, yellowish-green, hardly paler beneath and sometimes with a violet midrib, with short scattered simple eglandular hairs on the upper surface, with few to dense arachnoid and short curly hairs beneath; basal 50–120 × 10–40 mm, obovate or obovate-lanceolate in outline, acute at apex, pinnatisect, the terminal segment oblanceolate, acute at apex and toothed, the lateral segments linear or oblong, acute at apex and entire or with an occasional tooth, cuneate at base to a narrow, arachnoid petiole; lower cauline similar to basal; median and upper cauline gradually becoming smaller, pinnatisect, with all segments linear or oblong, sometimes very narrowly so, the lateral subparallel and forwardly directed, more or less sessile, the lowest lobes semiamplexicaul; all leaves with more or less revolute margins. Capitula 10–22 mm in diameter, in terminal and axillary corymbs; peduncles yellowish-green, sparsely arachnoid-hairy, with 1–3 narrowly linear bracts. Involucral bracts in 2 rows, the outer 4–8 about half as long as the inner and narrowly linear and acute at apex, the inner 6–8 × 0.9–1.1 mm, green, narrowly scarious in upper half, linear-lanceolate and acute at apex. Flowers of 2 kinds, the outer 12–15, ligulate, the ligules yellow, narrow, discrete and entire or emarginate at apex, the inner tubular, yellow to treacly-yellow and shortly 5-lobed at apex. Receptacle flat, without scales. Achenes about 2 mm, cylindrical, all with hairy ribs; pappus about 6 mm, whitish, of simple hairs. Flowers 7–9. Visited by flies and bees. 2n = 40. (i) Var. viridulus (Martrin-Donos) Rouy S. viridulus Martrin-Donos; S. erucifolius forma glabra Druce Slender plants. Leaves lanceolate, oblong-lanceolate or ovate in outline, pinnatisect almost to midrib, the segments, both terminal and lateral oblong, acute or obtuse at apex and entire or with a few teeth, green and glabrous or with very few, very short hairs beneath and not arachnoid. Open capitula up to 17 mm in diameter.

(ii) Var. erucifolius S. tenuifolius Jacq., non Burm. fil.; S. erucifolius var. linnaeanus Rouy nom. illegit. Slender plants. Leaves lanceolate, oblong-lanceolate or ovate in outline, pinnatisect almost to midrib, the segments both terminal and lateral linear, acute or long acute at apex and often with small teeth, greyish or whitish hairy beneath, often densely so. Open capitula up to 15 mm in diameter. (iii) Var. communis Rouy Tall, robust plants. Leaves mostly broadly ovate in outline, pinnatisect, the segments often rather widely oblong, obtuse or acute at apex and usually toothed, greyish-hairy beneath. Open capitula up to 22 mm in diameter. (iv) Var. latilobus Boiss. Short, robust plants. Leaves obovate in outline, divided to about halfway to the midrib, with a large, dentate terminal lobe and broadly oblong lateral ones. Native. Grassy places, waysides, banks, downs and field margins particularly on base-rich soils. Common in most of England and Wales, rare in Scotland and very local in eastern Ireland. Central and southern Europe northwards to Denmark, southern Sweden and Lithuania; western Asia. A member of the Eurosiberian Temperate element. Var. erucifolius is a plant of open grassland. Var. communis occurs along hedge- and streambanks and is very like in appearance the late-flowering S. jacobaea subsp. jacobaea var. nemorosus with which it sometimes grows. Var. viridulus occurs in a few scattered localities. It appears to have been introduced with wild flower seed on the Gog Magog Hills near Cambridge where the native plant is var. erucifolius. In Cardiganshire it occurs on a boulder clay slope by the sea. Var. latilobus is a very distinct plant known to us only from a specimen collected in Southampton in 1836. All these varieties occur in Continental Europe. G. C. Druce described a var. subintegrifolius from the south coast with narrow leaves little divided, but we have seen no material of this. × jacobaea = × leichtensteinensis Murray S. × whitwellianus Lees ex Cheetham nom. nud. This hybrid has been recorded from several places but never confirmed. Large mixed colonies of these two species often occur. Section 9. Senecio Annual or short-lived perennial herbs, variously hairy. Leaves subentire to pinnately divided. Ligules yellow. Achenes glabrous or hairy. 16. S. squalidus L. Oxford Ragwort Annual, biennial or perennial herb with a tap-root and fibrous side-roots. Stems 20–50 cm, pale yellowishor glaucous-green, sometimes suffused brownish-purple, decumbent at base and sometimes woody, then erect, flexuous, more or less glabrous, branched and leafy. Leaves alternate, medium to dark yellowish- or glaucous-green on upper surface, paler beneath, flat and glabrous or with a few long hairs beneath; lower 2–12(–15) × 1.5–4.0 cm, oblong, oblanceolate, oblong-oblanceolate or lanceolate in outline,

82. Senecio obtuse or acute at apex, dentate or deeply pinnatisect, the lobes oblong or lanceolate and acute to obtuse at apex and entire or dentate, narrowed at base to a winged petiole; upper similar, gradually decreasing in size upwards, sessile, semiamplexicaul and sometimes with rounded auricles. Capitula few to numerous, 16–20 mm in diameter, bell-shaped in flower, in an irregular, lax, simple or compound corymb; peduncles glabrous. Involucral bracts in 2 rows, the inner 19–22, 6–8 × 0.8–1.0 mm, pale green, sometimes conspicuously black-tipped, linear-lanceolate, acute at apex, the outer 5–13, 1.5–3.0 mm, linear, acute at apex, black-tipped. Flowers of 2 kinds, the outer usually 12–15, ligulate, the ligules usually more than 8 mm, bright yellow, broad and entire or 3-lobed at apex, the inner tubular, yellow and with 5-lobed at apex. Stigmatic papillae 40–60, pollen grains 20–25 µm. Receptacle flat, without scales. Achenes 2.0– 3.0 mm, pale brownish, cylindrical, ribbed, usually hairy in the grooves; pappus whitish, of simple hairs. Flowers 3–12. 2n = 20. (a) Subsp. rupestris (Waldst. & Kit.) P. D. Sell S. rupestris Waldst. & Kit.; S. nebrodensis auct. Annual to perennial herb. Leaves medium to dark yellowish-green, at least some of them regularly pinnately lobed nearly to the midrib, or others regularly dentate, the uppermost often auriculate-semiamplexicaul. (i) Var. calabricus Fiori S. nebrodensis var. calabricus (Fiori) Fiori; S. rupestris var. calabricus (Fiori) Pign. Stem up to 30 cm. Leaves up to 6 × 2 cm, oblong in outline, regularly pinnately lobed, the lobes up to 7 mm, linear and sometimes denticulate. (ii) Var. pinnatifidus (Evers ex Hegi) P. D. Sell S. rupestris forma pinnatifidus Evers ex Hegi Stem up to 45 cm. Leaves up to 9 × 4 cm, regularly pinnately lobed nearly to midrib, the lobes up to 15 mm, oblong and dentate. (iii) Var. bipinnatifidus (Vis.) P. D. Sell S. rupestris var. bipinnatifidus Vis.; S. nebrodensis var. bipinnatifidus (Vis.) Hayek Stem up to 40 cm. Leaves up to 15 × 6 cm, bipinnately lobed, laciniate and dentate. (iv) Var. rupestris (Waldst. & Kit.) P. D. Sell Stem up to 40 cm. Leaves up to 10 × 4 cm, the lower broadly divided more than halfway, the upper regularly pinnately lobed and often toothed, the upper leaves markedly auriculate-semiamplexicaul. (v) Var. laciniatus (Vis.) P. D. Sell S. nebrodensis var. laciniatus Vis. Stem up to 45 cm. Leaves up to 12(–15) × 3(–4) cm, oblong or obovate in outline, some divided only to halfway, others laciniate to midrib, sometimes with acute linear lobes, the upper markedly auriculate-amplexicaul. (vi) Var. subinteger (Druce) Druce S. squalidus forma subinteger Druce; S. rupestris forma sinuatodentatus Evers ex Hegi; S. rupestris var. sinuatodentatus (Evers ex Hegi) E. I. Ny´arˆady

497

Stem up to 40 cm. Leaves up to 10 × 3.5 cm, oblong or oblong-lanceolate, sinuate-dentate, at least the upper broadly auriculate-semiamplexicaul. (b) Subsp. chrysanthemifolius (Poir.) Greuter S. chrysanthemifolius Poir.; Jacobaea incisa C. Presl; S. incisus (C. Presl) C. Presl, non Thunb. Perennial herb. Leaves deeply and irregularly pinnatisect almost to the midrib, the lobes 2–40 × 1–2 mm, with short, sharp irregular teeth on the side. Achenes 2.0–2.5 mm. (c) Subsp. squalidus S. glaber Ucria; S. squalidus subsp. glaber (Ucria) Nyman; S. squalidus var. glaucescens Spruce Perennial herb. Leaves 2–11 × 2–8 cm, glaucous-green, shallowly to deeply and irregularly pinnatisect with the lobes linear, oblong or lanceolate, acute or obtuse at apex, mostly more than 2 mm wide, irregularly dentate and showing a wide variation, or the leaves only dentate. (d) Subsp. aethnensis (Jan. ex DC.) Greuter S. aethnensis Jan ex DC.; S. chrysanthemifolius subsp. aethnensis (Jan ex DC.) Lambinon; S. squalidus var. aethnensis (Jan ex DC.) Fiori; S. carnosus (C. Presl) C. Presl, non Thunb.; Jacobaea carnosa C. Presl Perennial herb. Leaves 3–9 × 0.8–2.0 cm, glaucous-green, narrowly elliptical to lanceolate, obtuse to acute at apex, entire, semiamplexicaul. Achenes 2.5–3.0 mm. Introduced. It has always been assumed that the source of this aggregate was the Oxford Botanic Garden. Early material from Oxford is exactly like the Linnaean type which is thought to have come from Oxford. On Mount Etna in Sicily are three taxa which we include here as subspecies. Subsp. chrysanthemifolius occurs below 1,000 m and subsp. aethnensis above 2000 m. Those plants occurring in the area in between having a mixture of characters of the subspecies occurring above and below them and are thought to be the same as the plant introduced to Oxford and named by Linnaeus S. squalidus. Many plants from Oxford, but not all, do seem to be identical with these intermediate plants and one sheet in CGE contains two plants one of which is very near to subsp. chrysanthemifolius and the other very near to subsp. aethnensis, indeed Italian botanists might call it subsp. aethnensis as they allow that taxon to have toothed leaves. Plants from Yarmouth in Norfolk, Bideford and Teignmouth Docks in Devonshire, Cardiff, Co. Cork, Linley in Shropshire and Doncaster in Yorkshire also seem to be S. squalidus subsp. squalidus. These plants appear to be glaucous, as the plants of Sicily are said to be by the Italian botanists. The great bulk of the plants which later spread throughout Great Britain and Ireland are green, often yellowish-green, have more handsome flowers (a point noted by Druce) and often more auricled upper stem leaves and seem to better fit S. rupestris which is here also treated as a subspecies of S. squalidus. S. rupestris is a very variable plant and six distinct varieties are recognised here. All have names under S. rupestris in Continental floras, which seem to fit them. Similar names do not seem to be available for Etna plants and specimens have not been seen from there that match them. Var. calabricus has been seen from Cornwall, Kent,

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S. eboracensis R. J. Abbott & A. J. Lowe

S. cambrensis Rosser

subsp. squalidus

subsp. aethnensis (Jan. ex DC.) Greuter

subsp. chrysanthemifolius (Poir.) Greuter Senecio squalidus L. aggr.

82. Senecio

499

var. pinnatifidus (Evers ex Hegi) P. D. Sell

var. rupestris (Waldst. & Kit.) P. D. Sell

var. subinteger (Druce) Druce var. laciniatus (Vis.) P. D. Sell

var. calabricus Fiori

var. bipinnatifidus (Vis.) P. D. Sell Senecio squalidus L. subsp. rupestris (Waldst. & Kit.) P. D. Sell

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Oxford, Cambridgeshire, Glamorganshire and Midlothian. Var. pinnatifidus is common everywhere. Var. bipinnatifidus occurs in Bideford in Devonshire (1867), Oxford, Suffolk, Norfolk, Oxford, Huntingdonshire and Moray. It is common around Cambridge. Var. laciniatus is recorded from Exmouth Docks in Devonshire, Berkshire, Suffolk, Cambridgeshire, Cardiff Docks and Midlothian. Var. rupestris is recorded from Yarmouth and Snettisham in Norfolk, Belfast Docks and Ashburton, Co. Cork. Var. subinteger occurs at Exmouth in Devonshire, Didcot in Berkshire and Oxford. It is common around Cambridge. Var. laciniatus reproduced exactly from seed, and var. calabricus, var. pinnatifidus, var. bipinnatifidus and var. subinteger have been seen in uniform colonies suggesting they also might do so. A great deal of experimental work has been carried out on this group, much of which suggests all this variation comes from Etna plants. This account is based entirely on herbarium and field work. The distribution of the variation badly needs to be recorded. If other botanists believe all our plants originated from Etna the arrangement of infraspecific taxa is such that the same varietal names can be used under S. squalidus, and subsp. rupestris omitted. × viscosus = S. × subnebrodensis Simonk. S. × londinensis Lousley This hybrid is intermediate in leaf shape and number of glandular hairs between the parents and is highly sterile. 2n = 30. Introduced. In scattered localities on waste land. Great Britain mainly in the south. It occurs also in Continental Europe. If S. rupestris is not thought to be in Great Britain and Ireland, the correct name for this hybrid is S. × londinensis. × vulgaris = S. × baxteri Druce This hybrid is intermediate in leaf and capitulum and is highly sterile. Stigmatic papillae 0–30. Pollen 10–40µm; pores 2–6. 2n = 30. Some populations however, yield some seeds giving rise to fertile tetraploids, which when backcrossed with S. vulgaris show some S. squalidus-like characters in otherwise S. vulgaris-like offspring. S. × baxteri is very similar in appearance to S. cambrensis which is, however, fully fertile. For stable S. vulgaris variants which have ligules and are thought to be this hybrid see that species. S. × baxteri has been recorded in a few scattered localities in Great Britain and Ireland. (Over the past century, specimens of putative, semifertile hybrids between S. squalidus aggr. and S. vulgaris have been recorded. See Abbott and Lowe (2003) for list of records.) 17. S. cambrensis Rosser Welsh Groundsel Annual or short-lived perennial herb with fibrous roots. Stems up to 35(–60) cm, pale yellowish-green, sometimes tinted purplish, erect, ridged, somewhat woody at base, glabrous or with sparse arachnoid hairs, branched from the base or middle. Leaves alternate, 2–12 × 0. 3–4.0(–7.0) cm, dull, medium yellowish-green, paler beneath, all more or less cauline and gradually decreasing in size upwards, obovate, oblanceolate, oblong or linear in outline, obtuse to acute at apex, pinnatifid to pinnatisect, the lowest often less divided, the terminal lobe oblong to ovate and irregularly dentate, the lateral oblong to linear and dentate to incise-

dentate, all very sparsely arachnoid-hairy, the lower with winged petioles up to 3 cm, the upper sessile and auricled. Capitula up to 15 mm in diameter, cylindrical in flower, in corymbs of 4–7 at the ends of stems and branches; branches erecto-patent; peduncles 5–40 mm, with sparse arachnoid and sparse, short glandular hairs. Involucral bracts in 2 rows, yellowish-green with brownish-black tips, the inner 7–10 × 0.5–0.7 mm and linear, the outer few, 1.0–1.5 × 0.5– 0.7 mm, linear, and obtuse to subacute at apex. Flowers yellow; the inner tubular and 5-lobed at apex; the outer ligulate, the ligules (8–)13(–15), 4.0–7.0 × 2.0–2.5 mm, or absent, obovate, with 3 lobes at the apex; becoming revolute. Stigmatic papillae 30–50. Pollen grains 30–35 µm, mostly with 4 pores. Receptacle flat and pitted, without scales. Achenes 3.0–3.5 mm, pale brown, cylindrical to fusiform, with appressed, white hairs; pappus 4–7 mm, of pale grey simple hairs. Flowers 5–10. 2n = 60. Native. Waste ground, gardens and by footpaths and roadsides. First found in 1948 at Ffrith in Flintshire, where it still occurs at several sites and now known also in Denbighshire where it is much more widespread. Records from Shropshire require confirmation. In 1982 it was found at Leith, near Edinburgh in Midlothian, where it now appears to be extinct. Endemic. A naturally arisen alloploid from a cross between S. squalidus (possibly var. pinnatifidus) and S. vulgaris. 18. S. eboracensis R. J. Abbott & A. J. Lowe York Ragwort Annual herb with tap-root and fibrous side-roots, occasionally with adventitious roots on the horizontal base of the stem. Stems up to 40 cm, pale green, often purplish at base, erect or ascending, striate, arachnoid-hairy. Leaves alternate, up to 8 × 3 cm, medium to deep yellowish-green on upper surface, paler beneath, oblong, oblanceolate or narrowly elliptical in outline, obtuse or acute at apex, pinnately divided up to three-quarters of the way to the midrib, the lobes triangular, acute at apex, spreading or up-curved, sometimes denticulate, narrowed at base to a short petiole or the upper semiamplexicaul, auriculate, with few arachnoid hairs on upper surface and more numerous beneath. Capitula about 10 mm in diameter, broadly cylindrical in flower, becoming slightly campanulate, in a corymb of up to 4 at the ends of stems or branches, erecto-patent; peduncles 5–20 mm, elongating in fruit, pale green, sparsely hairy. Involucral bracts in 2 rows, pale yellowish green with or without black tips, the inner 20–22, 7–8 × 0.8–1.0 mm, linear, sharply acute at apex, the outer 7–8, 2.5–3.0 × 0.6–0.7 mm, linear, sharply acute at apex and denticulate; all glabrous or nearly so. Flowers yellow, the inner tubular and 5-lobed at apex, the outer (6–)8(–9), 5–6 × about 1.5 mm, occasionally becoming revolute, with 3 lobes at apex. Stigmatic papillae 10–30. Pollen grains in polar view about 30–35 µm when fully expanded, mostly 4-pored. Receptacle flat and pitted, without scales. Achenes 2.5–3.5 mm, pale brown, straight, cylindrical, ribs glabrous, grooves hairy; pappus white, of silky hairs, readily becoming detached from the fruit. Flowers 4–10. Self-pollination frequent. 2n = 40. Native. Disturbed ground, car parks, pavement cracks and other urban industrial sites. Known only from York.

82. Senecio Endemic. A hybrid product of S. squalidus (possibly var. subinteger) × S. vulgaris. 19. S. vulgaris L. Groundsel S. semperflorens Stokes nom. illegit. Annual or overwintering herb with fibrous roots. Stems 8–45(–50) cm, yellowish-green, often flushed brownishpurple especially towards the base, ascending or erect, often rather fleshy, ridged and channelled, nearly glabrous to arachnoid-hairy, leafy, simple to much-branched, the branches sometimes nearly to the base, erecto-patent. Leaves alternate, 10–50 × 3–25 mm, pale to medium yellowish- or bluish-green, paler and bluish- or greyishgreen beneath, sometimes tinted brownish-purple or purple, slightly fleshy to very fleshy, glabrous to densely arachnoid-hairy especially beneath, the lower sometimes forming a rosette or pseudorosette, oblanceolate to obovate, rounded and dentate at apex, dentate or serrate to pinnatisect and narrowed at base to a short or medium, sometimes narrowly winged, hairy petiole, the median and upper oblanceolate to oblong in outline, obtuse or acute at apex, irregularly lacerate-pinnatisect, the terminal lanceolate or oblong and often toothed, the lateral segments oblong or linear and sometimes toothed, usually sessile, more or less amplexicaul and auriculate at base. Capitula 4–5 mm wide when flowers all tubular, wider when ligulate flowers are present, in small terminal corymbs or sometimes solitary, the involucre cylindrical; peduncles yellowishgreen, glabrous or more or less arachnoid-hairy. Involucral bracts in 2 rows, the inner 5–8 × about 0.8 mm, yellowishgreen, with a narrow, scarious margin and usually becoming blackish-brown at apex, linear, and narrowing to an acute apex, the outer 6–10, 1–4 × 0.5–0.7 mm, yellowish-green with blackish-brown tips, and sometimes blackish throughout, linear or subulate, narrowed to an acute apex and more or less appressed, all glabrous or arachnoid-hairy. Flowers uniformly tubular or with an outer row which are ligulate, the tubular 5–6 mm, yellow, with 5 deltoid, recurved lobes at apex, the ligulate when present yellow, with 3 lobes at apex. Pollen grains with 3 pores. Receptacle flat or slightly convex, pitted, without scales. Achenes 1.5–2.5 mm, brown, cylindrical-fusiform, 8- to 10-ribbed, the apex expanding into a shallow cup, appressed-hairy between the ribs; pappus 5–6 mm, white, of scabridulous hairs. Flowers 1–12. Said to be little visited by insects and normally self-pollinated, but aphids are often present in the capitulum even in winter, and variation in large populations in arable land does not seem to suggest only self-pollination. Haskell (1953) believes there is a continual process of hybridisation, dissemination and inbreeding, depending to some extent on how many wild generations there are in one year. 2n = 40. (a) Subsp. vulgaris Short-lived annual. Stems 8–45(–50) cm, often strongly branched. Leaves fleshy or not, glabrous to moderately arachnoid-hairy, often deeply incise. Ligulate flowers absent or present with ligules 3.5–5.5 mm. 2n = 40. (i) Var. crassifolius (Rouy) P. D. Sell S. vulgaris subvar. crassifolius Rouy

501

Stems up to 15 cm, often branched nearly to base, arachnoidhairy in upper part of branches and peduncles. Leaves very fleshy, often arachnoid-hairy on under surface. Ligulate flowers absent. (ii) Var. vulgaris S. vulgaris var. erectus (Trow) Druce; S. erectus Trow; S. vulgaris var. praecox (Trow) Druce; S. praecox Trow; S. vulgaris var. latifolius (Trow) Druce; S. latifolius Trow; S. vulgaris var. multicaulis (Trow) Druce; S. multicaulis Trow; S. vulgaris var. rubricaulis Trow ex C. E. Salmon Stem up to 45(–50) cm, with few to numerous branched, glabrous to sparsely arachnoid-hairy. Leaves not or slightly fleshy, glabrous to sparsely arachnoid-hairy. Ligulate flowers absent. 2n = 40. (iii) Var. hibernicus Syme S. vulgaris var. radiatus auct.; S. × hybridus Burb., non Hort. ex Regel Stem up to 36 cm, with few to numerous branches, glabrous to sparsely arachnoid-hairy. Leaves not fleshy, glabrous to sparsely arachnoid-hairy. Ligulate flowers present, the ligules 3.5–5.5 mm. 2n = 40. (b) Subsp. denticulatus (O. F. M¨ull.) P. D. Sell S. denticulatus O. F. M¨ull.; S. sylvaticus var. denticulatus (O. F. M¨ull.) Wahlenb.; S. lividus var. denticulatus (O. F. M¨ull.) DC.; S. vulgaris var. radiatus W. D. J. Koch nom. illegit.; S. vulgaris var. dunensis Knuth; S. lanuginosus Trow, non Spreng.; S. vulgaris var. lanuginosus Druce; S. vulgaris forma radiatus Hegi; S. vulgaris var. lanifer Wilmott Stem up to 20(–30) cm, simple or weakly branched, densely arachnoid-hairy especially above. Leaves rather fleshy, not usually deeply divided, arachnoid-hairy. Ligulate flowers present, the ligules 6–12, 2.3–3.0 mm and becoming revolute after flowering. 2n = 40. Native. Cultivated, waste and other open ground, shingle, cliffs, dunes and walls. Common throughout Great Britain and Ireland reaching 530 m altitude in Scotland. Europe to 71◦ N; Asia; North Africa; widely introduced in other parts of the world. A member of the European Southerntemperate element, but total range is now Circumpolar Southern-temperate. Var. crassifolius is a plant of coastal shingle, dunes and cliffs. It probably occurs in many places round Great Britain and Ireland. It was described from France and is probably elsewhere on the coasts of western Europe. Var. vulgaris is very variable in habit, leaf shape and indumentum and is a ruderal which occurs throughout the range of the species. Var. hibernicus occurs in scattered populations in waste and cultivated ground, sometimes mixed with var. vulgaris, in Great Britain and Ireland. The origin of this plant is not certain. It was first recorded in Co. Cork in Ireland in 1853, but did not become frequent until the 1930s and did not become common until the 1950s. In Continental Europe it seems to be rare. Evidence has been put forward that it is a stabilised introgressant between S. vulgaris and S. squalidus, but with the exception of the ligulate flowers all its characters are those of S. vulgaris, of which it has been thought to be a mutation. Subsp. denticulatus occurs on dunes, sandy fields and cliffs near the sea. It occurs from

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south and east Sweden down the coast of western Europe to western France and in the Mediterranean region mainly on mountains in Spain and Sicily. In Great Britain it has only been seen recently from Ainsdale in Lancashire and in the Channel Islands. It is possible that var. crassifolius is more nearly related to subsp. denticulatus than to subsp. vulgaris and the discoid plants found by Comes (1995) in Guernsey could clearly not be separated from var. crassifolius. Comes also found plants with shorter ligules which he suggested were hybrids, but it may be there is a coastal ecotype more dwarf, fleshy and arachnoid-hairy with or without ligulate flowers. This coastal ecotype may have spread north after the last glaciation and from it the ruderal S. vulgaris subsp. vulgaris was derived (cf. Kadereit, 1984). 20. S. sylvaticus L. Heath Groundsel S. montanus L. Annual herb with a tap-root and fibrous side-roots. Stems 5– 70 cm, pale yellowish-green, sometimes suffused brownishpurple towards the base, erect, slender, furrowed, more or less arachnoid-hairy and with short glandular hairs, but not viscid, branched, the branches ascending, leafy. Leaves alternate, yellowish-green on upper surface, paler beneath, arachnoid-hairy at first becoming glabrous; lower 3–9 × 3–5 cm, oblanceolate, oblong-oblanceolate or obovate in outline, more or less acute at apex, deeply and irregularly pinnatifid, the lobes unequal, mostly obtuse and more or less lobed, sessile or with enlarged auricle-like basal lobes which clasp the stem. Capitula numerous, cylindrical, 5–6 mm in diameter, in a large, flat-topped terminal corymb; peduncle with short glandular hairs. Involucral bracts in 2 rows, the inner 6.5–7.5 × 0.7–0.8 mm, green, linear-lanceolate and acute at apex, the outer 1.0–1.5 mm, linear and acute at apex; all with short glandular hairs. Flowers of 2 kinds, the outer 8–14 ligulate, the ligules bright yellow with 3 lobes at apex and very short and revolute, the inner tubular, darker yellow and 5-lobed at apex. Receptacle flat, without scales. Achenes 2.0–2.5 mm, dark green, cylindrical, stiffly hairy on the ribs; pappus 4–5 mm, whitish, of simple hairs. Flowers 7–9. Visited by flies. 2n = 40. (i) Var. nanus Rouy Stems 5–20 cm. Capitula solitary or in a small corymb at the top of the stem. (ii) Var. sylvaticus Stems up to 70 cm. Capitula numerous in a large, flat-topped terminal corymb. Native. Open ground on sandy heaths, roadsides, waste places, dunes and shingle beaches. Widespread throughout Great Britain and Ireland but absent from large areas where there are no sandy soils. Central Europe from Portugal, central Italy and the northern Balkans to 66◦ N in Scandinavia; western Asia. A member of the European Temperate element. The common plant is var. sylvaticus. Var. nanus is the plant of sand and shingle by the sea. Its miniscule size makes it easily missed. It was described from the coast of France. × viscosus = S. × viscidulus Scheele This hybrid is intermediate between the parents in leaf colour and shape, capitulum shape and size, length of

involucral bracts and ligules and degree of viscid hairiness. It has lower seed and pollen fertility, but occasional achenes are set, being black or dark brown as in S. viscosus but hairy as in S. sylvaticus. 2n = 40. Native. Scattered localities in England and Scotland and Co. Cork in Ireland where the parents grow together. It also occurs widely through central Europe. 21. S. vernalis Waldst. & Kit. Eastern Groundsel S. leucanthemifolius Poir. subsp. vernalis (Waldst. & Kit.) C. Alexander; S. rapistoides DC. Annual to perennial herb with fibrous roots. Stems 10–50 cm, pale yellowish-green, ascending or erect, with arachnoid hair when young becoming nearly glabrous with age, simple or branching in upper half, leafy. Leaves alternate, 2–8 × 0.5–2.5 cm, yellowish-green, usually with arachnoid hairs, the basal and lower oblong, withering early, the cauline oblong or broadly linear, obtuse at apex, pinnately divided with spreading, obtuse lobes and sharply dentate, the lower long-petiolate, the rest sessile with amplexicaul auricles. Capitula 20–30 mm in diameter, bell-shaped in flower, forming a rather small, tight corymb at the apex of the stem; peduncles slightly arachnoid-hairy. Involucral bracts in 2 rows, the inner 8–10 × 1.0–1.5 mm, pale green, usually black-tipped, linear-lanceolate, acute at apex, arachnoid-hairy, the outer 5–13, about 3 mm. Flowers of 2 kinds, the outer 8–15, (6–)8–10 mm, ligulate, the ligules 3-lobed at apex, non-revolute and pale yellow, the inner tubular, 5-lobed at apex and yellow. Receptacle flat, pitted, without scales. Achenes 2–3 mm, cylindrical, ribbed, appressed-hairy; pappus of whitish simple hairs. Flowers 5–11. 2n = 20. Introduced. Roadside verges and newly landscaped areas as a seminaturalised grass-seed alien. Scattered records in Great Britain and Isle of Man. Native of Eastern Europe, western Asia and North Africa. It has been spreading westward during the last century and has become a troublesome weed in parts of central and western Europe. Alexander (1979) makes it a subspecies of the Mediterranean S. leucanthemifolius Poir. and gives good reasons for doing so. × vulgaris = S. × helwingii Begar ex Hegi S. × pseudovernalis Zabel ex Nyman nom. inval. Intermediate in leaf shape between the parents with ligules as large as in S. vernalis and deformed pollen. Introduced. Road verges newly sown with seed mixtures obtained from Denmark and Holland, near Market Harborough in Leicestershire in 1968 and 1969. It is also recorded from Germany, Switzerland, Poland and Romania. 22. S. viscosus L. Sticky Groundsel Foetid annual herb with the main root often thick and strong especially when growing in shingle. Stems (5–)10–60 cm, erect, yellowish-green, ridged and channelled, with dense, unequal, viscid, pale glandular hairs, leafy, branched sometimes to the base, the branches suberect, ascending or ascending-erect and undulate. Leaves alternate, 10–60 × 2– 30 mm, medium yellowish-green above, paler beneath, oblanceolate, obovate, oblong or linear in outline, obtuse at apex or the upper acute, pinnately divided, the terminal lobe often toothed, the lateral lobes oblanceolate, obovate,

83. Pericallis and toothed to linear and entire, narrowed at base, the lower often petiolate, the rest sessile, all densely covered with unequal, pale, viscid glandular hairs. Capitula 10–15 mm in diameter, rounded at base, few to numerous, in an irregular, compound corymb; peduncles up to 30 mm, pale green, furrowed, with dense, mostly short, pale, viscid glandular hairs and occasionally linear bracts. Involucral bracts in 2 rows, 8–9(–11) × 0.8–1.2 mm, pale yellowish-green, blackish-brown tipped, linear or narrowly linear-lanceolate, gradually narrowed to an acute apex, covered with numerous to dense, pale, unequal, viscid glandular hairs. Flowers of 2 kinds, the outer 6–7 mm, ligulate, the ligules yellow, soon revolute and shallowly 3-lobed at apex, the inner 5–6 mm, tubular, yellow, shortly lobed at apex. Receptacle flat, pitted, with a raised rim round the pits, without scales. Achenes 3–4 mm, the outer pale greenish-brown to black, the inner with medium brown ribs and pale brown between, glabrous; pappus 5–6 mm, white, of minutely denticulate hairs. Flowers 7–9. Visited by flies and bees. 2n = 40. The plants of coastal shingle are much branched, often to the base, and of dense habit. The inland weeds are often less branched and taller. Probably native in coastal localities, a weed inland. Shingle and dune, waste and rough ground, railway tracks, waysides and walls. Frequent in most of Britain, introduced and local in north and central Scotland and Ireland. Europe to 66◦ N in Scandinavia and eastwards to Karelia; Turkey; introduced in North America. A member of the European Temperate element. 23. S. elegans L. Purple Groundsel Jacobaea elegans (L.) Moench. Annual herb with a tap-root and fibrous side-roots. Stems up to 60 cm, pale green, ridged, erect, glabrous or nearly so, usually branched only in the inflorescence, leafy. Leaves alternate, 4–10(–20) × 1–3(–6) cm, green on upper surface, paler beneath, elliptic-oblong or oblong in outline, obtuse at apex, pinnatipartite, with 2–4 pairs of oblong, obtuse, crenate or shallowly lobulate segments, the lower petiolate, the upper sessile and auriculate-amplexicaul. Capitula few, 20–25 mm in diameter, grouped in a subcorymb; peduncles long and sparsely bracteate. Involucral bracts in 2 rows, the inner 6–10 × 0.6–0.8 mm, green, lanceolate and more or less acute at apex, the outer 8–15, 2–4 mm, more or less ovate, acute at apex. Flowers of 2 kinds, the outer about 13, 6–8 mm, ligulate, the ligules 3-lobed at apex and purple, the inner tubular, 5-lobed at apex and yellow. Receptacle flat, without scales. Achenes 2.5–3.0 mm, cylindrical, usually hairy between the ribs; pappus 5–8 mm whitish, of simple hairs. Flowers 7–8. 2n = 20. Introduced. A casual or persistent garden escape in Devonshire, the Channel Islands and Isles of Scilly. Native of South Africa. 24. S. minimus Poir. Toothed Fireweed Erechtites minima (Poir.) DC.; Erechtites prenanthoides DC.; Erechtites pumila DC. Annual herb with a tap-root and fibrous side-roots. Stems up to 1 m, pale yellowish-green, sometimes slightly tinted brownish-purple, stiffly erect, ridged, glabrous or with a few minute hairs, branched in upper part, leafy. Leaves alternate,

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2–13 × 0.2–2.5 cm, medium slightly bluish-green on upper surface, paler beneath, linear to elliptical, obtuse or acute at apex, sharply dentate to serrate, the lower narrowed at base, the upper auriculate and clasping the stem, glabrous or rugose. Capitula very numerous, 2–4 mm in diameter, in large corymbose panicles; peduncles glabrous or with a few minute hairs. Involucral bracts in 2 rows, the inner (6–)8(–10), 5–6 × 0.6–0.8 mm, greenish-brown with a narrow, pale margin, narrowly linear-lanceolate, obtuse to subacute at apex and glabrous, the outer few, about one-sixth as long as inner and linear. Flowers all tubular, 5-toothed at apex and yellow. Receptacle flat, without scales. Achenes 1.8–2.0 mm, brown, cylindrical, ribbed, with appressed hairs on the ribs; pappus 5–6 mm, white, of simple hairs. Flowers 8–9. 2n = 60. Introduced. A wool casual. Native of Australia. 25. S. quadridentatus Labill. Cotton Fireweed Erechtites quadridentata (Labill.) DC. Short-lived perennial herb with a stout root. Stem up to 60 cm, pale green, erect, ridged, arachnoid-woolly, branched, the branches ascending, leafy. Leaves alternate, 3–13 × 0.2– 0.5(–3.0) cm, dull medium green on upper surface, paler beneath, long-linear, gradually drawn out to an acute apex, entire, denticulate or coarsely toothed, revolute, gradually narrowed at base and sometimes auriculate, arachnoidtomentose especially beneath. Capitula numerous, 2–3 mm in diameter, in corymose corymbs; peduncles arachnoidtomentose. Involucral bracts in 2 rows, the inner 11–13, 6–8 × 0.5–0.6 mm, greenish-brown with a pale margin, linear, often tipped red at the obtuse apex and arachnoid-hairy or tomentose, the outer few, about one-quarter the length of the inner and linear. Flowers all tubular, 5-lobed at apex and yellow. Receptacle flat, without scales. Achenes 2.8– 3.0 mm, pale brown, cylindrical, ribbed, with appressed hairs on the ribs; pappus 5–6 mm, whitish, of simple hairs. Flowers 8–10. 2n = 40. Introduced. A wool casual. Native of Australia. 83. Pericallis D. Don Annual to perennial herbs. Stems erect, leafy. Leaves alternate, palmately lobed and veined. Capitula in more or less dense corymbose masses. Involucral bracts in 1 main row. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female. Corolla white to pink, red, maroon, deep purple, violet or blue, or bicoloured. Anthers without tails; apical appendage ovate-lanceolate to oblong, flat; filament collar dilated. Style with oblonglinear, truncate branches; stigmatic areas in 2 separate lines. Receptacle flat, without scales. Achenes more or less cylindrical but somewhat compressed, the outer without a pappus, the inner with a caducous pappus. Fifteen species in the Canary Islands, Madeira and the Azores. Nordenstam, B. (1978). Taxonomic studies in the tribe Senecioneae (Compositae). Opera Bot. 44: 1–83.

1. P. hybrida R. Nordenstam Cineraria Senecio hybridus Hyl. nom. inval.; Senecio cruentus auct. Annual to perennial herb in compact cushions or openly branched. Stems 20–80 cm, yellowish-green, sometimes

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purplish, erect, with simple eglandular hairs, branched above, leafy. Leaves alternate, 9–12 × 10–15 cm, medium yellowish-green on upper surface, paler beneath, ovate to subrotund, palmately veined, acute at apex, lobed, with shallow, triangular acute lobes, cordate at base, glabrous, long-petiolate. Inflorescence a more or less dense corymbose mass; peduncles striate, glabrous. Capitula 15–60 mm in diameter. Involucral bracts in 1 row, 5–6 × 1.0–1.5 mm, greyish-green, oblanceolate, abruptly narrowed to a long acute apex, glabrous, ribbed. Flowers of 2 kinds; outer 15– 30 mm, ligulate, the ligules from white to pink, red, maroon, deep purple, violet and blue, sometimes bicoloured with a white base and coloured apex, entire, undulate or 3-lobed at apex; inner 4–10 mm, tubular, coloured similar to outer, 5-lobed at apex. Receptacle flat, without scales. Achene cylindrical but somewhat compressed, ribbed, those of the outer flowers sparsely hairy and without a pappus, those of the inner flowers densely hairy with a caducous pappus. Flowers 6–9. Introduced. A very popular, but frost-sensitive pot plant well naturalised on open ground, walls and waysides in the Isles of Scilly and rarely mainland Cornwall. A variable complex of forms appearing to have originated in England as hybrids between P. cruenta (L’H´er.) Bolle and other species from the Canary Islands. Cineraria aspera Thunb., C. burkei Burtt Davy, C. geifolia L., C. lobata L’H´er. and C. lyrata DC. have been recorded as wool casuals. 84. Tephroseris (Rchb.) Rchb. Senecio section Tephroseris (Rchb.) DC.; Cineraria section Tephroseris Rchb. Biennial to perennial herbs. Stems erect, leafy. Leaves alternate, entire to coarsely dentate, woolly. Capitula in small, dense terminal and axillary corymbs. Involucral bracts in 1 main row. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female. Corolla yellow. Anthers without tails; apical appendages ovatelanceolate to oblong, flat; filament collar slender. Style with oblong-linear, truncate branches; stigmatic area continuous. Receptacle flat, without scales. Achenes more or less cylindrical; pappus of hairs. About 50 species in Eurasia and 1 species in North America. Nordenstam, B. (1978). Taxonomic studies in the tribe Senecioneae (Compositae). Opera Bot. 44: 1–83. Stewart, A., Pearman, D. A. & Preston, C. D. (1994). Scarce plants in Britain. Peterborough. [T. integrifolia.] Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough. [T. integrifolia subsp. maritima.] 1. Tall plants to 100 cm of fen ditches; achenes glabrous 2. palustris 1. Usually much shorter plant of dry grassland or coastal 2. cliffs; achenes hairy 2. Basal leaves entire or remotely denticulate; involucral 1(a). integrifolia subsp. integrifolia bracts 6–8 mm 2. Basal leaves rather coarsely crenate-dentate; involucral 1(b). integrifolia subsp. maritima bracts 8–12 mm

1. T. integrifolia (L.) Holub Field Fleawort Senecio integrifolius (L.) Clairv.; Cineraria alpina var. integrifolia L.; Cineraria campestris auct.; Senecio campestris auct.; Othonna integrifolia L.; Cineraria integrifolia (L.) Murr. Short-lived perennial herb with a short, more or less erect stock. Stem 5–50(–90) cm, pale green, erect, arachnoidlanate and minutely glandular hairy, becoming more or less glabrous, branching only in the inflorescence, sparingly leafy. Leaves alternate, greyish or whitish on both surfaces with woolly-arachnoid and minute glandular hairs; basal 2– 10 × 2–8 cm, oblong, ovate-lanceolate or ovate-spathulate or subrotund, obtuse to acute at apex, entire to coarsely crenate-dentate, narrowed, truncate or subcordate at base, with a winged petiole; cauline few to several, narrowly to broadly lanceolate, more or less acute at apex, entire or denticulate, sessile and the lower often semiamplexicaul, the lower sometimes shortly petiolate. Capitula (1–)3–15, 15– 25 mm in diameter, in a simple corymb; peduncles short, minutely glandular-hairy and arachnoid-hairy. Involucral bracts in 1 main row, 6–12 × 0.8–1.0 mm, linear-lanceolate, acute at apex, minutely glandular-hairy and arachnoid-hairy at base. Flowers of 2 kinds, the outer 12–15 ligulate, the ligules bright yellow and shortly 3-lobed at apex, the inner tubular, yellow and with 5 sharp lobes at apex. Receptacle flat, without scales. Achenes 2.5–4.0 mm, more or less cylindrical, usually shortly and densely hairy; pappus 6–7 mm, whitish, of numerous simple hairs. Flowers 5–7. In its total range T. integrifolia is extremely variable and it is difficult to distinguish infraspecific taxa, but in Great Britain the two races are easily distinguished. (a) Subsp. integrifolia Stems 5–30 cm. Basal leaves 2–5 × 0.7–2.0 cm, lanceolate, entire or remotely denticulate, the cauline oblonglanceolate to narrowly lanceolate. Involucral bracts 6–8 mm. 2n = 48. (b) Subsp. maritima (Syme) B. Nord. Senecio campestris var. maritimus Syme; Senecio integrifolius subsp. maritimus (Syme) Chater; Senecio integrifolius var. maritimus (Syme) A. R. Clapham comb. inval.; Senecio spathulifolius auct. Stems 25–60(–90) cm. Basal leaves 5–10 × 3–8 cm, broadly oblong to ovate-spathulate or subrotund, rather coarsely crenate-dentate, the cauline several and broadly lanceolate. Involucral bracts 8–12 mm. 2n = 48. Native. The species as a whole occurs in central and east Europe from England and Wales to the Urals, Roumania and Greece, northwards to 70◦ N in Norway and Arctic Russia, and in temperate Asia eastwards to Japan and in western North America. The T. integrifolia complex (including T. atropurpurea) is a member of the Circumpolar Wide-boreal element. It can be divided into many subspecies. The closely related T. helenitis (L.) B. Nord. is sometimes replacing. Subsp. integrifolia occurs in short, calcareous Festuca grassland, mainly on the chalk but also on the Cotswold oolite. It stretches across the East Anglian Heights to the Sussex and Wiltshire Downs where it is still frequent. It has decreased considerably over the last century, probably due to abandonment of sheep grazing and is

86. Brachyglottis nearly extinct in Kent and on the Cotswold oolite. Subsp. maritima is endemic to glacial drift on sea-cliffs on Holy Island, Anglesey. The only specimen seen from the Pennine area, where the species is now extinct, is that from Mickle Fell, which in all characters is subsp. integrifolia. Plants from the Sussex coast can have very broad leaves, but are not subsp. maritima.

2. T. palustris (L.) Fourr. Marsh Fleawort Othona palustris L.; Cineraria palustris (L.) L.; Senecio palustris (L.) Hook., non Vell.; Senecio arcticus Rupr. Biennial or sometimes perennial herb with a short, stout, erect stock and numerous fibrous roots. Stems 30–100 cm, pale green, furrowed, hollow, stout, long arachnoidhairy and minutely glandular-hairy, branched only above, very leafy. Leaves alternate, pale yellowish-green on upper surface, paler beneath; very shortly glandular-hairy and woolly; basal and lower cauline 7–12 × 2–3 cm, lanceolate to ovate-lanceolate, acute at apex, entire or sinuate-serrate, with large jagged teeth, narrowed into short, stalk-like base, the basal dying before flowering; medium and upper cauline similar to basal but gradually getting smaller, sessile with a broad semiamplexicaul base. Capitula numerous, 20– 30 mm in diameter, in small, dense terminal and axillary corymbs; peduncles short, long arachnoid-hairy and shortly glandular-hairy. Involucral bracts in 1 row, 8–10 × 1.0–1.5 mm, green, linear-lanceolate, acute at apex, long arachnoid and shortly glandular-hairy. Flowers of 2 kinds, the outer ligulate, the ligules sulphur yellow and 2–3 lobed at apex or entire, the inner tubular and deep yellow with 5 triangular lobes at apex. Receptacle flat, without scales. Achenes 1.5–2.0 mm, 10-ribbed, more or less cylindrical, glabrous; pappus 6–7 mm, whitish, of simple hairs. Flowers 6–7. 2n = 48. An extinct native. Formerly in fen ditches. Local in eastern England from Sussex to Yorkshire. Last recorded in 1899. A variable species found in the Arctic and Temperate regions throughout the northern hemisphere, extending north in Europe to central France and central Ukraine. It is a member of the Circumpolar Wide-boreal element. Our plant was subsp. congesta (R. Br.) Holub (Cineraria congesta R. Br.; Senecio congestus (R. Br.) DC.)

85. Delairea Lem. Woody perennial herbs. Stems trailing or climbing; woody below, succulent above. Leaves alternate, succulent, palmately lobed or angled. Capitula numerous in dense axillary and terminal panicles. Involucral bracts in 1 main row with short supplementary ones at the base of the capitulum. Flowers all tubular, 5-lobed at apex and bisexual. Corolla yellow. Anthers with tails, apical appendage ovatelanceolate to oblong, flat; filament collar dilated. Style with oblong-linear, truncate branches; stigmatic areas in 2 separate lines. Receptacle flat, without scales. Achenes subcylindrical; pappus of hairs. One species in South Africa.

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1. D. odorata Lem. German Ivy Senecio mikanioides Otto ex Walp.; Senecio scandens DC., non D. Don Perennial herb. Stems up to 3(–6) m, climbing or trailing, woody below, succulent above, glabrous, branched or not, leafy. Leaves alternate, 2.5–10.0 × 2–10 cm, medium yellowish-green on upper surface, paler beneath, more or less succulent, ovate, deltoid or more or less subrotund, acute at apex, with (3–)5–7(–11) acute angles or triangular lobes and dentate with flat or concave-sided teeth, cordate or truncate at base, sometimes with small, leaf-like auricles, palmately veined, glabrous or with a few hairs below, with a long, glabrous petiole. Capitula 5–7 mm in diameter, in dense axillary or terminal corymbose panicles; peduncles glabrous. Involucral bracts in 1 main row with short supplementary ones at base of capitulum, the main row 3–4 × 0.9– 1.2 mm, oblong, at apex and glabrous, the supplementary ones 2–4, 1.5–3.0 mm, linear, subacute at apex. Flowers all tubular, 5-lobed at apex, bright yellow, fragrant. Receptacle flat, without scales. Achenes 2.0–2.5 mm, subcylindrical, glabrous or with scattered hairs on the ribs; pappus of many fine hairs. Flowers 6–9. 2n = 20. Introduced. Climbing over hedges and walls. Naturalised in the Channel Isles and the Isles of Scilly, rarely in mainland Cornwall. Native of South Africa. 86. Brachyglottis J. R. Forster & G. Forster Urostemon B. Nord. Shrubs or small trees. Leaves alternate, entire to remotely dentate or sinuately lobed, densely white-felted. Capitula solitary on long peduncles, or numerous in panicles. Involucral bracts in 1 main row with short supplementary ones at the base of the capitulum. Inner flowers tubular, 5-lobed and bisexual, the outer ligulate, 3-lobed and female. Corolla of tubular flowers cream or yellow, of ligulate flowers yellow. Anthers with or without tails, apical appendage oblong and flat; filament collar slender. Style with oblong-linear branches with stigmatic areas confluent except basally. Receptacle flat, without scales. Achenes subcylindrical or linear-oblong; pappus of hairs. Thirty-one species in New Zealand and Tasmania. Nordenstam, B. (1978). Taxonomic studies in the tribe Senecioneae (Compositae). Opera Bot. 44: 1–83. 1. Many leaves exceeding 8 cm, sinuate-dentate; involucral 3. repanda bracts glabrous or hairy only at base 1. Leaves less than 8 cm, entire, denticulate or undulate-crenate; involucral bracts glandular or with 2. woolly hairs all over 2. Many leaves exceeding 4 cm, entire to remotely 1. × jubar denticulate 2. monroi 2. Leaves less than 4 cm; undulate-crenate

1. B. × jubar P. D. Sell Shrub Ragwort Probably B. compacta (Kirk) B. Nord. × laxifolia (Buchanan) B. Nord. Senecio greyi auct.; B. cv. Sunshine Shrub up to 1(–2) m. Branches numerous, spreading and somewhat straggling, grey-woolly; twigs rather densely leafy, white-woolly at first. Leaves alternate; lamina (1.5–)3–8 × 2–4 cm, dark green on upper surface, white

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beneath, ovate, obovate, elliptical or oblong-elliptical, obtuse at apex, entire to remotely denticulate, rounded, subtruncate or shortly cuneate at base, tomentose on upper surface when young, white-woolly beneath; petiole up to 30 mm, woolly. Capitula 20–45 mm in diameter, in a loose terminal corymb; peduncles white-woolly. Involucral bracts in 1 main row with short supplementary ones, 5–7 × 0.5–1.2 mm, greyish-green, linear-lanceolate or oblong, obtuse or subacute at apex, grey-woolly. Flowers of 2 kinds, the outer 10–15, 10–15 mm, ligulate, 3-lobed at apex and yellow, the inner 6–7 mm, tubular, 5-lobed at apex and yellow or orange, turning brownish. Receptacle flat, pitted, without scales. Achenes 2.0–2.5 mm, linear-oblong, sparsely short-hairy; pappus of slender, whitish hairs. Flowers 7–8. 2n = 60. Introduced. Much grown in gardens and often massplanted along roads. Persistent in rough ground and on dunes. Scattered records in England and Wales. First recorded in 1910–13 in Dunedin Botanic Garden, New Zealand and now a widespread plant of gardens. 2. B. monroi (Hook. fil.) B. Nord. Monro’s Ragwort Senecio monroi Hook. fil. Shrub up to 1 m. Branches numerous, spreading; twigs compact, densely leafy, short white-woolly. Leaves alternate; with lamina 2–4 × 0.5–1.5 cm, brownish-green on upper surface, white below, oblong, oblong-obovate or oblonglanceolate, obtuse at apex, thickened at margin and distinctly undulate-crenate, narrowed at base, thick, finely reticulate veined and slightly glandular-viscid on upper surface, shortly white-woolly beneath with the midrib raised but the venation not visible; petiole short. Capitula 18– 25 mm in diameter, in a lax corymb; peduncles slender, white-woolly and with short glandular hairs. Involucral bracts in 1 main row with short supplementary ones, 5– 10 × 1.0–1.2 mm, green, the inner with a pale margin, linear, acute at apex, with very short glandular hairs, the outer spreading. Flowers of 2 kinds, the outer 10–15, ligulate, the ligules yellow, 3-lobed at apex and spreading to recurved, the inner tubular, numerous, brownish and with 5 lobes at apex. Receptacle flat, pitted, without scales. Achenes 1.5– 2.0 mm, linear-oblong, grooved, shortly hairy; pappus up to 5 mm, of slender, whitish hairs. Flowers 7–8. 2n= 60. Introduced. Persistent and more or less naturalised on dunes near Llandudno in Caernarvonshire. Native of South Island, New Zealand. 3. B. repanda J. R. & G. Forst. Hedge Ragwort Cineraria repanda (J. R. & G. Forst.) Forst. fil.; Senecio georgii Endl.; Senecio forsteri Hook. fil.; B. rangina Buchanan Shrub or small tree up to 6 m. Branches spreading, stout, brittle; twigs densely clad in soft, white to buff, subappressed tomentum, leafy. Leaves alternate with lamina 5–25 × 5–20 cm, dark green on upper surface, whitish beneath, ovate, ovate-oblong or elliptic-oblong, obtuse to subacute at apex, sinuate to dentate-sinuate, obliquely cordate to truncate at base, membranous to subcoriaceous, glabrous on upper surface, tomentose beneath; petiole up to 10 cm, stout, grooved. Capitula 4–5 mm in diameter, numerous, in large dense, terminal, often drooping, pyramidal panicles; peduncles short, tomentose. Involucral bracts

in 1 main row with short supplementary ones, 3.0–3.5 × 0.2–0.5 mm, pale except for midrib, scarious, oblong, obtuse at apex, glabrous or with woolly hairs only at base. Flowers 8–12, of 2 kinds, the outer 2-lipped to ligulate, the upper lip short, broad and obscurely lobed to entire, the lower lip 1- to 2-lobed and white to creamy white, the inner tubular-campanulate. Receptacle flat, pitted, without scales. Achenes very small, subcylindrical, hispid-papillose; pappus of white hairs. Flowers 7–8. 2n = 60. Introduced. Used for hedging in the Isles of Scilly where it is often long-persistent after neglect. Native of New Zealand. 87. Sincalia H. Rob. & Brettell Perennial herbs with an underground tuber. Stems erect, leafy. Leaves alternate, pinnately lobed. Capitula numerous, in large terminal panicles. Involucral bracts in 1 main row, with small bracts some way below the base of the capitulum. Inner flowers tubular, 5-lobed and bisexual, the outer ligulate, 3-lobed and female. Corolla of all flowers yellow. Anthers without tails, apical appendage oblong and flat; filament collar slender. Style with oblong-linear branches with stigmatic areas mainly continuous but basally separate. Receptacle flat, without scales. Achenes subcylindrical; pappus of hairs. Four species in China. Nordenstam, B. (1978). Taxonomic studies in the Tribe Senecioneae (Compositae). Opera Bot. 44: 40–83.

1. S. tangutica (Maxim.) B. Nord. Chinese Ragwort Senecion tanguticus Maxim.; Ligularia tangutica (Maxim.) Mattf.; Cacalia tangutica (Maxim.) Hand.-Mazz.; Senecilis tangutica (Maxim.) Kitam.; Senecio henryi Hemsl.; Sincalia henryi (Hemsl.) H. Rob. & Brettell Perennial herb with an underground tuber. Stems 100– 200 cm, erect, robust, pale yellowish-green, often suffused reddish-purple, striate, glabrous or arachnoid-hairy, unbranched except in the inflorescence, leafy. Leaves alternate, gradually decreasing in size upwards, 4–20 × 3–15 cm, medium yellowish-green on upper surface, pale greyish-green beneath with prominent yellowish veins, glabrous, broadly ovate in outline, sharply acute at apex, pinnately divided nearly to the midrib, the lobes oblong to triangular-lanceolate, the lower dentate and the upper entire to denticulate, cuneate or truncate at base to a rather long petiole. Inflorescence a large, terminal, pyramidal panicle; rhachis and peduncles glabrous or with short simple eglandular hairs. Capitula very numerous, 10–20 mm in diameter, cylindrical. Involucral bracts in 1 row with small bracts some way below the capitulum, 6.5–7.5 × 0.8–1.0 mm, pale yellowish-green, with a brownish tip, linear, more or less obtuse at apex, glabrous. Flowers of 2 kinds, the outer 3– 4, 5–9 mm, ligulate and female, the ligules yellow and 3lobed at apex, the inner 3–4, tubular and bisexual, orange and with 5 revolute lobes at apex. Receptacle flat, without scales. Achenes 2.0–2.5 mm, pale brown, sub-cylindrical, ribbed; pappus 4–5 mm, of whitish simple hairs. Flowers 7–9.

89. Doronicum Introduced. Grown in gardens, escaping and becoming naturalised in damp shady places. Scattered records in northern Wales and northern England and Renfrewshire in Scotland; naturalised for more than 30 years by the River North Esk in Forfarshire. Native of China. 88. Ligularia Cass. Perennial herbs with creeping rhizomes. Stems erect, leafy. Leaves alternate, reniform, dentate, palmately veined, with sheathing petioles. Capitula few to numerous, in a terminal, corymbose cluster. Involucral bracts all in 1 row. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female. Corolla yellow to orange. Anthers without tails, apical appendge oblong and flat; filament collar slender. Style with oblong-linear branches, stigmatic areas continuous. Receptacle convex without scales. Achenes subcylindrical; pappus of hairs. About 125 species in temperate Eurasia. 1. Basal leaves dentate; capitula 40–100 mm in diameter; 1. dentata ligules orange 1. Basal leaves deeply palmately lobed; capitula 15–30 mm 2. przewalskii in diameter; ligules yellow

1. L. dentata (A. Gray) H. Hara Leopard-plant Erythrochaete dentata A. Gray; L. clivorum Maxim.; Senecio clivorum (Maxim.) Maxim. Perennial herb with creeping rhizome forming large patches. Stems 45–120 cm, erect, robust, fleshy, hollow, yellowish-green, often suffused brownish-purple, angled, slightly striate, glabrous, unbranched except in the inflorescence, sparingly leafy. Leaves alternate, dull medium yellowish-green on the upper surface and with impressed veins, pale greyish-green beneath with markedly prominent, purplish, palmate veins, glabrous; basal numerous 20–50 × 20–50 cm, subrotund, broadly ovate or reniform, broadly rounded to subacute at apex, dentate or crenatedentate, the teeth more or less mammiform, widely cordate at base, the petioles up to 75 cm, usually brownishpurple, rounded above, flattened beneath, glabrous; cauline much smaller but similar in shape and toothing, the petioles broadly winged and amplexicaul. Inflorescence a small, subcorymbose cluster; peduncles often brownish-purple with short, rather crisp or subarachnoid hairs; bracts large, ovate, acute at apex, entire, slightly clasping. Capitula 40– 100 mm in diameter. Involucral bracts in 1 row, 12–16 × 4–6 mm, dull medium green with broad scarious margins, tinted red but the margins cannot be seen because the phyllaries overlap, ovate, acuminate at apex, with short, crisped or subarachnoid hairs. Flowers of 2 kinds, the outer 15–50 mm, ligulate and female, the ligules orange and shortly 3lobed at apex, the inner 10–12 mm, tubular and bisexual, brown and narrowly 5-lobed at apex. Receptacle slightly convex, pitted, without scales. Achenes 11–12 mm, reddishpurple, subcylindrical, ribbed; pappus 10–12 mm, brown, of rather stiff hairs. Flowers 7–11. 2n = 60. Introduced. Grown in gardens from which it escapes and becomes naturalised in damp and shady places. Very scattered records in England and Scotland, especially in the north. Native of China and Japan.

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2. L. przewalskii (Maxim.) Diels Przewalski’s Leopard-plant Senecio przewalskii Maxim. Perennial herb with long rhizomes. Stems up to 1.8 m, usually suffused deep brownish-purple, stout, hollow, markedly ridged and channelled, glabrous. Leaves alternate, dull dark to medium green on upper surface, glaucous with marked venation beneath; basal 15–40 × 15–50 cm, broadly ovate in outline, acute at apex, deeply palmately lobed, the lobes triangular or oblong and toothed or lobulate, both lobes and lobules acute at apex, deeply cordate at base, with very long, brownish-purple, striate petioles; cauline often similar to basal and petiolate or the upper only deeply toothed and the uppermost sometimes sessile; all glabrous. Inflorescence a long, narrow, raceform panicle; peduncles 2–5 mm, slender, glabrous. Capitula numerous, 15–30 mm in diameter. Involucral bracts in 1 row, 9–11 × 1.5–2.0 mm, medium green, tinted red near the apex, linear, shortly narrowed to a subacute apex, glabrous. Flowers of 2 kinds, the 2 outer 6–15 mm, ligulate and female, the ligules 3-lobed at apex and yellow, the 3 inner tubular 5-lobed and bisexual, brownish-yellow. Receptacle slightly convex, pitted, without scales. Achenes 10–12 mm, purplish, subcylindrical, ribbed; pappus of rather stiff hairs. Flowers 7–11. 2n = 60. Introduced. Grown in gardens and a persistent escape by the River Tyne in Northumberland and in a wild bog garden at Witherslack in Westmorland. Native of northern China. 89. Doronicum L. Rhizomatous perennial herbs. Stems erect, leafy. Leaves alternate, simple, entire to dentate; veins more or less palmate. Capitula solitary or in terminal corymbs. Involucral bracts in 2 rows of equal length. Inner flowers tubular, 5lobed and bisexual, the outer ligulate, 3-lobed and female and in 1 row. Corolla yellow. Anthers without tails, apical appendage oblong and flat; filament collar slender. Style with oblong-linear branches, stigmatic areas continuous. Receptacle convex, without scales. Achenes oblanceolate; pappus of 1–2 rows of simple hairs, sometimes absent in outer flowers. About 40 species in Europe, Asia and Africa, especially in the mountains of temperate Asia. 3. plantagineum 1. Basal leaves all cuneate at base 1. Basal leaves mostly or all truncate, rounded or cordate at 2. base 2. Petioles of basal leaves with many flexuous or patent hairs longer than 1 mm; capitula usually 3–6 per stem 1. pardalianches 2. Petioles of basal leaves without hairs longer than 1 mm or with very few hairs longer than 1 mm; capitula 1–2(–3) 3. per stem 3. Basal leaves deeply cordate at base; all hairs on the stem 5. columnae glandular and less than 1 mm 3. Basal leaves shallowly cordate, truncate or rounded at base; stems usually with a few, long simple eglandular 4. hairs as well as short glandular ones 4. Basal leaves obtuse to rounded at apex, mainly rounded to truncate at base, with teeth less than 2 mm 2. × willdenowii 4. Basal leaves acute at apex, mainly shallowly cordate at 4. × excelsum base, with prominent teeth more than 2 mm

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1. D. pardalianches L. Leopard’s-bane D. cordatum Lam.; D. cordifolium Stokes Perennial herb with a rhizome with tufts of sericeous hairs. Stems up to 90 cm, erect, yellowish-green, with numerous, short, yellowish glandular hairs intermixed with few to numerous, long, pale simple eglandular hairs especially in the lower half. Leaves alternate, dull yellowish-green on upper surface, paler beneath; basal (6–)7–12(–15) × (5–)7– 11(–14) cm, ovate to ovate-subrotund, rounded at apex, subentire to very shallowly undulate-dentate or undulatedenticulate, the teeth broad and flat, cordate at base, the petioles up to 10 cm and pale yellowish-green; lower cauline ovate, cordate and petiolate, the middle cauline often panduriform and the upper cauline ovate-lanceolate or lanceolate and more or less amplexicaul; all with numerous, short, pale glandular and few to numerous, pale simple eglandular hairs on both surfaces and the margins, the petioles with numerous, short glandular hairs and more or less numerous, medium and long, pale simple eglandular hairs. Capitula (2–)3–6(–8), in a terminal corymb, 30–50(–60) mm in diameter; peduncles very long, with numerous, short, pale glandular hairs and few, longer, pale simple eglandular hairs. Involucral bracts (8–)12–18 × 1.8–2.0 mm, yellowish-green, linear-lanceolate, gradually narrowed to a sharp apex, with dense, short, pale glandular hairs. Flowers of 2 kinds, outer in 1 row, ligulate and female, the ligules 11–14 × 2.5–3.0 mm, yellow, linear and with 2–3 lobes at apex, the inner 5–6 mm, tubular and bisexual, 5-lobed at apex and yellow. Receptacle convex, with dense, short, pale glandular and simple eglandular hairs. Achenes 2.0–2.5 mm, those of outer flowers without a pappus and glabrous, pale yellowish-green with a darker green band at the apex, oblanceolate, with dense, pale, appressed simple eglandular hairs; pappus 4.0–4.5 mm, of whitish denticulate hairs. Flowers 5–7. Visited by flies, beetles and Lepidoptera. 2n = 60. Introduced. Well naturalised in woods and shady places. Frequent throughout Great Britain and Ireland. West Europe eastwards to Italy, western Switzerland and western Germany. 2. D. × willdenowii (Rouy) A. W. Hill Willdenow’s Leopard’s-bane D. pardalianches × plantagineum D. plantagineum var. willdenowii (Rouy) A. B. Jacks.; D. plantagineum forma willdenowii Rouy Perennial herb with a rhizome with hairs at the nodes. Stems up to 1 m, erect, yellowish-green, markedly striate, with numerous, short, colourless glandular hairs and usually a few, longer simple eglandular hairs especially in the lower half. Leaves alternate, dull yellowish-green on upper surface, paler beneath; basal 5–12(–17) × 5–10 cm, more or less ovate, obtuse to rounded at apex, entire to shallowly undulate-dentate, rounded, truncate or shortly cuneate at base, the petioles up to 17 cm and pale yellowish-green; lower cauline ovate to oblong-ovate, obtuse at apex, entire to shallowly undulate-dentate especially towards the base, the upper gradually getting smaller, more or less ovate, obtuse to acute at apex, entire or shallowly dentate; all with short, colourless glandular hairs on both surfaces and the margins

and some longer, pale simple eglandular hairs particularly on the veins beneath and the petioles. Capitula 1–2(–3) per stem, 45–80 mm in diameter; peduncles with numerous, short but unequal, colourless glandular hairs and few, much longer, pale simple eglandular hairs. Involucral bracts 20– 25 × 2.0–2.5 mm, yellowish-green, linear-lanceolate, with a long-drawn-out acute apex, with numerous to dense, short, pale glandular hairs. Flowers of 2 kinds, the outer in 1 row, ligulate and female, the ligules 28–32 × 2.5–3.0 mm, yellow, linear and with 2–3 lobes at apex, the inner 5–6 mm, tubular, bisexual, 5-lobed at apex and yellow. Receptacle convex, with numerous, pale, rather stiff hairs. Achenes poorly developed, 1.5–2.5 mm, pale brown, oblanceolate, those of the tubular flowers appressed-hairy, those of the ligulate flowers glabrous; pappus of 1–2 rows of whitish, denticulate hairs, absent in the ligulate flowers. Flowers 4–6. Introduced. Grown in gardens and naturalised in woods and shady places. Scattered records in Great Britain. Probably under-recorded for D. plantagineum. Western Europe or of garden origin. 3. D. plantagineum L. Plantain-leaved Leopard’s-bane Perennial herb with a rhizome with tufts of sericeous hairs at the nodes. Stems up to 1 m, erect, yellowish-green, markedly striate, with numerous, very short glandular hairs. Leaves alternate, dull yellowish-green on upper surface, paler beneath; basal 4.5–11.0 × 2.5–5.0(–6.0) cm, more or less ovate, more or less obtuse at apex, entire to shallowly undulate-dentate, all cuneate at base, with petioles up to 15 cm; cauline 2–4, ovate to ovate-elliptical, obtuse to acute at apex, entire or shallowly undulate-dentate, the lower often with a winged petiole, the upper sessile and semiamplexicaul; all with few to numerous, colourless glandular hairs and a very occasional simple eglandular hair. Capitula usually solitary, 30–60 mm in diameter; peduncles with numerous, short glandular hairs. Involucral bracts 14– 20 × 1.5–2.0 mm, yellowish-green, linear-lanceolate, with a long-drawn-out acute apex, with numerous to dense, short, pale glandular hairs and longer, simple eglandular ones along the margin. Flowers of 2 kinds, outer in 1 row, 15– 30 × 2.0–2.5 mm, ligulate and female, yellow, linear and with 2–3 lobes at apex, the inner 4.5–5.5 mm, tubular and bisexual, 5-lobed at apex and yellow. Receptacle convex, with long, pale hairs. Achenes 2.0–2.5 mm, pale brown, oblanceolate, those of the tubular flowers appressed hairy, those of the ligulate flowers glabrous; pappus of 1–2 rows of whitish, denticulate hairs, absent in the ligulate flowers. Flowers 6–7. 2n = 120. Introduced. Grown in gardens, escaping and becoming naturalised in woods and shady places. Scattered records in Great Britain, but over-recorded for its hybrids. Native of western Europe northwards to northern France. 4. D. × excelsum (N. E. Br.) Stace Harper-Crewe’s Leopard’s-bane ?D. columnae × pardalianches × plantagineum D. plantagineum var. excelsum N. E. Br.; D. × draytonense hort.; D. × hybridum hort. Perennial herb with rhizomes. Stems up to 1 m, pale yellowish-green, erect, with numerous very short, pale

90. Tussilago glandular hairs and scattered pale, medium to long simple eglandular hairs. Leaves alternate, dull, medium yellowishgreen on upper surface, paler beneath; basal 4–8 × 3–7 cm, ovate, more or less acute at apex, sinuate-dentate with rather shallow teeth, mostly more or less cordate but sometimes rounded or truncate at base, the petioles up to 14 cm, pale yellowish-green, channelled; cauline similar to basal but more closely toothed, sometimes panduriform and more or less amplexicaul at base or the lower petiolate; all with numerous, short, pale glandular and scattered short to medium, pale, simple eglandular hairs on both surfaces and the petiole, and more numerous on the margins, arachnoid-hairy in the channel on the petioles. Capitula 1–2(–3) per stem, 60–90 mm in diameter; peduncles very long, with numerous, unequal, very short to short, pale glandular and occasional, slightly longer, pale simple eglandular hairs. Involucral bracts in 2–3 rows, 15–20 × 2.0–2.5 mm, yellowish-green, narrowly linear-lanceolate, gradually narrowed to a sharp point at apex, with dense, very short and short, pale eglandular and few to numerous, slightly longer, pale simple eglandular hairs. Flowers of 2 kinds, the outer in 1 row, ligulate and female, the ligules 25–30 × 3.5–4.0 mm, yellow, linear, with 2–3 lobes at apex and glabrous, the inner 5–6 mm, tubular, bisexual, 5-lobed at apex and yellow. Receptacle convex, with dense, short, colourless glandular and simple eglandular hairs, without scales. Achenes not well-formed, 2.5–3.5 mm, yellowish-green, with a darker green rim at apex, oblanceolate, with dense, short simple eglandular hairs; pappus 5–6 mm, of 1–2 rows of denticulate hairs. Flowers 5–6. Introduced. Grown in gardens and escaping and becoming naturalised in woods and shady places. Scattered records in Great Britain. Garden origin or western Europe. 5. D. columnae Ten. Eastern Leopard’s-bane D. cordatum auct. Perennial herb with glabrous or sparsely hairy rhizomes. Stems 12–60 cm, erect, yellowish-green, with a few, short, pale glandular hairs. Leaves alternate, dull medium yellowish-green on upper surface, paler beneath; basal 3–8(–9) × 3–7(–9) cm, ovate to ovate-subrotund, more or less acute at apex, undulate-dentate the teeth subobtuse, cordate at base, the petioles up to 22 cm, pale green; cauline 3–4, similar to basal but more or less amplexicaul; all with very short, pale glandular and few simple eglandular hairs on both surfaces, the margins and petioles. Capitula 1(–3), (20–)25–50(–60) mm in diameter; peduncles very long, with numerous, short and very short glandular hairs. Involucral bracts in 2–3 rows, 8–14(–18) × 1.5–2.0 mm, narrowly linear-lanceolate, gradually narrowed to a sharply pointed apex, with dense, short, pale glandular hairs. Flowers of 2 kinds, the outer in 1 row, ligulate 3-lobed at apex and female, yellow, 16–28(–36) mm and glabrous, the inner 4.0–4.5 mm, tubular, 5-lobed at apex, bisexual and yellow. Receptacle convex, with dense, short, pale glandular and simple eglandular hairs, without scales. Achenes 1.4– 1.5 mm, the marginal without a pappus and glabrous, pale yellowish-green with a darker green band near the apex, oblanceolate, with dense, pale, appressed simple eglandular hairs; pappus 3.5–4.0 mm, of whitish, denticulate hairs. Flowers 5. 2n = 60.

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Introduced. Naturalised on the bank of a reservoir in Surrey and in a Cambridgeshire churchyard. Perhaps overlooked elsewhere. Native of south-east Europe. 90. Tussilago L. Rhizomatous perennial herbs. Stems erect, scaly, but not leafy. Leaves all basal, appearing after the flowers, cordate at base, dentate, arachnoid-hairy beneath. Capitulum solitary, terminal. Involucral bracts all in 1 row. Inner flowers tubular, 5-lobed, few, and functionally male, the outer ligulate, numerous, 3-lobed at apex and female. Corolla yellow. Anthers without tails, apical appendage oblong and flat; filament collar slender. Style with oblong-linear branches, stigmatic areas contiguous. Receptacle slightly concave, without scales. Achenes narrowly cylindrical; pappus of 1 row of numerous denticulate hairs. One species in Eurasia and North Africa. 1. T. farfara L. Colt’s-foot T. vulgaris Lam. nom. illegit.; T. ruderalis Salisb. nom. illegit Perennial herb with long, rather stout, white, scaly stolons, these and their short branches terminating in rosettes of leaves in whose axils arise the flowering shoots of the following season. Leaves appearing after the flowers, all basal, numerous, 8–20 × 10–30 cm, at first white-felted on both surfaces, later upper surface becoming glabrous and yellowish-green, broadly ovate or subrotund-polygonal, shortly acute at apex, very shallowly 5- to 12-lobed, the lobes broadly triangular and acute at apex, and with small, distant, blackish teeth, deeply cordate at base; petiole 10– 30 cm, pale green often tinted purplish, rounded beneath, broadly furrowed above, with arachnoid hairs. Flower stem 5–15 cm, elongating up to 30 cm in fruit, erect at first, after flowering drooping, then becoming erect again when fruit is ripe, pale green, sometimes flushed brownish-red, whitewoolly, covered throughout with scales, the scales 10–15 × 3–7 mm, appressed to stem, dull green, suffused brownisholive or purplish, tipped red, lanceolate, linear-lanceolate or linear, obtuse at apex, entire. Capitulum solitary, terminal, 15–35 mm in diameter. Involucral bracts numerous, mostly in 1 row, 7–15 × 1.5–2.5 mm, pale yellowish-green, the outer with narrow, pale margins, linear, slightly redtipped at the more or less acute apices, the inner longer and broader with wide, reddish-suffused margins and broadly rounded apices, curved outwards at the top when the flowers are open. Flowers up to 300, of 2 kinds, the outer numerous, ligulate and female, the ligule 12–17 mm, 3-lobed at apex and bright pale yellow, sometimes tipped darker, the inner few, tubular and male, 9–11 mm, 5-lobed at apex and yellow. Receptacle slightly concave, without scales. Achenes 5–10 mm, pale, narrowly cylindrical, 5-ribbed, glabrous; pappus 10–14 mm, of many rows of whitish simple eglandular hairs. Flowers 3–4. Visited chiefly by flies and bees, the flowers closing at night. 2n = 60. Native. Waste places, banks, landslides, boulder-clay cliffs, dunes, screes, streamside shingle and seepage-fens on hillsides; formerly a troublesome weed of agriculture now mostly controlled with herbicides. Throughout Great Britain and Ireland reaching, 1,070 m on Ben Lawers in Perthshire. Europe northwards to 71◦ N in Norway; western

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and northern Asia; North Africa; introduced in North America. A member of the Eurosiberian Boreo-temperate element. 91. Petasites Mill. Dioecious, rhizomatous perennial herbs. Stems erect, with few to many bracts. Leaves all basal, cordate at base, dentate, more or less palmately veined, arachnoid-hairy on underside. Capitula numerous, in a terminal raceme or panicle. Involucral bracts in 1–2 rows. Male capitula with numerous tubular, functionally male flowers, usually with a peripheral ring of 1–5(–10) sterile ligulate or tubular female flowers. Female capitula with numerous fertile ligulate or tubular female flowers and 0–5(–8) sterile tubular flowers in the centre. Corolla white, cream or purple. Anthers without tails, apical appendage oblong and flat; filament collar slender. Style with oblong-linear branches, stigmatic areas contiguous. Receptacle flat, without scales. Achenes cylindrical; pappus of simple hairs, numerous in female, few in male flowers. Nineteen species in Eurasia and North America. Grime, J. P. et al. (1988). Comparative plant ecology. London. [P. hybridus.] Toman, J. (1972). A taxonomic survey of the genera Petasites and Endocellion. Folia Geobot. Phytotax. 7: 381–406. Valentine, D. H. (1947). The distribution of the sexes in Butterbur. North Western Nat. 1947: 111–114. 1. Leaves present at anthesis, regularly denticulate, the small teeth all of 1 size; flowering from November to February, plants always male with outer female flowers ligulate 4. fragrans 1. Leaves absent at anthesis, irregularly dentate, with large teeth or shallow lobes dispersed amongst small teeth; flowering from February to May, plants male or female, 2. flowers all tubular 2. Leaves usually less than 30 cm in diameter, sinus of basal with parallel or divergent sides bordered by 0–1 veins on 3. albus each side; flowers pure white 2. At least some leaves usually more than 30 cm in diameter, sinus of basal with convergent sides and more than 2 veins on each side; flowers cream or with a purplish tinge 3. 3. Leaves distinctly, but very shallowly lobed; upper part of flowering stem with bracts less than 10 mm wide; inflorescence more or less cylindrical; involucral bracts 1. hybridus subsp. hybridus and/or flowers tinted purplish 3. Leaves scarcely or not lobed; upper part of flowering stem with bracts more than 10 mm wide; inflorescence more or less hemispherical when mature; involucral bracts greenish, flowers creamy-white 2. japonicus subsp. giganteus

Subgenus 1. Petasites Corolla tube of female flowers with a truncate or laciniate mouth, the laciniae being shorter than the double diameter of the corolla tube. In the capitula of andromorphous (‘male’) individuals female flowers occur only in small numbers and form only 1 marginal row which is usually incomplete, and they are mostly sterile or are absent. In gynomorphous (‘female’) individuals, female, fully fertile flowers form the whole capitulum with the exception of a

few, central, regular pseudobisexual agamic flowers which are only rarely absent. 1. P. hybridus (L.) P. Gaertn., B. Mey. & Scherb. Butterbur Tussilago hybrida L.; Tussilago petasites L.; P. ovatus Hill; P. elatior Hill nom. illegit.; P. officinalis Moench nom. illegit.; P. vulgaris Desf. nom. illegit.; P. petasites (L.) Karst. nom. illegit. Perennial herb with a stout, more or less horizontal rhizome. Leaves 10–90 cm in diameter, all basal, medium yellowishgreen on upper surface, grey beneath, subrotund, broadly ovate or reniform, broadly rounded at apex, obscurely lobed, with larger distant teeth and smaller intervening teeth, cordate at base, each basal lobe bordered by 2–5 lateral veins, at first downy on both surfaces, but later much less hairy above and green, grey arachnoid-hairy beneath; petioles up to 1.5 m, stout, hollow, channelled above, rounded beneath, arachnoid-hairy. Flowering stems 10–40 cm, the female to 1 m in fruit, appearing before the leaves, yellowish-green, often suffused brownish-purple, stout, hollow, with dense arachnoid hair mixed with numerous, pale, short glandular hairs; scale leaves numerous, sometimes with a rudimentary lamina, 0.5–9.0 × 0.2–1.5 cm, yellowish-green, ovateoblong to narrowly linear-lanceolate, gradually narrowed to a more or less obtuse apex, glabrous to arachnoid-hairy. Capitula numerous, 3–12 mm in diameter, in a loose, terminal corymb, cylindrical when mature, the male 7–12 mm in diameter, with very short peduncles, 0–3 female flowers and 20–40, sterile ‘bisexual’ flowers, the female 3–6 mm in diameter, lengthening in fruit, with longer peduncles, about 100 female flowers and 1–3 sterile flowers. Involucral bracts irregularly 2- to 3-rowed, 5–7 × 1.0–1.5 mm, green, tinted purplish with a pale margin, linear to narrowly linear-oblanceolate, obtuse to acute at apex, glabrous. Corollas all tubular, 5-lobed at apex, white with a purplish tinge; stigma of male flowers 0.5–1.3 mm, divided only at apex. Receptacle flat, without scales. Achenes 2–3 mm, cylindrical, glabrous; pappus of long, white, slender simple eglandular hairs. Flowers 3–5. Visited chiefly by bees; only the sterile flowers secrete nectar. 2n = 60. Native. Wet meadows, copses and by streams up to 460 m in Scotland. The male plant is locally common throughout Great Britain and Ireland. The female plant is not uncommon in Lancashire, Yorkshire, Cheshire, Derbyshire and Lincolnshire but rare or absent elsewhere, though extending north to Perthshire. Europe to 63◦ 26 N in Scandinavia, north and west Asia. A member of the European Temperate element. Our plant is subsp. hybridus which occurs throughout Europe except for the southern part of the Balkan peninsula. It is introduced in Iceland and North America. 2. P. japonicus (Siebold & Zucc.) Maxim. Great Butterbur Nardosmia japonica Siebold & Zucc. Perennial herb with long rhizomes and fibrous roots. Leaves 30–100 cm in diameter, all basal, medium yellowish-green on upper surface, greyish-green beneath, subrotund to reniform, broadly rounded at apex, not angled, sharply and irregularly dentate, cordate at base, the sinuses convergent

92. Homogyne and with about 5 veins on each side, glabrous on upper surface, arachnoid-hairy beneath; petiole long, hollow, flat above, rounded beneath, arachnoid-hairy. Flowering stems 30–120 cm, appearing before the leaves, pale yellowishgreen, robust, hollow, angled, with numerous to dense, short to long, slender, unequal, pale glandular hairs; scaleleaves numerous, only sometimes with a fully developed lamina, 5–12 × 3–6 cm, yellowish-green, ovate to oblong-ovate, rounded-obtuse at apex, entire, with prominent longitudinal veins. Capitula numerous, 10–15 mm in diameter, in a loose terminal corymb; more or less hemispherical when mature, peduncles short, yellowish-green, ridged, glandular-hairy, the top of the flowering stem looking like a small cauliflower. Involucral bracts in 1–2 rows, 13–15 × 1.5–2.0 mm, pale green with darker longitudinal lines and scarious margins, oblong, rounded-obtuse at apex, glandular-hairy. Corollas all tubular, creamy-white, fragrant; stigma of male flowers 1.6–1.8 mm. Receptacle flat, without scales. Achenes 3–4 mm, cylindrical, glabrous; pappus of long, white, slender simple eglandular hairs. Flowers 3–4. 2n = 84–87. Introduced. Naturalised by rivers and in damp places. Scattered localities throughout most of Great Britain; the female plant rarely or never naturalised here. Our plant is subsp. giganteus Kitam. (P. amplus Kitam.) which is native of northern Japan, Sakhalin and the Kurile Islands. 3. P. albus (L.) Gaertn. White Butterbur Tussilago alba L. Perennial herb with long rhizomes, not thickened at the nodes and with fibrous roots. Leaves 15–30 cm in diameter, all basal, dark green on upper surface, white beneath, subrotund or very broadly ovate, acute at apex, with large, sharp teeth or shallow lobes at regular intervals terminating the main veins, the intervening spaces denticulate, cordate at base, the sinus with parallel or divergent sides and bordered by 0–1 veins on each side, glabrous on upper surface, white arachnoid-hairy beneath; petioles up to 30 cm long, arachnoid-hairy. Flowering stems 30–70 cm, appearing before the leaves, pale yellowish-green, with long, slender glandular hairs and slightly arachnoid-hairy; scale leaves (5–)20–40 × 1.8–8.0 mm, only sometimes with a fully developed lamina, pale yellowish-green, linear-lanceolate, acute at apex, entire, arachnoid-hairy. Capitula 10–15 mm in diameter, in a loose, terminal corymb, male and female inflorescences separate; peduncles short, yellowish-green, glandular-hairy. Involucral bracts in 1–2 rows, 6–12 × 1.4– 1.5 mm, pale yellowish-green, with scarious margin, linearlanceolate, acute at apex, glandular-hairy. Corollas all tubular, pure white; stigma of male flowers 1.5–2.8 mm, divided almost to base. Receptacle flat, without scales. Achenes 2–3 mm, cylindrical; glabrous; pappus of long, white, slender simple eglandular hairs. Flowers 3–5. Visited by various insects. 2n = 60. Introduced. Naturalised in rough ground, waysides and woods. Throughout Great Britain but rare in the south and common in the north; northern Ireland. Native of Europe and the Caucasus. The female plant is much less common than the male in Great Britain and Ireland.

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Subgenus 2. Nardosmia (Cass.) Peterm. Nardosmia Cass. Corolla of female flowers distinctly ligulate, the ligules 1–9 mm, at least twice as long as the diameter of the throat of the corolla tube. In the capitula of andromorphous (‘male’) plants female flowers usually form several outer rows. In gynomorphous (‘female’) plants these flowers with ligulate corollas form the whole capitulum and are fully fertile with the exception of several agamous central flowers which are rarely absent. 4. P. fragrans (Vill.) C. Presl Winter Heliotrope Tussilago fragrans Vill.; Nardosmia fragrans (Vill.) Rchb.; Nardosmia denticulata Cass. nom. illegit. Perennial herb with far-creeping rhizome and fibrous sideroots. Leaves 10–20 cm in diameter, all basal, medium yellowish-green on upper surface, greyish-green beneath, subrotund to reniform, broadly rounded at apex, regularly denticulate, cordate at base, the lobes near the base divergent, with 2–5 lateral veins bordering the broad sinus, glabrous on upper surface, hairy beneath; petiole short to rather long, often brownish-purple tinted, channelled above, rounded beneath, at first arachnoid-hairy; the leaves appear in spring and remain green through the following winter until the new leaves appear. Flowering stem 10–25 cm, pale yellowish-green, often suffused brownish-purple, slender, hollow, with numerous to dense, short to long, slender, dark glandular hairs and arachnoid-hairy; scale leaves 2– 7, 2–7 × 1–3 cm, commonly with a fully developed lamina like basal and with a broadly winged, sheathing base. Capitula 12–15 mm in diameter, 6–20 in a loose terminal column; peduncles pale green, short, glandular-hairy. Involucral bracts 8–10 × 1.5–2.0 mm, green, often tinted purplish, oblong, obtuse at apex, glandular-hairy. Flowers in ‘male’ plants of 2 kinds, the outer ligulate, 3-lobed at apex and female, the ligules pale lilac, the inner tubular, 5-lobed at apex, pale lilac and vanilla scented, the ‘female’ plants with flowers all ligulate, the ligules 3-lobed at apex, pale lilac, with the exception of several central flowers; stigma of male flowers 1.5–2.5 mm. Receptacle flat, without scales. Achenes 2–3 mm, cylindrical, glabrous; pappus of long, white, slender simple eglandular hairs. Flowers 11–2. Visited by flies and hive-bees. 2n = 60. Introduced. Naturalised on waste and rough ground and waysides. Throughout Great Britain and Ireland, common in southern England, local in the north. Native of Libya, Tunisia and Algeria; naturalised in southern Europe, occasionally in central Europe. ‘Female’ plants not recorded in Great Britain and Ireland. 92. Homogyne Cass. Rhizomatous perennial herbs. Stems erect, not leafy, but with small scales. Leaves cordate, dentate, palmately veined, sparsely hairy beneath. Capitulum solitary, terminal. Involucral bracts in more or less 1 row. Inner flowers tubular, numerous, bisexual, the outer row tubular, numerous, bisexual. Corolla 5-lobed at apex, purple. Receptacle flat, without scales. Achenes narrowly cylindrical; pappus of 1 row of numerous, denticulate hairs.

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Three species in central and southern Europe. Don, G. (1813). Account of the native plants in the County of Forfar, and the animals to be found there in Headrick, J. General view of the agriculture of the County of Angus, or Forfarshire. Edinburgh. Druce, G. C. (1920). The extinct and dubious plants of Britain. Rep. Bot. Soc. Exch. Cl. Brit. Isles 5: 731–799. Ribbons, B. W. (1952). Homogyne alpina in Scotland. Watsonia 2: 237–238. Wigginton, M. J. (Edit.) (1999). British red data books. Vol. 1. Vascular plants. Peterborough.

1. H. alpina (L.) Cass. Purple Colt’s-foot Tussilago alpina L. Perennial herb with slender rhizomes covered with woolly scales. Leaves all basal, 1.5–4.5 × 1.5–5.0 cm, shining yellowish-green on upper surface, paler beneath, and often suffused brownish-purple, subrotund to reniform, broadly rounded at apex, shallowly crenate-dentate, cordate at base, glabrous on upper surface, with short, pale simple eglandular hairs beneath, especially on the veins, and the margins; petiole 1–8 cm, pale green, often suffused brownish-purple, with sparse to dense, short to long, pale simple eglandular hairs. Flowering stem 10–40 cm, pale green, often suffused purplish, striate, woolly-arachnoid-hairy mixed with glands above; scales 5–20 × 2–10 mm, lanceolate or ovate, acute at apex, entire, the lower sometimes with a small lamina and a sheathing base. Capitula 10–20 mm in diameter, solitary. Involucral bracts in more or less 1 row, 8–10 × 1.5–2.0 mm, yellowish-green, flushed rosy-purple especially at the scarious margins and tips, oblong, obtuse at apex, arachnoidhairy at base. Flowers all tubular, purple, the outer obliquely truncate, the inner 5-lobed at apex. Receptacle flat, without scales. Achenes 4–5 mm, cylindrical, slightly ribbed; pappus of several rows of pure white simple hairs. Flowers 5–8. 2n = 120–140. Probably always planted. First recorded as a native plant of the Clova Mountains, Forfarshire in 1813 by George Don. Not seen again until found in the same mountains by A. A. P. Slack in 1951 (cf. B. W. Ribbons). There is not doubt that specimens collected by Don are correctly named, but it has been suggested that he planted some of his finds. It is not known if the Slack locality is the same as that of Don, but is restricted to a very small area. It was recorded from Beinn Mhor, South Uist in 1957, but had been planted there by students as a hoax about 1955. It is native of the mountains of central Europe from the Pyrenees to the north Balkan peninsula, and has a European Boreo-arctic Montane element. Tribe 8. Calendulae Cass. Annual to perennial herbs. Leaves alternate, simple, more or less sessile. Capitula with ligulate outer flowers and tubular inner ones. Flowers yellow to orange, sometimes flore pleno. Involucral bracts in 1–2 rows, herbaceous with scarious margins. Receptacular scales absent. Achenes in each capitula varying in size, degree of curving and wartiness. Pappus absent.

93. Calendula L. Annual to perennial aromatic herbs. Stems procumbent to erect, leafy, sometimes woody at base. Leaves alternate, simple, entire or slightly toothed, more or less sessile. Capitula 1–3 at ends of stems or branches. Involucral bracts in 1–2 rows, with narrow, scarious margins. Inner flowers tubular, 5-lobed at apex and functionally male, the outer ligulate, 3-lobed at apex, female and fertile. Corolla yellow to orange or rarely white. Anthers tailed, apical appendage triangular-ovate. Style undivided with an acute conical tip; surrounded by a ring of hairs. Receptacle flat, without scales. Achenes varying in 1 capitulum in size, degree of curving and wartiness; pappus absent. About 10–25 species, depending on taxonomic opinion; with a wide distribution in the Mediterranean region and western Asia. 1. Capitula 20–70 mm in diameter; flowers at least twice as 1. officinalis long as involucral bracts 1. Capitula 10–20(–35) mm in diameter; flowers less than 2. arvensis twice as long as involucral bracts

1. C. officinalis L. Pot Marigold Annual to perennial herb with a tap-root. Stems (17–)20– 50(–70) cm, pale yellowish-green, erect, diffuse or procumbent, much branched, leafy to apex, slightly angled, striate, with dense, short glandular hairs. Leaves (3–)7–15(–17) × 1.0–4.5(–6.0) cm, alternate, dull medium green on upper surface, slightly paler beneath, oblanceolate, narrowly obovate, oblong or spathulate, shortly acute or obtuse at apex, subentire to repand-dentate or denticulate, narrowed at base, the lower sometimes shortly petiolate, the upper sometimes semiamplexicaul, with dense, very short, yellow glandular hairs and sparsely arachnoid-hairy. Capitula 20–70 mm in diameter, solitary at the end of branches. Involucral bracts in 2 rows, 7–11 × 3.5–4.0 mm, pale to medium greyishgreen with narrow scarious margins; lanceolate or oblonglanceolate, acute at apex, with white and red glandular and simple eglandular hairs. Flowers of 2 kinds, strong smelling, the outer 15–25 mm, ligulate and female, the ligules cream, pale yellow to golden-yellow or orange, with 3 shallow lobes at apex, the inner 3–5 mm, tubular, pale creamyyellow at base, golden yellow at apex. Receptacle slightly convex, pitted and spongy, without scales. Achenes with or without an outer row which are incurved or rarely patent, narrowly beaked and 20–25 mm, alternating with much shorter, cymbiform, rarely 3-winged ones; pappus absent. Flowers 6–8. 2n = 32. Very variable in flower colour and achenes. Introduced. Much grown in gardens and a frequent escape or throw-out on tips and waste ground. In scattered localities in Great Britain and the Channel Islands and sometimes more or less naturalised. Origin unknown, perhaps in cultivation. It is cultivated world wide and is frequently a casual and sometimes naturalised in Europe. 2. C. arvensis L. Field Marigold Annual strong-smelling herb with a slender tap-root. Stem (5–)15–25(–30) cm, pale yellowish-green, erect to diffuse, usually much-branched, with more or less numerous, very

94. Ambrosia short glandular hairs and usually some longer arachnoid hairs. Leaves (1–)3–8(–10) × 0.4–1.4(–2.0) cm, alternate, rather pale yellowish-green, with short glandular, eglandular and branched hairs; basal and lower cauline oblong to oblanceolate, obtuse at apex, entire or remotely denticulate, rounded and semiamplexicaul at base. Capitula 10–20(–35) mm in diameter, solitary at the ends of branches; peduncles up to 5 cm, erect, with dense, short glandular and some longer simple eglandular hairs. Involucral bracts 9–12 × 1.0–1.5 mm, yellowish-green, tipped reddish-purple, with narrow scarious margins, linearlanceolate, subacute at apex, with numerous to dense, short glandular, few longer simple eglandular and some minutely forked hairs. Flowers of 2 kinds, the outer ligulate, female, 3-lobed at apex, and fertile, the ligules yellow or orange, usually less than 18 mm and often exceeding the involucral bracts, but not up to twice as long as them, the inner tubular, yellow, orange, brown or violet-purple, 5-lobed at apex and functionally male. Receptacle flat, without scales. Achenes of 2 kinds, the outer 13–20 mm, incurved and narrowly beaked, the inner 6–10(–25) mm, broadly cymbiform and beaked, or rarely wholly with cymbiform or annular achenes; pappus absent. Flowers 5–8. 2n = 44. Introduced. Naturalised in cultivated ground in Guernsey in the Channel Islands and in the Isles of Scilly; an uncommon casual elsewhere in southern England. Tribe 9. Heliantheae Cass. Tribe Helenieae; Tribe Trageteae Annual to perennial herbs. Leaves often opposite, sometimes alternate. Capitula with all tubular flowers or ligulate outer and tubular inner ones. Tubular flowers yellow to brown, ray flowers yellow to brown, orange, red, purple or white. Involucral bracts in 1–several rows, all herbaceous or the inner scarious. Receptacular scales usually present, sometimes absent. Pappus usually absent or minute, sometimes of barbed bristles. 94. Ambrosia L. Franseria Cav.; Gaertneria Medik.; Xanthidium Delpino Annual or perennial herbs. Stems erect, leafy. Leaves mostly in opposite pairs, but those above alternate, usually pinnately or palmately lobed or dissected. Capitula inconspicuous and unisexual, the plants usually monoecious. Male capitula in terminal racemes or spike-like racemes, hemispherical, drooping, each with 7–12, proximally fused involucral bracts enclosing 5–20 flowers, the stamens with anthers separate or nearly so, being borne on the greenish tubular corolla of each flower. Female capitula usually in the axils of leaves or bracts at the base of the male inflorescence, each with a single flower lacking a pappus, corolla and stamens, but enclosed in the involucre which is united with the ovary below, bears a ring of 4–8 spines or tubercles at or above midway and then tapers into a weakly bifid beak through the top of which the stigmatic arms protrude; the ripe achene is nut-like but with the involucre assuming the role of a sclerocarp. Anthers without tails, appendage ovate and acuminate. Style with linear-oblong branches; stigmatic areas in 2 separate lines. Receptacle flat, usually with chaffy scales.

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About 43 species, chiefly in the New World, but several cosmopolitan weeds. In North America pollen from the species of this genus is one of the commonest causes of hay fever. Payne, W. W. (1964). A re-evaluation of the genus Ambrosia (Compositae). Jour. Arnold Arbor. 45(4): 401–430. Rich, T. C. G. (1994). Ragweeds (Ambrosia L.) in Britain. Grana 33: 38–43. 1. Lower and middle leaves palmately 3- to 5-lobed, the 3. trifida upper ones not lobed, all in opposite pairs. 1. Leaves pinnately divided nearly to middle, the lower and 2. middle ones in opposite pairs, the upper alternate 2. Annual with a tap-root and fibrous side roots; involucre in 1. artemisiifolia fruit with 3–7 spines or sharp tubercles 2. Perennial with a deep rootstock and creeping roots, rarely annual; involucre in fruit unarmed or with blunt tubercles 2. psilostachys

1. A. artemisiifolia L. Ragweed A. elatior L.; A. longistylis Nutt.; A. media Rydb.; Franseria artemisioides Willd.; Gaertneria artemisioides (Willd.) Kuntze; Xanthideum artemisioides (Willd.) Delpino Annual herb with a tap-root and fibrous side-roots. Stem 30–100 cm, pale green, erect, furrowed and bluntly ridged, with more or less dense, short, rough, appressed simple eglandular hairs, branched above, leafy. Leaves mostly in opposite pairs, but the uppermost often alternate, 4–10 × 1–7 cm, dark green on upper surface, greyish-green beneath, ovate to lanceolate in outline, acute at apex, deeply pinnatisect into narrow, acuminate segments which are mostly pinnatifid to coarsely and sharply toothed, the uppermost sometimes entire, more or less glabrous on upper surface, with short, pale simple eglandular hairs beneath; petioles 10–30 mm, narrowly winged, the uppermost leaves sessile. Inflorescences of 2 kinds, the male capitula in spike-like, ebracteate racemes and shortly stalked, the female capitula in axillary clusters immediately below the male. Involucre of male capitula 2–3 mm, shortly or scarcely lobed, hispid or with coarser, longer hairs, the flowers with a greenish, tubular corolla. Involucre of female capitula 3–5 mm at maturity, with a single ring of 3–7 spines or sharp tubercles up to 1 mm near to or above the middle, overtopped by the slender apical beak, the flower without a corolla or pappus, the ripe achene nut-like. Receptacle flat with chaffy scales. Flowers 8–10. Anemophilous. 2n = 36. Introduced. Bird-seed, grain, oil-seed and agricultural seed alien on tips and waste ground. A frequent casual in England, Wales, southern Scotland and the Channel Islands since the 1860s, sometimes semi-naturalised particularly in the south-east. Native of North America; widely naturalised in Europe. 2. A. psilostachya DC. Perennial Ragweed A. coronopifolia Torr. & A. Gray Usually a perennial herb with a deep rootstock and creeping roots, rarely annual. Stem 30–60(–100) cm, pale green, erect, bluntly ridged, with more or less dense, short, ascending, stiff simple eglandular hairs. Leaves mostly in opposite

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pairs, but the uppermost often alternate, 4–8(–10) × 1–5 cm, dark green on upper surface, greyish-green beneath, ovate to lanceolate in outline, acute at apex, pinnatisect, the segments oblong, acute at apex and more or less entire to incise-serrate, scabrid on the upper surface, with short, pale simple eglandular hairs beneath, sometimes weakly sticky-pubescent, all subsessile. Inflorescence of 2 kinds, the male capitula in spike-like, ebracteate racemes and shortly stalked, the female capitula in axillary clusters immediately below the male. Involucre of male capitula 2–3 mm, shallowly and inconspicuously lobed, with sparse to numerous, stiff hairs, the flowers with greenish, tubular corolla. Involucre of female capitula 4–6 mm at maturity, with a single ring of 4, short, blunt tubercles above the middle, or without tubercles, the beak 0.6–1.0 mm, the flower without a corolla or pappus, the ripe achene nut-like. Receptacle flat with chaffy scales. Flowers 8–10. Anemophilous. Probably mostly reproduces vegetatively. 2n = 36, 72, 108. Introduced. Grain alien occurring on rough ground and dunes and in waste places. In scattered localities in Great Britain; naturalised in Lancashire since the 1880s and known for at least ten years in Ayrshire. Native of North America. 3. A. trifida L. Giant Ragweed A. striata Rydb.; A. variabilis Rydb.; A. integrifolia Muhlenb. ex Willd.; A. aptera DC. Annual herb with a tap-root and fibrous side-roots. Stem 100–300(–400) cm, pale green, robust, erect, striate, glabrous below, with more or less numerous, spreading, stiff, pale simple eglandular hairs above, simple or sparingly branched, leafy. Leaves mostly in opposite pairs, 10– 20 × 10–20 cm, dark green on upper surface, much paler beneath, ovate to subrotund in outline some or all of the lower and median deeply, palmately 3- to 5-lobed, the lobes ovate or ovate-lanceolate, acute or acuminate at apex and serrate or rarely lobulate, the upper smaller and only 3cleft or subentire, occasional plants may occur with all the leaves simple and subentire, all scabrid on both surfaces and petiolate. Inflorescence of 2 kinds, the male capitula forming raceme-like, branching panicles, the female capitula in clusters in the axils of bracts subtending the male inflorescence. Involucre of male capitula wide open and 2– 4 mm in diameter, becoming almost rotate, shallowly or obscurely lobed, with 3, conspicuous, dark, vein-like straps extending from the centre to margin, with 10–15 flowers with a greenish, tubular corolla. Involucre of female capitula 5–10 mm, with several ribs, each rib ending in a short spine, the spines forming a circle round the short, stout beak 1–4 mm, the flower without a corolla and pappus, the ripe achene nut-like. Receptacle flat, usually with chaffy scales. Flowers 7–9. Anemophilous. 2n = 24. Introduced. A semi-persistent oil-seed, soya-bean or grain alien, casual on tips, docks and in waste places since about 1900. It was recorded at Par Harbour, Cornwall from at least 1909 to 1926. It sometimes reproduces from seed on the beaches and walls of the River Thames in the Dartford– Greenhythe area of Kent. Native of North America.

95. Iva L. Annual herb. Stems erect, leafy. Leaves mostly opposite, but those above alternate, simple, ovate, petiolate. Capitula small, subsessile, numerous, in axillary and terminal, leafless spikes or panicles. Involucral bracts 5, in 1 row. Flowers all tubular, 5 of the outer female, the 8–20 inner male. Corolla 5-lobed at apex, greenish. Anthers without tails, appendage ovate. Styles with linear-oblong branches; stigmatic areas in 2 separate lines. Receptacle flat, with scales. Achenes cuneate-obovate, somewhat compressed; pappus absent. Fifteen species in North America. Jackson, R. C. (1960). A revision of the Genus Iva L. Univ. Kansas Sci. Bull. 41: 793–875.

1. I. xanthifolia Nutt. Marsh Elder Cyclachaena xanthifolia (Nutt.) Fresen.; Euphrosyne xanthifolia (Nutt.) A. Gray. Annual herb with tap-root and fibrous side-roots. Stems 40– 200 cm, pale green, erect, robust, glabrous or with short to long, glandular hairs in the upper part, branched in the upper part, leafy. Leaves mostly opposite, but those above alternate, 5–30 × 2.5–15 cm, dark green on upper surface, greyish beneath, narrowly to broadly ovate or rhombic, acuminate at apex, coarsely serrate and sometimes 3- to 5-lobed, the teeth unequal and often cusped, subcordate at base, strigose-scabrid on the upper surface, more or less sericeous beneath; 3-veined, long-petiolate. Inflorescence a large paniculate cluster or spike of subsessile capitula not subtended by bracts. Capitula 4–5 mm in diameter, turbinate. Involucral bracts 5, in 1 row, 1.5–3.0 × 0.5–2.5 mm, greenish-white, broadly ovate, shortly acuminate at apex, stiffly hairy. Flowers all tubular, the outer 5 female, up to 0.5 mm with the corolla greatly reduced and often only a mere crown at the base of the style, the inner 8– 20 male, about 2.5 mm with a fusiform, 5-lobed, greenish corolla. Receptacle flat, with chaffy scales, the outer partly enclosing the achenes. Achenes 2.0–2.5 mm, blackish or dark brown, cuneate-obovate, somewhat compressed at right angles to the radius of the capitulum, muricate, glabrous or hispid distally; pappus absent. Flowers 7–9. Anemophilous. 2n = 36. Introduced. A persistent bird-seed, grain, carrot-seed and wool alien, seminaturalised in some places, elsewhere a casual. Scattered records in southern England, mainly south-east England and East Anglia. Native of North America. 96. Xanthium L. Annual, sometimes spiny herbs. Stems decumbent to erect, leafy. Leaves all alternate, slightly to very deeply pinnately or palmately lobed. Capitula solitary or in axillary clusters, unisexual, the male above the female. Male capitula subglobose, involucral bracts in 1 row; receptacle conical with scales; flowers numerous. Female capitula ovoid, involucral bracts in 2 rows, the outer small and free, the inner connate, coriaceous, prickly, ending in 2 (rarely 1) beaks and forming a 2-locular structure containing 2 flowers. Corolla absent. Anthers without tails, appendage ovate; filaments

96. Xanthium connate. Style with linear-oblong branches, protruding from the beaks of the receptacle; stigmatic areas in 2 separate lines. Achenes ovoid; pappus absent. About 3 species which are widespread weeds in the warmer parts of the world. L¨ove, D. & Dansereau, P. (1959). Biosystematic studies on Xanthium: taxonomic appraisal and ecological status. Canad. Jour. Bot. 37: 173–208. Millspaugh, C. F. & Sherff, E. E. (1919). Revision of the North American species of Xanthium. Publ. Field Columbian Mus. Bot. Ser. 4: 9–54. 2. 1. Stems without spines at base of leaves 3. 1. Stems with spines at base of leaves 2. Plant not aromatic; stems and branches green; fruiting involucre 10–15 × 5–10 mm 1(a). strumarium subsp. strumarium 2. Plant aromatic; stems and branches often with violet or brownish dots or stripes; fruiting involucre 15–35 × 6–25 mm, yellow or brown when ripe 1(b). strumarium subsp. italicum 3. Leaves sessile or with petioles under 10 mm, simple or 2. spinosum with 1–2 pairs of lobes 3. Leaves with some petioles more than 20 mm, divided nearly to midrib and larger lobes toothed or with lobules 3. ambrosioides

Section 1. Xanthium Stems without spines at base of petiole. Leaves green beneath, often cordate at base. Fruiting involucre usually with 2 strong beaks. 1. X. strumarium L. Rough Cocklebur Annual herb with fibrous roots, sometimes aromatic. Stems 20–120 cm, pale greyish-green, sometimes with brownish or violet lines or dots, decumbent to erect, variously hairy, sometimes glandular, much branched, leafy. Leaves alternate, 4–15 × 5–16 cm, medium yellowish-green, paler beneath, ovate in outline, obtuse or acute at apex, entire or shallowly palmately 3- to 7-lobed, with rather wide basal sinuses, shallowly to coarsely and irregularly serratedentate, usually more or less cordate at base, rarely cuneate, with short, white bristles and sessile, yellow glands; petiole 1–10 cm. Capitula unisexual, in axillary clusters, the male above the female. Male capitula 5–8 mm in diameter, subglobose to conical; involucral bracts in 1 row, 2–3 mm, linear-lanceolate, obtuse at apex, with simple eglandular and glandular hairs; flowers about 3 mm, glandular in the upper half; receptacle conical, with scales about 3 mm, obovate. Female capitula with a basal involucre of numerous, uniseriate, free, lanceolate, involucral bracts about 3 × 1 mm with simple and glandular hairs and 2 erect, divergent or incurved apical beaks. Fruiting involucre 10–35 × 5–25 mm, green or greyish-green to yellow or brown, oblongellipsoid or ovoid, shortly glandular-hairy, with hooked spines 4–6 mm and glandular-hairy at base. Achenes 9– 15 × 4–5 mm, flattened-hemispherical in transverse section, longitudinally ribbed and sometimes with 2 narrow wings; style persisting as a beak-like appendage about 2.5 mm. Flowers 7–10.

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The following treatment into 2 subspecies follows D. L¨ove & P. M. Dansereau (1959). Very variable populations in Eurasia and North America are believed to have been derived from hybrids between the two subspecies. They have been named X. albinum (Widder) H. Scholz, X. orientale L., X. macrocarpum DC., X. basilicum Vell., X. saccharatum Wallr. and X. riparium Itzigs. & Hertsch. There is obviously a rich gene pool and predominant inbreeding has favoured the formation of local races. The best account of this variation is to be found in C. F. Millspaugh & E. E. Sherff (1919). (a) Subsp. strumarium Plant not aromatic. Stems and branches green. Fruiting involucre 10–15 × 5–10 mm, covered with dense but slender spines. 2n = 36. (b) Subsp. italicum (Moretti) D. L¨ove X. italicum Moretti; X. californicum Greene; X. echinatum Murray; X. cavanillesii Schouw & Didr Plant aromatic. Stems and branches often with violet or brownish dots or stripes. Fruiting involucre 15–35 × 6–25 mm, covered with stout spines, yellow or brown when ripe. 2n = 36. Introduced. Casual on tips and waste ground and sometimes naturalised for a short period, mainly from wool and soya-bean waste. Scattered records in England and Wales and frequent on the estuarine shores of the River Thames from the London border as far as the east coast of the Isle of Grain in Kent. Occurs as a weed throughout the warmer parts of the world. Both subspecies occur. Subsp. strumarium is thought to be native of east, central and south Europe, subsp. italicum of North and South America. Section 2. Acanthoxanthum DC. Stems with 3-fid spines at the base of each leaf. Leaves greyish-white beneath, narrowed at base. Fruiting involucre with 1 or 2 short, straight beaks or beakless. 2. X. spinosum L. Spiny Cocklebur Annual herb with fibrous roots. Stems 20–100 cm, yellowish-green, sometimes tinted brownish-purple, decumbent to erect, much branched, striate, glabrous or with few to dense, short, pale simple eglandular hairs, leafy, with 1–2, 3-branched, yellow spines 10–30 mm at the base of each leaf. Leaves alternate 2–12 × 1.0–3.5 cm, medium to dark green on upper surface, greyish-white beneath, lanceolate to ovate, long-acute at apex, the upper often entire, the lower generally with 1–2 pairs of entire lobes, the terminal lobe usually longer than the lateral ones, cuneate at base, with short, appressed, pale simple eglandular hairs on the upper surface, particularly along the midrib, densely felted beneath; sessile or with petioles up to 10 mm. Capitula unisexual, in axillary clusters, the male above the female. Male capitula 4–5 mm in diameter, subglobose; involucral bracts in 1 row, about 2 × 1 mm, oblanceolate, obtuse at apex, with simple eglandular hairs; flowers glandular in the upper half; receptacle conical, with scales about 2.5 mm and oblanceolate. Female capitula often solitary, at first subglobose, about 2 mm in

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diameter, without basal involucre, with slender, straight apical beaks, one usually developed more than the other. Fruiting involucre 8–13 × 3.5–8.0 mm, oblong-ellipsoid, with hooked spines 4–6 mm. Achenes 6–7 × 0.8–1.0 mm, fuscous, narrowly oblong-cylindrical, slightly compressed. Flowers 7–10. 2n = 36. Introduced. Casual on farms manured by shoddy, tips, railway sidings, sewage-works and waste ground, sometimes more or less naturalised for a while, mainly from wool-waste and bird-seed. Scattered records in England and Wales. Native of South America. 3. X. ambrosioides Hook. & Arn. Argentine Cocklebur X. leucocarpon Wallr.; X. eriocarpon Wallr.; X. catharticum Coste & Sennen Annual herb with fibrous roots. Stems 20–80 cm, pale green often suffused brownish-purple, decumbent to erect, striate, angled, with short, appressed, white simple eglandular hairs, branched, leafy, with 3-branched, orange spines 6–11 mm at the base of each leaf. Leaves alternate, 2–7 × 1–3 cm, dark green on upper surface, greyish-white beneath, ovate in outline, more or less obtuse at apex, divided nearly to midrib, more or less obtuse at apex, divided nearly to midrib, the larger lobes toothed or with lobules, cuneate at base, with dense, short, appressed simple eglandular hairs on both surfaces and felted beneath; petioles up to 30 mm, slender, shortly hairy. Capitula unisexual, in axillary clusters, the male above the female. Male capitula 4–5 mm in diameter, subglobose; involucral bracts in 1 row, oblanceolate, obtuse at apex, with simple eglandular hairs; receptacle conical with oblanceolate scales. Female capitula often solitary, at first subglobose, with a single, straight apical beak. Fruiting involucre 7–8 × 3–5 mm, ellipsoid, more or less sparsely covered with yellow, hooked spines 2–4 mm. Achenes fuscous, narrowly oblong-cylindrical, slightly compressed. Flowers 7–10. Introduced. Casual on tips and waste ground mainly from wool and soya-bean waste, and sometimes more or less naturalised for a short period. A few scattered localities in England. Native of Argentina. 97. Guizotia Cass. Annual herbs. Stems erect, divaricately branched, leafy. Leaves opposite below, alternate above, simple, subentire to serrate, amplexicaul. Capitula numerous, in a loose panicle. Involucral bracts in 2 dissimilar rows, the outer longer and wider, herbaceous, the inner scarious. Inner flowers tubular 5-lobed and bisexual, the outer ligulate, minutely 3-lobed and female. Corolla yellow. Anthers without tails, appendages ovate. Styles with branches linear-oblong; stigmatic areas in 2 separate lines. Receptacle convex, outer flowers subtended by scales resembling the inner involucral bracts. Achenes obpyramidal, compressed, 3- to 4-angled; pappus absent. Named after Fran¸cois Pierre Guillaume Guizot (1787–1874), a French statesman and historian. Six species in tropical Africa. Baagøe, J. (1974). The Genus Guizotia (Compositae). A taxonomic revision. Bot. Tidsskr. 69: 1–39.

1. G. abyssinica (L. fil.) Cass. Niger Polymnia abyssinica L. fil.; Tetragonotheca abyssinica (L. fil.) Ledeb.; Ramtilia oleifera DC.; G. oleifera (DC.) DC.; Jaegeria abyssinica (L. fil.) Spreng.; Polymnia frondosa Bruce ex Murray; Verbesina sativa Roxb. ex Sims.; Heliopsis platyglossa Cass. Annual herb with fibrous roots. Stems 30–50(–200) cm, pale green, usually purple-streaked, erect, striate, glabrous or with few to numerous glandular hairs above, divaricately branched, leafy. Leaves 3–5 × 2–6 cm, opposite below, alternate above, medium green on upper surface, paler beneath, lanceolate to oblong-ovate, acuminate at apex, subentire to serrate, semiamplexicaul, glabrous on upper surface, with short, pale simple eglandular hairs beneath, especially on the veins. Capitula numerous, 20–50(–60) mm in diameter, in a loose panicle; peduncles short to long, with short, pale simple eglandular hairs or glabrous. Involucral bracts in 2 rows, the outer 5, 7–10(–30) × (2.3–)4– 6(–11) mm, green and ovate or broadly elliptical, the inner narrower, and membranous, all hairy. Flowers of 2 kinds, the outer about 8, 11–15 mm, ligulate and female, the ligules yellow with a short tube and 3 deeply cut lobes at apex, the inner tubular, bisexual, yellow and with 5 lobes at apex. Receptacle convex, the outer flowers subtended by broadly elliptical scales resembling the inner involucral bracts. Achenes 3–6 mm, brown to black, obpyramidal, compressed, outer 3-angled, inner more or less 4-angled; pappus absent. Flowers 8–9. 2n = 30. Introduced. Bird-seed casual on tips and waste ground. Scattered records in Great Britain and Channel Islands. Native of East Africa. Grown in warm countries as oil-seed. 98. Sigesbeckia L. Annual herbs. Stems erect, dichotomously branched, leafy. Leaves opposite, simple, irregularly toothed or lobed, with winged petioles. Capitula small, in lax corymbs or rarely solitary. Involucral bracts of 2 kinds, the outer linear to linear-spathulate, patent and with glandular hairs, the inner much shorter, enfolding the outer achenes. Inner flowers tubular, 5-lobed and bisexual, the outer shortly ligulate, 2- to 3-lobed and female. Corolla yellow. Anthers without tails, appendages ovate. Styles with branches linearoblong, stigmatic areas in 2 separate lines. Receptacle flat, the inner flowers subtended by scales. Achenes elongateobovate, 3- to 4-angled; pappus absent. Named after Johann Georg Siegesbeck (1686–1755). S. microcephala DC. has been recorded as a wool casual. Brummit, R. K. (1967). Three species of Sigesbeckia in Britain. Proc. B.S.B.I. 7: 19–21. Henker, H. (1965). Die Gattung Sigesbeckia L. in Europa unter besonderer Ber¨ucksichtund von Deutschland. Arch. Freunde Nat. Gesch. Mecklenb. 11: 7–54.

About 3 species in tropical Africa and Asia and secondarily widespread in other tropical and subtropical areas. 1. Petioles winged distally, the wings tapering proximally and more or less absent at base, not amplexicaul 1. orientalis 1. Petioles winged to base, the wings tapering proximally 2. serrata but widened at base and semiamplexicaul

99. Rudbeckia 1. S. orientalis L. Eastern St Paul’s-wort S. iberica Willd.; S. caspica Fisch. & C. A. Mey. Aromatic annual herb with fibrous roots. Stems 20–120 cm, pale yellowish-green, sometimes tinted purplish, erect, striate, usually with numerous to dense glandular hairs, but sometimes with only sparse hairs and becoming glabrous below, much dichotomously branched, often to the base, leafy. Leaves opposite, yellowish-green on upper surface, paler and slightly bluish beneath and sometimes tinted reddish, lower and medium 3–10(–17) × 1–12 cm, triangular or triangular-ovate, sometimes ovate-elliptical, more or less acute at apex, more or less entire or with irregular teeth which are larger towards the base, or slightly lobed, shortly cuneate, rounded or subcordate at base, the petioles winged distally, the wings tapering proximally and more or less absent at base; upper leaves gradually becoming smaller, narrowly elliptical or lanceolate, more or less acute at apex, less distinctly toothed than lower, shortly cuneate or rounded at base, shortly petiolate or sessile; hairy on both surfaces and margins with short hairs and sometimes glandular hairs. Capitula 6–10 mm in diameter, in small corymbs; peduncles slender, with short, dark glandular hairs. Involucral bracts in 2 rows, the outer 5–10 mm, pale yellowish-green, linear to linear-spathulate, obtuse at apex, patent and with numerous short glandular hairs and sometimes some simple eglandular ones, the inner 2.8–3.2 mm, pale green, obovate, obtuse at apex, with sparse simple eglandular hairs and sometimes few to numerous glandular hairs and enfolding the outer achenes. Flowers of 2 kinds, the outer 3–5, ligulate and female, the ligules yellow, with 2–3 lobes at apex, the inner 4–20, tubular, 5-lobed, bisexual and yellow. Receptacle flat, the scales like the inner involucral bracts and with simple eglandular hairs near the apex only. Achenes 2–3 mm, dark brownish-black, elongateobovate, irregularly 3- to 4-angled, the outer curved; pappus absent. Flowers 7–9. Introduced. An occasional wool alien not usually naturalised. Scattered records in central and southern England and probably over-recorded for S. serrata. Native of tropical Asia. 2. S. serrata DC. Western St Paul’s-wort S. jorullensis auct.; S. cordifolia auct. Aromatic annual herb with fibrous roots. Stems 20–120 cm, pale yellowish-green, often suffused brownish-purple, erect, striate, usually with numerous to dense, short, dark glandular hairs, sometimes nearly glabrous, much branched, often to the base, leafy. Leaves opposite, yellowish-green on upper surface, paler and slightly bluish beneath; lower and medium 3–10(–17) × 1–12 cm, ovate or triangular-ovate, acute or acuminate at apex, more or less entire to shallowly crenate-serrate, shortly cuneate, rounded, truncate or subcordate at base, the petioles winged to base, the wings tapering proximally but widened at base and clasping the stem; upper leaves gradually becoming smaller, lanceolate or linear-lanceolate, acute at apex, entire or minutely toothed, narrowed, but rounded and semiamplexicaul at base; all with short, appressed, pale, stiff, bulbous-based simple eglandular hairs on both surfaces and

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sometimes with small glands. Capitula 5–8 mm in diameter, in small corymbs; peduncles slender, with short, dark glandular hairs. Involucral bracts in 2 rows, the outer 5– 10 mm, pale yellowish-green, linear to linear-spathulate, obtuse at apex, patent and with numerous, short, dark glandular hairs, the inner 2.8–3.2 mm, pale green, obovate, obtuse at apex, with numerous, short, dark glandular hairs and enfolding the outer achenes. Flowers of 2 kinds, the outer 3–5, ligulate, female and with 2–3 lobes at apex, the inner 4–20, tubular, 5-lobed at apex, bisexual and yellow. Receptacle flat, the scales like the inner involucral bracts and with simple eglandular hairs near the apex only. Achenes 2–3 mm, dark brownish-black, elongate-obovate, irregularly 3- to 4-angled, the outer curved; pappus absent. Flowers 7–9. Introduced. Weed of waste and cultivated ground. Naturalised at Freshfield in Lancashire since 1928 and occasionally casual elsewhere. Native of Central and South America. (Verbesina encelioides (Cav.) Benth. & Hook. fil. ex A. Gray (Ximenesia encelioides Cav.) has been recorded as a wool casual and Chrysogonum virginianum L. as a casual garden escape.) 99. Rudbeckia L. Biennial or perennial herbs. Stems erect, leafy, branched. Leaves alternate, simple, entire to deeply lobed. Capitula solitary at the ends of branches on long peduncles. Involucral bracts in 2 or more rows. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, obscurely 3-lobed at apex and sterile. Corolla yellow to orange. Anthers without tails, appendages ovate. Styles with branches linear-oblong; stigmatic areas in 2 separate lines. Receptacle conical, with scales which partly enclose the achenes. Achenes more or less 4-angled; pappus a short corona or absent. About 16 species in North America. Named after Olaus Johannis Rudbeck (1630–1702). R. amplexicaulis Vahl, R. fulgida Aiton and R. trilobata L. occur as casual garden escapes. 1. Stem and leaves with numerous spreading simple eglandular hairs; leaves on upper surface, subentire to 1. hirta remotely denticulate 1. Stem glabrous or with an occasional simple hair, leaves with only rough minute hairs on upper surface; leaves 2. laciniata 1- to 2-pinnatifid

1. R. hirta L. Black-eyed Susan R. serotina Nutt. Biennial or short-lived perennial herb. Stems 30–100 cm, yellowish-green, often suffused brownish-purple, ridged and channelled, angled, erect, with numerous, spreading pale, medium, dark-tubercular-based simple eglandular hairs, leafy, branched in upper part. Leaves alternate, dull medium green on upper surface, paler beneath, with numerous, soft, medium, pale simple eglandular hairs on both surfaces and the margins; basal and lower cauline 10–20 × 4–8 cm, elliptical, lanceolate or oblong-lanceolate, acute or obtuse at apex, subentire or remotely denticulate, long-attenuate at base; veins depressed above and prominent beneath; petioles up to 150 mm, pale green, narrowly

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winged, channelled above and beneath, with numerous, medium to long, pale simple eglandular hairs; middle and upper cauline smaller and broader but similar to lower, sometimes panduriform and more or less sessile and semiamplexicaul. Capitula 40–150 mm in diameter, solitary; peduncles very long, green, angled, with more or less numerous, medium or short, pale, spreading simple eglandular hairs. Involucral bracts in numerous rows, lax, spreading or curved downwards, 9–16 × 2–4 mm, medium green, lanceolate or narrowly elliptical, obtuse at apex, glabrous on the surfaces, with short, pale, spreading simple eglandular hairs round and near the margin. Flowers of 2 kinds, the outer 20–35 mm, ligulate and sterile, the ligules golden yellow, and sometimes brownish-red in the lower half, rounded, retuse or 3-lobed at apex and softly and minutely hairy below and on the margin, the inner 7–8 mm, tubular and bisexual, brownish-black and shallowly 5-lobed at apex. Receptacle conical, with purplish, spathulate, obtuse scales partly enclosing the achenes. Achenes about 4 mm, blackish, more or less 4-angled, glabrous; pappus absent. Flowers 7–10. Visited by bees and butterflies. 2n = 38. Very variable. The so-called Rustic Dwarfs are fairly short with large bicoloured ligules. The cv. Terra Gloriosa is a tall plant with very large capitula with bicoloured ligules. Introduced. Grown in gardens and occasionally persistent in rough and waste ground. Scattered localities in southern Great Britain. Native of North America. 2. R. laciniata L. Coneflower Perennial herb with a branched, creeping rhizome. Stems 50–250 cm, pale green, erect, smooth, angled, glabrous or with an occasional hair, leafy, branched in the upper part. Leaves alternate, dull dark green on upper surface, paler beneath, rough with minute simple eglandular hairs on upper surface, with numerous, short, pale simple eglandular hairs beneath and with short, rigid, upwardly curved hairs on the margin; basal and lower cauline, 14–20 × 15–17 cm, broadly ovate in outline, acute at apex, 1–2 pinnatifid, sometimes almost to middle, the terminal segment ovate, often asymmetrical, often incisely lobed and the lobes often dentate, the lateral segments usually asymmetrical, ovate or oblong, sometimes asymmetrically lobed and the outer serrate-dentate, the inter-segment area winged, the petioles up to 150 mm, with a pale channel above, narrowly winged, angled on the lower surface and rough with minute hairs; cauline gradually decreasing in size upwards, the middle ones similar to basal but with shorter petioles and smaller, the upper ovate, acute at apex, irregularly serrate-dentate and sessile, the uppermost ovate or lanceolate, acute at apex and entire. Capitula (50–)70–140 mm in diameter, solitary; peduncles very long, green, with a few, minute hairs. Involucral bracts 12–16 × 4–5 mm, green, ovate-oblong, acute and reflexed at apex, minutely hairy on the margin and with a few hairs on the surface. Flowers of 2 kinds, the outer 30–60 mm, ligulate and sterile, the ligules yellow to orange and deeply emarginate or 3-lobed at apex, the inner 7–8 mm, tubular, bisexual, yellowish-green and 5-lobed at apex. Receptacle conical or elongate, with green, rigid, oblanceolate, acute scales partly enclosing the achenes. Achenes 4.8–5.2 mm, brownish, more or less 4-angled, glabrous;

pappus a short, toothed corona. Flowers 7–10. Visited by bees and hoverflies. 2n = 36, 38, 54, 72, 76, 102. Introduced. Widely grown in gardens and naturalised on waste and rough ground. In scattered localities mainly in south and central Great Britain. Native of North America. (Ratibida columnifera (Nutt.) Wooton & Standley has been recorded as a wool casual.) 100. Helianthus L. Annual to perennial herbs. Stems erect, leafy. Leaves simple, toothed. Capitula 1–several, pedunculate. Involucral bracts in 2 or more rows, herbaceous. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and sterile, often flore pleno. Corolla yellow. Anthers without tails, appendages ovate; filaments surrounded by a collar. Style with branches linear-oblong; stigmatic areas in 2 separate lines. Receptacle flat or slightly convex, with scales which partly enclose the achenes. Achenes slightly compressed and angled; pappus of 2 awns, soon deciduous, sometimes with some shorter extra awns. About 50 species in North America. In North America species boundaries are indistinct because of hybridisation and intergradation. Widespread cultivation has compounded the situation. Heiser, C. B. (1969). The North American sunflower. Mem. Torrey Bot. Club 22(3): 1–218. Phillips, R. & Rix, M. (1993). Vegetables. London. 2. 1. Plant annual with a simple tap-root 3. 1. Plant perennial, with rhizomes and clump-forming 2. Lower leaves cordate at base; involucral bracts acuminate to long cuspidate at apex; receptacular scales 1. annuus inconspicuously hairy 2. Lower leaves cuneate to truncate at base; involucral bracts acuminate at apex; central receptacular scales with 2. petiolaris a dense white-hairy apex 5. tuberosus 3. Rhizomes with swollen tubers 4. 3. Rhizomes without tubers 4. Involucral bracts lax or more less appressed 4. × laetiflorus 4. Outer involucral bracts spreading or reflexed 3. × multiflorus

1. H. annuus L. Sunflower Annual herb with a tap-root and thick, fibrous side-roots. Stems 75–300 cm, pale yellowish-green, erect, robust, markedly ridged, solid, roughly hairy, densely so above, leafy, rarely branched. Leaves 10–40 × 5–35 cm, dull medium green with paler veins on upper surface, paler beneath, broadly ovate, more or less acuminate at apex, regularly and shallowly mammiform-dentate, mostly cordate at base, the upper sometimes cuneate or truncate, shortly, unequally and roughly hairy on both surfaces and the margins; veins very prominent beneath; petioles long, pale green, densely and roughly hairy. Capitula 100–300 mm in diameter, usually solitary, nodding; peduncle pale green, with dense, unequal, short to long, subrigid, pale simple eglandular hairs. Involucral bracts 20–80 × 15–40 mm, medium yellowish-green, ovate, acuminate to long cuspidate, minutely dentate, with 1–3 prominent veins, densely covered with short to long, pale simple eglandular hairs,

100. Helianthus the bracts often subtended by several upper cauline leaves, in fact there is a gradual sequence from leaves to inner involucral bracts. Flowers of 2 kinds, the outer 25–35 mm, ligulate and sterile, the ligules golden yellow, elliptical and acute or bifid at apex, the inner tubular, bisexual yellow and with a brown, 5-lobed apex. Receptacle flat, with scales which are whitish at base and brown at apex with 3 teeth, the outer 2 short, the middle hooked at apex, slightly hairy, partly enclosing the achenes at maturity. Achenes 5–17 mm, variable in colour but often white with black streaks, compressed, shortly hairy; pappus of 2(–4) deciduous bristles, sometimes also with small scales. Flowers 8–10. Visited by many bees and long-tongued flies. 2n = 34. Introduced. Grown for ornament in gardens, in fields as oil-seed and for Pheasant cover. Common casual on tips and in waste places, also a bird-seed alien. Throughout much of the British Isles. Widely cultivated and grown in gardens in temperate regions. Probably native of Mexico. 2. H. petiolaris Nutt. Lesser Sunflower H. patens Nutt.; H. integrifolius Nutt.; H. aridus Rydb. Annual herb with a tap-root and fibrous side-roots. Stems 50–120 cm, pale green, erect, robust, ridged, glabrous or strigose-hairy, simple or branched, leafy. Leaves alternate, 4–15 × 1–8 cm, medium green becoming bluish-green with age on upper surface, paler beneath, lanceolate to ovate or ovate-oblong, acute at apex, entire to obscurely serrate, cuneate to truncate at base, strigose-hairy on both surfaces; petiole short to medium, hairy. Capitula few, 50– 75 mm in diameter; peduncles long. Involucral bracts in 2 or more rows, 12–15 × 2–4 mm, green, lanceolate to ovatelanceolate, acuminate at apex, with short, stiff simple eglandular hairs on the outer face and shortly ciliate. Flowers of 2 kinds, the outer 15–30, ligulate and sterile, the ligules 15–30 mm, 3-lobed at apex and yellow, the inner tubular, 5-lobed at apex and reddish-purple or rarely yellow. Receptacle flat, with 3-fid scales, the middle cusp exceeding the tubular flowers and those in the centre of the capitulum densely white-hairy at the apex. Achenes 3.5–4.5 mm, pale brown, compressed, sparsely appressed-hairy; pappus of 2 early-deciduous awns. Flowers 8–10. 2n = 34. Introduced. Casual, especially from soya-bean waste, frequent in the London area, infrequent elsewhere. Native of North America. 3. H. × multiflorus L. Thin-leaved Sunflower H. annuus × decapetalus L. H. decapetalus auct. Perennial herb with short rhizomes forming clumps. Stems 50–220 cm, pale yellow-green, sometimes suffused brownish-purple, erect, markedly ridged, glabrous below becoming scabrid with short, pale rigid hairs above, branched, leafy. Leaves 3–10 × 1–8 cm, dull medium green on upper surface, paler beneath, ovate to rhomboid-ovate, acute or acuminate at apex, shallowly serrate or biserrate, cuneate at base, scabrid on both surfaces with very short, rough hairs; petioles short, pale green, roughly hairy. Capitula 30–120 mm in diameter, solitary at the ends of stems or branches; peduncles pale green, with short, pale, rigid hairs. Involucral bracts in several rows, 10–15 × 2.5–3.0 mm,

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olive-green, lanceolate, long drawn out to an acute apex, the outer lax or reflexed, the inner erect, with very short simple eglandular hairs on the surface and longer, white ones on the margins. Flowers of 2 kinds, the outer 30–50 mm, ligulate and sterile, the ligules golden yellow and shortly 3lobed at apex, the inner tubular, 5-lobed at apex and deeper yellow, sometimes all flowers ligulate. Receptacle convex, with scales which partly embrace the achene. Achenes 5–6 mm, somewhat flattened, slightly angled; pappus of 2 early deciduous awns. Flowers 8–10. 2n = 102. Introduced. A rather infrequent escape or throw-out from gardens. Scattered records in Great Britain, mainly the south and centre. Of garden origin. 4. H. × laetiflorus Pers. Perennial Sunflower H. rigidus (Cass.) Desf. × tuberosus H. pauciflorus Nutt. Perennial herb with rhizomes. Stems up to 200 cm, yellowish-green, sometimes suffused brownish-purple, striate, solid, erect, rough with medium to long, pale, rigid, curved simple eglandular hairs, leafy, branched in the upper half. Leaves all cauline, gradually decreasing in size upwards, 4–30 × 1.5–11.0 cm, dull medium green on upper surface, paler beneath, the lower ovate, the upper lanceolate, long-acute or acuminate at apex, shallowly and bluntly serrate, cuneate or gradually narrowed at base, very rough on both surfaces with very short, rigid simple eglandular hairs and longer ones on the prominent veins beneath; petioles very short and winged. Capitula 10–60 mm in diameter, solitary at the end of long, erect branches or sometimes 2 to a branch; peduncles very long, pale green, striate, with numerous, short simple eglandular hairs. Involucral bracts in several rows, lax or loosely appressed, 8–15 × 2–4 mm, the outer yellowish-green, the inner dark or blackish-green, lanceolate to ovate, gradually narrowed to an acute apex, the inner sometimes obtuse, glabrous on the surfaces but markedly ciliate, the hairs pale, rather stiff and patent. Flowers usually of 2 kinds, the outer 30–35 mm, ligulate and sterile, the ligules yellow, acute at apex, the inner 10–15 mm, tubular and bisexual, yellow or brownish-yellow and with 5 short triangular lobes at apex, sometimes all ligulate. Receptacle convex, pitted, with pale, lanceolate scales which partly enclose the achenes. Achenes 5–6 mm, oblong, somewhat flattened, glabrous; pappus of 2 scarious scales which soon fall. Flowers 9–10. 2n = 102. Introduced. The commonest garden perennial sunflower which is a frequent escape or throw-out. Throughout most of Great Britain and Ireland. Probably of garden origin. 5. H. tuberosus L. Jerusalem Artichoke Perennial herb with large irregular tubers developing on the tips of rhizomes. Stems 100–280 cm, pale yellowish-green, often suffused brownish-purple, erect, strongly ridged, solid, with dense medium to long, pale, stiff simple eglandular hairs with brownish-purple tuberous bases, very leafy, sometimes branched at apex. Leaves all cauline, smaller ones often developing in the axils of larger ones, 10–25 × 7– 15 cm, dull medium green on upper surface, paler beneath, ovate, sometimes broadly so, acute or acuminate at apex, subentire to shallowly serrate, cuneate at base, rough with

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small rigid hairs on upper surface, rough with similar hairs beneath but more densely so on the very prominent veins; petioles up to 80 mm, pale green, flat, with narrow, darker wings above, rounded beneath, with dense, stiff, pale simple eglandular hairs thickened towards their bases. Capitula often only 1, sometimes several, solitary, 40–80 mm in diameter, peduncles rather dark green, with numerous hairs similar to those on the stem. Involucral bracts in several lax rows, 10–17 × 3–4 mm, dark green, linear-lanceolate, long-acute or acuminate at apex, with long, pale, stiff simple eglandular hairs on the margins and midrib and very short ones on the surfaces. Flowers of 2 kinds, the outer 30–40 mm, ligulate and sterile, the ligules yellow and acute or retuse or 3-lobed at apex, the inner very short, tubular, 5-lobed at apex, bisexual and yellow. Receptacle convex, with scales which partly enclose the achenes. Achenes 5–6 mm, somewhat flattened, slightly angled, glabrous or hairy; pappus of 1–4 ciliate awns. Flowers 10–11, but plants are often killed by frost before they can develop. Visited by many bees and Diptera. 2n = 102. Introduced. Grown as a minor crop and commonly, at least in East Anglia, as Pheasant food and cover. Escapes and is often very persistent in waste places. Scattered throughout much of Great Britain and the Channel Islands. Native of North America. Jerusalem is said to be a corruption of the French Girasol, a sunflower, but is more likely to be a corruption of Terneuzen, the place in Holland from which it was first introduced into England. Jerusalem Artichokes are grown for their fleshy, tuberous rhizomes, which are similar to knobbly potatoes. 101. Heliopsis Pers. Perennial herbs. Stems erect, simple or branched, leafy. Leaves opposite, coarsely toothed. Capitula solitary. Involucral bracts in 2 or 3 rows. Inner flowers tubular, 4- to 5-lobed at apex and bisexual, the outer row ligulate, 3-lobed at apex and female. Corolla yellow. Anthers without tails, apical appendage ovate; filament collar dilated. Style with linearoblong branches; stigmatic areas in 2 separate lines. Receptacle conical, with numerous scales. Achenes obtusely 3to 4-angled; pappus absent or a small, more or less toothed rim. Fifteen species in North and Central America and northern to central South America. Fischer, T. R. (1957). Taxonomy of the genus Heliopsis (Compositae). Ohio Jour. Sci. 57: 171–191.

1. H. helianthoides (L.) Sweet Rough Oxeye Bupthalmum helianthoides L.; H. laevis (L.) Pers.; Helianthus laevis L. Perennial herb with a stout rootstock. Stem 80–150 cm, pale yellowish-green, erect, rigid, ridged, glabrous or scabrid, simple or branched above, sparingly leafy. Leaves 5–15 × 3–8 cm, yellowish-green on upper surface, paler beneath, opposite, broadly lanceolate to ovate, acute at apex, serrate, shortly cuneate to subtruncate at base, petiolate, glabrous or scabrous. Capitula 50–80 mm in diameter, solitary at the ends of branches; peduncles long, pale yellowishgreen, ridged, glabrous or scabrous. Involucral bracts in 2

or 3 rows, subequal, 10–12 × 4–5 mm, green, ovate, bluntly acuminate at apex, minutely scabrid. Flowers of 2 kinds, the outer 25–40 mm, ligulate, yellow and female, the ligule 3lobed at apex, the inner tubular and bisexual, dull yellow and 5-lobed at apex. Receptacle conical, elongating at maturity, with concave and clasping scales. Achenes of ligulate flowers often 3-angled, those of tubular flowers 4-angled, glabrous; pappus absent or a short, obscure, more or less toothed crown. Flowers 8–10. 2n = 28. Var. helianthoides is glabrous and var. scabra (Dunal) Fern. (H. scabra Dunal) is scabrid. They occupy different geographical areas. It is not known to which the Kent plants belong. Introduced. A garden escape established on waste ground near Swanscombe in Kent. Native of North America. 102. Galinsoga Ruiz & Pavon Wiborgia Roth nom. rejic.; Adventina Raf. Annual herbs. Stems erect or ascending, leafy. Leaves opposite and simple, ovate or lanceolate, crenate-serrate. Capitula small, in dichasial cymes. Involucral bracts in 2 rows, largely herbaceous, the outer much shorter, the inner with a membranous margin. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female. Corolla of tubular flowers yellow, of ligulate flower white or pinkish. Anthers without tails, apical appendage ovate; filament collar dilated. Style with oblong-linear branches; stigmatic areas in 2 separate lines. Receptacle conical, with scales. Achenes obovoid-prismatic, the outer somewhat compressed dorsally; pappus of several scales. Thirteen species in North, Central and South America. Canne, J. M. (1977). A revision of the Genus Galinsoga [Compositae: Heliantheae]. Rhodora 79: 319–389. 1. Peduncles with short eglandular hairs and an occasional short glandular hair; receptacular scales trifid; pappus 1. parviflora scales fimbriate, but not awned 1. Peduncles with numerous, long, slender glandular hairs, a few simple hairs and minute glandular hairs; receptacular scales acute, but rarely shallowly trifid; pappus scales 2. quadriradiata fimbriate and with a fine terminal awn

1. G. parviflora Cav. Gallant Soldier G. quinqueradiata Ruiz & Pavon; Wiborgia parviflora (Cav.) Kunth; Adventina parviflora (Cav.) Raf.; G. hirsuta Baker; Wiborgia acmella Roth nom. illegit. Annual herb with fibrous roots. Stems 10–80 cm, medium green, erect, ascending or flopping, striate, flexuous, glabrous below, with short, pale, appressed, simple eglandular hairs, sparsely leafy, much branched throughout. Leaves opposite, 1–5(–9) × 0.5–2.0 cm, dull medium green on upper surface, slightly paler beneath, ovate or lanceolate, acute at apex, shallowly undulate-crenate-serrate, rounded to cuneate at base; with glabrous upper surface and short pale simple eglandular hairs on the margins and prominent veins beneath; petiole up to 10 mm, pale green, slender, with short, pale simple eglandular hairs. Capitula 4–7 mm in diameter, subglobose, in dichasial cymes at the ends of stems and branches; peduncles up to 25 mm, pale green, with short, pale, appressed simple eglandular hairs and

103. Bidens an occasional short glandular hair. Involucral bracts 3– 4 × 1.5–2.0 mm, medium green, the outer with narrow, scarious margins, broadly ovate, narrowed to an obtuse apex, glabrous. Flowers of 2 kinds, the outer usually 5, about 4 mm, ligulate, the ligules white and 3-lobed at apex, the inner about 3 mm, yellow and 5-lobed at apex. Receptacle shortly conical, with obovate, 3-fid scales finely toothed above, those subtending the ligulate flowers about 0.4 mm, those subtending the tubular flowers at least 1 mm. Achenes 1.0–1.5 mm; those of the tubular flowers narrowly obovoidprismatic, their faces with more or less sparse, short ascending rigid hairs, their pappus of 15–20, silvery, lanceolate to spathulate, deeply fimbriate, obtuse to acute but not awned scales about 1.5 mm; those of the ligulate flowers 3-angled, flattened, more or less curved, with rigid hairs above and their pappus scales about 0.4 mm or absent. Flowers 5–10. 2n = 16. Introduced. A well-naturalised weed in cultivated and waste ground, first recorded in 1860. Locally frequent in the Channel Islands and Great Britain north to central Scotland. Native of South America, but now a cosmopolitan weed. 2. G. quadriradiata Ruiz & Pavon Shaggy Soldier G. ciliata (Raf.) S. F. Blake; Adventina ciliata Raf.; Wiborgia urticaefolia Kunth; Jaegeria urticaefolia (Kunth) Spreng.; G. aristulata E. P. Bicknell.; Sabazia urticaefolia (Kunth) DC.; Baziasa urticaefolia (Kunth) Steud.; G. urticaefolia (Kunth) Benth. Annual herb with fibrous roots. Stem 10–80 cm, medium green, erect, ascending or flopping, striate, sometimes flexuous, with more or less numerous, short to medium, pale, thickened-based simple eglandular hairs and numerous, minute glandular hairs throughout, much branched, leafy. Leaves opposite, 1.0–9.5 × 1.0–5.5 cm, dull medium green on upper surface, slightly pale beneath, lanceolate or ovate, more or less acute at apex, shallowly undulate-crenateserrate, rounded to cuneate at base; glabrous or with a few, pale simple eglandular hairs on both surfaces and the margin, especially on the prominent veins; petioles up to 40 mm, pale green, with few to numerous, pale simple eglandular hairs and those of the upper leaves with numerous, minute glandular hairs. Capitula 5–7 mm in diameter, in dichasial cymes at the ends of stems and branches; peduncles up to 20 mm, with numerous, long, slender glandular hairs, a few simple eglandular hairs and numerous, minute glandular hairs. Involucral bracts 2.5–4.0 × 1.5–2.7 mm, medium green, the inner only with a narrow scarious margin, broadly ovate, narrowed to an obtuse apex, with few to numerous, long, slender glandular hairs. Flowers of 2 kinds, the outer (4–)5, 3.5–4.0 mm, ligulate and female, the ligules white and 3-lobed at apex, the inner 2.5– 3.0 mm, tubular and bisexual, dirty yellow and 5-lobed at apex, hairy. Receptacle shortly conical, with lanceolate, acute scales, rarely shallowly 3-fid at apex. Achenes 1.0– 1.6 mm, brown, obovate-prismatic, with pale ascendingappressed hairs; pappus scales 0.5–1.5 mm, fimbriate and with a fine terminal awn. Flowers 5–10. 2n = 16, 24, 32. Introduced. Naturalised weed in cultivated and waste ground, first recorded in 1909. Frequent in the Channel Islands and Great Britain north to central Scotland. Native

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of Mexico, but widely established as a weed elsewhere in the New World, Europe, Nepal, Japan, the Philippines and Africa. (Hemizonia pungens Torrey & A. Gray has been recorded as a grain casual and Flaveria bidentis (L.) Kunze as a casual at Avonmouth Docks.) 103. Bidens L. Annual herbs. Stems erect, branched, leafy. Leaves opposite, simple and toothed to pinnate. Capitula solitary, pedunculate. Involucral bracts in 2 dissimilar rows, the outer herbaceous, the inner more or less membranous with a usually scarious margin. Flowers all tubular, 5-lobed at apex and bisexual, or the outer row ligulate, 3-lobed at apex and sterile. Corolla yellow or rarely white. Receptacle flat or slightly convex, with scales. Anthers without tails, appendages ovate, often with a conspicuous resin duct; filament collar dilated. Style with oblong-linear branches; stigmatic areas in 2 lines. Achenes obovoid-oblong or linear, compressed or somewhat 4-angled; pappus of 2–5, strong, barbed bristles. About 240 species, cosmopolitan, but mostly native in the New World, though widely introduced as weeds. The American plants of the genus Bidens form a complex of intergrading species plus many local variants and it is difficult to find constant characters to distinguish them. Even our native populations include some plants which are difficult to allocate to the two recognised species, which have been thought to be hybrids. Some of the achene characters such as bristle number and direction of barbs, which have been thought to be important, seem to be unreliable. The presence of ligulate outer flowers in some populations adds to the difficulty. Identification should be based on a survey of a population rather than an individual plant. Harriman N. A. (1998). Proposal to conserve the name Bidens (Asteraceae) with a conserved gender. Taxon 47: 485–486. Kay, G. M. (1998). A tale of two Bidens. B.S.B.I. News 78: 64–65. Sherff, E. E. (1937). The Genus Bidens. Publ. Field Columbian Mus. Botanical Series 16(1): 1–346; 16(2): 347–709.

B. aristosa (Michx) Britton, B. biternata (Lour.) Merr. & Sherff; B. camporum (Hutch.) Mesfin, B. subalternans DC. and B. tenuisecta A. Gray have been recorded as casuals. 1. Leaves not lobed or lobed less than halfway to the midrib 2. 1. Leaves at least in the lower part of the stem pinnate or 5. lobed nearly to the midrib 2. Achenes with the faces between the 4 midribs warty, with 3. connata large, rounded, raised areas 3. 2. Achenes with the faces between the midribs smooth 3. Receptacular scales 8–10 mm, broadly linear and acute at 2. tripartita apex 3. Receptacular scales 6–8 mm, broadly oblanceolate and 4. subacute at apex 1(i). cernua var. cernua 4. All flowers tubular 1(ii). cernua var. radiata 4. Outer flowers ligulate 5. Leaflets lobed again to midrib or more or less so 7. bipinnata 6. 5. Leaflets unlobed

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7. 6. Achenes more or less linear 8. 6. Achenes narrowly obovoid-oblong 7. All flowers tubular and yellowish-orange 6(i). pilosa var. pilosa 7. Outer flowers ligulate, and white, inner tubular and 6(ii). pilosa var. radiata yellowish-orange 5. vulgata 8. Outer involucral bracts 10–16 9. 8. Outer involucral bracts 5–10 9. Lower leaves 3(–5)-partite; achenes downwardly barbed 2. tripartita on the angles 9. Lower leaves pinnate, the central segment petiolulate; 4. frondosa achenes upwardly barbed on the angles

1. B. cernua L. Nodding Bur-marigold Annual herb with a tap-root and long, fleshy side-roots. Stems 8–80 cm, erect, pale yellowish-green, sometimes flushed brownish-purple, striate, glabrous or with sparse simple eglandular hairs, usually branched, sometimes to the base, sometimes tall and little branched, rarely very small and slender and unbranched, leafy. Leaves opposite, sometimes large all the way up, sometimes the lower smaller, sometimes a mixture of small and large, 2–17 × 0.5–4.0 cm, yellowish-green on upper surface, paler and slightly greyish or bluish beneath, linear-lanceolate, lanceolate, ovate, narrowly elliptical or narrowly oblong-elliptical, mostly long drawn out to an acute or acuminate apex, mostly serrate or crenate-serrate, the smaller ones often more or less entire, narrowed at base and often slightly connate, glabrous or with a few simple eglandular hairs, scabridulous on the margin, sometimes with a short, winged petiole. Capitula 15–25 mm in diameter, usually solitary at the ends of stems or branches, drooping; peduncles long, pale green, striate, glabrous. Involucral bracts in 2 rows, the outer lax, 11– 26 × 2–7 mm, green, lanceolate, drawn out to an acute apex and with tiny, hooked teeth along the margin, the inner 6– 11 × 3–8 mm, green with dense, black, longitudinal lines in the middle and broad, scarious margins, broadly obovate, rounded at apex and with an apiculate tip, glabrous. Flowers usually all tubular, bisexual and yellow, sometimes of 2 kinds, the outer 10–12 mm, ligulate and sterile, the ligules yellow, 3-lobed at apex and spreading, the inner tubular, 5-lobed at apex, bisexual and yellow. Receptacle flat or slightly convex, the scales 6–8 mm, scarious, broadly oblanceolate and subacute at apex. Achenes 5–6 mm, narrowly obovoid, compressed, but made strongly 4-angled by the stout midribs, the outer with 3 and the inner with 4 barbed angles and smooth flat sides, the angles continuing upwards as 4 (rarely 3), downwardly barbed awns. Flowers 7–9. Visited sparingly by hive-bees and flies. 2n = 24. (i) Var. cernua B. cernua forma discoidea DC. nom. illegit.; B. minima Hill.; B. cernua forma minima (Hill.) F. N. Williams; B. cernua forma rugosa Druce Flowers all tubular, yellow. (ii) Var. radiata DC. Coreopsis bidens L. Flowers of 2 kinds, the outer 10–12 mm, ligulate, the ligules yellow, the inner tubular, yellow.

Native. Ponds and streamsides, especially places with standing water in winter but not during the growing season. Locally common in central and southern England, Wales and Ireland, rare and only in scattered localities further north. Europe northwards to 60◦ 40 N; northern Asia; North America. A member of the Circumpolar Temperate element. The common plant is var. cernua. Var. radiata is frequent in north-west England and rare elsewhere. It is widespread in Continental Europe. The plant called forma minima grows with var. cernua with intermediates and would appear to be a growth form. 2. B. tripartita L. Trifid Bur-marigold B. peacockii Druce Annual herb with fibrous roots. Stems 15–75 cm, pale yellowish-green, often suffused brownish-purple; striate, slender to robust, glabrous or with short hairs below, simple or branched, sometimes to base, leafy. Leaves opposite, gradually decreasing in size upwards or sometimes the upper larger, 5–15 × 0.5–8.0 cm, deep green on upper surface, paler beneath, lanceolate to broadly ovate in outline, acute at apex, usually 3(–5)-partite but sometimes undivided, some leaflets lanceolate to ovate-lanceolate, acute at apex and coarsely serrate, the terminal leaflet usually larger and sometimes 3-lobed, glabrous or minutely ciliate; petioles short, winged and semiamplexicaul. Capitula 15– 25 mm in diameter, solitary at the ends of stems and branches, suberect; peduncles pale green, striate, glabrous. Involucral bracts in 2 dissimilar rows, the outer 5–10, 14–20 × 3–5 mm, leaf-like, linear or linear-oblanceolate, subacute at apex and spreading, the inner 8–10 × 2.5–3.0 mm, with a dense longitudinal black line in the centre and a wide scarious margin, oblong-ovate and subobtuse at apex, glabrous. Flowers all tubular, 5-lobed at apex and yellow. Receptacle flat or slightly convex, the scales 8– 10 mm, broadly linear and acute at apex. Achenes 7–8 mm, narrowly obovoid-oblong, much compressed, downwardly barbed on the angles, 2 of which continue upwards as downwardly barbed awns forming the pappus, with or without 1–2 awns on the other 2 angles, the faces glabrous and smooth. Flowers 7–9. Visited by bees and hoverflies. 2n = 48. Native. Ditches, streams and pond and lake margins. Locally common in central and southern England, Wales and Ireland, rare and only in scattered localities further north. Throughout Europe to 63◦ 45 N, and in western Asia; introduced in Australia and North America. A member of the Eurasian Temperate element. Plants included here with only two awns on the fruits and undivided leaves strongly approach B. radiata Thuill., but they do not seem to have more than eight outer involucral bracts. They badly need study in the field. 3. B. connata Muhlenb. ex Willd. London Bur-marigold Annual herb with tap-root and side-roots. Stems 40–150 cm, pale yellowish-green, sometimes suffused brownishpurple, erect, glabrous, much branched, leafy. Leaves opposite, 5–15 × 2–8 cm, medium green on upper surface, paler beneath, lanceolate to elliptical, acute at apex, mostly

103. Bidens simple or with 2–4 divergent lobes not reaching more than halfway to midrib, coarsely serrate or dentate to inciseserrate, narrowed at base into a short, winged petiole or sessile, glabrous or slightly hairy. Capitula 10–15 mm in diameter, 1–few at ends of stems or branches; peduncles 1–6 cm, more or less erect, striate, glabrous. Involucral bracts in 2 rows, the outer 10–20(–40) mm, somewhat leafy, but narrow and linear and spreading or loosely ascending, the inner 6–10 mm, green with black lines and scarious margins, elliptical, ovate or ovate-lanceolate and obtuse at apex, glabrous. Flowers all tubular, bisexual, and orange or reddish-yellow, or of 2 kinds, the outer ligulate and sterile, the ligules small, 3-lobed at apex, and yellow and the inner tubular, 5-lobed at apex and bisexual. Receptacle flat or slightly convex, with broadly linear, acute scales. Achenes 3.5–8.0 mm, black or purplish, obovoid-oblong, compressed, made strongly 4-angled by the stout midribs, the marginal with 3, the central with 3–4 awns which are upwardly barbed, the faces verrucose with large, rounded, raised areas, rarely with downwardly directed barbs. Flowers 7–9. 2n = 72. Introduced. Seems to prefer cracks in stonework. Well established by the Grand Union Canal in Essex, Middlesex, Hertfordshire and Buckinghamshire, the Trent and Mersey Canal in Cheshire and by the River Thames near Cross Ness in Kent. Native of eastern North America. 4. B. frondosa L. Beggar’s-ticks B. melanocarpa Wieg. Annual herb with a tap-root and side-roots. Stems 50–120 cm, pale yellowish-green, sometimes flushed brownishpurple, slender, erect, glabrous or with scattered simple eglandular hairs, branched, leafy. Leaves opposite, 5–15(–20) × 2–10 cm, medium green or glaucous on upper surface, paler beneath, ovate in outline, acute at apex, pinnate, with 3–5 lanceolate or lanceolate-ovate, acute or acuminate, neatly serrate segments, the middle segment petiolulate, the others sessile, glabrous or with a few minute hairs on the surfaces, shortly ciliate; petioles 1–6 cm, green, glabrous. Capitula 8–20 mm in diameter, solitary or few at the ends of stems and branches; peduncles slender, with short, whitish hairs. Involucral bracts in 2 rows, the outer 5–10 often very long, 30–50 mm, leafy, linear-spathulate and loosely ascending, the inner 5–7 mm, brownish with scarious margins, lanceolate to ovate and obtuse at apex, glabrous. Flowers all tubular and bisexual, orange-yellow and usually 5-lobed at apex, or of 2 kinds, the outer rudimentary, ligulate and sterile, the ligules 2–3 mm, 3-lobed at apex and golden yellow, the inner tubular, 5-lobed at apex and bisexual, orange-yellow. Receptacle flat or slightly convex, with linear-lanceolate, subobtuse scales. Achenes 6–10 mm, dark brown or black, narrowly obovoid-oblong, tapered from apex to base, slightly compressed, with a midrib and papillae on each face, glabrous or upwardly barbed on the angles, 2 of which continue upwards as downwardly barbed awns forming the pappus. Flowers 7–9. 2n = 48. Introduced. Naturalised by canals and rivers, on damp ground and in waste places. Scattered records in England and Wales, frequent around Birmingham and London and

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in Cheshire and south Wales. Native of North and South America. 5. B. vulgata Greene Eastern Beggar’s-ticks Annual herb with a tap-root and side-roots. Stems 30–150 cm, pale yellowish-green, often purplish or reddish, erect, angled, striate, glabrous to hairy, branched, leafy. Leaves opposite, 5–15 × 2–8 cm, acute at apex, pinnatifid or pinnate, the 3–5 segments lanceolate, acute at apex, serrate, cuneate at base, sessile or petiolulate, glabrous or rarely shortly hairy; petiole 5–15 cm, glabrous or shortly hairy. Capitula 15–28 mm in diameter, at the ends of stems and branches; peduncles 4–23 mm, slender, glabrous or hairy. Involucral bracts in 2 rows, the outer 10–16, 10–20(–40) mm, leaf-like and loosely ascending, the inner 8–18, 7–9 mm, olive-green to reddish-brown, ovate-lanceolate, subacute at apex. Flowers all tubular and bisexual, pale yellow and 5-lobed at apex, or of 2 kinds, the outer ligulate and sterile, the ligules up to 3.5 mm, 3-lobed at apex and yellow, the inner tubular, bisexual and pale yellow. Receptacle flat or slightly convex, with linear, obtuse scales. Achenes 6–12 mm, olive to brownish, narrowly obovoid-oblong, compressed, with slender midribs, with 2 downwardly barbed awns forming the pappus. Flowers 7–9. 2n = 24. Introduced. A rather rare wool casual. Recorded from a few localities, but may have been overlooked for B. frondosa. Native of North America from Nova Scotia to Quebec and Alberta southwards to North Carolina, Missouri, Nevada and California. 6. B. pilosa L. Black-jack B. californica DC. Annual to short-lived perennial herb. Stem 10–100 cm, pale green, often suffused brownish-purple, erect, ridged, glabrous or with a few simple eglandular hairs, muchbranched, leafy. Leaves opposite, medium yellowish-green on upper surface, paler beneath, sometimes tinted purplish, ovate in outline, 3- to 7-foliolate or sometimes simple and not lobed, or 3-lobed; leaflets (1–)2–10(–25) × 2–5(–10) cm, the terminal largest, lanceolate to ovate, acute to acuminate at apex, evenly serrate, cuneate or rounded at base, sessile or shortly petiolulate; glabrous to hairy on both surfaces and the margins, especially beneath, petiolate. Capitula 7–20 mm in diameter, in irregular cymes; peduncles pale green, short to long, channelled, glabrous. Involucral bracts in 2 dissimilar rows, the outer 3–6 mm, green, linear to narrowly oblong-obovate and acute at apex, the inner 6– 10 mm, with dark-lined centre and pale margins, lanceolate to oblong-lanceolate, obtuse to subacute at apex. Flowers all tubular, 5-lobed at apex, bisexual and yellowish-orange, or of 2 kinds, the outer ligulate, 3-lobed at apex and sterile and the ligules white, the inner tubular, bisexual and yellowish-orange. Receptacle flat or slightly convex, with scales similar to inner involucral bracts but narrower. Achenes 5–9 mm, dark brown, more or less linear, flattened or 3(–4)-angled, ribbed, usually glabrous except for the ribs ciliate near the apex, sometimes with scattered hairs elsewhere; pappus of mostly 3 awns, sometimes 2, rarely 4, sometimes unequal, (0.5–)1.5–2.5 mm. Flowers 6–9. 2n = 24, 36, 48, 72.

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(i) Var. pilosa Flowers all tubular, yellowish-orange.

About 114 species in North, Central and South America and Africa.

(ii) Var. radiata DC. Outer flowers ligulate, the ligules white, inner flowers tubular and yellowish-orange.

Sherff, E. E. (1936). Revision of the genus Coreopsis. Publ. Field Columbian Mus. Botanical Series 11: 279–475.

Introduced. Wool, cotton and bird-seed alien. Scattered records in England, both varieties occur. Native of South America and has become a cosmopolitan weed. 7. B. bipinnata L. Spanish Needles B. fervida Hort. ex Colla; B. myrrhidifolia Tausch; B. cicutaefolia Tausch; B. elongata Tausch; B. tenuifolia Tausch; B. decomposita Wall. ex DC.; Coreopsis corymbifolia Wall. ex DC.; B. kotschyi Sch. Bip.; B. kotschyana Sch. Bip.; Kerneria bipinnata (L.) Gren. & Godr.; B. pilosa var. bipinnata (L.) Hook. fil. Annual herb with a tap-root and side-roots. Stems 30– 120(–170) cm, erect, pale yellowish-green, striate, glabrous or with minute simple eglandular hairs, branched, leafy. Leaves 4–12 × 2–10 cm, opposite, medium green on upper surface, paler beneath, broadly ovate in outline, acute at apex, 2- to 3-pinnate, the segments lanceolate or rhomboid in outline and subacute at apex, the lobes linear to lanceolate and sometimes dentate, glabrous on surfaces, minutely ciliate; petiole 2–5 cm, striate, glabrous. Capitula 4–6 mm in diameter, at the ends of stems and branches; peduncles 1–10 cm, striate, glabrous. Involucral bracts in 2 rows, the outer 3–5 mm, green with a dark base, linear and acute at apex, the inner with dark lines in centre, linear-lanceolate and obtuse at apex, shortly hairy at base. Flowers of 2 kinds, the outer ligulate and sterile, the ligules yellowish-white and 2- to 3-lobed at apex, the inner tubular, 5-lobed at apex, bisexual and yellow. Receptacle flat or slightly convex, with narrowly linear-lanceolate scales. Achenes 2–4 mm, blackish, linear, 4-angled, with 3–4 awns which are downwardly barbed. Flowers 7–9. 2n = 72. Introduced. A characteristic wool alien and bird-seed casual, and formerly a cotton alien. Scattered records in England and Scotland. Probably native in recent times only in the eastern United States and eastern Asia; introduced elsewhere in North America, South America, Europe and Australia. 104. Coreopsis L. Annual to perennial herbs. Stems erect, branched and leafy. Leaves mostly pinnately divided to the midrib or more or less so, the primary divisions often divided again. Capitula on long, slender peduncles. Involucral bracts in 2 dissimilar rows, the outer narrower, the inner very broad. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female or sterile. Corolla of tubular flowers yellow or dark red, of ligulate flowers yellow, sometimes with a dark basal blotch. Anthers without tails, appendages ovate, often with a conspicuous resin duct. Style with branches linear-oblong; stigmatic areas in 2 separate lines. Receptacle flat or slightly convex, with scales. Achenes flattened, sometimes with wings; pappus absent or of very few teeth or bristles or an obscure crown.

1. Outer involucral bracts 7–10 mm; ligulate and tubular 1. grandiflora flowers yellow, 5-lobed 1. Outer involucral bracts 2–3 mm; ligulate flowers yellow with a prominent reddish-brown basal blotch, tubular 2. tinctoria flowers dark red, 4-lobed

1. C. grandiflora Hogg ex Sweet Large-flowered Tickseed C. boykiniana Nutt.; C. heterophylla Nutt. Tufted perennial herb with a short stock. Stems 30–100 cm, usually clustered, pale yellowish-green, erect or ascending, fairly robust, ridged, glabrous or with short simple eglandular hairs, branched above, leafy. Leaves evenly scattered up the stem, 5–15 × 0.3–6.0 cm, yellowish-green on upper surface, paler beneath, broadly ovate in outline, mostly pinnatifid, the segments linear or linear-lanceolate and obtuse to acute at apex, narrowed below to a semiamplexicaul petiole, glabrous or nearly so, the lowermost leaves sometimes entire. Capitula few or solitary 30–60 mm in diameter; peduncles up to 15 cm, pale yellowish-green, ridged, naked, glabrous. Involucral bracts in 2 dissimilar rows, the outer 7–10 × 2.5–3.0 mm, olive-green with pale margins, ovatelanceolate and obtuse at apex, the inner about as long but much broader, olive-green, dark lined and with a pale margin, broadly ovate and obtuse at apex. Flowers of 2 kinds, the outer 13–25 mm, ligulate and sterile, the ligules golden yellow and 3-lobed at apex, the inner tubular, bisexual, yellow and 5-lobed at apex. Receptacle flat, the scales 6–7 mm, narrowly oblong. Achenes 2.0–2.5 mm, flattened, with thin, flat entire wings; pappus absent or of a few short teeth or bristles. Flowers 7–9. 2n = 26, with 0–4 B chromosomes. Introduced. Grown in gardens and occurring in waste places as a throw-out or escape. Infrequently naturalised in south-east England. Native of the United States. 2. C. tinctoria Nutt. Garden Tickseed Calliopsis bicolor Rchb.; Diplosastera tinctoria (Nutt.) Tausch; Calliopsis tinctoria (Nutt.) DC.; C. elegans hort. Annual herb with fibrous roots. Stems 30–100 cm, usually arising singly, pale yellowish-green, erect, slender to robust, ridged, glabrous, branched above, leafy. Leaves becoming gradually smaller upwards, 3–10 × 3–6 cm, yellowishgreen on upper surface, paler beneath, broadly ovate in outline, pinnatifid, the segments linear or linear-lanceolate and obtuse to acute at apex, narrowed below to a short petiole, glabrous. Capitula numerous, 25–40 mm in diameter; peduncles pale yellowish-green, ridged, sometimes with narrow linear bracts, glabrous. Involucral bracts in 2 dissimilar rows, the outer 2–3 × 1.5–2.0 mm, green to brown with a membranous margin, ovate or triangular and obtuse at apex, the inner much larger, with a dark-lined centre and membranous margins, broadly ovate and obtuse at apex. Flowers of 2 kinds, the outer 7–15 mm, ligulate and sterile, the ligule yellow with a prominent reddishbrown basal blotch, the inner tubular and bisexual, dark red

107. Tagetes and 4- to 5-lobed at apex. Receptacle flat, the scales 4.0– 4.5 mm, linear-lanceolate. Achenes 3–4 mm, linear-oblong, flattened, often wingless, but sometimes with narrow to prominently broad, thin wings; pappus of minute awns or reduced to an obscure crown. Flowers 8–10. 2n = 24. Introduced. Garden escape and wool casual. A few records in southern England. Native of North America. 105. Cosmos Cav. Annual herbs. Stems erect, leafy. Leaves opposite 2- to 3pinnate with linear to filiform segments. Capitula on long peduncles in loose cymes. Involucral bracts in 2 dissimilar rows, the outer narrower and herbaceous, with darker lines, the inner membranous with a lined centre. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and sterile. Corolla of tubular flowers yellow, those of ligulate flowers pinkish to purple or rarely white. Anthers without tails, appendage ovate; filaments hairy. Style with branches linear-oblong; stigmatic areas in 2 separate lines. Receptacle flat, with scales. Achenes fusiform with an attenuate beak; pappus of (0–)2(–3) bristles which have backwardly directed barbs. Twenty-six species in North, Central and South America.

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or less herbaceous. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and female or sterile, but most often flore pleno. Corolla of tubular flowers yellow, of ligulate flowers yellow, white, pink or purple. Receptacle flat or slightly convex, with scales. Achenes linear-oblong or wider at apex; pappus absent, or of 2 obscure awn-like teeth. Twenty-eight species in Mexico and Central and northern South America. Sorensen, P. D. (1969). Revision of the genus Dahlia (Compositae, Heliantheae – Coreopsidinae). Rhodora 71: 309–416.

1. C. bipinnatus Cav. Mexican Aster Annual herb with fibrous roots. Stem (15–)30–200 cm, pale yellowish-green, often suffused brownish-purple, erect, markedly striate, rigid, with sparse to dense, short simple eglandular hairs, sometimes becoming glabrous in part, branched, leafy. Leaves (2–)3–12 × 3–7 cm, opposite, greyish-green, triangular-ovate in outline, (1–)2- to 3-pinnate, the segments usually linear or filiform, rarely narrowly lanceolate, glabrous or with a few, short simple eglandular hairs, shortly petiolate or more or less sessile. Capitula (25–)45–90 mm in diameter, in loose cymes; peduncles long, pale green, sometimes suffused purple, glabrous or with numerous, short simple eglandular hairs. Involucral bracts in 2 rows, the outer 7–10 × 3.5–4.5 mm, green with darker lines, ovate-oblong, acuminate-caudate at apex and glabrous, the inner 12–15 × 5–6 mm, membranous, pale green to brown with dark lined centre and pale margins, triangular-ovate, obtuse to acute at apex and glabrous. Flowers of 2 kinds, the outer ligulate, 3-lobed at apex and sterile, the ligules pale to deep mauve-pink, pink or white, the inner tubular, 5-lobed at apex, bisexual and yellow. Receptacle flat, with acuminate, membranous scales. Achenes 5–12 mm, brown, all similar, fusiform, (3–)4-angled, with an attenuate beak, glabrous except for short, antrorse hairs towards the apex; pappus absent or with (0–)2(–3) bristles with backwardly directed barbs. Flowers 7–9. 2n = 24. Introduced. Much grown in gardens and a frequent escape on tips and in waste places. Scattered localities in Great Britain, but mostly the centre and south. Native of Mexico and the southern United States.

1. D. × hortensis Guillaumot Garden Dahlia Perennial herb with tuberous roots, but killed by the first frosts. Stems 50–200 cm, pale green, erect, smooth, fleshy, glabrous or sparsely hairy, branched above, leafy. Leaves 10–30 cm, medium yellowish-green on upper surface, paler beneath; lower pinnate or ternate, the leaflets ovate to elliptical, acute to acuminate at apex, simple or 2- to 3-lobed, coarsely serrate and sessile; the upper simple or ternate; all glabrous or sparsely clothed with short simple eglandular hairs. Capitula 5–30 cm in diameter, solitary; peduncles long, pale green, glabrous. Involucral bracts in 2 dissimilar rows, the outer 15–25 mm, pale green, ellipticobovate to spathulate, acute at apex, recurved at anthesis and glabrous, the inner 15–25 mm, usually membranous at margin, triangular-ovate and acute at apex. Flowers of 2 kinds, the outer ligulate and sterile the ligules 3-lobed at apex, yellow, orange, pink to deep red and pale to deep purple, the inner tubular, 5-lobed at apex and yellow, orange or deep purple, often partially or completely replaced by ligulate flowers. Receptacle flat or nearly so, with ovatetriangular, obtuse to acute scales. Achenes 12–15 mm, linear-oblong or slightly broader towards the apex; pappus absent or of 2 reduced awns less than 0.5 mm. Flowers 7–10. 2n = 32, 64. The plants included here under D. × hortensis probably all originate as hybrids between D. pinnata Cav. (D. variabilis (Willd.) Desf.) and D. coccinia Cav., although some may belong to these species. The genus has a history in cultivation long pre-dating the Spanish conquest of South and Central America in the sixteenth century, and was probably an important crop for the Aztecs. Dahlias were first introduced into Europe to the Botanic Garden at Madrid in 1789, and from there distributed throughout Europe, and reached Kew in 1798. Nurserymen took full advantage of the natural proclivity for hybridisation, deriving new varieties by crossing and the chance seedlings which gave rise to the first doubles which were called glebe dahlias. Introduced. Much grown for ornament and a frequent throw-out on tips and waste ground. Scattered localities mainly in south-east England. The parents come from Mexico.

106. Dahlia Cav. Perennial herbs, but killed by the first frosts. Stems erect, rather fleshy, leafy. Leaves pinnate or ternate, the upper simple. Capitula solitary on long peduncles. Involucral bracts in 2 dissimilar rows, the inner membranous, the outer more

107. Tagetes L. Foetid annual herbs. Stems erect, leafy. Leaves alternate above, pinnate, glandular punctate. Capitula solitary at the ends of branches or numerous, in dense, terminal corymbs. Involucral bracts in 1 row, fused for most of their length

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to form a sheath round the capitulum. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, 3lobed at apex and female, or often flore pleno. Corolla yellow to orange, green or brownish. Anthers without tails, appendage ovate; filament collar dilated. Style with oblonglinear branches; stigmatic areas in 2 separate lines. Receptacle flat, without scales. Achenes linear, 3- to 4-angled, slightly compressed; pappus of unequal, often connate scales. About 50 species in North, Central and South America and widely introduced as weeds elsewhere. Named after Tages, an Etruscan deity, grandson of Jupiter. Species of this genus produce insecticidal and/or allelopathic herbicidal chemicals at the root. T. erythrocephala Rusby and T. gracilis DC. have been recorded as wool casuals while T. tenuifolia Cav. has occurred as a casual garden escape. 1. Capitulum 30–70(–120) mm in diameter; involucral 1. erecta bracts 4–8 mm wide 1. Capitulum, 3–45 mm in diameter; involucral bracts less 2. than 4 mm wide 2. Involucral bracts 8–12 mm; capitulum 3–7 mm in 2. minuta diameter; pappus scales 2.5–3.0 mm 2. Involucral bracts 12–18(–20) mm; capitulum 20–45 mm 3. patula in diameter; pappus scales 7–12 mm

1. T. erecta L. African Marigold Aromatic annual herb with a tap-root and fibrous sideroots. Stem 20–50(–100) cm, pale yellowish-green, robust, angled, striate, erect, glabrous or nearly so, branched, leafy. Leaves 1–12 × 0.5–7.0 cm, medium greyish-green on upper surface, paler beneath, pinnately divided, the segments 11– 17, 1–5 cm, linear, oblong or linear-lanceolate, obtuse to acute at apex, sharply dentate and with the teeth often bristle-tipped, hairless, but with scattered sessile glands, sessile. Capitula 30–70(–120) mm in diameter, solitary at the ends of stems and branches; peduncles long, swollen below the capitula, striate, glabrous. Involucral bracts 5–8 in 1 fused row, 10–20 × 4–8 mm, lanceolate or ovatelanceolate, acute at apex, glabrous. Flowers of 2 kinds, the outer ligulate, the ligules lemon yellow to orange or rarely white or cream, the inner tubular and lemon yellow to orange, often most or all flowers ligulate. Receptacle flat, without scales. Achenes 7–8 mm, linear, grooved; pappus of unequal scales. Flowers 5–10. 2n = 24. Introduced. Much grown in gardens and a frequent escape in waste places or on tips. Scattered records in England. Native of Mexico. 2. T. minuta L. Southern Marigold Foetid annual herb with a tap-root and fibrous side-roots. Stem 80–200 cm, pale green, sometimes suffused purplish, erect, ridged and angled, glabrous, branched, leafy. Leaves 2–15 × 1.5–8.0 cm, imparipinnate, medium green, paler beneath, broadly oblong in outline, the leaflets 3–8 pairs, linear to lanceolate, acute at apex, sharply serrate, shortly petiolate and glabrous, with large, brown glands especially on the lower surface. Capitula 3–7 mm in diameter, in dense corymbs; peduncles pale green and glabrous. Involu-

cral bracts in a single row, 8–12 mm, fused for most of their length, pale green, oblong, forming an acute, triangular tooth at apex, glabrous and fleshy. Flowers of 2 kinds, the outer 1–3 mm, ligulate, the ligules yellowish-green or cream and irregularly lobed at apex, the inner few, tubular, green, irregularly lobed at apex. Receptacle flat, pitted, without scales. Achenes 5.5–7.0 mm, dark brown, linear, 3to 4-angled, densely clothed in short, antrorse hairs; pappus 2.5–3.0 mm, of linear, unequal connate scales. Flowers 7–9. 2n = 48. Introduced. Wool, grain, cotton and bird-seed casual. A few, scattered records in Great Britain. Native of South America. 3. T. patula L. French Marigold Foetid annual herb with a tap-root and fibrous side-roots. Stems 10–40(–50) cm, pale green, sometimes suffused purplish, erect, markedly ridged and angled, glabrous, branched from the base, the branches widely angled and often as wide as the plant is high, sparingly leafy. Leaves 2–10 × 1.5–4.0 cm, the lower quickly dying, medium green, paler beneath, broadly oblong in outline, acute at apex, imparipinnate, the leaflets 2–5(–7) pairs, lanceolate, acute at apex and serrate, sessile or shortly petiolate, glabrous, with brown glands beneath. Capitula 20–45 mm in diameter, solitary at the end of branches forming a wide open inflorescence; peduncles long, swollen beneath the capitulum, pale green and glabrous. Involucral bracts in 1 row, 12–18(–20) mm, fused for most of their length, pale green below, cream-coloured above, oblong, forming an acute, triangular tooth at apex, glabrous and fleshy. Flowers of 2 kinds, the outer 25–40 mm, ligulate, the ligules variable in colour even on the same plant, deep orange with a narrow paler margin and sometimes a deeper colour towards the base, pale orange or yellow with an orange base or reddishbrown and irregularly lobed at apex, the inner 20–25 mm, very narrow, tubular, yellow or orange or reddish-brown, irregularly lobed at apex. Receptacle flat, pitted, with scales. Achenes 9–11 mm, dark brown or black, linear, 3- to 4angled, slightly compressed, glabrous; pappus 7–12 mm, of linear, unequal, whitish, connate scales. Flowers 7–9. 2n = 20, 24, 48. Introduced. Much grown in gardens and a frequent casual throw-out or escape. Scattered records in England. Native of Mexico. 108. Schkuhria Roth Annual herbs. Stems erect, branched, leafy. Leaves pinnatifid, glandular-punctate. Capitula very numerous in a more or less corymbose cyme. Involucral bracts few in 1 overlapping row, herbaceous, with membranous tips. Flowers inner tubular, 5-lobed at apex and bisexual, the outer ligulate, the ligules 3-lobed at apex and female. Anthers without tails, apical appendage ovate; filament collar dilated. Style with oblong-linear branches; stigmatic areas in 2 lines. Corolla yellow. Receptacle concave, without scales. Achenes obpyramidal 4-angled; pappus of 8 oblanceolate scales.

110. Helenium Six species in North and South America, but one species widely introduced elsewhere as a weed. Named after Christian Schkuhr (1741–1811). Heiser, C. B. (1945). A revision of the genus Schkuhria. Ann. Missouri Bot. Gard. 32: 265–278.

1. S. pinnata (Lam.) Thell. Dwarf Marigold Pectis pinnata Lam.; S. abrotanoides Roth; S. advena Thell.; S. bonariensis Hook. & Arn.; S. isopappa Benth.; S. pedata A. Gray Annual herb with a tap-root and fibrous side-roots. Stems 20–60 cm, pale yellowish-green, sometimes suffused brownish-purple, erect, ridged, with minute, pale glandular hairs, branched above, densely leafy. Leaves below opposite, above alternate, 1–6 × 2–6 cm, pale greyish-green, broadly ovate in outline, pinnate, the lobes narrowly linear to filiform and obtuse at apex, covered with sunken glands, sessile. Capitula numerous, 6–10 mm in diameter, in a corymbose cyme; peduncles thin, with short, pale glandular hairs and sunken glands. Involucral bracts in 1 overlapping row, 5–6 × 2.5–3.0 mm, pale yellowish-green, oblanceolate, with a rounded, scarious apex, covered with sunken glands. Flowers of 2 kinds, the outer up to 5, ligulate and female, the ligules 3-lobed at apex and yellow, the inner tubular, 5-lobed at apex and bisexual, yellow tinged red. Receptacle concave, without scales. Achenes 3.5–4.0 mm, brown, obpyramidal, 4-angled; pappus of 8 oblanceolate fringed scales. Flowers 8–10. 2n = 22. Introduced. A characteristic wool alien. Scattered records in England. Native of Central and South America. 109. Gaillardia Foug. Annual to short-lived perennial herbs. Stems erect, leafy. Leaves alternate, simple, entire to coarsely dentate. Capitula few, with long peduncles. Involucral bracts in 2–3 rows, more or less herbaceous, becoming reflexed before fruiting. Flowers the inner tubular, 5-lobed at apex and bisexual, the outer ligulate, 3-lobed at apex and usually female. Corolla of tubular flowers orange-red to purple or brownishpurple, of ligulate flowers yellow, often with a purplish base. Anthers without tails, apical appendage ovate; filament collar dilated. Style with oblong-linear branches; stigmatic areas in 2 lines. Receptacle strongly convex to subglobose, with bristle-like scales. Achenes obovoid, covered with a basal tuft of long hairs; pappus of scales with an apical bristle. Twenty-eight species in North and South America; often cultivated elsewhere. Biddulph, S. F. (1944). A revision of the genus Gaillardia. Res. Studies State Coll. Washington 12: 195–256.

1. G. × grandiflora Van Houtte Blanket Flower G. aristata Pursh × G. pulchella Foug. Annual to short-lived perennial herb with a slender tap-root. Stems 20–70 cm, yellowish-green, sometimes tinged purple, erect, slightly striate, with pale, medium simple eglandular hairs, dense at the base becoming sparser upwards, branched, leafy. Leaves medium dull yellowish-green with pale midrib on upper surface, paler beneath, with dense,

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short, felted hairs and scattered longer simple eglandular hairs; basal in a rosette, few to numerous, dying off when in flower, 3–15 × 1.0–6.5 cm, oblanceolate to obovate, rounded at apex, entire to jaggedly toothed, long attenuate at base to a winged petiole 20–100 mm; cauline 3–10 × 0.4– 3.0 cm, alternate, oblanceolate, linear-oblanceolate or panduriform, obtuse to more or less acute at apex, narrowed to a rounded, sessile, amplexicaul base. Capitula 20–60 mm in diameter, solitary at the end of branches; peduncles long, straight, greyish-green, slightly striate, with dense, short and some long, ascending, subappressed simple eglandular hairs. Involucral bracts in 2–3 rows, 3–17 × 2–6 mm, yellowish-green, linear-lanceolate to lanceolate, acute at apex, lax, with long, wavy, pale simple eglandular hairs. Flowers strong-smelling, of 2 kinds, the outer 15–30 mm, ligulate and usually female, the ligules yellow with dark purplish-red base, obtriangular-ovate with 3 triangular lobes and hairy, the inner 5–7 mm, tubular and bisexual, pale creamy-green at base and orange-red to purple or brownishpurple above, sharply jagged at apex and hairy. Receptacle strongly convex to subglobose, with 2 kinds of scales intermixed, triangular-ovate and long-aristate and linear and shortly aristate. Achenes 3–4 mm, obovoid, with appressed hairs; pappus of scales, white and 2–3 mm; plus long awns. Flowers 7–9. 2n = 72. Introduced. Much grown in gardens, thrown out on tips and waste ground and sometimes naturalised especially on sand and shingle by the sea. Scattered records in south-east England. Of garden origin. Most or all of the records of the parents are errors for the hybrid. 110. Helenium L. Tufted perennial herbs. Stems erect, very leafy, winged. Leaves alternate, simple, entire to shallowly dentate, decurrent on stems. Capitula in compact, corymbose cymes at the ends of stems or branches. Involucral bracts in 2–3 rows, herbaceous, becoming reflexed before flowering. Inner flowers tubular, 5-lobed at apex and bisexual, the outer ligulate, the ligules 3-lobed at apex and female. Corolla of outer flowers yellow to mahogany-red, those of inner flowers brownish-yellow. Anthers without tails, apical appendage ovate; filament collar dilated. Style with oblong-linear branches; stigmatic areas in 2 lines. Receptacle convex, pitted, without scales or hairs. Achenes oblong, strongly ribbed, hairy; pappus of scarious scales with apical bristles. Forty species in North, Central and South America, mainly the southern United States and Mexico. H. quadridentatum Labill. from Central America has been recorded as a wool casual. 1. H. autumnale L. Sneezeweed Tufted perennial herb. Stems up to 150 cm, markedly ridged to narrowly winged, pale yellowish-green with darker green ridges, erect, rigid, hollow, glabrous, very leafy, with long, more or less erect branches. Leaves all cauline, 2–8 × 1–2 cm, gradually decreasing in size upwards, dull medium yellowish-green on upper surface, slightly paler beneath, alternate, narrowly elliptical to oblong-elliptical or lanceolate, narrowed to a subacute apex, entire or to shallowly

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1 4 8 . A S T E R AC E A E

dentate, gradually narrowed below, sessile, decurrent on the stems and branches, hairless but densely gland-dotted on both surfaces, veins prominent on both surfaces. Inflorescence of compact corymbose cymes at the end of each stem and branch; peduncles pale green, angled, with darker green ridges, minutely appressed-hairy. Capitula 25–40 mm in diameter, truncate-based. Involucral bracts in 2–3 rows, 7–9 × 1.0–1.5 mm, pale green, narrowly linear-lanceolate, more or less acute at apex, curved upwards, becoming reflexed, shallowly channelled, minutely hairy. Flowers of 2 kinds, the outer up to 20 mm, ligulate and female, the ligules golden yellow, orange, bronze-red or mahogany-red and with a 3-lobed apex, often becoming reflexed, the inner 10–12 mm, tubular and bisexual, brownish-yellow and 5lobed at apex. Receptacle convex, pitted, without scales or hairs. Achenes 2.5–3.0 mm, brown, oblong, strongly ribbed, appressed-hairy; pappus about 0.5 mm, of scarious scales each with an apical bristle. Flowers 7–11. 2n = 34. Cultivars are very variable in height and colour of flowers; some may be hybrids with other species. Introduced. Much grown in gardens and occurring as a throw-out or relic. Scattered localities in south-east England. Native of North America. Tribe 10. Eupatorieae Cass. Annual to perennial herbs. Leaves opposite. Capitula in terminal corymbs or panicles. Flowers all tubular, blue or pinkish-purple, sometimes white. Involucral bracts in several rows, herbaceous. Receptacular scales absent. Pappus of 1 row of hairs or pointed scales. 111. Eupatorium L. Perennial herbs. Stems erect, leafy. Leaves opposite, palmately lobed, the lobes coarsely serrate. Capitula in terminal corymbs. Involucral bracts in 2–3 rows. Flowers all tubular, 5-lobed at apex and bisexual. Corolla pinkishpurple, rarely white. Anthers without tails, apical appendage ovate-lanceolate, flat; filament collar elongate-oblong. Style branches with long filiform-linear sterile branches; stigmatic areas in 2 short, widely separated marginal lines at the base of each style branch. Receptacle flat or convex, without scales. Achenes oblong or fusiform, 5-angled; pappus of 1 row of denticulate hairs. Forty-eight species in the narrowly defined genus of R. M. King & H. Robinson in eastern North America, West Indies, south and east Asia and one species reaching North Africa and Europe. 1. E. cannabinum L. Hemp Agrimony Perennial herb with a woody rootstock. Stems 30–175 cm, pale yellowish-green, often suffused reddish-purple, erect, robust, with numerous to dense, short, crisp, simple eglandular and glandular hairs, simple or with short branches above, leafy. Leaves opposite, dark green on upper surface, paler beneath, with numerous, short, pale, stiff simple eglandular hairs on both surface and the margins; basal 8– 12 × 8–12 cm, broadly ovate in outline, 3- to 5-palmately divided almost to base, the segments lanceolate to ovate or elliptical, acuminate at apex and coarsely serrate, cuneate at base, petiolate; lower cauline gradually becoming smaller,

similar to basal but sessile; upper cauline simple, lanceolate or ovate, long acute or acuminate at apex, serrate, cuneate at base, sessile. Inflorescence of dense terminal corymbs; peduncles densely, short, white hairy. Capitula very numerous, 2–5 mm in diameter. Involucral bracts in 2–3 rows, 2–6 × 0.9–1.1 mm, pale green with a rose-purple flush at the tips, the inner with broad scarious margins, oblong or ovate, obtuse at apex, very shortly hairy down the centre and on the margin. Flowers all tubular and bisexual, 3.0–3.5 mm, pinkish-purple or rarely whitish, 5-lobed at apex. Receptacle flat or convex, without scales. Achenes 2.5–3.0 mm, blackish, oblong or fusiform, 5-angled, glanddotted; pappus 4–5 mm, whitish, of 1 row of denticulate hairs. Flowers 7–9. Protandrous. Visited mainly by Lepidoptera and by some flies and bees. Automatic pollination between different flowers in the same capitulum occurs. 2n = 20. Native. Gregarious in marshes, fens, stream banks and other damp places and sometimes dry grassland or rough ground. Common in England and Wales, frequent in Ireland and the Channel Islands, local and mainly coastal in Scotland. Europe; west and central Asia; North Africa. A member of the European Temperate element. Our plant is subsp. cannabinum which is common throughout the range of the species except Corsica. 112. Ageratum L. Annual herbs. Stems ascending to erect, leafy. Leaves opposite, simple, truncate to cordate at base. Capitula numerous, in compound, subcorymbose panicles. Involucral bracts in 2–3 rows. Flowers all tubular, 5-lobed at apex and bisexual. Corolla usually blue, rarely pink or white. Anthers without tails, apical appendages large. Style branch appendages papillose; stigmatic areas in 2 short, widely separated margin lines at the base of each style branch. Receptacle flat, without scales. Achenes ellipsoid-cylindrical or obpyramical, 5-angled; pappus of narrow, aristate, fimbriate scales. Forty-four species in Central and South America and West Indies. Johnson, M. F. (1971). A monograph of the genus Ageratum L. [Compositae: Helenieae]. Ann. Missouri Bot. Gard. 58: 6–88.

1. A. houstonianum Mill. Flossflower A. mexicanum Sims; A. conyzoides var. mexicanum (Sims) DC. Annual herb with fibrous roots. Stems 30–100 cm, pale green, sometime suffused brownish-purple, ascending or erect, with dense, short glandular and long, pale simple eglandular hairs intermixed, becoming almost glabrous and somewhat woody towards the base, leafy. Leaves (1.2–)2.5– 6.0(–10) × (1.2–)1.5–5.0(–7.0) cm, opposite, medium green on upper surface, paler beneath, ovate or triangularovate, acute at apex, crenate-serrate, truncate to cordate at base, with numerous glandular and simple eglandular hairs especially on the lower surface; petiole 0.5–2.0(–5.0) cm, with glandular and simple eglandular hairs. Inflorescence of dense, compound, subcorymbose panicles; peduncles with glandular and simple eglandular hairs. Capitula numerous,

112. Ageratum 3–10 mm in diameter. Involucral bracts in 2–3 rows, more or less equal, sometimes with some supplementary bracts outside, 4–5(–6) mm, mostly chartaceous, triangular-oblong to lanceolate, acute at apex, with glandular and simple eglandular hairs. Flowers all tubular, 5-lobed at apex and bisexual, usually blue with conspicuous blue stigmas, rarely pink or white, fragrant. Receptacle conical, without scales. Achenes 1.4–1.8 mm, black, ellipsoid-cylindrical to obpyramidal, 5-angled, with sparse simple eglandular hairs;

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pappus of 5(–6), narrow, aristate, fimbriate scales. Flowers 6–9. 2n = 20. Introduced. Grown in gardens and escaping on to waste ground and tips. A few scattered records in south-east England. Native of Mexico and West Indies. Named after William Houstoun (1695–1733). Liatris ligulistylis (A. Nelson) A. Nelson (Lacinaria ligulistylis A. Nelson) has occurred as a casual garden escape.

NEW TAXA AND COMBINATIONS

The new taxa and combinations are to be attributed to the authors whose names follow the names of the taxa and who have supplied the supporting information. Campanula rotundifolia L., Sp. Pl. 163 (1753). subsp. montana (Syme) P. D. Sell stat. nov. C. rotundifolia var. montana Syme, Eng. Bot. ed. 3, 6: 13 (1866). Phyteuma orbiculare L., Sp. Pl. 170 (1753). subsp. tenerum (R. Schulz) P. D. Sell stat. nov. P. tenerum R. Schulz, Monogr. Bearb. Phyteuma 122 (1904). Galium × pomeranicum Retz., Fl. Scand. Prodr. ed. 2, 34 (1795). nothosubsp. hillardiae (Druce) P. D. Sell comb. et stat. nov. G. × hillardiae Druce in Rep. Bot. Exch. Cl. Brit. Isles 8: 869 (1929) pro Rep. Bot. Exch. Cl. Brit. Isles 5: 32 (1918). = G. mollugo subsp. mollugo × verum subsp. maritimum Galium pumilum Murray, Prodr. Stirp. Gott. 44 (1770). subsp. fleurotii (Jord.) P. D. Sell stat. nov. G. fleurotii Jord., Cat. Jard. Grenoble 1849: 2 (1849). Galium aparine L., Sp. Pl. 108, 1231 (1753). subsp. agreste P. D. Sell subsp. nov. Holotype: Margin of potato crop, Hall Close, Bassingbourn, Cambs, v.c. 29, 52/324437, 12 August 1994, P. D. Sell no. 94/282 (CGE). Caules ad 25 cm, prostrati, tegetem circularem humi applanatam facientes vel effusi; internodia ad 35 mm. Folia ad 25 × 3 mm. Flores saepe viridiusculi. Nuculae 2.5–4.5 mm. var. agreste (P. D. Sell) P. D. Sell stat. nov. Holotype as for subsp. agreste. Planta non succulenta, saepe tegetem circularem humi applanatam faciens. Folia plerumque viridia remanentia. Nuculae 3.0–4.5 mm. Floret 8–9(–3). Lonicera periclymenum L., Sp. Pl. 173 (1753). var. hirsuta (Rouy) P. D. Sell stat. nov. L. periclymenum subvar. hirsuta Rouy, Fl. France 8: 75 (1903). Viburnum opulus L., Sp. Pl. 268 (1753). forma flavum (Horwood) P. D. Sell stat. nov. V. opulus var. flavum Horwood, Watson Bot. Exch. Cl. Rep. 2: 540 (1916). Symphoricarpos albus (L.) S. F. Blake, Rhodora 16: 118 (1914). subsp. laevigatus (Fern.) P. D. Sell stat. nov. S. racemosus var. laevigatus Fern., Rhodora 7: 167 (1905).

Valeriana officinalis L., Sp. Pl. 31 (1753). subsp. dunensis P. D. Sell subsp. nov. Holotype: Dune hollow between Birkdale and Ainsdale, South Lancashire, v.c. 59, June 1951. Dwarf form, uniform over several miles of dunes. D. E. Allen (CGE). Plantae stolonibus praeditae. Caules pilosi vel glabri, ad 50 cm. Folia caulina media foliolis 7–13 anguste lanceolatis denticulatis praedita, foliolo terminali quam medio laterali aliquantum latiore. Inflorescentia ramis inferioribus raro praedita. Corolla 5–6 mm. Fructus 3–4 mm. Succisa pratensis Moench, Methodus Pl. 489 (1794). var. subacaulis (N. Bernadin) P. D. Sell comb. nov. ´ Succisa subacaulis N. Bernadin in Cariot, Etude Fl. ed. 5, 2: 301 (1872). Scabiosa succisa var. subacaulis (N. Bernadin) Rouy, Fl. France 8: 115 (1903). var. arenaria (Rouy) P. D. Sell comb. nov. Scabiosa succisa var. arenaria Rouy, Fl. France 8: 115 (1903). var. ovalis (Rouy) P. D. Sell comb. nov. Scabiosa succisa var. ovalis Rouy, Fl. France 8: 115 (1903). var. grandifolia (Rouy) P. D. Sell comb. nov. Scabiosa succisa var. grandifolia Rouy, Fl. France 8: 115 (1903). Carlina vulgaris L., Sp. Pl. 828, 1231 (1753). var. maritima (Rouy) P. D. Sell stat. nov. C. vulgaris forme maritima Rouy, Fl. France 8: 366 (1903). Arctium lappa L., Sp. Pl. 816 (1753). subsp. pubens (Bab.) P. D. Sell comb. nov. A. pubens Bab. in Ann. Mag. Nat. Hist. Ser. 2, 17: 376 (1856). A. minus subsp. pubens (Bab.) Ar`enes in Bull. Jard. Bot. ´ Bruxelles 20: 89 (1950). Etat subsp. nemorosum (Lej.) P. D. Sell comb. nov. A. nemorosum Lej. in Courtois Mag. Hort. (Li`ege) 1: 289 (1833). A. minus subsp. nemorosum (Lej.) Syme in Sowerby, Engl. Bot. ed. 3, 5: 25 (1866). Carduus crispus L., Sp. Pl. 821 (1753). var. glareicola P. D. Sell var. nov. Holotype: Shingle, Snettisham Scalp, W. Norfolk, v.c. 28, 53/652342, 23 July 1991, P. D. Sell no. 91/180 (CGE). Caules 30–45 cm. Capitula pauca, congesta. Carduus nutans L., Sp. Pl. 821 (1753). var. litoralis P. D. Sell var. nov.

New taxa and combinations Holotype: Frequent along the shingle beach, Snettisham Scalp, W. Norfolk, v.c. 28, 53/649339, 15 July 1987, P. D. Sell no. 87/213 and L. C. Nicol (CGE). Caules plerumque breviores quam 50 cm. Capitula 1–3, in ramis brevibus portata. Cirsium vulgare (Savi) Ten., Fl. Napol. 5: 209 (1835–1836). var. litorale P. D. Sell var. nov. Holotype: Frequent along the shingle beach, Snettisham Scalp, W. Norfolk, v.c. 28, 53/649339, 15 July 1987, P. D. Sell no. 87/211 and L. C. Nicol (CGE). Caules ad 50 cm. Capitula pauca, in ramis brevibus congesta. Centaurea scabiosa L., Sp. Pl. 913 (1753). var. nana P. D. Sell var. nov. Holotype: Exposed grassy bank above the sea, near Port Eynon, Gower Peninsula, Glamorganshire, v.c. 41, 21/467844, 2 May 1976, P. D. Sell no. 76/48 (CGE). Planta ad 20 cm. Folia profunde pinnatisecta. Capitula pauca, in pedunculis brevibus portata. Centaurea debeauxii Gren. & Godr., Fl. France 2: 243 (1850). subsp. nemoralis (Jord.) Dost´al in Bot. Jour. Linn. Soc. 71: 207 (1976). var. minima (C. E. Britton) P. D. Sell comb. nov. C. nemoralis var. minima C. E. Britton in Rep. Bot. Soc. Exch. Cl. Brit. Isles 6: 414 (1922). var. nemoralis (Jord.) P. D. Sell comb. nov. C. nemoralis Jord., M´em. Acad. Sci. Lyon 1 (1851) (Pug. Pl. Nov. 104 (1852)). C. nigra var. nemoralis (Jord.) Gremli, Exkursionsfl. Schweiz ed. 6, 254 (1889). subsp. thuillieri Dost´al in Bot. Jour. Linn. Soc. 71: 207 (1976). var. pumila P. D. Sell var. nov. Holotype: Freshwater Downs, Isle of Wight, August 1890, J. H. A. Stewart (CGE). Planta ad 30 cm. Capitula pauca, in pedunculis brevibus portata. Flores exteriores interioribus multo longiores. var. thuillieri (Dost´al) P. D. Sell stat. nov. C. debeauxii subsp. thuillieri Dost´al in Bot. Jour. Linn. Soc. 71: 207 (1976). C. pratensis Thuill., Fl. Env. Paris ed. 2, 444 (1799), non Salisb., Prod. Stirp. Chap. Allerton 208 (1796). Scorzoneroides autumnalis (L.) Moench, Methodus Pl. 549 (1794). Leontodon autumnalis L., Sp. Pl. 798 (1753). var. salina (Aspegren) P. D. Sell comb. nov. Apargia autumnalis var. salina Aspegren, Blekingsk Fl. 57 (1823). var. simplex (Duby) P. D. Sell comb. nov. Leontodon autumnalis var. simplex Duby, Bot. Gall. ed. 2, 1: 308 (1828) pro Sowerby, Eng. Bot. tab. 830 (1801). var. nigrolanata (Fr.) P. D. Sell comb. nov. Leontodon autumnalis var. nigrolanatus Fr., Summa Veg. Scand. 8, 187 (1846). var. alpina (Gaudin) P. D. Sell comb. nov.

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Apargia autumnalis var. alpina Gaudin, Fl. Helv. 5: 59 (1829). var. pratensis (Hornem.) P. D. Sell comb. et stat. nov. Apargia pratensis Hornem., Hort. Bot. Hafn. 2: 785 (1815). var. pinnatifida (Schur) P. D. Sell comb. nov. L. autumnalis var. pinnatifidus Schur, Enum. Pl. Transsilv. 356 (1866). var. coronopifolia (Lange) P. D. Sell comb. nov. L. autumnalis var. coronopifolius Lange, Haandb. Danske Fl. ed. 4, 399 (1886). var. cinerascens (Briq.) P. D. Sell comb. nov. L. autumnalis var. cinerascens Briq. in Annu. Cons. Jard. Bot. Gen`eve 3: 124 (1899). var. dentata (Holuby ex Schur) P. D. Sell comb. nov. L. autumnalis var. dentatus Holuby ex Schur in Verhandl. Naturf. Ver. Br¨unn 15: 186 (1877). var. latifolia (Schur) P. D. Sell comb. nov. L. autumnalis var. latifolius Schur, Enum. Pl. Transsilv. 356 (1866). Leontodon × vegetus Finch & P. D. Sell nothospecies nova = L. hispidus × saxatilis Holotype: Origin: Parish Church (St James the Less), North Lancing, W. Sussex, v.c. 13, 51/182055, 5 July 1996, P. D. Finch. Growing with both parental species nearby, in turf just outside the kerb at east end of graves of the Dabbs family, to the south of church, west of path down to Vicarage. Cult. in Lancing soil with coco-peat at bottom of pots till pressed on 29 August 1997, by R. A. Finch. 2n = 11. F1 . (CGE). Herba perennis variabilis, praesertim magnitudine, indumento, foliorum figura, papporum auctu. Caules ad 45 cm, graciles. Folia ad 20.0 × 2.5 cm, pallide luteoviridia, oblongooblanceolata, ad apicem obtusum angustata, sinuatodentata, pilis diverse furcatis vestita, ad basin in petiolum gracilem longe attenuata. Involucri bracteae ad 11 × 2 mm, pallide virides, glabrae vel pilis numerosis diverse furcatis pallidis vestitae. Cypselae pleraeque steriles maleque effectae, exteriores plerumque pappo ad instar coronae, interiores pilorum plumosorum pappo praeditae. Pollinis grana magnitudine variabillima. Leontodon saxatilis Lam., Fl. Franc¸. 2: 115 (1779). var. pristis (Druce) P. D. Sell comb. nov. L. nudicaulis var. pristis Druce in Jour. Bot. (London) 45: 423 (1907). var. arenarius (Duby) P. D. Sell comb. nov. L. hastilis var. arenarius Duby, Bot. Gall. ed. 2, 1: 302 (1828). Picris hieracioides L., Sp. Pl. 792 (1753). var. villarsii (Jord.) P. D. Sell comb. nov. P. villarsii Jord., Cat. Dijon 29 (1848). var. leteae (E. I. Ny´ar.) P. D. Sell comb. nov. P. sonchoides var. leteae E. I. Ny´ar. in Fl. Rep. Pop. Rom. 10: 719 (1965). var. arenaria (E. I. Ny´ar.) P. D. Sell comb. nov. P. sonchoides var. arenaria E. I. Ny´ar. in Fl. Rep. Pop. Rom. 10: 720 (1965).

532

New taxa and combinations

Helminthotheca echioides (L.) Holub, Folia Geobot. Phytotax. 8: 176 (1973). var. mollis (Duby) P. D. Sell comb. nov. Helminthia echioides var. mollis Duby, Bot. Gall. 1: 300 (1828). var. pratensis (Chevall.) P. D. Sell comb. nov. Helminthia echioides var. pratensis Chevall., Fl. G´en. Env. Par. 2: 352 (1827). var. incisa P. D. Sell var. nov. Holotype: Garden of 54, Cambridge Road, Impington, Cambs, v.c. 29, 52/436619, 15 July 1998, P. D. Sell no. 98/92 (CGE) (1 plant on 6 sheets). Planta erecta, pilis longioribus rigidis aeque cum brevibus vestita. Folia magna, costam versus ad partem dimidiam lobis latis dentatis incisa. Involucri bracteae exteriores ad basin cordatae. Sonchus oleraceus L., Sp. Pl. 794 (1753). var. litoralis P. D. Sell var. nov. Holotype: Shingle beach between Cley and Salthouse, E. Norfolk, v.c. 27, 63/059451, 21 September 1967, P. D. Sell no. 67/1775 and N. Marchant (CGE). Caulis 10–50 cm. Folia saepe rubripurpureotincta. Capitula pauca, in fasciculo denso congesta. Sonchus asper (L.) Hill, Herb. Brit. 1: 47 (1769). var. sabulosus P. D. Sell var. nov. Holotype: Stabilised shingle at back of beach, north of Dunwich, E. Suffolk, v.c. 25, 62/479709, 26 June 1990, P. D. Sell no. 90/157 and J. G. Murrell (CGE). Caulis ad 50 cm. Folia plerumque divisa, aculeatissima. Capitula pauca, in fasciculo denso congesta. Lactuca virosa L., Sp. Pl. 795 (1753). forma lactucarii (Lamotte) P. D. Sell stat. nov. L. lactucarii Lamotte, Prodr. Fl. Plat. Centr. 2: 467 (1881). L. scariola forme virosa var. lactucarii (Lamotte) Rouy, Fl. France 9: 199 (1905). Crepis vesicaria L., Sp. Pl. 805 (1753). var. rutilans (Lacaita) P. D. Sell comb. et stat. nov. C. rutilans Lacaita, Jour. Linn. Soc. Bot. 44: 128 (1918). Pilosella × stoloniflora (Waldst. & Kit.) F. W. Sch. & Sch. Bip. in Flora (Regensb.) 45: 423 (1862). nothosubsp. schuriana (N¨ageli & Peter) P. D. Sell comb. nov. Hieracium × stoloniflorum subsp. schurianum N¨ageli & Peter, Hier. Mittel-Europas: Piloselloiden 326 (1885). Probabiliter = P. aurantiaca subsp. carpathicola × officinarum subsp. euronota Hieracium L. In Section Sabauda (Fr.) Arv.-Touv. Hieracium vagum Jord., Cat. Grenoble 21 (1849). forma croceostylum (Pugsley) P. D. Sell comb. et stat. nov. H. croceostylum Pugsley in Jour. Linn. Soc. London (Bot.) 54: 310 (1948).

Hieracium prominentidens P. D. Sell nom. et stat. nov. H. dispalatum var. macrodon Sudre, Hier. Centr. France 24 (1902), non H. macrodon N¨ageli & Peter, Hier. MittelEuropas: Archieracien 84 (1886). Hieracium sabaudum L., Sp. Pl. 804 (1753). forma bladonii (Pugsley) P. D. Sell comb. et stat. nov. H. bladonii Pugsley in Jour. Linn. Soc. London (Bot.) 54: 299 (1948). In Section Hieracioides Dumort. Hieracium umbellatum L., Sp. Pl. 804 (1753). var. sarniense P. D. Sell var. nov. Holotype: Granite cliffs, north coast of Jersey, Channel Islands, 17 July 1901, L. V. Lester-Garland as E. F. & W. R. Linton, Set of British Hieracia no. 185 (CGE). Caulis 10–30 cm, robustus, succulentus. Folia crassa, succulenta, anguste vel late elliptica, ad apicem obtusa, inferiora ad c. 5 cm lata, prope caulis basin fasciculata, sursum minora. var. latifolium (W. R. Linton) P. D. Sell stat. nov. H. umbellatum forma latifolium W. R. Linton, Brit. Hier. 98 (1905). In Section Foliosa (Fr.) Arv.-Touv. Hieracium reticulatiforme P. D. Sell nom. et stat. nov. H. reticulatum var. angustifolium Pugsley in Jour. Linn. Soc. London (Bot.) 54: 280 (1948), non H. angustifolium Hoppe, Bot. Taschenb. 1799: 129 (1799). Hieracium reayense (Pugsley) P. D. Sell stat. nov. H. reticulatum var. reayense Pugsley in Jour. Linn. Soc. London (Bot.) 54: 280 (1948). Hieracium melanoglochin (E. F. Linton) P. D. Sell stat. nov. H. corymbosum var. melanoglochin E. F. Linton in W. R. Linton, Brit. Hier. 88 (1905). In Section Tridentata (Fr.) Arv.-Touv. Hieracium charitodon P. D. Sell sp. nov. Holotype: Waterfall at the head of Glyn Tarell, Brecon, v.c. 42, 25 July 1900, A. Ley (CGE). Herba perennis aphyllopoda vel hypophyllopoda, caudice ramoso. Caulis 35–100 cm, pallide luteoviridis, ad basin saepe purpurascens, gracilis vel robustus, infra pilis simplicibus eglandulosis densis longis pallidis, sursum paucioribus brevioribusque, et in parte superiore pilis stellatis numerosis vestitus. Folia in pagina superiore pallide vel mediocriter viridia, in pagina inferiore pallidiora, 5–15, sursum gradatim decrescentia; folia basalia et caulina infera lamina 6–11 × 1.5–4.0 cm, anguste elliptica vel oblongoelliptica vel lanceolatooblonga, ad apicem obtusomucronatum vel acutum gradatim angustata, dentibus mammiformibus vel longiusculis angustisque dentata, ad basin in petiolum alatum ad 4 cm longum attenuata; caulina mediana lamina 4–9 × 1.5–4.0 cm, linearielliptica

New taxa and combinations vel oblongoelliptica, ad apicem magis minusve acutum gradatim angustata, dentibus longis angustis ad 14 mm dentata, ad basin gradatim attenuata, sessilia vel petiolo brevi praedita; caulina supera lamina lineari vel linearilanceolata, ad apicem longe acuta, integra vel dentibus angustis dentata, sessilia; omnia in pagina superiore glabra vel pilis nonnullis brevissimis vestita, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis vestita. Inflorescentia capitulis 1–21, paniculatocorymbosa, saepe ramis inferioribus longis praedita; pedunculi graciles erecti, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis sed interdum base obscura pilisque stellatis densis vestiti. Capitula 25–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 5–12 × 1.2–1.5 mm, olivaceae vel nigrivirides, interiores ad marginem pallidiores, omnes linearilanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis vel densis brevibus vel mediocribus pallidis sed saepe base obscura pilisque glanduliferis paucis brevibus obscuris pilisque stellatis paucis vestitae. Ligulae flavae, glabrae. Styli obscuri. Receptaculi alveoli margine fibrillosodentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 7–8. Apomicticum. In Section Alpestria (Fr.) Arv.-Touv. Hieracium klingrahoolense Walter Scott & R. C. Palmer sp. nov. Holotype: Steep rocks by pool, lower end of the Burn of Skelladale, near Brae, Shetland Islands, v.c. 112, 15 August 1989, W. Scott no. 3286 (CGE). Herba perennis plerumque aphyllopoda, caudice ramoso. Caulis 20–70 cm, pallide luteoviridis, saepe colore brunneopurpureo suffusus, flexuosus, striatus, glaber vel fere glaber vel pilis paucis stellatis vel minutis vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora, interdum purpureotincta; folia basalia sub anthesi plerumque absentia sed interdum lamina parva elliptica, ad apicem obtusa, ad basin cuneata, praedita; folia caulina 10–19, in caulis dimidio superiore abrupte decrescentia, infera lamina 4–11 × 1–2 cm, anguste linearielliptica vel oblonga, ad apicem plerumque acutum attenuata, subintegra vel dentibus minutis praedita, ad basin gradatim angustata et magis minusve amplexicaulia, supera parva, linearia vel linearilanceolata, ad apicem acuta, integra et amplexicaulia; omnia glabra vel pilis paucis minutis, praesertim in margine, vestita. Inflorescentia capitulis 1–10; pedunculi pilis stellatis numerosis vestiti, plerumque sine pilis simplicibus eglandulosis pilisque glanduliferis manifestis, sed pilis glanduliferis minutissimis vestiti. Capitula 25–30 mm diametro, ad basin rotundata vel subtruncata. Involucri squamae ante anthesin incumbentes, 5–11 × 1.0– 1.2 mm, nigrivirides, interiores ad marginem pallidiores, omnes linearilanceolatae, ad apicem obtusae, pilis glanduliferis paucis vel numerosis brevibus vel mediocribus obscuris et interdum pilis simplicibus eglandulosis solitariis et basin versus pilis stellatis paucis vestitae. Ligulae flavae, ad apicem glabrae. Styli flavi. Receptaculi alveoli margine acute dentato. Cypselae 3.5–4.0 mm, intense rubrae. Floret 7–8. Apomicticum.

533

Hieracium amaurostictum Walter Scott & R. C. Palmer sp. nov. Holotype: Low sea-banks east of the north end of the Loch of Semblister, Shetland Islands, v.c. 112, 3 August 1993, L. A. Inkster and W. Scott no. 3257 (CGE). Herba perennis phyllopoda vel aphyllopoda, caudice ramoso. Caulis 20–45 cm, pallide luteoviridis, saepe colore brunneopurpureo suffusus, gracilis, flexuosus, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis, saepe densis, longis sinuatis pallidis, sursum aliquantum paucioribus, et in parte superiore pilis stellatis numerosis et raro pilis glanduliferis solitariis brevibus obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, colore brunneopurpureo dilute maculata et guttata, in pagina inferiore pallida, saepe purpureosuffusa; folia basalia sub anthesi pauca vel absentia, lamina 2– 7 × 2.0–2.5 cm, anguste elliptica vel oblongoelliptica, ad apicem acuta, integra vel denticulationibus minutis praedita, ad basin cuneata vel attenuata, petiolo ad 4 cm, saepe purpurascente, pilis simplicibus eglandulosis densis longis vestito; folia caulina 3–6, 1.5–7.0 × 0.2–2.5 cm, sursum gradatim minora, infera basalibus similia petiolataque, mediana elliptica vel lanceolata, ad apicem acuta, integra vel denticulata, ad basin sessilem cuneata, supera linearilanceolata vel linearia, ad apicem acuta, integra et sessilia, aut bracteiformia; omnia in paginis ambabus inque margine pilis simplicibus eglandulosis numerosis longis pallidis vestita. Inflorescentia capitulis 1–3; pedunculi pilis stellatis densis et pilis simplicibus eglandulosis numerosis brevibus mediocribusque base obscura et pilis glanduliferis paucis brevibus obscuris vestita. Capitula 30–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 6–12 × 1.3–1.6 mm, nigrivirides, interiores ad marginem pallidiores, omnes linearilanceolatae, ad apicem obtusae vel subacutae, pilis simplicibus eglandulosis numerosis brevibus mediocribusque base obscura et pilis glanduliferis paucis brevibus obscuris vestitae, pilis stellatis fere omnino absentibus. Ligulae flavae, glabrae vel ad apicem pilis paucis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine acute dentato. Cypselae 4–5 mm, rubrinigrae. Floret 7–8. Apomicticum. In Section Alpina (Griseb.) Gremli Hieracium deargicola P. D. Sell & D. J. Tennant sp. nov. Holotype: Cliff ledges, Ben Dearg, E. Ross, v.c. 106, August 1982, D. J. Tennant (CGE). Inter Hieracium kennethii et H. subglobosum aliquantum intermedium, a specie prima foliis minus dentatis, involucri squamis acutioribus, stylis flavis, a specie secunda foliis pilosioribus, in involucri squamis pilis glanduliferis paucioribus manifestis, stylis flavis differt. Herba perennis phyllopoda, caudice ramoso. Caulis 8– 23(–28) cm, pallide viridis, mediocris vel aliquantum gracilis, ubique pilis simplicibus eglandulosis dispersis vel numerosis brevibus mediocribusque longisque sinuatis vel rectis pallidis sed plerumque base obscura (in parte superiore saepe cinerascentibus base longa nigra) pilisque glanduliferis dispersis brevissimis obscuris (in parte superiore numerosioribus vel satis densis, brevissimis brevibusque

534

New taxa and combinations

mediocribusque (ad 1.0 mm), nigris) pilisque stellatis dispersis vel numerosis (in parte superiore numerosioribus et ubique pilos glanduliferos breves occulentibus) vestitus. Folia in pagina superiore pallidiuscule vel mediocriter vel interdum tristiuscule viridia pauloque nitida, in pagina inferiore aliquantum pallidiora, plerumque non colore rubriviolaceo tincta, aspectu manifeste graniculata; folia basalia magis minusve numerosa, interdum in rosula satis nitida, tenuia vel minime coriacea, paulo undulata, haud profunde canaliculata vel plana, exteriora lamina 5–35 × 4–25 mm, ovata vel obovata vel subrotunda, ad apicem magis minusve late rotundatoobtusa (pauca leviter retusa), subintegra, remote denticulata vel in margine dentibus paucis inaequalibus latis obtusiusculis praedita, ad basin in petiolum mediocre vel satis longum cuneata vel cuneatotruncata, interiora lamina 8–50 × 5–25 mm, plerumque elliptica (aliquot anguste elliptica vel ovata vel obovata, intima saepe oblanceolata), ad apicem plerumque magis minusve late obtusa (aliquot subacuta vel acuta leviterque torta), in margine subintegra vel denticulata vel serratodentata, dentibus haud profundis vel anguste mammiformibus vel late acutis, magis minusve cuspidatis, interdum ad acumen rubellis, ad basin plerumque cuneata (intima ad basin in petiolum mediocrem vel longum fere attenuata), exteriora lamina saepe subglabra, pilis simplicibus eglandulosis in margine numerosissimis mediocribus vel longiusculis inque pagina inferiore dispersis vel numerosis plerumque mediocribus (saepe in costa densioribus) et pilis glanduliferis plerumque in margine aliquot vel paucis brevissimis flavidis vestita, sine pilis stellatis, interiora lamina pilis simplicibus eglandulosis numerosis (vel dispersis) rigidiusculis sinuatis mediocribus longisque albis, ad basin pallidis, uniformiter distributis vel interdum in pagina superiore marginem versus limitatis, vestita, petiolis plerisque pilis simplicibus eglandulosis numerosis longis sinuatis vestitis, plerumque sine colore rubriviolaceo; folia caulina 1–4, infimum rigide suberectum vel subpatens, lamina 10–40 × 2–12 mm, lineariellipticum vel linearilanceolatum vel oblanceolatum, plerumque acutum, saepissime ad apicem acumine longo tenui praeditum, subintegrum vel interdum in margine aliquantum dentatum undulatumque, ad basin sessilem attenuatum vel petiolo breviusculo saepe alato praeditum, secundum linearioblanceolatum, ad apicem subtiliter acuminatum, integrum, vel infimo simile, cetera bracteiformia suberecta longiuscula, angustissime linearia, subtiliter acuta, omnia in pagina superiore pilis simplicibus eglandulosis paucis vel multis mediocribus vel longis, in margine paginaque inferiore pilis simplicibus eglandulosis numerosis sinuatis, ad basin magis minusve nigris, pilisque stellatis dispersis vel multis, aut nullis, vestita. Capitulum saepe solitarium, ad 3 (4 in cultura), 30–45 mm diametro, involucro aspectu obscurissime olivaceogriseo vel olivaceonigro, latiuscule campanulato, rotundato vel in pedunculum minime angustato, in medio plerumque constricto; pedunculus (quando adest) aliquantum furve dilute viridis, ut caulis superus ita vestitus. Involucri squamae ante anthesin incumbentes, nigrivirides, pilis simplicibus eglandulosis numerosis vel densis mediocribus longisque (ad 4(–5) mm) sinuatissimis albidis vel argentigriseis base nigra, pilos glanduliferos brevissimos vel breves (ad 0.6

mm) nigellos fere omnino occulentibus (vel interdum paucis visibilibus), vestitae, sine pilis stellatis sed ad apicem pilorum brevium ramosorum et simplicium caespite implicato praeditae, interiores inaequales, longissimae 11–15 × 1.0– 1.6 mm, appressae vel laxe appressae, anguste lanceolatosubulatae, acutissimae vel acuminatae (vel paucae subacutae), intimae ad apicem subtiliter acuminatae, exteriores multo breviores, laxae, magis minusve erectae, acutae vel subacutae. Ligulae mediocriter (vel intensiuscule) flavae, centraliter aliquantum densae, exteriores ad 3.5(–4.0) mm latae, satis discretae, rectae vel leviter sursum versae, lobis longiusculis angustis ad apicem praeditae et pilis simplicibus eglandulosis in pagina inferiore paucissimis vel aliquot brevibus vel mediocribus, saepe aliquantum versicoloribus (pallide aureobrunneis), aut nullis, in lobis ad apicem paucissimis vel multis brevissimis vel brevibus, ultra loborum margines vix procurrentibus, vestitae. Styli pure flavi. Receptaculi alveoli margine breviter dentato. Cypselae 3.3–4.4 mm, purpureonigrae. Floret 7–8. Apomicticum. Hieracium einichense P. D. Sell & D. J. Tennant sp. nov. Holotype: West side of Glen Einich, E. Inverness, v.c. 96, 6 August 1975, D. J. Tennant (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 8– 25 cm, mediocris vel raro aliquantum gracilis, ubique pilis simplicibus eglandulosis dispersis vel multis brevibus mediocribusque vel longiusculis sinuatissimis (in parte superiore plerumque nigrigriseis, base nigra) pilisque glanduliferis dispersis brevissimis brevibusque obscuris (in parte superiore numerosis vel densis, plerumque brevibus vel mediocribus, nigris) pilisque stellatis dispersis vel multis (in parte superiore magis minusve densis) vestitus. Folia in pagina superiore saturate viridia, fere hebetata, colore obscure tristeque rubriviolaceo magnopere vel omnino suffusa, in pagina inferiore pallidiora, dilute glaucoviridia, costa prominenti, saepe purpurascenti; folia basalia pauca vel numerosa, in rosula satis nitida aliquantum patenti, rigidiuscula, aliquantum coriacea, exteriora lamina 7–40 × 4–25 mm, obovata vel late elliptica vel subrotunda, ad apicem late rotundatoobtusa, subintegra, denticulata vel in margine dentibus nonnullis parvis praedita, ad basin in petiolum saepe breviusculum paulo alatum magis minusve cuneata vel rotundata, interiora lamina 20– 55 × 5–25 mm, plerumque elliptica vel oblongoelliptica (aliquot obovata, aliquando pauca ovata, intima saepe magis minusve oblanceolata), ad apicem obtusa, saepe mucronata (multa late rotundatoobtusa, intima subacuta, interdum apiculata), denticulata vel basin versus dentibus nonnullis satis regularibus parviusculis acutissimis, aliquando cuspidatis vel anguste mammiformibus, praedita, ad basin cuneata vel anguste cuneata (intima ad basin in petiolum late alatum vel aliquantum gracilem brevem vel longiusculum magis minusve attenuata), exteriora lamina magis minusve subglabra vel in margine pilis simplicibus eglandulosis numerosissimis plerumque mediocribus (sed aliis longioribus, aliis brevioribus) magis minusve rigidis inque pagina inferiore dispersis vel numerosissimis (in costa aliquantum densioribus) et pilis glanduliferis brevissimis, in margine multis inque paginis ambabus aliquot, et interdum

New taxa and combinations in pagina inferiore pilis stellatis nonnullis vestita, interiora lamina in pagina superiore pilis simplicibus eglandulosis magis minusve numerosis, plerumque mediocribus, subrigidis, albidis, base pallide vel obscure violacea, aut moderate uniformiterque distributis aut saepius marginem versus limitatis (in foliis intimis interdum numerosissimis et longiusculis), vestita, petiolis in margine pilis simplicibus eglandulosis mediocribus longisque sinuatis sparsim vel modice vestitis; folia caulina (0–)3–5(–6), infimum suberectum vel rigide subpatens, lamina 10–50 × 1– 9 mm, plerumque linearioblanceolatum, interdum anguste lineare et bracteiforme, ad apicem magis minusve acutum, integrum vel saepius denticulatum vel in margine interdum undulato dentibus aliquot parviusculis acutissimis praeditum, sessile et semiamplexicaule vel interdum petiolatum, in pagina inferiore saepe pilis stellatis multis, cetera magis minusve linearia, integra, summum bracteiforme. Capitulum plerumque solitarium (raro 2), 38–48 mm diametro (aliquando caule secundo a rosula basali exorienti, capitulum ferenti, saepe in cultura), involucro aspectu nigricanti, campanulato, ad basin rotundato et in medio constricto; pedunculus (quando adest) ut caulis superus ita vestitus. Involucri squamae ante anthesin incumbentes, nigrivirides, pilis simplicibus eglandulosis multis vel numerosis plerumque mediocribus nonnullisque longiusculis (ad 2(–2.5) mm), sinuatis, plerumque cinerascentibus vel obscurissimis base nigra et pilis glanduliferis satis densis brevissimis brevibusque mediocribusque (ad 1.5 mm) nigris, a pilis simplicibus eglandulosis vix celatis, interiores pilorum brevium ramosorum simpliciumque caespite definito ad apicem praeditae, exteriores pilis nonnullis stellatis ad basin praeditae, interiores inequales, longissimae (11.5–)13–15 × 1.1–1.6 mm, aliquantum appressae, anguste linearilanceolatae, plerumque abrupte acutae (nonnullae subacutae acuminataeque angusteque obtusae), intimae ad apicem acutissimae, exteriores breviores, magis minusve appressae. Ligulae intensiuscule (vel mediocriter) flavae, centraliter aliquantum densae, exteriores ad 3.6(–4.0) mm latae, discretae, ad apicem leviter vel valde sursum versae, lobis longiusculis angustis praeditae et in pagina inferiore pilis simplicibus eglandulosis paucissimis vel aliquot brevissimis (vel brevibus), interdum aliquantum appressis, et plerumque nonnullis ultra apicem brevissime (0.1–0.2 mm) procurrentibus (vel magis minusve nullis) vestitae. Styli mediocriter vel intense obscuri, uniformiter fuscate olivaceogrisei. Cypselae 3.2–3.8 mm, purpureonigrae. Floret 7–8. Apomicticum. Hieracium subtenuifrons P. D. Sell & D. J. Tennant sp. nov. Holotype: Granite slabs at about 2,250 ft, south side of Coire Garbhlach, Glen Feshie, E. Inverness, v.c. 96, 27/872945, 10 August 1966, R. W. Jones and B. A. Miles no. 66/203 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 10– 25(–32) cm, graciliusculus vel aliquantum robustus, interdum striatus, raro flexuosus, ad basin plerumque colore rubriviolaceo tinctus, in parte inferiore pilis simplicibus eglandulosis dispersis vel multis, plerumque mediocribus vel longis, sinuatisssimis albidis et pilis glanduliferis

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plerumque satis paucis brevissimis pallidis vel obscuris et pilis stellatis dispersis, in parte superiore pilis simplicibus eglandulosis multis vel numerosissimis (raro paucis) mediocribus longisque, fere strictis vel sinuatisssimis (aliquando irregulariter tortis), albidis vel saepe cinerascentibus base nigra, et pilis glanduliferis aliquot vel multis brevissimis (nonnullis brevibus, ad 0.7 mm) pallidis obscurisque et pilis stellatis magis minusve numerosis vestitus. Folia in pagina superiore mediocriter vel intensiuscule, clare viridia (raro triste dilutissimeque caeruleoviridia), omnia (vel saltem intima) aliquantum nitida, in pagina inferiore aliquantum pallidiora et leviter nitida vel subglauca, plerumque, praesertim in pagina inferiore, colore rubriviolaceo tincta vel saepe valde suffusa; folia basalia numerosa, exteriora lamina 8–45 × 3–28 mm, obovata vel elliptica vel oblanceolata, ad apicem plerumque obtusa vel late rotundatoobtusa (et saepe torta), subintegra vel denticulata vel dentibus nonnullis majoribus praedita, ad basin cuneata vel attenuata, interiora lamina 12–70 × 4–30 mm, rigida, interdum aliquantum coriacea, saepe sinuata et aliquantum canaliculata, plerumque anguste vel late elliptica (interdum oblanceolata), ad apicem plerumque latiuscule acuta vel subacuta et saepe mucronata vel apiculata (interdum obtusa, saepe valde torta), plerumque dentibus haud profunde vel anguste mammiformibus vel aliquando aquilinomammiformibus dentata vel serratodentata (raro subintegra), ad basin cuneata vel attenuata, omnia pilis simplicibus eglandulosis in pagina superiore aliquot vel multis, mediocribus vel longis, gracilibus vel crassiusculis, subrigidis, sinuatis vel ad apicem curvatis, base pallida (raro base nigra), plerumque prope marginem numerosioribus et saepe circa costam nullis (raro uniformiter distributis vel absentibus, in foliis intimis interdum numerosissimis longissimisque), in margine numerosis vel numerosissimis, saepe longis, rectis vel sinuatis, in pagina inferiore paucis vel multis, plerumque mediocribus vel longiusculis, base pallida vel nigra (aut nullis), et pilis glanduliferis aliquantum paucis vel dispersis brevissimis flavidis (in margine manifestioribus) vestita, plerumque sine pilis stellatis, petiolis breviusculis vel longiusculis, gracilibus vel aliquantum alatis, plerumque valde colore rubriviolaceo tinctis (saltem prope basin), pilis simplicibus eglandulosis dispersis vel numerosis vel numerosissimis, mediocribus vel longis, interdum sinuatissimis, vestitis; folia caulina (3–)4–5(–9), infimum subpatens, saepe undulatum, lamina 6–30(–40) × 0.5–7.0 mm, lineare vel linearioblanceolatum, ad apicem acutissimum vel acuminatum vel subacutum, integrum vel denticulatum vel dentibus paucis parvis acutis vel cuspidatis praeditum, sessile vel petiolatum, pilis simplicibus eglandulosis in pagina superiore paucis vel aliquot, in margine paginaque inferiore numerosis, saepe base nigra, vestitum, cetera similia, minora et semierecta, summum bracteiforme. Capitula solitaria vel pauca (raro ad 7), 35–55 mm diametro, involucro aspectu mediocriter olivaceogriseo vel olivaceonigro, campanulato vel late campanulato, ad basin late vel anguste rotundato. Involucri squamae ante anthesin incumbentes, nigrivirides; interiores 12–16 × 1.0–1.6 mm, appressae vel semiappressae, linearilanceolatae vel saepe lanceolatosubulatae, ad apicem magis minusve acutae vel acuminatae, exteriores breviores, aliquantum laxae, suberectae, ad apicem

536

New taxa and combinations

subacutae vel acuminatae, omnes pilis simplicibus eglandulosis numerosissimis vel densis, mediocribus vel longis (ad 4(–5) mm), fere rectis vel sinuatissimis, saepe magis minusve griseis sed acumine pallido et base nigra, et pilis glanduliferis numerosis brevissimis nonnullisque brevibus, pallidis obscurisque, fere semper a pilis simplicibus eglandulosis omnino occultis, vestitae, sine pilis stellatis, sed interiores ad apicem pilis nonnullis brevibus pallidis caespitem indistinctum formantibus praeditae. Ligulae clare et pallide flavae, exteriores ad 3.5–4.3 mm latae, in parte apicali aliquantum vel modice discretae, rectae vel leviter sursum curvatae, in pagina inferiore sine pilis simplicibus eglandulosis aut nonnullis (vel multis) plerumque brevibus pallidis (saepe nonnullis griseotinctis vel brunneotinctis) vestitae, ad apicem absentibus vel ultra apicem paucis vel aliquot brevissime (ad 0.2 mm) procurrentibus. Styli magis minusve obscuri (mediocriter olivaceogrisei vel fuscate grisei, stigmatibus interdum paulo pallidioribus). Receptaculi alveoli margine breviter dentato. Cypselae 3.0–4.1 mm, purpureonigrae. Floret 7–8. Apomicticum. Hieracium pentaploideum P. D. Sell & D. J. Tennant sp. nov. Holotype: North side of An Teallach Range, W. Ross, v.c. 105, 22 July 1986, D. J. Tennant (CGE). Species sola pentaploidea in genere Hieracio (Pilosella exclusa) nota, H. pentaploideum, notas characteristicas aliquot easdem habens ac H. holosericeum, quod illud comitatur, ex hoc hybridogenum fortasse exortum est. Herba perennis phyllopoda, caudice ramoso. Caulis 7–22 cm, aliquantum pallide viridis, mediocris vel aliquantum gracilis, saepe flexuosus, prope involucrum valde incrassatus, pilis simplicibus eglandulosis ubique numerosis, saepe densis, mediocribus et longis vel longissimis (ad 5 mm) sinuatissimis albis base nigra, et pilis glanduliferis tenuibus, in parte inferiore paucis vel multis brevissimis, in parte superiore magis minusve numerosis brevissimis brevibusque (ad 0.5 mm) obscuris, et pilis stellatis brevissimis, in parte inferiore sparsis, in parte superiore numerosis vel satis densis, vestitus. Folia in pagina superiore mediocriter vel intensiuscule, saepe tristiuscule viridia (griseiolivacea) aliquantumque nitida, in pagina inferiore pallidiora, saepe paulo nitida, extima (omniumque apices) aliquando colore triste intenseque rubriviolaceo tincta; folia basalia pauca vel numerosa, exteriora lamina 5–30 × 3– 16 mm, plerumque obovata vel late elliptica, ad apicem late rotundatoobtusa, saepe mucronata, interdum aliquantum retusa, integra vel denticulata vel interdum dentibus paucis parvis (raro magnis) praedita, ad basin cuneata vel rotundata vel in petiolum saepe breviusculum, interdum alatum, breviter angustata, interiora lamina 10–70 × 2– 15 mm, saepe paulo coriacea, plerumque anguste elliptica (interdum oblanceolata vel lanceolata et intima saepe linearielliptica vel linearia), ad apicem plerumque acuta vel subacuta (media interiora aliquando obtusomucronata), margine saepe undulatissimo, interdum torto, praesertim apicem versus, subintegra vel denticulata vel dentibus paucis haud profundis praedita (interdum dentibus nonnullis acutis, leviter vel angustissime mammiformibus, serratodentata), ad basin in petiolum longum gracilem (interdum brevem et alatum), aliquando colore rubriviolaceo

tinctum, cuneata vel magis minusve attenuata, omnia in pagina superiore pilis simplicibus eglandulosis plerumque numerosis longis erectis sinuatis albis base pallida, interdum uniformiter distributis sed saepius marginem versus limitatis (extima in margine interdum paucis, saepe satis densis sinuatissimisque) inque pagina inferiore magis minusve uniformiter dispersis vel numerosissimis mollioribus, plerumque base pallida sed raro base nigra, omnia in paginis et margine pilis glanduliferis dispersis brevissimis flavidis vestita, sine pilis stellatis, petiolis in margine pilis simplicibus eglandulosis numerosis, saepe densis, longis vel longissimis sinuatissimis vestitis; folia caulina 1–2(–4), infimum semipatens vel suberectum, lamina 8– 30 × 1–7 mm, linearioblanceolatum vel lineariellipticum vel lineare et bracteiforme, ad apicem plerumque acutissimum (aliquando magis minusve obtusum), integrum vel dentibus paucis acutis praeditum, ad basin in petiolum plerumque brevem attenuatum, pilis simplicibus eglandulosis in pagina superiore dispersis vel numerosis, in margine paginaque inferiore numerosis (saepe base nigra), ad basin densioribus longioribusque (ad 5 mm) vestitum, cetera linearia bracteiformia, plerumque magis minusve appressa. Capitulum solitarium (ad 3 in cultura), 35–50 mm diametro (raro caule secundo a rosula basali exorienti, capitulum ferenti), involucro aspectu obscurissime argentiolivaceogriseo, anguste campanulato, ad basin anguste rotundatam vel anguste subtruncatam contracto; pedunculus (quando adest) ut caulis superus ita vestitus. Involucri squamae ante anthesin incumbentes, nigrivirides; interiores inaequales, quarum longissimae (11–)13–15 ×1.0– 1.6 mm, linearilanceolatae, pleraeque ad apicem acutissimae vel acuminatae, intimae longae, subtiliter acuminatae, exteriores multo breviores, laxae erectae, ad apicem acutae, omnes pilis simplicibus eglandulosis longis vel longissimis (ad 6 mm) gracilibus sinuatissimis implexis (lanuginosis) albis vel pallide argentigriseis (basibus longis incrassatis nigrescentibus), ad basin interdum longioribus (ad 9 mm), et pilis glanduliferis brevissimis brevibusque (paucis ad 0.6 mm) tenuibus pallidiusculis obscurisque (a pilis simplicibus eglandulosis magis minusve omnino celatis) densissime vestitae, sine pilis stellatis sed pilorum brevium ramosorum caespite ad apicem praeditae. Ligulae intensiuscule (vel mediocriter) flavae, interdum dilute viriditinctae, interiores numerosae, exteriores ad 4.0(–4.3) mm latae, leviter vel valde extrorsae, paucae, fere semper discretissimae, ad apicem saepe lobis angustis praeditae, pilis simplicibus eglandulosis in pagina inferiore dispersis vel multis, brevibus vel longiusculis, pallidis vel obscuriusculis, et ultra apicem procurrentibus magis minusve numerosis brevissimis brevibusque et paucis mediocribus vestitae. Styli obscuri, mediocriter vel intensiuscule olivaceogrisei. Receptaculi alveoli non visi. Cypselae (3.7–)4.0–4.5 mm, purpureonigrae. Floret 7–8. Apomicticum. In Section Subalpina Pugsley Hieracium vorlichense P. D. Sell sp. nov. Holotype: Rock above a streamlet (2,000 ft), east side of Ben Vorlich, Dunbarton, v.c. 99, 10 July 1914, E. S. Marshall no. 4008 (CGE).

New taxa and combinations In Hieracio anfractiformi, quo aspectu generali simile est, antea inclusum, foliis hirsutioribus aliquantum aliter dentatis, involucri squamis longioribus angustioribus acutis differt. Herba perennis phyllopoda, caudice ramoso. Caulis 40–50 cm, pallide luteoviridis, infra interdum aliquantum colore brunneorubro tinctus, gracilis, striatus, in parte inferiore pilis simplicibus eglandulosis paucis vel numerosis mediocribus vel longis pallidis, sursum sparsioribus, in parte superiore pilis stellatis numerosis pilisque glanduliferis paucis vel numerosis brevibus obscuris vestitus. Folia in pagina superiore clare mediocriter viridia, in pagina inferiore pallidiora, interdum rubritincta; folia basalia pauca vel numerosa, exteriora lamina 2.0–5.5 × 1.5–2.5 cm, elliptica vel ovata, ad apicem rotundatomucronulata, in dimidio inferiore dentibus mammiformibus dentata, ad basin abrupte contracta, interiora lamina anguste elliptica vel oblonga vel lanceolata vel oblongolanceolata, ad apicem acuta vel longe acuta, in dimidio superiore integra vel denticulata, in dimidio inferiore dentibus mammiformibus vel anguste mammiformibus dentata, ad basin in petiolum longiusculum gracilem longipilosum attenuata, dentibus interdum in petiolum extensis; folia caulina 0–2, inferum quando adest basalibus interioribus simile, breviter petiolatum, superum bracteiforme; omnia in paginis ambabus et in margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis pallidis pilisque glanduliferis sparsis minutis vestita. Inflorescentia capitulis 2–3, furcatocorymbosa; pedunculi suberecti, pilis stellatis numerosis pilisque glanduliferis numerosis inaequalibus brevissimis vel brevibus vel mediocribus gracilibus obscuris pilisque simplicibus eglandulosis sparsis mediocribus pallidis base obscura vestiti. Capitula 35–40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 5–13 × 1.0–1.5 mm, nigrivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem angustum acutum gradatim extensae, interiores fere filamentosae, omnes pilis glanduliferis numerosis inaequalibus brevissimis brevibusque gracilibus obscuris et pilis simplicibus eglandulosis numerosis mediocribus vel longiusculis pallidis base obscura et in margine basinque versus pilis stellatis satis numerosis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis numerosis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 7–8. Apomicticum. Hieracium breacense P. D. Sell sp. nov. Holotype: Meal Breac, Ross-shire, v.c. 105, 28/225755, 17 July 1977, A. G. Kenneth no. 3477 (CGE). Hieracio centripetali proxime affine, capitulis cinerascentibus (non nigris), pilis in involucri squamis stellatis numerosioribus, foliis majoribus et dentibus plerumque majoribus praeditis distinguitur. Herba perennis phyllopoda, caudice ramoso. Caulis 25– 50 cm, pallide luteoviridis, ad basin aliquantum rubellus, gracilis vel robustus, striatus, ubique pilis simplicibus eglandulosis paucis vel satis numerosis mediocribus vel longis pallidis et in parte superiore pilis stellatis numerosis pilisque glanduliferis paucis vel numerosis brevibus

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obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora; folia basalia pauca vel numerosa, lamina 3–10 × 1.0–3.5 cm, elliptica vel ellipticooblonga, raro subrotunda, ad apicem obtusomucronata vel acuta, dentibus mammiformibus vel aquilinomammiformibus regulariter dentata, ad basin cuneata, petiolis ad 7 cm, pilis simplicibus eglandulosis densis longis pallidis vestitis; folium caulinum plerumque 1, linearilanceolatum, longe acutum, acute dentatum, breviter petiolatum (interdum superioribus 1–2 bracteiformibus); omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis vestita. Inflorescentia capitulis 1–11, furcatocorymbosa; pedunculi mediocres vel breves, suberecti, pilis stellatis densis et pilis glanduliferis numerosis brevissimis brevibusque obscuris et pilis simplicibus eglandulosis solitariis pallidis vestiti. Capitula 30– 35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–14 × 1.0–1.2 mm, cinereivirides, interiores margine pallidiore, omnes anguste linearilanceolatae, ad apicem longum gracilem acutum gradatim angustatae, pilis glanduliferis numerosis brevissimis brevibusque mediocribusque gracilibus obscuris et pilis simplicibus eglandulosis numerosis brevibus mediocribusque pallidis base obscura et secus marginem adque apicem pilis stellatis vestitae. Ligulae intense flavae, ad apicem pilis satis numerosis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 7. Apomicticum. Hieracium kingshousense P. D. Sell sp. nov. Holotype: Rocky stream, Allt nan Guibhas, Argyll, v.c. 98, 27/264522, 20 July 1978, P. D. Sell no. 78/134 and A. G. Kenneth (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 30–50 cm, pallide luteoviridis, basin versus saepe purpurascens, gracilis flexuosus striatus, in parte inferiore pilis simplicibus eglandulosis magis minusve numerosis longis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis stellatis paucis vel numerosis et interdum pilis glanduliferis solitariis brevibus obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora; folia basalia lamina 4–12 × 2.5–4.0 cm, elliptica vel late elliptica vel ovatolanceolata, ad apicem acuta vel longe acuminata (raro obtusa), denticulata vel haud profunde dentata (unum folium saepe dentibus longioribus aquilinomammiformibus praeditum), ad basin abrupte contracta vel cuneata, petiolis semper longis, ad 8 cm, longipilosis; folium caulinum 0–1, quando adest lanceolatum, ad apicem longe acuminatum, acute dentatum, ad basin cuneatum, petiolatum; omnia in paginis ambabus et in margine pilis simplicibus eglandulosis magis minusve numerosis brevibus mediocribusque vestita. Inflorescentia capitulis 4– 7, furcatocorymbosa; pedunculi breves vel mediocres, pilis stellatis densis et pilis simplicibus eglandulosis numerosis brevibus mediocribusque pallidis base obscura et pilis glanduliferis brevibus obscuris vestiti. Capitula 20–30 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–11 × 1.2–1.5 mm, nigrivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis

538

New taxa and combinations

vel densis, brevibus vel mediocribus, obscuris vel base obscura, et pilis glanduliferis paucis vel numerosis, brevibus vel brevissimis, obscuris, et secus marginem pilis stellatis numerosis vestitae. Ligulae intense flavae, glabrae vel ad apicem pilis paucis brevissimis vestitae. Styli flavidi, postea obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–8. Apomicticum. Hieracium nigrifactum P. D. Sell sp. nov. Holotype: Cam Creag, Killin, Perthshire, v.c. 88, 25 July 1891, E. S. Marshall no. 237 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 15–40 cm, pallide luteoviridis, ad basin saepe roseotinctus vel purpureotinctus, gracilis, aliquantum flexuosus, striatus, in parte inferiore pilis simplicibus eglandulosis paucis vel satis numerosis, mediocribus vel longis, pallidis, sursum paucis brevioribusque, in parte superiore pilis stellatis numerosis et pilis glanduliferis paucis brevissimis obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora; folia basalia pauca vel numerosa, exteriora lamina 2–7 × 1.5–3.5 cm, subrotunda vel elliptica vel late elliptica, ad apicem obtusomucronata, subintegra vel denticulata vel haud profunde dentata, dentibus mamillatis, ad basin cuneata vel subtruncata, interiora lamina 3–8(–10) × 1–3 cm, lanceolata vel anguste elliptica, ad apicem magis minusve acuta, dentata praesertim in dimidio inferiore, dentibus mammiformibus vel aquilinomammiformibus, ad basin saepe asymmetricam cuneata vel abrupte contracta, petiolis mediocribus vel longis, pilis simplicibus eglandulosis densis longis pallidis vestitis; folium caulinum 0–1, quando adest lanceolatum vel raro lineare, ad apicem longe acutum, ad basin plerumque acute dentatum, dentibus apice longe cuspidato aquilinomammiformibus, sessile vel petiolatum; omnia in paginis ambabus et in margine pilis simplicibus eglandulosis brevibus vel mediocribus pallidis et pilis glanduliferis paucis brevissimis flavidis vestita. Inflorescentia capitulis 1–6, furcatocorymbosa; pedunculi breves vel mediocres vel longi, pilis stellatis magis minusve densis et pilis glanduliferis numerosis brevissimis brevibusque obscuris et rarissime pilis simplicibus eglandulosis mediocribus vestiti. Capitula 25–30 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 11–13 × 1.2–1.5 mm, nigrivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem obtusae, pilis glanduliferis numerosis vel densis, brevissimis vel brevibus vel mediocribus, obscuris, et secus marginem pilis stellatis paucis et rarissime pilis simplicibus eglandulosis solitariis vestitae. Ligulae intense flavae, ad apicem pilis paucis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 6–8. Apomicticum. Hieracium atriglandulosum P. D. Sell sp. nov. Holotype: Ben More, 27/4425, Perthshire, v.c. 88, 1980, A. G. Kenneth no. 2780 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 25–40 cm, pallide luteoviridis, in parte inferiore saepe rubellus vel purpurascens, aliquantum gracilis flexuosusque, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis vel

densis longis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis stellatis magis minusve densis et pilis glanduliferis paucis vel numerosis brevibus obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora; folia basalia lamina 4–9 × 2– 4 cm, anguste vel late elliptica (interdum ovata), ad apicem exteriora obtusa, interiora acuta, subintegra vel denticulata vel nequaquam profunde dentata, dentibus mamillatis, ad basin magis minusve cuneata, petiolis plerumque longis, pilis simplicibus eglandulosis densis longis pallidis vestitis; folium caulinum 0–1, quando adest lanceolatum, ad apicem acutum, breviter dentatum, ad basin cuneatum, petiolatum; omnia in paginis ambabus et in margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis vestita. Inflorescentia capitulis 2–6 (ad 9 in cultura), furcatocorymbosa; pedunculi longi, suberecti, pilis stellatis numerosis vel densis et pilis glanduliferis numerosis brevissimis brevibusque mediocribusque obscuris et raro pilis simplicibus eglandulosis solitariis vestiti. Capitula 30–40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 5–14 × 1.0–1.4 mm, nigrivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem magis minusve acutae, pilis glanduliferis numerosis vel densis brevissimis brevibusque mediocribusque gracilibus obscuris et pilis simplicibus eglandulosis solitariis (interdum absentibus) et in margine adque apicem pilis stellatis paucis vestitae. Ligulae intense flavae, ad apicem pilis simplicibus eglandulosis numerosis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–8. Apomicticum. Hieracium ochthophilum P. D. Sell sp. nov. Holotype: Rocky bank, Allt nan Giubhas, near Kingshouse, Argyll, v.c. 98, 27/263520, 1 August 1969, C. West (CGE). Hieracio westii (H. submurorum auct.), quocum invenitur, confusum, foliorum dentibus paucioribus obtusioribus minus profundis, capitulis latioribus minus glandulopilosis distinguitur. Herba perennis phyllopoda, caudice ramoso. Caulis 24–37 cm, pallide luteoviridis, ad basin interdum aliquantum purpurascens, gracilis flexuosus striatus, pilis simplicibus eglandulosis paucis dispersis brevibus vel mediocribus pallidis sed ad basin obscuris et in parte superiore pilis stellatis paucis vel numerosis vestitus. Folia in pagina superiore pallide cinereiviridia, in pagina inferiore pallidiora; folia basalia plerumque pauca, lamina 2–6 × 1.5–3.0 cm, elliptica vel lanceolata vel ovata vel subrotunda, ad apicem obtusomucronata vel magis minusve acuta, subintegra vel saepius in dimidio inferiore dentibus haud profundis patentibus praedita, ad basin in petiolum mediocrem gracilem glabrum vel satis hirsutum cuneata; folium caulinum 0–1, quando adest lamina lanceolata, ad apicem acutum, in dimidio inferiore dentatum, petiolatum; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis paucis vel numerosis mediocribus vel longis pallidis vestita, pilis glanduliferis dispersis minutis flavidis praedita. Inflorescentia capitulis 2–4, furcatocorymbosa; pedunculi longiusculi suberecti, pilis stellatis numerosis vel densis pilisque glanduliferis paucis vel numerosis brevibus obscuris pilisque simplicibus

New taxa and combinations eglandulosis paucis brevibus vel mediocribus pallidis sed ad basin obscuris vestiti. Capitula 30–35 mm diametro, ad basin late rotundata vel subtruncata. Involucri squamae ante anthesin incumbentes, 6–14 × 1.5–2.0 mm, nigrivirides, late linearilanceolatae, ad apicem obtusum gradatim angustatae, pilis simplicibus eglandulosis numerosis vel satis densis brevibus vel mediocribus pallidis sed ad basin obscuris et pilis glanduliferis paucis vel satis numerosis brevibus obscuris et ad basin apicemque pilis stellatis nonnullis vestitae. Ligulae aureoflavae, ad apicem glabrae vel fere glabrae. Styli obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 7–8. Apomicticum. Hieracium raveniorum P. D. Sell sp. nov. Holotype: North-east-facing cliffs below summit of Beinn na Cille, Morven, Argyll, v.c. 97, 17/854544, 2 July 1970, P. D. Sell no. 70/626a and N. Spring (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20– 35 cm, pallide luteoviridis, saepe colore brunneopurpureo suffusus, gracilis sed rigidus, flexuosus, striatus, pilis simplicibus eglandulosis prope basin numerosis longis pallidis (sursum paucioribus brevioribus, base obscura) et in parte superiore pilis stellatis paucis vel numerosis et interdum pilis glanduliferis solitariis brevibus vestitus. Folia in pagina superiore obscure viridia, in pagina inferiore pallidiora, interdum rubritincta; folia basalia numerosa, lamina 2–7 × 1.5–2.5 cm, elliptica vel oblanceolata vel obovata vel lanceolata, ad apicem plerumque omnia obtusa (raro magis minusve acuta), dentibus parvis acutisque regulariter sinuatoserrulata, ad basin in petiolum brevem vel mediocrem dense longipilosum cuneata vel attenuata; folia caulina 0– 2, quando adsunt basalibus similia sed minora petiolataque aut bracteiformia; omnia in paginis ambabus et in margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis pallidis (in pagina superiore rigidiusculis) pilisque glanduliferis dispersis minutis flavidis vestita. Inflorescentia capitulis 1–3, furcatocorymbosa; pedunculi breviusculi, suberecti, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis paucioribus brevibus obscuris vestiti. Capitula 30–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–10 × 0.8–1.2 mm, obscure griseivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem subobtusae vel magis minusve acutae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis pallidis base obscura et pilis glanduliferis paucis vel satis numerosis brevibus obscuris et ad apicem pilorum caespite parvo vestitae. Ligulae flavae, glabrae. Styli flavidi. Receptaculi alveoli margine breviter dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 6–8. Apomicticum. In memoriam Caroli Earle Raven (1885–1964) et Joannis Earle Raven (1914–80) nominatur. Hieracium lakelandicum P. D. Sell sp. nov. Holotype: Keppel Cove, Helvellyn, Westmorland, v.c. 69, 7 July 1954, J. E. Raven (CGE).

539

Herba perennis phyllopoda, caudice ramoso. Caulis 15–35 cm, pallide luteoviridis, basin versus saepe brunneopurpureus, gracilis flexuosus striatus, pilis simplicibus eglandulosis prope basin numerosis longis pallidis (interdum sursum dispersis) et pilis stellatis paucis et in parte superiore pilis glanduliferis sparsis brevibus obscuris vestitus. Folia in pagina superiore caeruleoviridia, saepe colore brunneopurpureo guttata et maculata, in pagina inferiore pallidiora, interdum rubritincta; folia basalia saepe numerosa, lamina 1–6 × 1–3 cm, subrotunda vel late elliptica vel lanceolata vel ovata, ad apicem saepe omnia obtusa (interdum 1–2 magis minusve acuta), dentibus parvis denticulata vel dentata, ad basin in petiolum brevem longipilosum rotundata vel subtruncata vel abrupte contracta; folium caulinum 0–1, quando adest lamina lanceolata, ad apicem acutum vel acuminatum, dentatum, petiolatum; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore (praesertim in costa inque margine) pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis pallidis vestita, pilis glanduliferis paucis dispersis minutis flavidis praedita. Inflorescentia capitulis 1–4, furcatocorymbosa; pedunculi breviusculi, suberecti, pilis stellatis densis pilisque glanduliferis numerosis inaequalibus brevissimis brevibusque mediocribusque obscuris vestiti. Capitula 30– 40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–12 × 1.0–1.3 mm, obscure virides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem magis minusve acutum gradatim extensae, pilis glanduliferis densis brevissimis vel brevibus vel mediocribus obscuris et in margine et base pilis stellatis numerosis vestitae, sine pilis simplicibus eglandulosis. Ligulae flavae, glabrae. Styli flavi. Receptaculi alveoli margine subulatodentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 7–8. Apomicticum. Hieracium aristidens P. D. Sell sp. nov. Holotype: Carn Dearg, Sutherland, v.c. 107, 29 July 1975, A. G. Kenneth no. 8975. Herba perennis phyllopoda, caudice ramoso. Caulis 20– 40 cm, pallide viridis, in parte inferiore saepe purpurascens, flexuosus striatus, in parte inferiore pilis simplicibus eglandulosis numerosis longis sinuatis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis stellatis paucis vel numerosis et pilis glanduliferis paucis brevissimis obscuris vestitus. Folia in pagina superiore mediocriter viridia, in pagina inferiore pallidiora; folia basalia satis pauca, exteriora lamina 1.5–4.5 × 1.0–2.5 cm, elliptica, ad apicem rotundatomucronulata, sparsim denticulata, ad basin angustata vel rotundata, interiora lamina 6– 8 × 2–3 cm, anguste elliptica vel elliptica vel lanceolata, ad apicem acuta, dentibus parvis acute spinulosis serratodentata, ad basin cuneata, petiolis ad 5 cm, pilis simplicibus eglandulosis numerosis longis pallidis vestitis; folium caulinum 0–1, quando adest lamina lanceolata, ad apicem acutum, acute dentatum, ad basin angustatum, semiamplexicaule, aut bracteiforme; omnia pilis simplicibus eglandulosis in paginis ambabus inque margine paucis vel numerosis, brevibus vel mediocribus, et pilis glanduliferis nonnullis minutis vestita. Inflorescentia capitulis 1–5, interdum ramo inferiore longo praedita; pedunculi longi, graciles, pilis

540

New taxa and combinations

stellatis densis pilisque glanduliferis numerosis brevissimis obscuris pilisque simplicibus eglandulosis paucis longioribus pallidis vestiti. Capitula 30–40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–15 × 1.0–1.3 mm, nigrivirides, interiores margine angusto pallido, omnes linearilanceolatae, ad apicem magis minusve acutum angustatae, pilis simplicibus eglandulosis numerosis vel densis brevibus vel mediocribus, base obscura, et pilis glanduliferis paucis vel satis numerosis, sed non manifestis, brevibus obscuris et basin versus pilis stellatis paucis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis numerosis brevibus vestitae. Styli obscuri. Receptaculi alveoli margine acute dentato. Cypselae 2.5–3.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium oenophyllum P. D. Sell sp. nov. Holotype: Creag an Sturra, Kilmelford Glen, Argyll, v.c. 98, 7 June 1967, A. G. Kenneth (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20–35 cm, pallide luteoviridis, in parte inferiore plerumque valde vinicolor, gracilis vel satis robustus, striatus, in parte inferiore pilis simplicibus eglandulosis paucis vel numerosis mediocribus vel longis pallidis (sursum sparsis vel nullis), in parte superiore pilis stellatis paucis vel numerosis vestitus. Folia in pagina superiore mediocriter luteoviridia vel aliquantum caeruleoviridia (interdum colore brunneopurpureo suffusa vel maculis paucis notata), in pagina inferiore plerumque saturate purpureosuffusa; folia basalia pauca vel satis numerosa, lamina 1.5–6.0 × 1–3 cm, subrotunda vel elliptica vel ellipticooblonga vel lanceolata, ad apicem plerumque rotundatoobtuse mucronata (interdum interiorum aliquot magis minusve acuta), subintegra vel denticulata vel dentibus paucis haud profundis ad basin praedita, ad basin rotundata vel abrupte contracta vel breviter cuneata, petiolo brevi purpurascenti longipiloso praedita; folium caulinum 0–1, quando adest lamina lanceolata, ad apicem acutum, integrum vel denticulatum, ad basin cuneatum, breviter petiolatum (aut interdum bracteiforme); omnia in pagina superiore glabra vel pilis simplicibus eglandulosis paucis vel numerosis brevibus vel mediocribus pallidis vestita, in pagina inferiore et in margine et praesertim in costa pilis simplicibus eglandulosis numerosis mediocribus vel longis pallidis vestita. Inflorescentia capitulis 2–7, furcatocorymbosa, interdum ramo inferiore praedita; pedunculi pilis stellatis magis minusve densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis pilisque glanduliferis paucis brevibus obscuris vestiti. Capitula 30–40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin porrectae vel leviter incumbentes, 5–12 × 1.0–1.3 mm, obscure cinereivirides, interiores margine pallidiore, omnes anguste linearilanceolatae, ad apicem acutum gradatim angustatae, interiores angustissimae, omnes pilis stellatis numerosis (in margine saepe densis) pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis paucis brevibus obscuris vestitae. Ligulae mediocriter vel saturate flavae, ad apicem pilis simplicibus eglandulosis numerosis brevissimis vestitae. Styli flavi vel aliquantum obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 5–7. Apomicticum.

Hieracium sannoxense P. D. Sell sp. nov. Holotype: Sannox Bridge, Arran, v.c. 100, 26/015515, 1 June 1974, A. G. Kenneth no. 25/74 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20– 55 cm, pallide viridis, basin versus aliquantum purpurascens, gracilis striatus, in parte inferiore pilis simplicibus eglandulosis numerosis longis sinuatis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis glanduliferis numerosis minutis pilisque stellatis vestitus. Folia in pagina superiore mediocriter viridia, in pagina inferiore pallidiora; folia basalia numerosa, omnia similia, lamina 3.0–6.5 × 1.5–3.5 cm, ovata vel elliptica, ad apicem rotundatomucronulata vel subacuta, denticulata vel dentibus paucis angustis acutis praeditum, ad basin abrupte contracta vel subtruncata, petiolis ad 4 cm, pilis simplicibus eglandulosis densis longis pallidis vestitis; folium caulinum 0–1, quando adest parvum lanceolatum, ad apicem acutum, semiamplexicaule; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis mediocribus vel longis pallidis vestita. Inflorescentia capitulis 1–15, interdum ramo inferiore longo praedita; pedunculi longi graciles, pilis stellatis numerosis vel densis et pilis glanduliferis numerosis inaequalibus brevissimis brevibusque obscuris et pilis simplicibus eglandulosis paucis brevibus mediocribusque pallidis vestiti. Capitula 30–40 mm diametro, ad basin rotundata vel subtruncata. Involucri squamae ante anthesin incumbentes, 5–13 × 1.0–1.2 mm, nigrivirides, interiores margine pallidiore, omnes anguste linearilanceolatae, ad apicem acutae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis satis numerosis brevibus obscuris et secus marginem adque basin pilis stellatis paucis vestitae. Ligulae ad apicem pilis simplicibus eglandulosis numerosis brevibus vestitae. Styli obscuri. Receptaculi alveoli margine acute dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 5– 6. Apomicticum. Hieracium fissuricola P. D. Sell sp. nov. Holotype: Fisher’s Ghyll gully, Helvellyn, Cumberland, v.c. 70, 8 July 1954, J. E. Raven (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20–40 cm, pallide luteoviridis, in parte inferiore saepe purpurascens, gracilis striatus, in parte inferiore pilis simplicibus eglandulosis paucis vel numerosis longis pallidis (sursum brevioribus paucioribusque vel nullis), in parte superiore pilis stellatis densis vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora, interdum aliquantum purpureotincta; folia basalia magis minusve numerosa, lamina 2–10 × 1–4 cm, elliptica vel lanceolata vel oblongolanceolata vel oblongoelliptica vel subrotunda, ad apicem pleraque obtusa, sed alia late rotundata, alia angustata raroque acuta, integra vel remote denticulata, ad basin rotundata vel cuneata, petiolis ad 6 cm, pilis simplicibus eglandulosis numerosis longis flexuosis pallidis vestitis; folium caulinum 0–1, quando adest lamina anguste lineari, integrum, sessile; omnia in pagina superiore glabra vel pilis simplicibus eglandulosis dispersis brevibus rigidis vestita, in pagina inferiore et in margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus

New taxa and combinations rigidis, in pagina inferiore pilis stellatis dispersis vestita. Inflorescentia capitulis 1–11, furcatocorymbosa, interdum ramo inferiore longo praedita; pedunculi aliquantum breves rectique, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis pilisque glanduliferis paucis vel numerosis brevibus obscuris vestiti. Capitula 25–30 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 5–12 × 1.0– 1.2 mm, obscure olivaceae, interiores margine pallidiore, omnes anguste linearilanceolatae, ad apicem obtusum gradatim angustatae, pilis simplicibus eglandulosis numerosis brevibus mediocribusque base obscura et pilis glanduliferis paucis vel satis numerosis brevibus obscuris et basin versus pilis stellatis nonnullis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis brevissimis vestitae. Styli flavidi vel obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 6– 7. Apomicticum. Hieracium ovaliforme P. D. Sell sp. nov. Holotype: Meallan Liath, W. Sutherland, v.c. 108, 29/ 355388, 1974, A. G. Kenneth no. 124/74 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 15– 25 cm, pallide viridis, basin versus interdum purpurascens, gracilis, in parte inferiore pilis simplicibus eglandulosis longis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis stellatis pilisque glanduliferis minutis paucis vestitus. Folia in pagina superiore pallidiuscule viridia, interdum maculis dilutis, in pagina inferiore pallidiora; folia basalia numerosa, lamina 1.5–5.0 × 1.0–3.0 cm, subrotunda vel elliptica, ad apicem rotundata (raro subacuta), integra vel remote denticulata, ad basin rotundata vel abrupte contracta, petiolis ad 2 cm, pilis simplicibus eglandulosis numerosis longis flexuosis pallidis vestitis; folium caulinum 0–1, quando adest lamina parva, linearilanceolatum vel lanceolatum, integrum acutum semiamplexicaule, aut bracteiforme; omnia in pagina superiore pilis simplicibus eglandulosis paucis vestita vel glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis mediocribus vel longis pallidis vestita. Inflorescentia capitulis 1–2; pedunculi graciles, pilis stellatis densis pilisque glanduliferis paucis vel numerosis brevibus obscuris pilisque simplicibus eglandulosis sparsis vestiti. Capitula 35– 40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 5–12 × 1.2–1.5 mm, obscure griseivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem angustatae sed obtusae, pilis simplicibus eglandulosis numerosis mediocribus base obscura et pilis glanduliferis paucis vel satis numerosis brevibus obscuris et secus marginem pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis brevibus vestitae. Styli obscuri. Receptaculi alveoli margine acute dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium arranense P. D. Sell sp. nov. Holotype: Near King’s Cave, Arran, Clyde Islands, v.c. 100, 16/885310, 2 June 1974, A. G. Kenneth no. 3374 and A. McG. Stirling (CGE). Hieracio petrocharidi proxime affine, a quo foliis latioribus, plerumque magis ovatis, petiolis breviusculis, involucri squamarum indumento breviore differt.

541

Herba perennis phyllopoda, caudice ramoso. Caulis 10– 40 cm, pallide luteoviridis, ad basin interdum aliquantum rubellus, gracilis vel aliquantum robustus, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis longis pallidis (sursum paucis vel nullis), in parte superiore pilis stellatis paucis vel numerosis vestitus. Folia in pagina superiore luteoviridia, colore brunneopurpureo maculata et guttata, in pagina inferiore pallidiora, interdum colore violaceopurpureo suffusa; folia basalia lamina 1.5–5.5 × 1– 3 cm, plerumque ovata, interdum elliptica vel lanceolata, ad apicem pleraque obtusomucronata, sed interiora interdum magis minusve acuta, integra vel minute denticulata, ad basin in petiolum breviusculum, saepe rubellum, dense longipilosum magis minusve rotundata; folia caulina nulla aut linearia, bracteiformia; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis mediocribus vel longis pallidis, in margine pilis stellatis paucis minutis vestita. Inflorescentia capitulis 1–5, furcatocorymbosa; pedunculi pilis stellatis numerosis vel densis, interdum pilis simplicibus eglandulosis paucis brevibus vel mediocribus pallidis, raro pilis glanduliferis paucis brevibus obscuris vestiti. Capitula 30–40 mm diametro, ad basin late rotundata. Involucri squamae ante anthesin magis minusve porrectae, 4–12 × 1.2–1.5 mm, obscure cinereivirides, interiores margine pallidiore, omnes linearilanceolatae (interiores angustissimae), ad apicem magis minusve acutae, pilis stellatis ubique (sed praesertim in margine) numerosis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis pilisque glanduliferis paucis brevibus obscuris vestitae. Ligulae intense flavae, glabrae. Styli obscuri. Receptaculi alveoli margine fimbriatodentato. Cypselae c. 3.0 mm, rubrinigrae. Floret 5–6. Apomicticum. Hieracium thalassinum P. D. Sell sp. nov. Holotype: Cliff ledges by the sea, Ballygroggan, Kintyre, v.c. 101, 16/624197, 28 June 1968, P. D. Sell no. 68/369 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 25–50 cm, pallide luteoviridis, basin versus saepe purpurascentisuffusus, in parte inferiore pilis simplicibus eglandulosis numerosis longis pallidis, sursum brevioribus, et in parte superiore pilis stellatis numerosis vestitus. Folia in pagina superiore mediocriter viridia, in pagina inferiore pallidiora, interdum rubellitincta vel purpurascentitincta; folia basalia pauca, lamina 1.5–8.0(–10) × 1.0– 3.5(–5) cm, ovata vel obovata vel elliptica, ad apicem obtusomucronata vel acuta, subintegra vel sparse denticulata vel dentibus sparsis praedita, ad basin rotundata vel cuneata vel attenuata, petiolis ad 7 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folia caulina 1–3, infera basalibus similia, petiolo alato praedita, semiamplexicaulia, supera angustiora, magis minusve sessilia; omnia in paginis ambabus inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis pallidis vestita. Inflorescentia capitulis 2–6, furcatocorymbosa, interdum ramo inferiore praedita; pedunculi aliquantum longi gracilesque, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura vestiti. Capitula 25–35 mm diametro, ad basin rotundata. Involucri squamae ante

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New taxa and combinations

anthesin incumbentes, 4–12 × 1.0–1.2 mm, obscure virides, interiores ad marginem pallidiores, omnes oblongolanceolatae, ad apicem acutae, pilis simplicibus eglandulosis densis brevibus vel mediocribus vel longis pallidis base obscura et in margine pilis stellatis nonnullis et pilis glanduliferis sparsissimis brevibus obscuris vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis numerosis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine incisodentato. Cypselae 2.5–3.0 mm, rubribrunneae. Floret 6–7. Apomicticum. Hieracium hyparcticoides Pugsley ex P. D. Sell sp. nov. H. hyparcticoides Pugsley, Jour. Bot. (London) 79: 179 (1941) sine diagn. lat. Holotype: On limestone, by the Traligill and Allt-na-Glaise Moire; scarce; near Inchnadamph, W. Sutherland, v.c. 108, 15 July 1908, E. S. Marshall no. 3267 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20–50 cm, pallide luteoviridis, in parte inferiore aliquando rubellus, gracilis vel robustus, striatus, pilis simplicibus eglandulosis dispersis mediocribus vel longis pallidis, interdum base obscura, et in parte superiore pilis stellatis paucis glanduliferisque paucis brevibus obscuris vestitus. Folia in pagina superiore luteoviridia, in pagina inferiore pallidiora; folia basalia pauca, exteriora lamina 2–7 × 1.5–3.0 cm, obovata vel oblongoobovata vel elliptica, ad apicem obtusomucronata, integra vel denticulata vel raro haud profunde dentata, ad basin in petiolum longiusculum dense longipilosum abrupte contracta vel cuneata, interiora lamina anguste elliptica vel lanceolata, ad apicem magis minusve acuta, plerumque denticulata, ad basin in petiolum satis longum cuneata; folia caulina 0–2, infera basalibus interioribus similia petiolataque, cetera linearia bracteiformiaque; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis paucis vel numerosis brevibus vel mediocribus pallidis et in pagina inferiore interdumque in pagina superiore pilis stellatis numerosis vel densis, cum pilis glanduliferis paucis minutis flavidis, vestita. Inflorescentia capitulis 2–9, furcatocorymbosa; pedunculi longi suberecti, pilis stellatis numerosis vel densis et pilis gladuliferis numerosis brevibus obscuris et aliquando pilis simplicibus eglandulosis paucis brevibus vel mediocribus pallidis, base obscura, vestiti. Capitula 30–35 mm diametro, ad basin angustata. Involucri squamae ante anthesin porrectae, 5–13 × 1.0–1.5 mm, nigrivirides, linearilanceolatae, ad apicem obtusum gradatim angustatae, pilis glanduliferis numerosis inaequalibus brevissimis brevibusque obscuris et pilis simplicibus eglandulosis paucis mediocribus nigriusculis et praecipue in margine adque apicem pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis numerosis brevissimis vestitae. Styli flavidae vel obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.5– 4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. In Section Cerinthoidea Monnier Hieracium oxybeles P. D. Sell nom. et stat. nov. H. cerinthoides var. acutifolium Backh. fil., Hier. Mon. 37 (1856), non H. acutifolium Vill., Pr´ec. Voy. Bot. 59 (1812).

Hieracium cyclicum P. D. Sell sp. nov. Holotype: About 2,000 ft on east-facing micaceous cliffs above Loch an Fhuarain, Ben Klibreck, Sutherland, v.c. 108, 29/579279, 19 July 1966, R. W. Jones and B. A. Miles no. 66/24 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20– 50 cm, pallide luteoviridis, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis longis pallidis, sursum paucioribus brevioribusque, et in parte superiore pilis stellatis paucis vel numerosis pilisque glanduliferis paucis brevibus obscuris vestitus. Folia in pagina superiore pallidiuscule caeruleoviridia, in pagina inferiore pallidiora; folia basalia numerosa, exteriora lamina 2–8 × 2–5 cm, subrotunda vel late ovata, ad apicem rotundatomucronata, in dimidio superiore subintegra vel denticulata, infra dentata, saepe dentibus mammiformibus vel aquilinomammiformibus, ad basin subtruncata vel late cuneata, interiora lamina 5–10 × 3.0–5.5 cm, lanceolata vel ovata vel elliptica, ad apicem plerumque acuta, in dimidio superiore integra vel denticulata, infra plerumque dentata, dentibus angustis, ad basin late cuneata vel subtruncata, petiolis ad 5 cm, pilis simplicibus eglandulosis densis longis pallidis vestitis; folia caulina 1–2, inferum basalibus interioribus simile, petiolo semiamplexicauli praeditum, superum bracteiforme; omnia in paginis ambabus pilis simplicibus eglandulosis paucis mediocribus pallidis vestita. Inflorescentia capitulis 1–9, furcatocorymbosa; pedunculi longi, pilis stellatis densis pilisque glanduliferis numerosis brevibus obscuris pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura vestiti. Capitula 30–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin porrecti, 5–13 × 1.0–1.2 mm, nigrivirides, interiores margine angusto pallidiore, omnes linearilanceolatae, ad apicem magis minusve acutum gradatim angustatae, pilis simplicibus eglandulosis numerosis mediocribus pallidis base obscura et pilis glanduliferis paucis vel satis numerosis brevibus obscuris et ad basin pilis stellatis paucis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis numerosis brevibus vestitae. Styli flavi vel aliquantum obscuri. Receptaculi alveoli margine fimbriatodentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium ampliatiforme P. D. Sell nom. et stat. nov. H. ampliatum var. gracilius Pugsley in Jour. Linn. Soc. London (Bot.) 54: 75 (1948).

Hieracium flocculosiforme P. D. Sell nom. et stat. nov. H. flocculosum var. denticulatum Pugsley in Jour. Linn. Soc. London (Bot.) 54: 78 (1948), non H. denticulatum Sm., Eng. Bot. t. 2122 (1810).

Hieracium flocculipubens P. D. Sell nom. et stat. nov. H. flocculosum var. angustifolium Pugsley in Jour. Linn. Soc. London (Bot.) 54: 78 (1948), non H. angustifolium Hoppe, Bot. Taschenb. 1799: 129 (1799).

New taxa and combinations In Section Oreadea (Fr.) Dahlst. Hieracium subscoticum P. D. Sell sp. nov. Holotype: Crags at seashore, north-east side of Aith Hope, South Waas, Hoy, Orkney, v.c. 111, 21 July 1913, H. H. Johnston (CGE). Hujus speciei plantae Hieracium scoticum antea nominatae sunt, sed ab illa specie foliis angustioribus, ad basin attenuatis, differunt. Herba perennis phyllopoda, caudice ramoso. Caulis 25–60 cm, pallide luteoviridis, in parte inferiore saepe rubellitinctus, gracilis vel magis minusve robustus, in parte inferiore pilis simplicibus eglandulosis numerosis mediocribus vel longis sinuatis pallidis (interdum ubique, interdum sursum paucioribus brevioribusque), in parte superiore pilis stellatis magis minusve numerosis vestitus. Folia in pagina superiore obscure viridia vel glauciuscula, in pagina inferiore pallidiora; folia basalia pauca, sub anthesi saepe siccata, lamina 5–10 × 1–3 cm, elliptica vel oblongoelliptica vel lanceolatoelliptica, ad apicem obtusum vel acutum angustata, subintegra vel denticulata, ad basin attenuata, petiolis ad 5 cm, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folia caulina 2–6, sursum gradatim magnitudine decrescentia, infera lamina lanceolata vel elliptica vel oblongoelliptica vel late elliptica, plerumque magis minusve acuta, dentibus parvis acutis dentata, ad basin attenuata, petiolo alato ad 6 cm longo praedita, supera lamina lanceolata vel ovata vel elliptica, ad apicem longe acuta, dentibus parvis acutis praedita, ad basin sessilem rotundata vel cuneata; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis subrigidis pallidis vestita. Inflorescentia capitulis 1–5(–7), furcatocorymbosa; pedunculi longi recti, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis paucis brevissimis obscuris vestiti. Capitula 30–40 mm diametro, ad basin subtruncata. Involucri squamae ante anthesin incumbentes, 5–13 × 1.2– 1.7 mm, obscure virides, interiores ad marginem pallidiores, omnes linearilanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis brevibus mediocribusque pallidis base obscura et pilis glanduliferis paucis brevissimis obscuris et basin versus pilis stellatis paucis vestitae. Ligulae citrinae, ad apicem glabrae. Styli triste fulvi. Receptaculi alveoli margine breviter dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium apheles P. D. Sell nom. et stat. nov. H. septentrionale var. simplex Ley in Jour. Bot. (London) 47: 50 (1909), non H. simplex Viv., Fl. Lybicae Spec. 50 (1824). Hieracium sowadeense P. D. Sell sp. nov. Holotype: Grassy, heathery, rocky crags at burnside, Burn of Sowa Dee, Sandwick, Mainland, Orkney, v.c. 111, 23 July 1925, H. H. Johnston no. 3107. Herba perennis phyllopoda, caudice ramoso. Caulis 20– 50 cm, pallide luteoviridis, plerumque colore brunneopurpureo multum suffusus, gracilis flexuosusque aut robustus, striatus, in parte inferiore pilis simplicibus eglandu-

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losis numerosis longis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis stellatis densis vestitus. Folia in pagina superiore mediocriter viridia, in pagina inferiore pallidiora, saepe purpurascentia; folia basalia pauca, sub anthesi interdum siccata, lamina 2–8 × 1.5–4.0 cm, elliptica vel lanceolata vel ovata, ad apicem obtusa vel acuta, integra vel remote denticulata, ad basin cuneata vel attenuata, petiolis ad 3 cm, pilis simplicibus eglandulosis densis longis sinuatis vestitis; folia caulina 3–6, infera basalibus similia, sessilia vel breviter petiolata, cetera sursum gradatim minora, supera lamina lineari vel bracteiformia; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis subrigidis pallidis vestita. Inflorescentia capitulis 1–6, furcatocorymbosa; pedunculi longi recti, pilis stellatis numerosis pilisque simplicibus eglandulosis satis numerosis brevibus vel mediocribus vel longis pallidis vestiti. Capitula 30–40 mm diametro, ad basin subtruncata. Involucri squamae ante anthesin incumbentes, 5–13 × 1.5–2.0 mm, obscure olivaceae, interiores ad marginem pallidiores, omnes lanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis et secus marginem pilis stellatis paucis dispersis vestitae. Ligulae flavae, ad apicem glabrae. Styli flavi. Receptaculi alveoli margine breviter dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium pusillifolium P. D. Sell nom. et stat. nov. H. scoticum var. microphyllum Pugsley in Jour. Linn. Soc. London (Bot.) 54: 99 (1948), non H. microphyllum Hyl. in Sym. Bot. Upsal. 7(1): 179 (1941). Hieracium aggregatifolium P. D. Sell nom. et stat. nov. H. scoticum var. rosulare Pugsley in Jour. Linn. Soc. London (Bot.) 54: 99 (1948), non H. rosulare Brenner in Acta Soc. Fauna Fl. Fenn. 9(5): 20 (1893). Hieracium breconicola P. D. Sell sp. nov. Holotype: Sandstone rocks, Fan Nedd, Brecon Beacons, 7 August 1906, A. Ley (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 35– 55 cm, pallide luteoviridis, infra interdum rubellus, gracilis striatus, in parte inferiore pilis simplicibus eglandulosis numerosis mediocribus vel longis pallidis, sursum paucioribus vel interdum nullis, et in parte superiore pilis stellatis numerosis vestitus, sine pilis glanduliferis. Folia in pagina superiore mediocriter viridia, in pagina inferiore pallidiora, saepe rubripurpureosuffusa, subcoriacea; folia basalia pauca vel satis numerosa, lamina 3–10 × 2–4 cm, elliptica vel lanceolata vel ovata vel oblongoelliptica, ad apicem angustata sed obtusomucronata, integra vel remote denticulata (aliquando dentibus sparsis praedita), ad basin cuneata vel attenuata, petiolis ad 4 cm, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folia caulina 2–5, infera basalibus interioribus similia, longipetiolata, supera lamina ovata, acuta, denticulata, sessilia; omnia in pagina superiore inque margine pilis simplicibus eglandulosis brevibus vel mediocribus rigidis pallidis vestita (aut in pagina superiore glabra vel fere glabra) et in

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New taxa and combinations

pagina inferiore (praesertim in costa) pilis numerosioribus longioribus vestita. Inflorescentia capitulis 1–7, furcatocorymbosa, interdum ramo inferiore longo praedita; pedunculi longi rectique, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus mediocribusque pallidis pilisque glanduliferis paucis minutis obscuris vestiti. Capitula 30–40 mm diametro, ad basin subtruncata. Involucri squamae ante anthesin porrectae, 3–12 × 1.0–1.5 mm, nigrivirides, interiores ad marginem pallidiores, omnes late linearilanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis brevibus mediocribusque pallidis base obscura et pilis glanduliferis paucissimis brevibus obscuris et in margine pilis stellatis paucis vestitae. Ligulae flavae, ad apicem glabrae. Styli flavi. Receptaculi alveoli margine subulatodentato. Cypselae 4.0–4.5 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium scottii P. D. Sell sp. nov. Holotype: Rocky geos below Windy Scord, Voe of Snarraness, Walls and Sandness, Zetland, v.c. 112, 41/2256, 5 August 1964, W. Scott (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 15–30 cm, pallide luteoviridis, ad basin saepe purpurascens, gracilis flexuosusque, striatus, pilis simplicibus eglandulosis numerosis mediocribus vel longis sinuatis pallidis, sursum paucioribus brevioribusque, et in parte superiore pilis stellatis densis pilisque glanduliferis paucis minutis obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora, interdum rubellitincta; folia basalia pauca, lamina 4–9 × 1.5–3.5 cm, lanceolata vel elliptica, rotundatomucronulata vel acuta, remote denticulata, ad basin cuneata vel attenuata, petiolis ad 4 cm, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folia caulina 2–4, sursum magnitudine abrupte decrescentia, infera basalibus similia petiolataque, supera lamina lanceolata vel lineari, ad apicem acuta, denticulata vel integra, sessilia; omnia in paginis ambabus inque margine pilis simplicibus eglandulosis mediocribus vel longis subrigidis pallidis et in pagina inferiore pilis stellatis paucis vestita. Inflorescentia capitulis 1–8, furcata; pedunculi erecti, recti, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis pilisque glanduliferis numerosis minutis obscuris vestiti. Capitula 30–40 (ad 50 in cultura) mm diametro, ad basin late rotundata vel subtruncata. Involucri squamae ante anthesin incumbentes, 5–14 × 1.5–2.0 mm, obscure virides, interiores ad marginem pallidiores, omnes late lanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis numerosis brevissimis brevibusque obscuris et praesertim in margine pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem glabrae. Styli obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 7–8. Apomicticum. Honori Walteri Scott ob studium florae zetlandicae praelongum nominatur. Hieracium ascendentidens P. D. Sell sp. nov. Holotype: Lingadale, North Harris, v.c. 110, July 1955, C. West (CGE).

Herba perennis phyllopoda, caudice ramoso. Caulis 30– 50 cm, pallide luteoviridis, ad basin raro aliquantum purpurascens, gracilis flexuosus, pilis simplicibus eglandulosis paucis vel numerosis mediocribus vel longis pallidis et pilis stellatis numerosis et in parte superiore pilis glanduliferis numerosis brevibus obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora; folia basalia numerosa, lamina 1.5–9.0 × 1.0– 3.5 cm, lanceolata vel ovata, ad apicem acuta vel acuminata (saepe acumine longo), acute dentata, dentibus apiculatis ascendentibusque (plerisque angustissimis sed nonnullis mammiformibus) praedita, ad basin cuneata vel attenuata, petiolis ad 6 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folia caulina 0–2, inferum quando adest basalibus simile, petiolatum, superum lamina anguste lanceolata, acuta, integra, aut bracteiforme; omnia in paginis ambabus inque margine pilis simplicibus eglandulosis numerosis mediocribus vel longis rigidis pallidis vestita. Inflorescentia capitulis 2–6, furcatocorymbosa; pedunculi longi rectique, pilis stellatis densis pilisque glanduliferis numerosis brevissimis brevibusque (0.2–0.7 mm) obscuris pilisque simplicibus eglandulosis sparsis mediocribus pallidis vestiti. Capitula 30–35 mm diametro, ad basin subtruncata. Involucri squamae 5–12 × 1.0–1.5 mm, olivaceae, interiores ad marginem pallidiores, omnes linearilanceolatae, ad apicem tenuem magis minusve acutum gradatim angustatae, pilis stellatis numerosis (praesertim in margine), pilisque glanduliferis numerosis inaequalibus brevibus vel mediocribus (0.3–1.0 mm) obscuris pilisque simplicibus eglandulosis magis minusve numerosis brevibus vel mediocribus pallidis base obscura vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis brevissimis vestitae. Styli flavi. Receptaculi alveoli margine incisodentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6– 7. Apomicticum. Hieracium filisquamum P. D. Sell sp. nov. Holotype: Rigg Head, near Seatoller, Cumberland, v.c. 70, July 1958, C. West (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 30–40 cm, pallide luteoviridis, basin versus saepe purpurascens, gracilis flexuosusque, striatus, infra pilis simplicibus eglandulosis densis longis sinuatis pallidis, sursum paucioribus brevioribusque, et in parte superiore pilis stellatis densis vestitus, sine pilis glanduliferis. Folia in pagina superiore aliquantum caeruleoviridia, in pagina inferiore pallidiora, interdum aliquantum purpurascentitincta; folia basalia pauca, lamina 6–10 × 2–3 cm, anguste elliptica vel lanceolata, ad apicem acuta vel obtusomucronata, subintegra vel sinuatodenticulata, ad basin cuneata vel attenuata, petiolis ad 3 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folia caulina 2–3, infera basalibus similia et saepe magna, petiolo alato praedita, supera lamina angustiore, longe acuta, denticulata, ad basin angustata et semiamplexicaulia, vel bracteiformia; omnia in pagina superiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis rigidis pallidis, in pagina inferiore numerosioribus longioribus minus rigidis vestita. Inflorescentia capitulis 2–3, corymbosa; pedunculi graciles erecti, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis interdumque

New taxa and combinations paulo sub capitulo pilis glanduliferis paucis minutis vestiti. Capitula 15–20 mm diametro, ad basin rotundata. Involucri squamae ante anthesin porrectae, 3–10 × 0.6–1.0 mm, nigrivirides, interiores ad marginem pallidiores, omnes lineares vel linearilanceolatae, ad apicem filamentosum protractae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis paucis brevissimis obscuris et basin versus pilis stellatis paucis vestitae. Ligulae flavae, ad apicem glabrae, brevissimae et debiliter effectae. Styli flavi. Receptaculi alveoli et cypselae non visi. Floret 7. Apomicticum. Hieracium chloranthum Pugsley ex P. D. Sell sp. nov. H. chloranthum Pugsley in Jour. Bot. (London) 79: 180 (1941) sine diagn. lat. Holotype: Uig, Isle of Skye, v.c. 104, 13 August 1888, E. F. Linton, Set of British Hieracia no. 12 (BM). Hieraciorum in Magna Britannia et Hibernia sola species ligulis viridiflavis, debiliter effectis. Herba perennis phyllopoda, caudice ramoso. Caulis 20–55 cm, pallide luteoviridis, basin versus saepe purpurascens, plerumque gracilis flexuosusque, striatus, infra pilis simplicibus eglandulosis numerosis longis sinuatis pallidis, sursum paucioribus brevioribusque, et ubique pilis stellatis numerosis vestitus. Folia in pagina superiore mediocriter caeruleoviridia, in pagina inferiore pallidiora; folia basalia pauca, exteriora lamina 2–6 × 1.0–2.5 cm, plerumque elliptica, raro ovata, ad apicem obtusomucronata, subintegra vel denticulata vel haud profunde dentata, ad basin rotundata vel cuneata, interiora lamina 7–11 × 1.5–2.5 cm, anguste elliptica vel oblongoelliptica, a media parte ad apicem acutum gradatim angustata, denticulata vel haud profunde dentata, interdum dentibus majoribus paucis praedita, ad basin gradatim angustata, petiolis ad 4 cm, alatis, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folia caulina 2–4, infera basalibus interioribus similia, petiolata, supera lanceolata vel linearia, ad apicem acutum gradatim angustata, dentata, sessilia; omnia in pagina superiore glabra vel fere glabra, raro pilis dispersis rigidis, in pagina inferiore pilis simplicibus eglandulosis numerosis sed dispersis brevibus vel mediocribus rigidis pallidis, in margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis magis minusve setiformibus vestita, in pagina inferiore, praecipue in foliis caulinis, pilis stellatis quoque saepe praesentibus. Inflorescentia capitulis (1–)3–5(–8), furcatosubcorymbosa, interdum ramo inferiore longo praedita; pedunculi satis longi gracilesque, pilis stellatis densis et pilis simplicibus eglandulosis numerosis brevibus mediocribusque et interdum pilis glanduliferis sparsis brevibus obscuris vestiti. Capitula 25–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin magis minusve porrectae, 4–12 × 0.8– 1.2 mm, obscure virides, interiores ad marginem pallidiores, omnes linearilanceolatae, ad apicem acutae, pilis stellatis numerosis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis pilisque glanduliferis paucis minutis inconspicuis obscuris vestitae. Ligulae triste viridiflavae, glabrae vel ad apicem pilis simplicibus eglandulosis brevissimis vestitae, plerumque debiliter effectae vel magis minusve tubulares. Styli triste flavi. Receptaculi

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alveoli margine fimbriatodentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. In Section Stelligera Zahn Hieracium hesperium P. D. Sell nom. et stat. nov. H. scoticum var. occidentale F. Hanb. in Jour. Bot. (London) 31: 18 (1893) et 32: 228 (1894). Hieracium dentulum (E. F. Linton) P. D. Sell stat. nov. H. holophyllum var. dentulum E. F. Linton in Jour. Bot. (London) 49: 356 (1911). Hieracium siluriense (F. Hanb.) P. D. Sell stat. nov. H. nitidum var. siluriense F. Hanb. in Jour. Bot. (London) 32: 228 (1894). Hieracium hypophalacrum P. D. Sell nom. et stat. nov. H. oreades var. subglabratum F. Hanb. in Jour. Bot. (London) 32: 227 (1894). Hieracium argentatum (Pugsley) P. D. Sell stat. nov. H. orimeles var. argentatum Pugsley in Jour. Linn. Soc. London (Bot.) 54: 102 (1948). Hieracium argentiforme P. D. Sell nom. et stat. nov. H. argenteum var. septentrionale F. Hanb. in Jour. Bot. (London) 32: 227 (1894), non H. septentrionale (Norrl.) Norrl. in Mela, Suomen Kasvio 653 (1906). Hieracium amydrostictum P. D. Sell sp. nov. Holotype: Rocks at about 2,000 ft, Coire Choimhlidh, below Beinn na Socaigh, Lochaber Mountains, W. Inverness, v.c. 97, 27/236744, 24 July 1966, R. W. Jones and B. A. Miles no. 66/65 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20– 40 cm, pallide luteoviridis, ad basin saepe purpurascens, gracilis flexuosus striatus, in parte inferiore pilis simplicibus eglandulosis paucis vel satis numerosis mediocribus vel longis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis stellatis aliquantum paucis pilisque glanduliferis sparsis brevibus obscuris vestitus. Folia in pagina superiore aliquantum glaucoviridia, colore brunneopurpureo maculata et guttata, in pagina inferiore pallidiora, interdum purpureotincta; folia basalia pauca vel numerosa, lamina 2–8 × 1.0–2.5 cm, plerumque lanceolata vel ovata, interdum oblongoelliptica, ad apicem obtusomucronata vel magis minusve acuta, denticulata vel interdum basin versus dentibus paucis majoribus praedita, ad basin cuneata, petiolis ad 4 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folium caulinum 0–1, quando adest saepe basalibus simile, petiolatum, vel lamina linearilanceolata, ad apicem acuta, integra vel dentata; omnia in pagina superiore pilis simplicibus eglandulosis numerosis brevibus vel mediocribus subrigidis, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosioribus brevibus vel mediocribus vel longis pallidis vestita. Inflorescentia capitulis 1–4, furcatocorymbosa; pedunculi longi graciles erecti, pilis stellatis magis minusve densis pilisque glanduliferis satis numerosis brevibus obscuris pilisque simplicibus eglandulosis paucis vel satis numerosis brevibus vel mediocribus base obscura vestiti. Capitula

546

New taxa and combinations

30–35 mm diametro, ad basin subtruncata. Involucri squamae ante anthesin porrectae, 5–13 × 1.2–1.5 mm, nigrivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem acutum gradatim angustatae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus base obscura et pilis glanduliferis numerosis brevibus obscuris et secus marginem pilis stellatis aliquot vestitae. Ligulae flavae, ad apicem pilis paucis brevissimis vestitae. Styli flavi. Receptaculi alveoli margine incisodentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium fucatifolium P. D. Sell sp. nov. Holotype: Riverside rocks, Linn of Quoich, near Braemar, S. Aberdeen, v.c. 92, 37/117912, 22 July 1965, R. W. Jones and B. A. Miles no. 65/106 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 30–60 cm, pallide luteoviridis, ad basin saepe brunneopurpureus, magis minusve robustus, striatus, prope basin pilis simplicibus eglandulosis paucis vel numerosis longis sinuatis pallidis vestitus, in parte superiore glaber vel fere glaber vel pilis stellatis paucis vestitus. Folia in pagina superiore caeruleoviridia, maculis punctisque obscure brunneopurpureis fucata, in pagina inferiore pallidiora; folia basalia pauca, lamina magna, 4–12 × 2–4 cm, elliptica vel lanceolatoelliptica vel oblongoelliptica, ad apicem rotundatomucronata vel acuta, denticulata vel dentata, interdum dentibus magnis uncinatis praedita, ad basin cuneata vel attenuata, petiolis longis, ad 10 cm, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folia caulina 0–2, unum plerumque magnum, basalibus simile, longipetiolatum, sed interdum bracteiforme; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore (praesertim in costa) inque margine pilis simplicibus eglandulosis paucis vel numerosis mediocribus vel longis pallidis vestita. Inflorescentia capitulis 2–6, furcatocorymbosa; pedunculi longi graciles erecti, pilis stellatis numerosis pilisque glanduliferis satis numerosis brevissimis vel brevibus obscuris pilisque simplicibus eglandulosis sparsis mediocribus pallidis vestiti. Capitula 30–40 mm diametro, ad basin rotundata vel subtruncata. Involucri squamae ante anthesin porrectae, 5–14 × 1.0–1.5 mm, obscure olivaceae, interiores in margine pallidiores, omnes linearilanceolatae, ad apicem angustum acutum gradatim attenuatae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis magis minusve numerosis brevibus obscuris et secus marginem pilis stellatis satis numerosis vestitae. Ligulae flavae, ad apicem glabrae. Styli flavi vel aliquantum obscuri. Receptaculi alveoli margine incisodentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6– 7. Apomicticum. Hieracium inaequilaterum P. D. Sell nom. et stat. nov. H. silvaticum var. asymmetricum Ley in W. R. Linton, Brit. Hier. 40 (1905). Hieracium promontoriale P. D. Sell sp. nov. Holotype: Limestone sea-cliffs, Humphrey Head, Westmorland, v.c. 69, 22 July 1962, C. West (CGE).

Herba perennis phyllopoda, caudice ramoso. Caulis 30– 50 cm, pallide luteoviridis, ad basin interdum aliquantum purpurascens, gracilis vel magis minusve robustus, striatus, in parte inferiore pilis simplicibus eglandulosis paucis mediocribus vel longis pallidis, sursum absentibus vel fere absentibus, in parte superiore pilis stellatis satis numerosis vestitus. Folia in pagina superiore pallide viridia, raro dilute guttata, in pagina inferiore pallidiora; folia basalia numerosa, lamina 4–9 × 1.0–3.0 cm, elliptica vel oblongoelliptica vel lanceolata, ad apicem obtusomucronata vel acuta vel acuminata vel cuspidata, dentibus angustis apiculatissimisque acutissime dentata vel serrata, ad basin plerumque cuneata, petiolis ad 2 cm, pilis simplicibus eglandulosis numerosis longis pallidis vestitis; folia caulina 0– 2, quando adsunt lamina magis minusve lineari vel linearilanceolata, apice longo angusto acuto, plerumque dentibus longis angustis praedita, breviter petiolata, interdum bracteiformia; omnia in pagina superiore magis minusve glabra vel pilis paucis prope marginem vestita, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis pallidis vestita. Inflorescentia capitulis 3–5 (vel in cultu numerosis), furcatocorymbosa; pedunculi recti erecti, pilis stellatis densis pilisque glanduliferis paucis vel numerosis brevibus obscuris pilisque simplicibus eglandulosis sparsis brevibus vel mediocribus pallidis vestiti. Capitula 30–40 mm diametro, ad basin subtruncata. Involucri squamae ante anthesin incumbentes, 5.0–13.5 × 1.0–1.2 mm, griseivirides, interiores margine pallidiore, omnes anguste linearilanceolatae, ad apicem angustum subacutum gradatim angustatae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis paucis brevibus obscuris et in margine basinque versus pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem glabrae. Styli flavidi vel aliquantum obscuri. Receptaculi alveoli margine subulatodentato. Cypselae 4.0–4.4 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium arvonense P. D. Sell sp. nov. Holotype: Cwm Idwal, Caernarvonshire, v.c. 49, 13 July 1912, E. S. Marshall no. 3725 (CGE). Hieracio leyi (quocum saepe invenitur) et H. saxorum et H. petrocharite antea confusum, ad H. leyi proximum est, sed foliis pallide vel mediocriter viridibus guttatis minoribus, ad basin magis contractis, in agro statim agnoscitur. Herba perennis phyllopoda, caudice ramoso. Caulis 15–35 cm, pallide luteoviridis, in parte inferiore saepe colore brunneopurpureo suffusus, gracilis flexuosus striatus, in parte inferiore pilis simplicibus eglandulosis paucis vel satis numerosis longis pallidis, sursum paucioribus brevioribusque vel nullis, in parte superiore pilis stellatis numerosis pilisque glanduliferis sparsis brevissimis obscuris vestitus. Folia in pagina superiore pallide vel mediocriter viridia, guttis purpureobrunneis praedita, in pagina inferiore pallidiora, interdum purpurascentitincta; folia basalia plerumque numerosa, lamina 1.5–7.0 × 1–3 cm, lanceolata vel elliptica, ad apicem rotundatoobtusa vel acutissima, sinuatodenticulata, ad basin abrupte contractam vel subtruncatam saepe asymmetricam plerumque dentibus paucis majoribus praedita, petiolis ad 4 cm, pilis

New taxa and combinations simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folium caulinum 0–1, quando adest lamina ovata, apice longo angusto acuto, integrum vel in dimidio inferiore dentatum, ad basin cuneatum, plerumque petiolatum; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis pallidis (in margine subrigidis) vestita. Inflorescentia capitulis 1–5, furcatocorymbosa; pedunculi graciles, erecti, pilis stellatis numerosis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis numerosis brevibus tenuibus obscuris vestiti. Capitula 25–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin porrectae, 5–13 × 1.0–1.2 mm, griseivirides, interiores ad marginem pallidae, omnes anguste linearilanceolatae, ad apicem angustum magis minusve acutum gradatim angustatae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis numerosis brevibus obscuris et secus marginem pilis stellatis numerosis vestitae. Ligulae flavae, glabrae vel pilis sparsis brevissimis ad apicem vestitae. Styli flavi. Receptaculi alveoli margine subulatodentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium sublasiophyllum P. D. Sell nom. et stat. nov. H. leyi var. vestitum Ley ex E. F. Linton in Jour. Bot. (London) 49: 353 (1911), non H. vestitum Gren. & Godr., Fl. France 2: 369 (1850). Hieracium peroblongum P. D. Sell sp. nov. Holotype: Lower rocks, Snake Fell, Ambleside, Westmorland, v.c. 69, 12 July 1905, A. Ley (CGE). Hieracio argenteo, quocum invenitur, simillimum, foliis oblongis magis minusve integris, involucri squamis angustioribus, pilis simplicibus pluribus glanduliferisque paucioribus vestitis, ligulis ciliatis distinguitur. Herba perennis phyllopoda, caudice ramoso. Caulis 12–30 cm, pallide luteoviridis, ad basin purpurascens, gracilis, in parte superiore pilis simplicibus eglandulosis paucis vel satis numerosis vel fere nullis brevibus vel mediocribus vel longis pallidis pilisque stellatis aliquantum paucis pilisque glanduliferis sparsis brevibus obscuris vestitus. Folia in pagina superiore caeruleoviridia, in pagina inferiore pallidiora, saepe purpureosuffusa; folia basalia numerosa, lamina 2–8 × 1–2 cm, oblonga vel anguste elliptica vel anguste lanceolata, ad apicem obtusomucronata vel magis minusve acuta, integra vel remote denticulata, ad basin rotundata vel cuneata, petiolis ad 4 cm, pilis simplicibus eglandulosis longis pallidis vestitis; folium caulinum 0– 1, quando adest bracteiforme; omnia in pagina superiore glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus aliquantum rigidis vestita. Inflorescentia capitulis 1–4, furcatocorymbosa; pedunculi recti graciles, pilis stellatis aliquantum paucis pilisque glanduliferis numerosis brevibus brevissimisque tenuibus obscuris pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus obscuris vel base obscura vestiti. Capitula 25–30 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–12 × 1.0–1.2 mm, nigrivirides,

547

interiores margine pallidiore, omnes anguste linearilanceolatae, ad apicem angustissimae acutae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis numerosis brevibus obscuris et secus marginem pilis stellatis paucis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis brevissimis vestitae. Styli flavi. Receptaculi alveoli margine fimbriatodentato. Cypselae 2.5–3.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium grampianum P. D. Sell sp. nov. Holotype: South-east-facing cliffs above Dubh Lochan, Beinn a’ Bhuird, S. Aberdeen, v.c. 92, 37/091993, 1 August 1966, R. W. Jones and B. A. Miles no. 66/120 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 15– 45(–50) cm, luteoviridis, ad basin interdum purpurascens, graciliusculus striatus, interdum flexuosus, pilis simplicibus eglandulosis paucis mediocribus vel longis pallidis ubique dispersis vel basin versus numerosioribus, in parte superiore pilis stellatis paucis pilisque glanduliferis paucis minutis obscuris vestitus. Folia magis minusve caeruleoviridia, praecipue in pagina inferiore, raro aliquantum guttata; folia basalia lamina 4–11 × 1.2–3.5 cm, exteriora elliptica, interiora magis minusve lanceolata vel suboblongolanceolata, ad apicem exteriora obtusa interioraque acuta, omnia denticulata vel dentata, dentibus triangularimucronatis, basin versus gradatim majoribus, interdum secus petioli latus unum angustissimis, ad basin abrupte angustata vel asymmetrice subtruncata, petiolis plerumque mediocribus, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folium caulinum 0–1, quando adest plerumque parvum et bracteiforme sed interdum basalibus interioribus simile; omnia in pagina superiore glabra vel pilis simplicibus eglandulosis sparsis pallidis, in pagina inferiore sicut ad et prope marginem pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis subrigidis pallidis, in pagina inferiore, praesertim caulina, saepe pilis stellatis numerosis vestita. Inflorescentia capitulis 1–6, furcatocorymbosa, ramo inferiore longo interdum praedita, acladio saepe satis longo; pedunculi longi recti erecti, pilis stellatis numerosis vel densis pilisque glanduliferis numerosis brevissimis obscuris pilisque simplicibus eglandulosis sparsis pallidis vestiti. Capitula 35–45 mm diametro, ad basin rotundata. Involucri squamae ante anthesin porrectae, 6–14 × 1.0–1.5 mm, griseivirides (interiores in margine pallidiores, ad apicem interdum rubellae), anguste linearilanceolatae (interiores fere filamentosae), ad apicem acutum gradatim angustatae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis paucis brevibus obscuris (interdum absentibus) et pilis stellatis in margine densis inque pagina paucis vestitae. Ligulae pallide flavae, glabrae vel ad apicem aliquantum pilosae. Styli magis minusve flavi. Receptaculi alveoli margine subulatodentato. Cypselae 3.0–3.5 mm, nigellae. Floret 7–8. Apomicticum. Hieracium bettyhillense P. D. Sell sp. nov. Holotype: Farr Bay, Bettyhill, W. Sutherland, v.c. 108, 19 July 1897, E. S. Marshall and W. A. Shoolbred as E. F. & W. R. Linton, Set of British Hieracia no. 106 (CGE).

548

New taxa and combinations

Herba perennis phyllopoda, caudice ramoso. Caulis 20–50 cm, pallide viridis, basin versus saepe purpurascens, gracilis, flexuosus vel satis robustus, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis longis pallidis, sursum paucioribus brevioribusque vel absentibus, in parte superiore pilis stellatis numerosis et interdum pilis glanduliferis sparsis brevibus obscuris vestitus. Folia in pagina superiore caeruleoviridia, in pagina inferiore pallidiora, crassa; folia basalia numerosa, lamina 4–10 × 1.5–3.0 cm, plerumque ovata, interdum oblongoovata vel elliptica, ad apicem plerumque acuta, acute dentata vel serratodentata, ad basin abrupte contracta vel cuneata, breviter petiolata; folium caulinum 0–1(–2), quando adest plerumque bracteiforme, raro lanceolatum, ad apicem acutissimum, serratum, ad basin ad petiolum brevem alatum cuneatum; omnia in pagina superiore glabra vel pilis paucis rigidis vestita, in margine pilis simplicibus eglandulosis numerosis mediocribus longisque rigidis pallidis, in pagina inferiore pilis numerosis mollioribus vestita. Inflorescentia capitulis 2–4(–8), furcatocorymbosa; pedunculi breviusculi recti, pilis stellatis densis pilisque glanduliferis numerosis brevissimis tenuibus pilisque simplicibus eglandulosis paucis brevibus vel mediocribus pallidis vestiti. Capitula 30–35 mm diametro, ad basin subtruncata. Involucri squamae 4–12 × 1.0–1.2 mm, nigrivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem magis minusve acutae, pilis simplicibus eglandulosis numerosis mediocribus pallidis base obscura et pilis glanduliferis magis minusve numerosis brevissimis brevibusque obscuris et pilis stellatis paucis vestitae. Ligulae flavae, ad apicem pilis paucis brevissimis vestitae. Styli flavi. Receptaculi alveoli margine dentatofimbriato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium stictum P. D. Sell nom. et stat. nov. H. hypochaeroides var. lancifolium W. R. Linton, Brit. Hier. 29 (1905), non H. lancifolium Vuk. in Rad. Jugos. Akad. Zagreb 58: 167 (1881). Hieracium dalense P. D. Sell sp. nov. Holotype: Rocks by roadside, Litton Dale, Derbyshire, v.c. 57, 43/156748, 21 June 1954, P. D. Sell no. 54/165 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20–45 cm, pallide luteoviridis, in parte inferiore saepe purpurascentitinctus, gracilis sed rigidus, striatus, in parte inferiore pilis simplicibus eglandulosis paucis longis pallidis, in parte superiore pilis stellatis numerosis pilisque glanduliferis numerosis brevibus obscuris vestitus. Folia in pagina superiore glaucoviridia, in pagina inferiore pallidiora, saepe rubellitincta; folia basalia plerumque numerosa, exteriora lamina 3–9 × 1.5–4.0 cm, late elliptica vel elliptica, plerumque ad apicem obtusomucronata, subintegra vel denticulata, ad basin rotundata vel breviter cuneata, interiora lamina 4–11 × 1.5–4.5 cm, elliptica vel ovata vel lanceolata, ad apicem magis minusve acuta, subintegra vel denticulata vel prope basin dentibus paucis parvis praedita, ad basin abrupte contracta vel breviter cuneata, petiolis brevibus vel mediocribus, pilis simplicibus eglandulosis densis longis pallidis vestitis; folium caulinum 0–1, quando

adest basalibus interioribus simile petiolatumque vel lamina lineari adque apicem acutum; omnia in pagina superiore glabra, in pagina inferiore pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis, in margine pilis rigidis vestita. Inflorescentia capitulis 2–6, furcatocorymbosa; pedunculi breviusculi, magis minusve erecti, pilis stellatis densis et pilis glanduliferis numerosis brevibus obscuris et raro pilis simplicibus eglandulosis solitariis vestiti. Capitula 30–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin magis minusve porrectae, 4–13 × 1.0–1.2 mm, nigrivirides (interiores in margine pallidiores), anguste linearilanceolatae (interiores filamentosae), omnes ad apicem magis minusve acutae, pilis glanduliferis numerosis maxime inaequalibus brevibus mediocribusque obscuris et pilis simplicibus eglandulosis paucis mediocribus base obscura et plerumque basin versus pilis stellatis paucis vestitae. Ligulae flavae, ad apicem glabrae. Styli flavi. Receptaculi alveoli margine acute dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 5–7. Apomicticum. Hieracium apiculatidens P. D. Sell sp. nov. Holotype: Grassy limestone bank at Swindon Quarry, near Threshfield, M. W. Yorkshire, v.c. 64, 25 May 1961, C. West (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 30–55 cm, pallide luteoviridis, basin versus saepe brunneopurpureus, gracilis vel satis robustus, striatus, in parte inferiore pilis simplicibus eglandulosis paucis longis pallidis, sursum paucioribus, in parte superiore pilis stellatis satis numerosis vestitus. Folia in pagina superiore obscure caeruleoviridia, in pagina inferiore pallidiora, crassa et rigida; folia basalia plerumque numerosa, lamina magna brassiciformi, 2– 10 × 1.5–6.0 cm, late ovata vel late elliptica, interdum fere subrotunda, ad apicem plerumque breviter acuta, raro rotundatomucronulata, regulariter dentata vel serratodentata, dentibus acutis apiculatis interdum curvatis, ad basin truncata vel abrupte contracta vel breviter cuneata, petiolis plerumque longis, praecipue in plantis mense Julio florentibus, ad 11 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folium caulinum 0–1, quando adest basalibus simile sed multo minus, petiolatum, aut angustius profundeque dentatum; omnia in pagina superiore glabra vel pilis simplicibus eglandulosis paucis brevibus vel mediocribus pallidis (plerumque prope marginem), in pagina inferiore inque margine pilis simplicibus eglandulosis magis minusve numerosis mediocribus vel longis pallidis vestita. Inflorescentia capitulis 4–7, furcatocorymbosa; pedunculi longi graciles erecti, pilis stellatis densis vestiti, sine pilis simplicibus eglandulosis vel glanduliferis. Capitula 25–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin porrectae, 4–11 × 1.0–1.5 mm, griseivirides, interiores ad marginem pallidiores, omnes linearilanceolatae, ad apicem gradatim angustatae sed obtusae, pilis stellatis densis (saltem in margine sed saepe ubique) pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis paucis brevibus obscuris vestitae. Ligulae flavae, ad apicem pilis paucis brevissimis vestitae. Styli obscuri. Receptaculi

New taxa and combinations

549

alveoli margine breviter dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 5–7. Apomicticum.

todentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 5–7. Apomicticum.

Hieracium breconense P. D. Sell nom. et stat. nov. H. britannicum var. ovale Ley in Jour. Bot. (London) 45: 109 (1907).

Hieracium vagicola P. D. Sell sp. nov. Holotype: Pen Moel Rocks, Chepstow, W. Gloucestershire, v.c. 34, 6 June 1900, A. Ley as E. F. & W. R. Linton, Set of British Hieracia no. 136 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 15– 40 cm, pallide viridis, ad basin interdum colore brunneopurpureo tinctus, striatus, magis minusve robustus, erectus, glaber vel in parte superiore pilis simplicibus eglandulosis sparsis pallidis pilisque stellatis paucis vestitus. Folia plerumque rigidiuscula, in pagina superiore aliquantum glaucoviridia, in pagina inferiore pallidiora glaucaque, interdum colore brunneopurpureo suffusa; folia basalia numerosa, lamina 2–11 × 2–6 cm, exteriora ovata vel late ovata vel late elliptica vel late oblonga vel subrotunda, ad apicem late rotundatomucronulata, passim praeter quadrantem summum profunde et irregulariter dentata, dentibus anguste vel late mammiformibus, saepe curvatis, ad basin truncatam plerumque dentibus duobus reflexis, interiora ovata vel lanceolata, ad apicem saepe tortum acuta vel acuminata, plerumque profunde et irregularissime dentata, dentibus quam foliorum exteriorum angustioribus longioribus irregularioribus, petiolis ad 6 cm, pallide viridibus, saepe colore brunneopurpureo tinctis, pilis simplicibus eglandulosis densis longissimis crispatis pallidis vestitis; folia caulina plerumque absentia sed, quando adsunt, lamina lineari bracteiformique; omnia in pagina superiore glabra vel pilis simplicibus eglandulosis sparsissimis vestita, in pagina inferiore glabra vel, praecipue in costa, pilis simplicibus eglandulosis paucis vel numerosis brevibus vel mediocribus vel longis pallidis, ad vel prope marginem pilis simplicibus eglandulosis numerosis mediocribus vel longis subrigidis pallidis vestita. Inflorescentia capitulis 2–4(–7), plerumque satis spisse cymosocorymbosa, ramo inferiore longo interdum praedita, acladio brevi; pedunculi 15– 30(–60) mm pilis stellatis densis pilisque glanduliferis paucis vel satis numerosis brevibus magis minusve aequalibus obscuris, interdum et pilis simplicibus eglandulosis paucis mediocribus, vestiti. Capitula 35–45 mm diametro, ad basin rotundata. Involucri squamae ante anthesin porrectae, 8– 16 × 1.0–1.5 mm, nigrivirides, interiores saltem in margine pallidiores, omnes anguste linearilanceolatae, apice longo angusto fere filamentoso acuto, pilis simplicibus eglandulosis paucis vel numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis paucis vel numerosis brevibus obscuris et in margine pilis stellatis densis vestitae. Ligulae pallide flavae, ad apicem glabrae. Styli flavi. Receptaculi alveoli subulatodentati. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 5–6(–7). Apomicticum.

Hieracium britannicoides P. D. Sell sp. nov. Holotype: Growing in cracks on south-facing limestone cliffs, Bryn Euryn, Colwyn Bay, Denbigh, v.c. 50, 23/832797, 15 June 1968, J. M. Brummitt (CGE). Inter Hieracium britannicum et H. britanniciforme intermedium, ad utrumque aliquando relatum est; illius folia indumentumque molliora, hujus inflorescentiam compactam, involucri squamas pilosiores interdum filamentosas, pedunculos glandulosiores habet. Herba perennis phyllopoda, caudice ramoso. Caulis 15–35 cm, pallide luteoviridis, in parte inferiore saepe purpurascentitinctus, gracilis vel satis robustus, erectus, striatus, pilis simplicibus eglandulosis sparsis brevibus vel mediocribus pallidis et in parte superiore pilis stellatis numerosis pilisque glanduliferis paucis vel numerosis brevibus obscuris vestitus. Folia plerumque flaccidiuscula, in pagina superiore magis minusve glaucoviridia, in pagina inferiore pallidiora glaucioraque, interdum purpurascentisuffusa; folia basalia numerosa, lamina 2–7(–9) × 1–4 cm, exteriora ovata vel lanceolata, ad apicem rotundatomucronata vel magis minusve acuta, ubique denticulata vel in parte inferiore dentata, dentibus mammiformibus, ad basin subtruncata et interdum dentibus retrorsis praedita, interiora omnia ovata, ad apicem acuta et saepe torta, ubique denticulata vel in parte inferiore dentata, dentibus mammiformibus, ad basin subtruncata et saepe dentibus retrorsis praedita, petiolis ad 5(–7) cm, pilis simplicibus eglandulosis densis mediocribus vel longis crispis pallidis vestitis; folia caulina saepe absentia, interdum lamina lineari, interdum basalibus interioribus similia petiolataque; omnia in pagina superiore glabra vel pilis simplicibus eglandulosis sparsis mediocribus pallidis vestita, in pagina inferiore (praesertim in costa) pilis simplicibus eglandulosis paucis vel satis numerosis brevibus vel mediocribus vel longis pallidis, ad et prope marginem pilis simplicibus eglandulosis numerosis vel densis rigidiusculis pallidis vestita. Inflorescentia capitulis 2–6, plerumque aliquantum spisse cymosocorymbosa, interdum ramo inferiore praedita, acladio plerumque breviusculo; pedunculi 5– 35 mm, pilis stellatis densis pilisque glanduliferis numerosis brevibus obscuris et interdum pilis simplicibus eglandulosis sparsis vestiti. Capitula 20–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin porrectae, 8–16 × 1.0–1.2 mm, nigrivirides, interiores in margine pallidiores, omnes anguste linearilanceolatae, ad apicem longum gracilem interdum filamentosum acutum in medio abrupte angustatae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus vel longis pallidis sed plerumque base obscura, cum pilis glanduliferis satis paucis brevibus obscuris intermixtis, pilisque stellatis paucis vestitae. Ligulae pallide flavae, ad apicem glabrae. Styli flavidi. Receptaculi alveoli margine subula-

In Section Vulgata (Griseb.) Willk. & Lange Hieracium vulgatum Fr., Novit. Fl. Suec. 76 (1819). forma subfasciculare (W. R. Linton) P. D. Sell stat. nov. H. vulgatum var. subfasciculare W. R. Linton, Brit. Hier. 64 (1905). forma pseudosubramosum (Pugsley) P. D. Sell stat. nov.

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New taxa and combinations

H. vulgatum var. pseudosubramosum Pugsley in Jour. Linn. Soc. London (Bot.) 54: 196 (1948). forma subravusculum (W. R. Linton) P. D. Sell stat. nov. H. vulgatum var. subravusculum W. R. Linton, Brit. Hier. 64 (1905). forma sejunctum (W. R. Linton) P. D. Sell stat. nov. H. vulgatum var. sejunctum W. R. Linton in Jour. Bot. (London) 39: 105 (1901). Hieracium peccense (W. R. Linton) P. D. Sell stat. nov. H. rubiginosum var. peccense W. R. Linton, Brit. Hier. 49 (1905). Hieracium robertsii P. D. Sell nom. et stat. nov. H. rubiginosum var. glabrescens Pugsley, non H. glabrescens Murr in Deutsch. Bot. Monatsschr. 15: 226 (1897). In memoriam Evan Roberts (1906–91) nominatur. Hieracium kintyricum P. D. Sell sp. nov. Holotype: Largybaan ravine, Kintyre, v.c. 101, 5 July 1969, A. G. Kenneth no. 56/69. Herba perennis phyllopoda, caudice ramoso. Caulis 30–60 cm, pallide luteoviridis, in dimidio inferiore colore brunneopurpureo suffusus, gracilis vel magis minusve robustus, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis longis pallidis, sursum sparsissimis multoque brevioribus vel absentibus, in parte superiore pilis stellatis paucis vel numerosis vestitus. Folia in pagina superiore mediocriter glaucoviridia, in pagina inferiore pallidiora; folia basalia pauca, lamina 4–10 × 2.0–4.5 cm, late ovata vel late elliptica vel elliptica, sursum attenuata sed ad apicem rotundatoobtusa, subintegra vel remote denticulata vel dentibus paucis brevibus praedita, ad basin abrupte contracta vel cuneata, petiolis ad 4 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folia caulina 3–5, infimum vel saepe duo infima basalibus similia sed interdum dentibus pluribus praedita et petiolata, supera lamina minore linearilanceolata longe acuta dentata sessilia, aut bracteiformia; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel longis pallidis vestita. Inflorescentia capitulis 2–7, paniculatocorymbosa; pedunculi aliquantum graciles, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis pilisque glanduliferis sparsissimis brevibus obscuris vestiti. Capitula 25–30 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–11 × 1.0–1.5 mm, plerumque pallide virides, in centro parum fuscatae, linearilanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis et pilis glanduliferis paucis vel satis numerosis brevibus obscuris, a pilis simplicibus celatis, et basin versus pilis stellatis paucis vestitae. Ligulae flavae, ad apicem glabrae. Styli flavi. Receptaculi alveoli et cypselae non visi. Floret 6–7. Apomicticum. Hieracium aequiserratum P. D. Sell sp. nov. Holotype: Barras, Westmorland, v.c. 69, July 1952, C. West (CGE).

Herba perennis phyllopoda, caudice ramoso. Caulis 20–55 cm, pallide luteoviridis, infra saepe purpureoruber, magis minusve gracilis, interdum flexuosus, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis longis pallidis, sursum gradatim paucioribus brevioribusque et supra interdum sparsissimis, et in parte superiore pilis stellatis densis vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora; folia basalia numerosa, lamina 3–8 × 1.5–3.5 cm, plerumque elliptica, interdum ovata vel lanceolata, ad apicem obtusa vel acuta, pariter sinuatoserrata, dentibus parvis et pro parte maxima prorsum flexis, apiculo parvo ad apicem munitis, ad basin cuneata vel attenuata, petiolis plerumque aliquantum brevibus sed ad 4 cm longis, pilis simplicibus eglandulosis densis longis pallidis vestitis; folia caulina 2–3, basalibus similia sed minora, angustiora, ad apicem acutissima et petiolo brevi alato praedita, summum plerumque bracteiforme; omnia in pagina superiore pilis simplicibus eglandulosis brevibus rigidis pallidis et in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis vestita. Inflorescentia capitulis 2–10, confertim paniculatocorymbosa; pedunculi aliquantum breves, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis pilisque glanduliferis paucis vel numerosis brevibus obscuris vestiti. Capitula 25–30 mm diametro, ad basin subtruncata. Involucri squamae ante anthesin incumbentes, 4–10 × 1.0– 1.2 mm, nigrivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis paucis brevibus obscuris et secus marginem adque apicem pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem pilis sparsis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine incisodentato. Cypselae 3.0–3.5 mm longae, rubrinigrae. Floret 6–7. Apomicticum. Hieracium vinifolium P. D. Sell sp. nov. Holotype: Rock ledges on cliffs by the sea, between Rudha Duin Bhain and Uamh Ropa, south-west of Campbeltown, Kintyre, v.c. 101, 16/594147, 26 June 1968, P. D. Sell no. 68/202 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 20–60 cm, pallide viridis, saepe colore purpureorubro suffusus, plerumque gracilis, raro robustus, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis longis sinuatis pallidis, sursum multo paucioribus brevioribusque, in parte superiore pilis stellatis paucis vestitus. Folia in pagina superiore caeruleogrisea, in pagina inferiore pallidiora et saepe colore rubripurpureo tincta, interdum tota colore rubripurpureo suffusa; folia basalia pauca vel numerosa, lamina 2– 10(–14) × 2–5(–6) cm, elliptica vel ovata vel lanceolata, ad apicem rotundatoobtusa vel subacuta, denticulata vel dentata, triente superiore interdum integro, ad basin plerumque cuneata, interdum attenuata vel subtruncata, petiolis ad 5 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folia caulina 2–5, infera medianaque basalibus similia, breviter petiolata vel sessilia, supera lamina lanceolata, ad apicem acuta, dentata sessilia, aut bracteiformia; omnia in pagina superiore glabra vel pilis simplicibus

New taxa and combinations eglandulosis brevibus pallidis vestita, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis mediocribus vel longis pallidis vestita. Inflorescentia capitulis 2–16, corymbosa, ramo inferiore longo interdum praedita; pedunculi breves, interdum brevissimi, saepe curvati, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus mediocribusque pallidis base obscura pilisque glanduliferis numerosis brevibus obscuris vestiti. Capitula 25–30 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 3–10 × 0.8–1.0 mm, obscure olivaceae, interiores margine pallidiore, omnes linearilanceolatae, gradatim ad apicem obtusum vel subacutum angustatae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis paucis brevibus obscuris pilisque stellatis sparsis vestitae. Ligulae intense flavae, ad apicem glabrae. Styli flavi. Receptaculi alveoli margine acute incisodentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium amnicola P. D. Sell nom. et stat. nov. H. rubiginosum var. rivulare Pugsley in Jour. Linn. Soc. London (Bot.) 54: 182 (1948), non H. rivulare Krocker, Fl. Siles. suppl. 2, 249 (1814). Hieracium subviolascens P. D. Sell nom. nov. H. irriguum subsp. violascens Almq. ex Dahlst. in Bot. Not. 1886: 164 (1886); H. violascens (Almq. ex Dahlst.) Dahlst. in Lindm., Svensk. Fanerogamfl. 626 (1926), non H. violascens Borbas, Math. Term´esz. 26 (1880). Hieracium mammidens P. D. Sell sp. nov. Holotype: Pembury, Kent, v.c. 16, 7 June 1956, C. West (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 30–85 cm, pallide luteoviridis, ad basin saepe colore brunneopurpureo suffusus, gracilis vel robustus, striatus, in parte inferiore pilis simplicibus eglandulosis numerosis vel densis longis sinuatis pallidis, sursum gradatim paucioribus brevioribusque, in parte superiore pilis stellatis numerosis vel densis pilisque glanduliferis sparsis brevissimis obscuris vestitus. Folia in pagina superiore luteoviridia, in pagina inferiore pallidiora, saepe purpureosuffusa; folia basalia pauca vel numerosa, lamina 4–10 × 2–4 cm, elliptica vel late elliptica vel obovata vel fere subrotunda, ad apicem rotundata vel obtusa, plerumque dentata, raro subintegra vel denticulata, dentibus mammiformibus (interdum anguste mammiformibus), ad basin cuneata vel subtruncata, petiolis ad 9 cm, pilis simplicibus eglandulosis densis longis pallidis vestitis; folia caulina 3–5, gradatim sursum decrescentia, infera basalibus similia petiolataque, supera interdum ovata vel lanceolata, saepe magis dentata; omnia pilis simplicibus eglandulosis in pagina superiore numerosis brevibus pallidis, in pagina inferiore (praesertim in costa) longioribus vestita, caulina in pagina inferiore pilis stellatis quantitate variantibus vestita. Inflorescentia capitulis 3–20, paniculatocorymbosa, ramis longis praedita; pedunculi breviusculi, graciliusculi, erecti, pilis stellatis densis pilisque glanduliferis numerosis brevibus obscuris pilisque simplicibus eglandulosis numerosis mediocribus pallidis vestiti. Capit-

551

ula 30–40 mm diametro. Involucri squamae ante anthesin incumbentes, 5.0–10.5 × 0.8–1.0 mm, olivaceae, interiores margine pallidiore, omnes anguste linearilanceolatae, ad apicem obtusae, pilis glanduliferis numerosis brevissimis brevibusque tenuibus obscuris et pilis simplicibus eglandulosis sparsis mediocribus pallidis base obscura et in margine pilis stellatis paucis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis brevissimis vestitae. Styli flavi. Receptaculi alveoli margine breviter dentato. Cypselae 2.5– 3.0 mm, rubrinigrae. Floret 5–6. Apomicticum. Hieracium lagganense P. D. Sell sp. nov. Holotype: River-banks at Crathie and Laggan Bridge, E. Inverness, v.c. 96, 4 August 1916, E. S. Marshall no. 4310 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 40–60 cm, pallide luteoviridis, basin versus saepe purpurascens, gracilis sed rigidus, striatus, basin versus pilis simplicibus eglandulosis numerosis longis sinuatis pallidis, sursum paucioribus vel absentibus, in parte superiore pilis stellatis et glanduliferis brevibus obscuris numerosis vestitus. Folia in pagina superiore mediocriter glaucoviridia, circa marginem saepe signo purpureo notata, in pagina inferiore pallidiora, saepe purpureosuffusa; folia basalia pauca vel satis numerosa, lamina 5.0–7.5 × 2–3 cm, anguste elliptica vel ovata, ad apicem subacuta, subintegra vel remote denticulata, ad basin cuneata vel attenuata, petiolis ad 5 cm, saepe purpureis, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folia caulina (1–)2(–3), infimum plerumque in caule humillime positum, lamina ovata, magis minusve acuta, denticulata, breviter petiolata, supera lanceolata, ad apicem acuta, integra et sessilia, aut bracteiformia; omnia in paginis ambabus inque margine pilis simplicibus eglandulosis brevibus rigidis, caulina in pagina inferiore pilis stellatis numerosis vestita. Inflorescentia capitulis 3–12, paniculatocorymbosa, ramo inferiore longo saepe praedita; pedunculi longi gracilesque, pilis stellatis densis pilisque glanduliferis numerosis brevibus obscuris vestiti. Capitula 25–30 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–10 × 0.8–1.2 mm, nigrivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem perangustae, acutae, pilis glanduliferis numerosis brevibus brevissimisque gracilibus obscuris et prope basin pilis stellatis paucis vestitae, sine pilis simplicibus eglandulosis. Ligulae flavae, ad apicem glabrae. Styli flavi. Receptaculi alveoli margine dentato. Cypselae 2.5–3.0 mm, rubrinigrae. Floret 7–8. Apomicticum. In Section Hieracium Hieracium irregularidens P. D. Sell sp. nov. Holotype: Grassy bank by the track to Dalmunzie Lodge, by the Spittal of Glen Shee, E. Perth, v.c. 89, 37/1070, 19 July 1959, P. D. Sell no. 59/37, N. D. Simpson and C. West (CGE). Ab Hieracio caesiomurorum foliorum dentibus numerosioribus obtusis petiolisque plerumque brevioribus et ab H. cuspidente (sectionis Subalpinorum) foliorum dentibus obtusioribus foliique base magis contracta differt.

552

New taxa and combinations

Herba perennis phyllopoda, caudice ramoso. Caulis 30–65 cm, pallide luteoviridis, infra saepe rubripurpureus, gracilis vel robustus, striatus, in parte inferiore pilis simplicibus eglandulosis satis numerosis longis pallidis, sursum paucioribus brevioribusque vel absentibus, in parte superiore pilis stellatis numerosis vestitus. Folia in pagina superiore magis minusve caeruleoviridia, in pagina inferiore pallidiora; folia basalia pauca vel numerosa, lamina saepe magna, 4–14 × 3.5–6.0 cm, ovatooblonga vel ovata vel lanceolatooblonga, ad apicem plerumque acuta sed raro obtusa, irregulariter dentata (saepe profunde), dentibus inferis saepe longis angustisque sed dentibus plurimis obtusis etsi apiculatis, ad basin abrupte contracta vel subtruncata, dentibus inferis interdum in petiolum extensis, petiolis plerumque breviusculis sed ad 9 cm, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folia caulina 0–2, unum saepe magnum basalibusque simile, petiolatum, superum (quando adest) angustum, dentibus longis angustis interdum praeditum; omnia in pagina superiore glabra vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosis brevibus vel longis pallidis vestita. Inflorescentia capitulis 2–12, paniculatocorymbosa, ramo inferiore longo saepe praedita; pedunculi satis graciles, erecti, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis paucis vel satis numerosis brevibus obscuris vestiti. Capitula 30–40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–12 × 1.2–1.5 mm, obscure olivaceae vel nigrivirides, interiores margine pallido, omnes linearilanceolatae, ad apicem gradatim angustatae sed obtusae, pilis simplicibus eglandulosis numerosissimis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis satis numerosis brevibus obscuris et secus marginem pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis brevibus vestitae. Styli flavi vel aliquantum obscuri. Receptaculi alveoli margine subulatodentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium armadalense P. D. Sell sp. nov. Holotype: Coastal sandhills, Armadale, W. Sutherland, v.c. 108, 16 July 1915, E. S. Marshall no. 4097 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 25–30 cm, pallide luteoviridis, basin versus brunneopurpureus, magis minusve robustus, in parte inferiore pilis simplicibus eglandulosis paucis longis pallidis, sursum paucis vel nullis, in parte superiore pilis stellatis numerosis et interdum pilis glanduliferis sparsis vestitus. Folia in pagina superiore mediocriter caeruleoviridia, in pagina inferiore pallidiora; folia basalia numerosa, lamina 2.5–10.0 × 1–6 cm, elliptica vel late elliptica vel ovata, plerumque magis minusve acuta, interdum rotundatomucronulata, denticulata vel haud profunde dentata vel dentibus acutis serratodentata, ad basin rotundata vel cuneata, petiolis brevibus, ad 2 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folium caulinum 0–1, quando adest lamina lineari vel linearilanceolata, ad apicem acutum, dentatum, sessile; omnia in pagina superiore glabra vel prope marginem pilis paucis simplicibus eglandulosis rigidis vestita, in pagina inferiore pilis simplicibus eglandulosis paucis brevibus vel mediocribus pallidis inque margine numero-

sioribus vestita. Inflorescentia capitulis 5–12, condensate paniculatocorymbosa, ramo inferiore longo saepe praedita; pedunculi plerumque breves rigidique, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis pilisque glanduliferis sparsis brevibus obscuris vestiti. Capitula 25–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–11 × 1.2–1.5 mm, nigrivirides, interiores margine pallidiore, omnes late linearilanceolatae, ad apicem subacutum angustatae, pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura et pilis glanduliferis sparsis brevibus obscuris et in margine basinque versus pilis stellatis nonnullis vestitae. Ligulae aureoflavae, apicem versus pilis simplicibus eglandulosis nonnullis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine acute dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 6– 7. Apomicticum. Hieracium inspissatum P. D. Sell nom. et stat. nov. H. silvaticum var. crassum Ley in Jour. Bot. (London) 48: 326 (1910), non H. crassum Almq. in Ark. Bot. (Stockh.) 17(17): 54 (1922). Hieracium sinuolatum P. D. Sell sp. nov. Holotype: Grassy, rocky banks of disused railway, near Threshfield, M. W. Yorkshire, v.c. 64, 26 May 1961, C. West (CGE). Ab Hieracio lintonii, quocum invenitur, foliis basalibus regulariter sinuatodentatis folioque nullo caulino magno differt. Herba perennis phyllopoda, caudice ramoso. Caulis 25–50 cm, pallide luteoviridis, in dimidio inferiore saepe brunneopurpureus, gracilis vel satis robustus, in parte inferiore pilis simplicibus eglandulosis numerosis longis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis stellatis numerosis pilisque glanduliferis paucis brevibus obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, colore brunneopurpureo guttata, in pagina inferiore pallida, saepe purpureosuffusa; folia basalia numerosa, lamina 2–11 × 1.5–4.5 cm, subrotundata vel late elliptica vel oblongoelliptica vel elliptica vel ovata, ad apicem rotundatoobtusa vel acuta, in trientibus duobus inferis dentibus brevibus apiculatis regulariter sinuatodentata, dentibus infimis interdum longis angustis curvatis, interdum folia aliquot margine modo denticulato, ad basin rotundata vel subtruncata, petiolis ad 5 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folium caulinum 0–1, etiam in plantis magnis saepe absens sed, quando adest, plerumque lamina lineari bracteiformique praeditum; omnia in paginis ambabus et margine pilis simplicibus eglandulosis brevibus vel mediocribus aliquantum rigidis pallidis vestita. Inflorescentia capitulis 2–11, paniculatocorymbosa; pedunculi breves recti, pilis stellatis densis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis numerosis brevibus tenuibus obscuris vestiti. Capitula 25–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–12 × 1.0–1.2 mm, nigrivirides, interiores margine pallido, omnes anguste linearilanceolatae, ad apicem obtusum vel acutum gradatim angustatae, pilis simplicibus eglandulosis numerosis

New taxa and combinations brevibus vel mediocribus pallidis base obscura et pilis glanduliferis numerosis brevibus obscuris et secus marginem, praecipue basin versus, pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem glabrae. Styli obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 5–7. Apomicticum. Hieracium triangularifolium P. D. Sell sp. nov. Holotype: Schistose cliff-ledges at c. 2,400 ft, Meall Buidhe, near Bridge of Orchy, Argyll, v.c. 98, 27/353433, 20 August 1966, R. W. Jones and B. A. Miles no. 66/264 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 35–45 cm, pallide luteoviridis, basin versus interdum purpurascentitinctus, in parte inferiore pilis simplicibus eglandulosis paucis vel numerosis longis pallidis, sursum multo brevioribus, in parte superiore pilis stellatis numerosis vestitus. Folia in pagina superiore pallide viridia, in pagina inferiore etiam pallidiora et saepe colore purpurascenti tincta vel suffusa; folia basalia pauca, lamina 6–9 × 3.0–5.5 cm, late triangulariovata, ad apicem rotundatoobtusa vel acuta, dentibus acutis apiculatisque dentata vel serratodentata, ad basin trucata vel subcordata, petiolis longis, ad 7 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folium caulinum 0–1, quando adest lamina parva ovata acuta, acute dentata, petiolatum; omnia in pagina superiore glabra vel pilis simplicibus eglandulosis sparsis pallidis vestita, in pagina inferiore pilis simplicibus eglandulosis dispersis brevibus vel mediocribus pallidis, in margine numerosioribus, vestita. Inflorescentia capitulis 2–4, furcatocorymbosa; pedunculi longi, pilis stellatis numerosis pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura pilisque glanduliferis paucis brevibus obscuris vestiti. Capitula 30–40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 5–14 × 1.2–1.5 mm, nigrivirides, interiores ad marginem pallidiores, omnes linearilanceolatae, ad apicem obtusae, pilis simplicibus eglandulosis numerosis brevibus mediocribusque pallidis base obscura et pilis glanduliferis paucis vel satis numerosis brevibus obscuris et secus marginem apicemque versus pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis paucis brevissimis vestitae. Styli obscuri. Receptaculi alveoli et cypselae non visi. Floret 7–8. Apomicticum. Hieracium elongatifolium P. D. Sell sp. nov. Holotype: Shaded rocks, above the River Pattack, about 21/2 miles east of Loch Laggan, W. Inverness, v.c. 97, 19 July 1916, E. S. Marshall no. 4339 (CGE). Hieracio pictorum perpropinquum, a quo foliis majoribus oblongioribusque et involucri squamis acutis differt. Herba perennis phyllopoda, caudice ramoso. Caulis 35–50 cm, pallide luteoviridis, infra interdum rubellitinctus, aliquantum gracilis flexuosusque, striatus, in parte superiore pilis simplicibus eglandulosis paucis mediocribus vel longis pallidis pilisque stellatis paucis pilisque glanduliferis paucis brevibus obscuris vestitus. Folia in pagina superiore mediocriter viridia, venis interdum pallidioribus, in pagina inferiore pallidiora et glauciuscula; folia basalia

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plerumque numerosa, exteriora lamina 2–9 × 2–4 cm, elliptica vel oblongoelliptica vel oblonga, ad apicem rotundatomucronulata, in dimidio inferiore subintegra vel denticulata vel dentata, dentibus inferis interdum mammiformibus curvatis, ad basin abrupte contracta vel subtruncata, interiora lamina 8–14 × 3.0–5.5 cm, oblongolanceolata vel oblonga, ad apicem acuta, in dimidio superiore denticulata vel haud profunde dentata, in parte inferiore dentata, dentibus obtuse vel acute mammiformibus, plerumque angustis et saepe curvatis, ad basin abrupte contracta vel subtruncata (interdum asymmetrice), dentibus angustissimis interdum in petiolum extensis, petiolis ad 8 cm, pilis simplicibus eglandulosis numerosis longis pallidis vestitis; folium caulinum absens aut lamina lineari bracteiformique praeditum; omnia in pagina superiore glabra vel fere glabra, sed pilis simplicibus eglandulosis paucis mediocribus vel longis pallidis trans paginam inferiorem dispersis, in costa et margine numerosioribus, vestita. Inflorescentia capitulis 2–9, aliquantum aperte paniculatocorymbosa; pedunculi satis longi curvatique, graciles, pilis stellatis densis pilisque glanduliferis numerosis brevissimis vel brevibus (0.2–0.5 mm) obscuris pilisque simplicibus eglandulosis sparsis brevibus vel mediocribus pallidis vestiti. Capitula 30–35 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 5–11 × 1.0–1.2 mm, triste cinereivirides, interiores margine pallidiore, omnes linearilanceolatae, ad apicem anguste acutae, pilis simplicibus eglandulosis numerosis brevibus mediocribusque pallidis base obscura, cum pilis glanduliferis brevissimis vel brevibus (0.2–0.5 mm) obscuris immixtis, et secus marginem adque apicem pilis stellatis numerosis vestitae. Ligulae flavae, interdum male evolutae, ad apicem glabrae. Styli flavi. Receptaculi alveoli margine breviter dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium mariae P. D. Sell sp. nov. Holotype: Small cliff in Eathie Burn gorge, Black Isle, E. Ross, v.c. 106, M. McCallum Webster no. 11,255b (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 30–60 cm, pallide luteoviridis, infra interdum purpurascentitinctus, gracilis vel magis minusve robustus, striatus, in parte inferiore pilis simplicibus eglandulosis satis numerosis longis pallidis, sursum paucioribus brevioribusque, in parte superiore pilis stellatis numerosis pilisque glanduliferis paucis brevibus obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora; folia basalia pauca vel numerosa, exteriora lamina 2.5– 9.0 × 1.5–4.5 cm, late elliptica vel fere subrotunda vel obovata, ad apicem rotundatomucronulata, subintegra vel sinuatodenticulata, ad basin rotundata vel subtruncata, interiora lamina 5–11 × 3.0–4.0 cm, anguste lateve elliptica vel ellipticooblonga vel oblonga, ad apicem magis minusve acuta, denticulata, interdum ad basin rotundatam vel subtruncatam dentibus paucis haud profundis praedita, petiolis ad 8 cm, pilis simplicibus eglandulosis numerosis longis sinuatis pallidis vestitis; folium caulinum 0–1, quando adest lamina linearilanceolata longe acuta denticulata, ad basin cuneata, petiolatum, aut bracteiforme; omnia in pagina superiore pilis simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis vestita, aut interdum fere glabra, in pagina inferiore, praecipue in costa, et in margine pilis

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New taxa and combinations

simplicibus eglandulosis numerosis mediocribus vel longis pallidis vestita. Inflorescentia capitulis 4–numerosis, paniculatocorymbosa; pedunculi longitudine variabiles, graciles, pilis stellatis densis pilisque glanduliferis numerosis brevissimis vel brevibus vel mediocribus obscuris pilisque simplicibus eglandulosis paucis brevibus vel mediocribus pallidis base obscura vestiti. Capitula 25–35 mm diametro, ad basin rotundatam angustata. Involucri squamae ante anthesin incumbentes, 4–11 × 1.0–1.2 mm, obscure olivaceae, interiores margine pallidiore, omnes linearilanceolatae, ad apicem obtusae, pilis glanduliferis numerosis brevissimis brevibusque obscuris et pilis simplicibus eglandulosis numerosis brevibus mediocribusque pallidis base obscura et ad apicem basinque versus pilis stellatis nonnullis vestitae. Ligulae flavae, ad apicem interdum pilis paucis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine incisodentato. Cypselae 3.0–3.5 mm, nigrirubrae. Floret 6–7. Apomicticum. In memoriam Mariae McCallum Webster (1906–85) ob contributionem phytologiae scoticae magnam nominatur. Hieracium boreoanglicum P. D. Sell sp. nov. Holotype: Buckden Glen, Wharfedale, M. W. Yorkshire, v.c. 64, 30 June 1904, A. Ley (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 30–55 cm, pallide luteoviridis, ad basin interdum purpureotinctus, gracilis vel robustus, striatus, in parte inferiore pilis simplicibus eglandulosis paucis longis pallidis, sursum numerosioribus sed brevioribus, in parte superiore pilis stellatis numerosis pilisque glanduliferis sparsis brevibus obscuris vestitus. Folia in pagina superiore triste caesioviridia, in pagina inferiore pallidiora et saepe rubellitincta, satis crassa; folia basalia pauca vel numerosa, exteriora lamina 2.5–9.0 × 2.0–3.5 cm, ovata vel elliptica vel oblongoelliptica, ad apicem rotundatomucronulata, subintegra vel denticulata vel dentata, dentibus haud profunde mammiformibus, ad basin subtruncata, interiora lamina 7–12 × 2.5–4.0 cm, plerumque ovatooblonga vel oblonga, interdum lanceolata, ad apicem acuta vel breviter caudata, dentata, infra saepe dentibus magnis mammiformibus vel aquilinomammiformibus in petiolum aliquando decurrentibus, ad basin subtruncata, petiolis ad 12 cm, pilis simplicibus eglandulosis densis longis sinuatis pallidis vestitis; folium caulinum 0–1, quando adest lamina lanceolata, ad apicem acuta, dentata, petiolatum; omnia in pagina superiore pilis simplicibus eglandulosis sparsis brevibus vel mediocribus pallidis vestita vel fere glabra, in pagina inferiore inque margine pilis simplicibus eglandulosis numerosioribus, in pagina inferiore saepe pilis stellatis numerosis vestita. Inflorescentia capitulis 3–10, furcatocorymbosa, ramo inferiore longo interdum praedita; pedunculi satis crassi, recti vel curvati, pilis stellatis densis pilisque glanduliferis numerosis brevissimis vel brevibus (0.2–0.5 mm) obscuris pilisque simplicibus eglandulosis numerosis brevibus vel mediocribus pallidis base obscura vestiti. Capitula 30–35 mm diametro, ad basin rotundata vel subtruncata. Involucri squamae ante anthesin porrectae, 5–12 × 0.8–1.0 mm, obscure virides, interiores margine pallidiore, omnes lanceolatooblongae, ad apicem magis minusve acutae, pilis glanduliferis numerosis maxime inaequalibus brevissimis

brevibusque mediocribusque (0.2–1.2 mm) obscuris et pilis simplicibus eglandulosis paucis vel numerosis brevibus vel mediocribus pallidis base obscura et in margine pilis stellatis numerosis vestitae. Ligulae flavae, ad apicem pilis simplicibus eglandulosis numerosis brevissimis vestitae. Styli obscuri. Receptaculi alveoli margine acute dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium neomicracladium P. D. Sell nom. nov. H. murorum subsp. micracladium Zahn in Engler, Pflanzenreich IV.280: 29 (1921); H. micracladium (Zahn) Omang, Vidensk. Skrift. I. Mat.-Naturv. Klasse 1924(2): 194 (1924), non (F. N. Williams) Ley, Rep. Bot. Soc. Exch. Cl. Brit. Isles 2: 29 (1905). Hieracium protentum P. D. Sell nom. et stat. nov. H. aggregatum var. prolongatum F. Hanb. in Jour. Bot. (London) 32: 228 (1894). Hieracium ronasii P. D. Sell sp. nov. Holotype: Among rocks high up on the south side of Ronas Voe, opposite Heylor, Shetland Islands, v.c. 112, HU/2981, 5 July 1964, C. J. Cadbury and W. Scott no. 20 (CGE). Herba perennis phyllopoda, caudice ramoso. Caulis 40– 60 cm, pallide luteoviridis, basin versus interdum purpurascens, magis minusve robustus, ad basin extremam pilis simplicibus eglandulosis numerosis longis sinuatis pallidis, sursum paucis vel nullis, in parte superiore pilis stellatis paucis pilisque glanduliferis paucis brevibus obscuris vestitus. Folia in pagina superiore aliquantum caeruleoviridia, in pagina inferiore pallidiora; folia basalia numerosa, lamina 4–10 × 2.5–4.0 cm, exteriora ovata vel elliptica, ad apicem rotundatoobtusa, denticulata vel dentibus paucis haud profundis praedita, ad basin cuneata, interiora elliptica vel obovatooblonga vel oblonga, ad apicem obtusa vel acuta, denticulata et basin versus plerumque dentibus paucis praedita, ad basin cuneata, petiolis ad 6 cm, pilis simplicibus eglandulosis numerosis longis pallidis vestitis; folia caulina 1–2, inferum basalibus simile, petiolatum, superum (quando adest) bracteiforme; omnia in paginis ambabus inque margine pilis simplicibus eglandulosis magis minusve numerosis brevibus vel mediocribus pallidis vestita. Inflorescentia capitulis numerosis, paniculatocorymbosa; pedunculi plerumque breves, pilis stellatis numerosis pilisque glanduliferis numerosis brevibus obscuris vestiti. Capitula 25–30 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–11 × 1.0–1.2 mm, aliquantum pallide virides, interiores margine pallidiore, omnes linearilanceolatae, magis minusve obtusae, pilis glanduliferis numerosis brevissimis vel brevibus (0.2– 0.5 mm) tenuibus obscuris et secus marginem pilis stellatis paucis vestitae, sine pilis simplicibus eglandulosis. Ligulae flavae, ad apicem glabrae. Styli obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.0–3.5 mm, rubrinigrae. Floret 6–7. Apomicticum. Hieracium adelphicum P. D. Sell nom. et stat. nov. H. praecox subsp. resensitum microgen. fraternum Sudre, Hier. Centre Fr. 80 (1902).

New taxa and combinations Hieracium elevatum P. D. Sell nom. et stat. nov. H. sparsidens var. elatius Ley in Jour. Bot. (London) 48: 326 (1910); H. elatius (Ley) Druce, Brit. Pl. List ed. 2, 68 (1928), non H. elatius Rehmann in Verh. Zool.-Bot. Ges. Wien 45: 332 (1895). Hieracium dowardense P. D. Sell sp. nov. Holotype: Great Doward, Herefordshire, v.c. 36, July 1956, C. West (CGE). Inter Hieracium stenstroemii et H. glevense, a speciebus ambabus foliis subrotundis vel ovatis vel late ellipticis (quorum unum saepe magnum pilaeque simile est), folio caulino (quando adest) plerumque magno, ovato, ad apicem anguste acuto, dentibus magnis praedito, capitulis obscurioribus differt. Herba perennis phyllopoda, caudice ramoso. Caulis 40–70 cm, pallide luteoviridis, basin versus saepe brunneopurpureus, gracilis vel robustus, striatus, in parte superiore pilis simplicibus eglandulosis paucis vel numerosis mediocribus vel longis pallidis pilisque stellatis numerosis pilisque glanduliferis numerosis brevibus obscuris vestitus. Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora, interdum rubritincta; folia basalia plerumque numerosa, exteriora lamina 3–10 × 2–6 cm, quorum unum saltem magnum subrotundumque vel latissime ellipticum, interdum ovatoellipticum, ad apicem rotundatomucronulata, in dimidio inferiore sinuatodentata, in dimidio superiore interdum denticulata, ad basin rotundata vel truncata vel subcordata, interiora lamina 6–9 × 2.5–6.0 cm, plerumque ovata vel elliptica, interdum ellipticoovata, ad apicem rotundatoobtusa vel magis minusve acuta, dentibus mammiformibus vel acute mammiformibus sinuatodentata, ad basin truncata vel subcordata, petiolis ad 9 cm, pilis simplicibus eglandulosis densis longis pallidis vestitis; folium caulinum 0–1, quando adest lamina lanceolata vel ovata, ad apicem acuta vel acuminata, magis minusve dentata, interdum dentibus magnis mammiformibusque praedita, ad basin truncata, petiolatum; omnia in paginis ambabus pilis simplicibus eglandulosis brevibus vel mediocribus pallidis, in margine petioloque longis, vestita. Inflorescentia plerumque capitulis numerosis, ramo inferiore longo saepe praedita, paniculatocorymbosa; pedunculi plerumque breviusculi, pilis stellatis densis pilisque glanduliferis numerosis brevibus (0.2–0.7 mm) obscuris vestiti, sine pilis simplicibus eglandulosis. Capitula 35–40 mm diametro, ad basin rotundata. Involucri squamae ante anthesin incumbentes, 4–13 × 0.8–1.5 mm, nigrivirides, interiores margine pallido, omnes linearilanceolatae, ad apicem obtusae vel acutae, pilis glanduliferis densis maxime inaequalibus brevissimis brevibusque mediocribusque (0.2–1.5 mm) nigris et in margine basinque versus pilis stellatis vestitae, sine pilis simplicibus eglandulosis. Ligulae flavae, ad apicem glabrae. Styli obscuri. Receptaculi alveoli margine breviter dentato. Cypselae 3.5–4.0 mm, rubrinigrae. Floret 5–7. Apomicticum. Hieracium neosparsum (Zahn) P. D. Sell comb. nov. H. sparsum Jord. ex Boreau, Fl. Centre Fr. ed. 3, 2: 415 (1857), non H. sparsum Friv. in Flora (Regensb.) 19: 436

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(1836); H. murorum subsp. neosparsum Zahn in Engler, Pflanzenreich IV.280: 312 (1921). Filago L., Sp. Pl. 972, 1199, 1230 (1753); Gen. Pl. ed. 5, 397 (1754) nom. conserv. subgenus Logfia (Cass.) P. D. Sell stat. nov. Logfia Cass. in Bull. Sci. Soc. Philom. Paris 1819: 143 (1819). Gnaphalium L., Sp. Pl. 850 (1753); Gen. Pl. ed. 5, 368 (1754). subgenus Laphangium (Hilliard & B. L. Burtt) P. D. Sell comb. nov. Pseudognaphalium subgenus Laphangium Hilliard & B. L. Burtt in Bot. Jour. Linn. Soc. 82: 205 (1981). Aster L., Sp. Pl. 872 (1753): Gen. Pl. ed. 5, 373 (1754). Subgenus Galatella (Cass.) P. D. Sell stat. nov. Galatella Cass. in Dict. Sci. Nat. 37: 463, 488 (1825). Subgenus Crinitaria (Cass.) P. D. Sell stat. nov. Crinitaria Cass. in Dict. Sci. Nat. 37: 460, 475 (1825). Subgenus Tripolium (Nees) P. D. Sell stat. nov. Tripolium Nees, Gen. Sp. Aster 10, 152 (1832). Aster tripolium L., Sp. Pl. 872 (1753). var. flosculosus (Gray) P. D. Sell comb. nov. Eurybia maritima var. flosculosa Gray, Nat. Arrang. Brit. Pl. 2: 465 (1821). Conyza floribunda Humb., Bonpl. & Kunth, Nov. Gen. Sp. 4: 73 (1820). var. linearifolia P. D. Sell var. nov. Holotype: Weed in garden of 54, Cambridge Road, Impington, Cambs, v.c. 29, 52/442619, 25 July 1999, P. D. Sell no. 99/269 (CGE) (1 plant on 4 sheets). Bracteae longae et lineares, ultra capitula extensae, ut inflorescentia capitula bracteaeque permixtae esse videatur. Conyza canadensis (L.) Cronquist in Bull. Torrey Bot. Club 70: 632 (1943). var. simplex P. D. Sell var. nov. Holotype: Derelict arable land planted with conifers, by road between Kneesworth and Meldreth, Cambs, v.c. 29, 52/353447, 10 July 1976, P. D. Sell no. 76/189 (CGE). Caulis ad 40 cm; inflorescentia longa angustaque, non magis quam 3 cm lata. Folia non magis quam 5 mm lata, linearia et integra. var. obovoidea P. D. Sell var. nov. Holotype: Car park, St George’s Way, Impington, Cambs, v.c. 29, 52/453635, 3 September 1999, P. D. Sell no. 99/334 and B. & R. Chapman (CGE). Caulis ad 60 cm; inflorescentia obovoidea, id est ad apicem ad 7 cm lata et ad basin angustata. Folia ad 7 mm lata, aliquando dentibus uno vel duobus praedita. var. incisa P. D. Sell var. nov. Holotype: Side of pavement, Melvin Way, Histon, Cambs, v.c. 29, 52/432636, 5 November 1999, P. D. Sell no. 99/407 and B. & R. Chapman (CGE). Caulis ad 100 cm; inflorescentia amplissima, ad 23 cm lata. Folia ad 2 cm lata, inferiora dentibus longis angustis, saepe cuspidatis, praedita.

556

New taxa and combinations

var. robusta P. D. Sell var. nov. Holotype: Grounds of Impington Village College, Cambs, v.c. 29, 52/447631, 3 September 1999, P. D. Sell no. 99/337 and B. & R. Chapman (CGE). Caulis ad 100 cm; inflorescentia amplissima, ad 23 cm lata. Folia ad 2 cm lata, subintegra vel haud profunde dentata. Bellis perennis L., Sp. Pl. 886 (1753). forma disciformis P. D. Sell nom. nov. B. perennis forma discoidea D. C. McClint. in Watsonia 15: 119 (1984), non B. perennis var. discoidea Hus in Amer. Naturalist 45: 648 (1911). Tanacetum balsamita L., Sp. Pl. 845 (1753). var. balsamitoides (Sch. Bip.) P. D. Sell stat. nov. T. balsamitoides Sch. Bip., Tanacet. 51 (1844). Achillea millefolium L., Sp. Pl. 899 (1753). var. densiloba P. D. Sell var. nov. Holotype: Shingle between Thorpeness and Aldeburgh, E. Suffolk, v.c. 25, 62/472592, 14 August 1996, P. D. Sell no. 96/113 and J. G. Murrell (CGE). Caules ad 60 cm, magis minusve hirsuti. Folia ad 7 cm, dense villosa, segmentis densis imbricatis praedita. Inflorescentia corymbis satis numerosis praedita. Chamaemelum nobile (L.) All., Fl. Pedem. 1: 185 (1785). forma duplicatum (Gray) P. D. Sell stat. nov. C. nobile var. duplicatum Gray, Nat. Arr. Brit. Pl. 2: 455 (1821). forma nudum (Gray) P. D. Sell stat. nov. C. nobile var. nudum Gray, Nat. Arr. Brit. Pl. 2: 455 (1821). Anthemis cotula L., Sp. Pl. 894 (1753). var. vectensis (F. N. Williams) P. D. Sell comb. nov. A. foetida var. vectensis F. N. Williams, Prodr. Fl. Brit. 27 (1901). × Tripleurothemis gruetterana (Asch.) P. D. Sell comb. nov. Anthemimatricaria gruetterana Asch. in Deutsch. Bot. Gesell. 9: 99 (1892). Xanthophthalmum coronarium (L.) P. D. Sell comb. nov. Chrysanthemum coronarium L., Sp. Pl. 890 (1753). var. discolor (d’Urv.) P. D. Sell comb. nov. Chrysanthemum coronarium var. discolor d’Urv., Enum. Pl. Ponti-Eux. 112 (1822). Leucanthemum vulgare Lam., Fl. Franc¸. 2: 137 (1778). subsp. ircutianum (DC.) P. D. Sell stat. nov. Leucanthemum ircutianum DC., Prodr. 6: 47 (1838). subsp. cantabricum (Sennen) P. D. Sell comb. nov. Leucanthemum cantabricum Sennen, Diagn. Nouv. Pl. Espagne Maroc. 1928–35: 50 (1936). Matricaria discoidea DC., Prodr. 6: 50 (1838). subsp. occidentalis (Greene) P. D. Sell stat. nov. M. occidentalis Greene in Bull. Calif. Acad. Sci. 2: 150 (1886). Tripleurospermum maritimum (L.) W. D. J. Koch, Synops. Fl. Germ. Helv. ed. 2, 1026 (1845).

subsp. nigriceps P. D. Sell subsp. nov. Holotype: Strip of waste ground on the south side of the cement wall just north of the sea, Walls pier, Walls, Mainland, Shetland, v.c. 112, 68/240492, 4 August 1996, C. D. Preston no. 96/185 and P. M. Hollingsworth (CGE). Herba biennis vel perennis. Caules luteovirides, interdum rubripurpureosuffusi, ascendentes vel erecti. Folia rosularia non quasi dentium peniculus, segmentis 0.6–1.0 mm latis praedita, succulenta. Capitula 45–60 mm diametro. Involucri bracteae late triangulares, marginibus conspicuis atrobrunneis scariosis ut minimum 0.4 mm latis praeditae. Cypselae 2.5–3.5 mm, costis in superficie adaxiali contiguis vel minime sejunctis; glandes oleosae magis quam duplo longiores quam latiores. subsp. vinicaule P. D. Sell subsp. nov. Holotype: Quarry face, Trevose Head, W. Cornwall, v.c. 1, 10/850763, 29 June 1989, P. D. Sell no. 89/180 and J. G. Murrell (CGE). Herba biennis vel perennis. Caules saltem ad medium purpureorubri, ascendentes vel prostrati. Folia rosularia segmentis coarctis 0.5–0.6 mm latis praedita ut quasi dentium peniculus videantur, succulenta. Capitula 30–45 mm diametro. Involucri bracteae oblongae vel anguste triangulares, marginibus pallide brunneis vel brunneis non magis quam 0.3 mm latis praeditae. Cypselae 1.8–2.5 mm, costis in superficie adaxiali contiguis vel minime sejunctis; glandes oleosae magis quam duplo longiores quam latiores. Senecio aquaticus Hill, Veg. Syst. 2: 120 (1761). subsp. ornatus (Druce) P. D. Sell stat. nov. S. aquaticus var. ornatus Druce in Rep. Bot. Soc. Exch. Cl. Brit. Isles 6: 24 (1921). Senecio squalidus L., Sp. Pl. 869 (1753). subsp. rupestris (Waldst. & Kit.) P. D. Sell comb. et stat. nov. S. rupestris Waldst. & Kit., Pl. Rar. Hung. 2: 136 (1803–4). var. pinnatifidus (Evers ex Hegi) P. D. Sell comb. et stat. nov. S. rupestris forma pinnatifidus Evers ex Hegi, Ill. Fl. Mitteleur. 6 (2): 780 (1928). var. bipinnatifidus (Vis.) P. D. Sell comb. nov. S. nebrodensis var. bipinnatifidus Vis., Fl. Dalmat. 2: 69 (1847). var. rupestris (Waldst. & Kit.) P. D. Sell comb. nov. S. rupestris Waldst. & Kit., Pl. Rar. Hung. 2: 136 (1803–4). var. laciniatus (Vis.) P. D. Sell comb. nov. S. nebrodensis var. laciniatus Vis., Fl. Dalmat. 2: 68 (1847). Senecio vulgaris L. Sp. Pl. 867 (1753). var. crassifolius (Rouy) P. D. Sell stat. nov. S. vulgaris subvar. crassifolius Rouy, Fl. France 8: 343 (1903). Brachyglottis × jubar P. D. Sell nothospecies nova Holotype: Garden in High Street, Bassingbourn, Cambs, v.c. 29, 52/332439, 2 July 1997, P. D. Sell no. 97/37 (CGE). Frutex ad 1(–2) m. Rami numerosi, patentes et aliquantum effusi, cinereilanuginosi, ramunculis aliquantum spisse foliosis, initio albilanuginosis, praediti. Folia alterna;

New taxa and combinations lamina (1.5–)3–8 × 2–4 cm, in pagina superiore obscure viridia, infra alba, ovata vel obovata vel elliptica vel oblongoelliptica, ad apicem obtusa, integra vel remote denticulata, ad basin rotundata vel subtruncata vel breviter cuneata, in pagina superiore quum juvenia tomentosa, infra albilanuginosa; petiolus ad 30 mm, lanuginosus. Capitula 20–45 mm diametro, in corymbo laxo terminali ordinata; pedunculi albilanuginosi. Involucri bracteae in serie una principali cum seriebus secundariis brevibus ordinatae, 5–7 × 0.5–1.2 mm, griseivirides, linearilanceolatae vel oblongae, ad apicem obtusae vel subacutae, cinereilanu-

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ginosae. Flores generum duorum, exteriores 10–15, 10– 15 mm, ligulati, ad apicem tridentati, lutei, interiores 6–7 mm, tubulares, ad apicem quinquedentati, lutei vel aurantiaci brunneolescentes. Receptaculum planum, foveatum, sine squamis. Cypselae 2.0–2.5 mm, linearioblongae, sparsim brevipilosae; pappus pilorum gracilium albidorum. Probabiliter = B. compacta (Kirk) B. Nord. × laxifolia (Buchanan) B. Nord. Widely known to gardeners as Senecio greyi cv. Sunshine. Jubar is Latin for sunbeam.

ABBREVIATIONS

Authors of taxa are consistent with the abbreviations in: Brummitt, R. & Powell, C. E. (1992). Authors of plant names. Kew. Abbreviations of journals in the references follow: Lawrence, H. M. et al. (1968). Botanico–Periodicum–Huntianum. Pittsburgh. Metric measurements follow standard abbreviations: µm = micrometre mm = millimetre cm = centimetre m = metre km = kilometre Infraspecific taxa are abbreviated: subsp. = subspecies var. = variety cv. = cultivar

GLOSSARY

abaxial Of a lateral organ, the side away from the axis, normally the lower side. accrescent Becoming larger after flowering, usually applied to the calyx. achene A dry, indehiscent, one-seeded fruit, more or less hard, with a papery to leathery wall. achene pits See receptacular pits. acicle A slender prickle with a scarcely widened base. acladium The first-opening capitulum in the Asteraceae inflorescence. actinomorphic Radially symmetrical, having more than one plane of symmetry. acumen The tip of an acuminate point. acuminate Curved inwards on both sides to a point. Often wrongly used for gradually narrowed to a point (see acute). acute With a point. Gradually narrowed to a point is longacute, but is often called acuminate. adaxial Of a lateral organ, the side towards the axis, normally the upper side. adherent Joined or fused. adnate Joined to another organ of a different kind. aerial Above ground or above water. agamosperm Species without sexual reproduction. alien Not native. Believed on good evidence to have been introduced by Man and now more or less naturalised. allohexaploid Having six sets of chromosomes and three distinct genomes. allopolyploid A polyploid derived by hybridisation between two different species with doubling of the chromosome number. alternate Lateral organs on an axis, one per node, successive ones on opposite sides. Commonly used also to include spiral arrangements. alveoles See receptacular pits. amphimixis Reproducing by seed resulting from normal sexual fusion (adjective amphimictic). amplexicaul Clasping the stem. anastomosing Joining up to form loops, usually referring to veins. anatropous (of an ovule) Bent over against the stalk. androecium The male parts of the flower, the stamens. andromonoecious Having male and bisexual flowers on the same plant. anemophilous Wind-pollinated. angustiseptate A fruit with the septum across the narrowest diameter. annual Completing its life in under 12 months, but often not within one calender year. annular Ring-shaped.

annulus Special thick-walled cells forming part of the opening mechanism of a fern sporangium, often forming a ring. anther The pollen-bearing part of a stamen, usually terminal on a filament. anthesis The time of flowering. aphyllopodous Without basal leaves. apiculate With an apiculus. apiculus A small, broad point at the apex. apogamous See apomictic. apomictic Reproducing by seed not formed by sexual fusion without pollen stimulation (apogamous), or by pollen stimulation (pseudogamous), or sterile and vegetatively over wide areas. appendage A small extra protrusion or extension such as on a petal, sepal or seed. appressed Pressed close to another organ, but not united with it. arachnoid Appearing as if covered with cobwebs. archegonium The structure containing the female sexual cell in many land plants. arcuate Curved so as to form a quarter of a circle or more. aril The succulent covering around a seed, outside the testa, but not the pericarp. arista An awn. aristate Extended into a long bristle. ascending Sloping or curving upwards. asperous Rough to the touch. attenuate Gradually tapering. auricles Small ear-like projections at the base of a leaf, especially in grasses. autogamous Self-fertilising. autopolyploid A polyploid derived from one diploid species, by multiplication of its chromosome sets. autotrophic Neither parasitic nor saprophytic. awn A stiff, bristle-like projection from the tip or back of the glumes and/or lemma in grasses, or from a fruit, usually the indurated style, e.g. Erodium, or less frequently the tip of a leaf. axil The angle between the main and lateral axis. axile Of a placenta formed by the central axis of an ovary, that is connected by septa to the wall. axillary Arising in the axil of a leaf or bract. base-rich Soils containing a relatively large amount of free basic ions, e.g. calcium, magnesium, etc. basic number See chromosomes. basifixed (of anthers) Joined by the base to the filament and not capable of independent movement. beak A narrow, usually apical projection, in this volume particularly at the top of an achene. berry A fleshy fruit, usually several-seeded, without a stony layer surrounding the seeds.

560

Glossary

leaf blade or lamina

axil

opposite perfoliate leaf or connate

petiole

axillary bud

stipule opposite leaves alternate leaves equitant alternate perfoliate leaf

linear

oblong

elliptical

lanceolate

ovate

sagittate

hastate

spathulate

lyrate

terminal leaflet lateral leaflet

lobes acute

long acute pinnate

acuminate

pinnatifid rounded

obtuse

obtuse-mucronate

lobes

rounded pinnatisect

leaf with 3 leaflets

cordate

bipinnate

palmate

palmatifid

truncate

palmatisect Leaves

cuneate

pedate

attenuate

peltate

Glossary

561

bract

raceme

monochasial cymes

spike

bracteole bract dichasial cyme

involucral bract receptacle

panicle capitulum

pedicel

rhachis peduncle

umbel corymb Inflorescences

562

Glossary

free carpels

dorsifixed anthers

stigma

receptacle

style

stamens

perianth segments petal (coloured)

basifixed anthers

ovary (3 united carpels) sepal (green)

connective

limb

anther cell claw

petal free

filament

tube basifixed

petals united

dorsifixed Stamens

Corollas

spur zygomorphic zygomorphic

actinomorphic

superior ovaries

inferior ovaries

half-inferior ovary

Glossary

apical

marginal

563

axile

free-central

parietal

basal with 1 ovule

basal with several ovules

Ovules and placentation

stigma androecium of stamens

anther

style

filament

ovary

gynoecium of more than 1 carpel

receptacle

Hypogynous flower ovary superior

corolla of petals

carpophore

calyx of sepals

hypanthium hypanthium Perigynous flower with cup-shaped hypanthium, ovary superior

Perigynous flower with flat hypanthium, ovary superior

hypanthium

Epigynous flower with tubular hypanthium ovary inferior

Epigynous flower, ovary inferior

564

Glossary

biennial Completing its life cycle in more than one year, but less than two years, not flowering in the first year. bifid Divided into two, usually deeply at the apex. bifurcate Dividing into two branches. biotype A genetically fixed variant of a taxon, particularly to some condition. bird-seed alien An alien introduced as a contaminant of bird-seed. bisexual Of a flower, bearing both sexes. blade The main part of a flat organ, e.g. petal, leaf. bloom A delicate, waxy, easily removed covering to fruit or leaves. bog A community on wet, very acid peat. bract Modified, often scale-like leaf subtending a flower, or less often a branch. bracteate With a bract or bracts. bracteole A supplementary or secondary bract, or a bract once removed. bud-scales Scales enclosing a bud before it expands. bulb A swollen, underground organ consisting of a short stem bearing a number of swollen, fleshy leaf-bases or scales with or without a tunic, the whole enclosing the next year’s bud. bulbil A small bulb or tuber arising in the axil of a leaf or in an inflorescence on the aerial part of the plant. bullate With the surface raised into blister-like swellings. caducous Falling off at an early stage. caesious Lavender blue. caespitose Tufted. calcicole More frequently found upon or confined to soils containing free calcium carbonate. calcifuge Not normally found on soils containing free calcium carbonate. callus A horny region at the base of the lemma in grasses. calyx The sepals as a whole, the outer whorl of the perianth if different from the inner, including the calyx-tube and calyx-lobes. campanulate Bell-shaped, widest at the mouth. campylotropous Having the ovule bent so that the stalk appears to be attached to the side midway between the micropyle and chalata. capillary Hair-like. capitate Head-like, such as in a tight inflorescence, a knoblike stigma or style, or a stalked gland. capitulescence Sometimes used for the inflorescence formed by capitula in Asteraceae. capitulum An aggregate head of sessile flowers in the Dipsacaceae and Asteraceae. capsule A dry, many-seeded dehiscent fruit formed from more than one carpel. carpel One of the units of which the gynoecium is composed; the basic reproductive unit of the Magnoliophyta. One to many per flower; if more than one then separate or fused. carpophore A stalk-like sterile part of a flower between the receptacle and the carpels as in the Apiaceae and Caryophyllaceae. cartilaginous Hard, not green, and easily cut with knife. caryopsis A fruit in the Poaceae with the ovary wall and seed coat united.

casual An alien plant not naturalised. catkin A dense spike of reduced flowers on a long axis, often wind-pollinated. cauline Pertaining to the stem. cell (of an ovary) The chambers into which the ovary may be divided (often each one corresponding to a carpel). chartaceous Of papery texture. chasmogamous Of flowers which open normally. chlorophyll The green colouring matter of leaves, etc. chromosomes Small deeply staining bodies, found in all nuclei, which determine most or all of the inheritable characters of organisms. Two similar sets of these are normally present in all vegetative cells, the number (diploid number 2n) usually being constant for a single species. The sexual reproductive cells normally contain half this number, or a multiple of a common basic number. cilia Small whip-like structures by means of which some sexual reproductive cells swim. ciliate With hairs projecting from the margin. circumscissile Dehiscing transversely, the top of the capsule coming off like a lid. cladode A green leaf-like lateral shoot in the Liliaceae which, as the leaves are reduced to scales, functionally acts as leaves. clavate Club-shaped, slender and thickened towards the apex. claw The narrow part of a flat organ of which the broader part is the blade or lamina. cleistogamous Of flowers that never open and are selfpollinated in the bud stage. Opposite of chasmogamous. clone An area of plant from vegetative spread, or from seeds from an apomictic plant. column A stout stalk formed by fusion of various floral parts in the Orchidaceae, Geraniaceae, etc. columnar Column-like. commissure The faces by which the two carpels are joined together in the Apiaceae. compound Of an inflorescence with the axis branched; of a leaf made up of several distinct leaflets. compressed Flattened. concolorous Of approximately the same colour throughout. cone A compact body composed of an axis with lateral organs bearing spores or seeds, as in the Lycopodiophyta and Pinophyta. cone-scales The lateral organs of a cone. connate Organs of the same kind growing together and becoming joined, though distinct in origin. connective Part of an anther connecting its two halves. connivent Of two or more organs with their bases wide apart, but with their apices approaching one another. contiguous Touching at the edge with no gap between. contorted With each lobe of the perianth overlapping the next with the same edge and appearing twisted. convergent Of more than two organs with their apices closer together than their bases. convolute Rolled together, coiled. cordate A flat object with incurved base on both sides. coriaceous Of a leathery texture.

Glossary corm A short, usually erect and tunicated, swollen underground stem of one year’s duration, that of the next year arising at the top of the old one and close to it. corolla The inner whorls of the perianth, if different from the outer; all the petals, corolla-tube and corollalobes. corona A long or short tube within the perianth segments in the Liliaceae. corymb A raceme with the pedicels becoming short towards the top, so that all the flowers are approximately at the same level. corymbose Corymb-like, but strictly not a corymb but a flat-topped cyme. cotyledon The first leaf or leaves of a plant already present in the seed and usually differing in shape from the other leaves. Cotyledons may remain within the testa or may be raised above the ground and become green during germination. crenate With round teeth on the margins of a flat organ. crenulate The diminutive of crenate. crisped Curled. culm The stem of a grass. cuneate Wedge-shaped, that is flat and narrowed at the base of an organ. cuneiform Wedge-shaped with the thin end at the base of a solid object. cupule A fruit that is a nut surrounded by a husk formed of fused scales in the Fagaceae and Corylaceae. cuspidate Abruptly narrowed to a point. cymbiform Boat-shaped. cyme An inflorescence to which each flower terminates the growth of a branch, the more distal flowers being produced by longer branches lateral to it. cymose In the form of a cyme. cypsela Name sometimes given to achenes in Asteraceae. decaploid See polyploid. deciduous Not persistent, the leaves falling in autumn or petals falling after anthesis. decumbent Lying on the ground but turning up at the ends. decurrent Having the base prolonged down the axis as in leaves where the blade continues down the petiole or stem as a wing. decussate Opposite, with successive pairs at right angles to each other. deflexed Bent sharply downwards. dehiscent Opening naturally to shed its seeds or pores. deltate Shaped like the Greek letter
. dentate With patent teeth at the margin of a flat organ. denticulate With small teeth. depressed-globose Like globose but wider than long. diadelphous With one stamen free, and the rest connate (Fabaceae). dichasium A cyme in which the branches are opposite and more or less equal. didymus Formed of two similar parts attached to each other by a small portion of their surface. digitate See palmate. dimorphic Occurring in two forms. dioecious Having the sexes on different plants. diploid Having two matching sets of chromosomes.

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disc (disk) Anything disc-shaped, such as the fleshy, sometimes nectar-secreting, portion of the receptacle, surrounding or surmounting the ovary. disc flower The central, eligulate flowers on an Asteraceae receptacle. dissected Deeply divided up into segments. distal At the end away from the point of attachment. distichous Arranged in two diametrically opposed rows. divaricate Divided into widely divergent branches. diverging, divergent With organs having their apices further apart than their bases. dominant The chief constituents of a particular plant community, e.g. oaks in an oak wood or heather on a moor. dorsifixed (of anthers) Attached by their back. dorsiventral With a distinct upper side and lower side. drupe A more or less fleshy fruit with one or more seeds each surrounded by a stony layer. dry Not succulent. e- Without, e.g. eglandular, ebracteate. ebracteate Without bracts. ectomycorrhizal An association of roots with a fungus which may form a layer outside the root. effuse Spreading widely. eglandular Without glands of any kind. elaiasome An oily appendage to seeds offering food-bodies to ants. ellipsoid A solid, elliptical in side view; broadly and narrowly can be used as in elliptical. elliptical A flat shape widest in the middle and 1.2–3.0 times as long as wide; if less, broadly so, if more, narrowly so. By some authors the latter shape is called lanceolate. emarginate Shallowly notched at the apex. embryo-sac See gametophyte. emersed Standing above. endemic Confined to one particular area, i.e. in this book to the British Isles. endomycorrhizal An association of roots with a fungus which may form a layer inside the root. endosperm In the Magnoliophyta the nutritive tissue for the embryo in the developing seed; it might or might not remain as the food-store in the mature seed. ensiform A flat, narrow shape broadest in the middle, both ends with straight sides but gradually narrowed. entire The margin of a flat shape not toothed or lobed. entomophilous Insect-pollinated. epicalyx Organs on the outside of a flower, calyx-like but outside and additional to the calyx. epicarpic On the carpels. epicormic Of new shoots, borne direct from the trunk of a tree. epigeal Above ground; in epigeal germination, the cotyledons are raised above the ground. epigynous Of a flower, with an inferior ovary. epipetalous Inserted upon the corolla. equitant Of distichous leaves, folded longitudinally and overlapping in their parts. erect Upright. erecto-patent Between erect and patent. erose Appearing as if gnawed.

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Glossary

escape A plant growing outside a garden, but having spread vegetatively or by seed from one. evergreen Retaining its leaves throughout the year. exceeding Longer than. exserted Protruding from. exstipulate Without stipules. extravaginal Of branches, breaking through the sheaths of the subtending leaves. extrorse (of anthers) Opening towards the outside of the flower. F1 The offspring from a simple cross. falcate Sickle-shaped. false fruit An apparent fruit actually formed from tissue in addition to the real fruit. fasciculate In tight bundles. fastigiate Having upright branches forming a narrow outline. fen A community on alkaline, neutral or slightly acid wet peat. fertile Producing seed capable of germination or (of anthers) containing viable pollen. fibrous roots A root system where there is no main axis. filament The stalk of an anther, the two together forming the stamen. filiform Thread-like. fimbriate With the margins divided into a fringe. fistular Hollow and cylindrical; tube-like. flexuous Wavy. floccose A short, whitish indumentum often of stellate hairs. flore pleno A double flower with many more petals than normal, usually due to conversion of the stamens to petals; in the Asteraceae, a double capitulum with all or many disc flowers converted to ray flowers. floret A small flower. flush Wet ground, often on hillsides, where water flows but not in a definite channel. foliaceous Leaf-like, of an organ not usually a leaf. foliolate With leaflets. follicle A dry, usually many-seeded fruit dehiscent along its side, formed from one carpel. free Separate, not fused to another organ or to one another except at the point of origin. free-central Of a placenta, formed by the central axis of any ovary which is not connected by septa to the wall. fruit The ripe seeds and structure surrounding them, whether fleshy or dry; strictly the ovary and seeds, but often used to include other associated parts such as the fleshy receptacle, as in the rose and strawberry. fugacious Withering or falling off very rapidly. funicle The stalk of the ovary. furcate Forked. fuscous Dull brown. fusiform Spindle-shaped. gametophyte The haploid generations of a plant that bears the true sex organs; in the Pteridophyta the prothallus, in the Pinophyta and Magnoliophyta the pollen grains (male) and embryo-sac (female). gamopetalous Having the petals joined into a tube, at least at the base.

geminate Condition in the genus Hieracium when the first opening capitulum is more or less sessile. geniculate Bent abruptly to make a knee. gibbous With a rounded swelling on one side as on the base of the calyx in Acinos. glabrescent Becoming glabrous; sometimes wrongly used for sparsely hairy. glabrous Hairless, including all types of glands and spines. gland A small globose or oblong vesicle often containing oil, resin or other liquid, sunk in on the surface of or protruding from any part of a plant. When furnished with stalks they are called glandular hairs (see also hairs). glandular Furnished with glands. glaucous Bluish-white or bluish-green. globose Spherical. glumaceous Resembling a glume. glume A flat bract-like organ(s) subtending a flower in Cyperaceae and Poaceae. grain alien Alien introduced as a contaminant of grain. graniculate Consisting of many small knobs. granulose With a fine, sand-like surface texture. gynodioecious Having female and bisexual flowers on the same plant. gynoecium The group of female parts of a flower made up of one or more ovaries, with their styles and stigmas. hairs Generally, hairs can be described in terms of pubescent, tomentose, lanate, hispid, strigose, etc. Usually it is indicated if the hairs are eglandular, glandular, stellate, tufted or plumose. If precise comparisons are made of indumentum it is often diagnostic of a taxon, but very difficult to put into few words. In some critical genera, where the indumentum is a very important character, a more detailed account of the hairs is given. There are two main types of hair, branched and simple. Branched hairs are either more or less stellate, or plumose or subplumose, that is pinnately branched with the branches projecting longer than the diameter of the hair. Simple hairs with a capitate tip are glandular, or when more or less the same throughout the hairs are eglandular. Eglandular hairs usually include those hairs with minute side-projections not longer than the diameter of the hairs, although these are sometimes called denticulate hairs. Two rarer types of hair are medifixed hairs in the form of a T and glochidiate hairs in the form of an anchor. Sessile glands should also be thought of as hairs and some glands are so minutely stalked they are difficult to see. Spines are really rigid hairs and there is every intermediate between a stiff hair, acicle and pricklet, all of which can be gland-tipped, to a large spine. The abundance of hairs is indicated by: few or sparse when the hairs in question form only a small proportion of the total indumentum or are scattered; numerous when the hairs are abundant but separate enough to be individually distinct; dense when they form a continuous indumentum. The length of the hairs is referred to as very short up to 0.3 mm, short 0.3–0.7 mm, medium 0.7–1.5 mm and long 1.5–4.0 mm. If more than 4 mm the length is given. Colour, rigidity, angle and waviness can be added. In those cases when an attempt has been

Glossary made to be very precise in a particular group of plants it is so stated after the generic description. half-epigynous Of a flower, with a semi-inferior ovary. haploid Having only one set of chromosomes, as in gametophytic tissue. hastate Of a leaf, having two spreading basal lobes and a much longer erect central lobe. heath A lowland community dominated by heath or ling, usually on sandy soils with a shallow layer of peat. hemiparasite See parasite. heptaploid See polyploid. herb A plant dying down to ground level each year. herbaceous Not woody, dying down each year; leaf-like as opposed to woody, horny, scarious or spongy. hermaphrodite Bisexual. heterochlamydeous Having the perianth segments in two distinct series which differ from one another. heterophyllous Having leaves of more than two distinct forms. heterosporous Having spores of two sorts, megaspores female and microspores male, as in all Pinophyta and Magnoliophyta and a few Pteridiophyta. heterostylous Having two forms, not sexes, of flowers on different plants, the two sorts with different styles and pollen. hexaploid See polyploid. hilum The scar on a seed where it left its point of attachment. hip The fruit of Rosa for which there appears to be no other name; it is a hypanthium. hirsute Clothed with long stiff hairs. hispid Coarsely and stiffly hairy. homochlamydeous or homolochlamydeous Having all the perianth segments similar. homogamous Having anthers and stigmas that mature simultaneously. homosporous Having spores of approximately the same size, as in most Pteridophyta. hyaline Thin and translucent. hybrid A plant originating by the fertilisation of one species by another. hybrid swarm A series of plants originating by hybridisation between two or more species and subsequently recrossing with the parents and between themselves, so that a continuous series of forms arises. hypanthium The extension of the receptacle above the base of the ovary in perigynous and epigynous flowers. hypogeal Below ground; in hypogeal germination, the cotyledons remain below ground. hypogynous Of a flower with a superior ovary, having the calyx, corolla and stamens inserted at the base of the ovary. imbricate Overlapping at the edges. imparipinnate Pinnate with an unpaired terminal leaflet. impressed Sunk below the surface. incise Cut. included Not exserted. incumbent Incurved. incurved Curved inwards. indehiscent Not dehiscent.

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indumentum The hairy coverings as a whole. indurated Hardened and toughened. indusium A small flap or pocket of tissue covering a group of sporangia in many Pteridophyta. inferior An ovary which is borne below the point of origin of the petals, sepals and stamens and is fused with the receptacle (hypanthium) surrounding it. inflated Of an organ which is dilated, leaving a gap between the organ and its contents. inflexed Bent inwards. inflorescence A group of flowers with their branching system and associated bracts and bracteoles. infundibuliform Funnel-shaped. insertion The position and form of the point of attachment of an organ. intercalary leaves The leaves on the main stem between the topmost branches and the lowest flowers of the terminal spike in Rhinanthus. internode The stem between adjacent nodes. interpetiolar Between the petioles. interrupted Not continuous. intrapetiolar Between the petiole and the stem. intravaginal Within the sheath. introduced A plant which owes its existence in our country to importation by Man, deliberate or not. introgression The acquiring of characteristics by one species from another by hybridisation followed by backcrossing. introrse Of anthers, opening towards the middle of the flower. involucel An involucre at the base of a flower in the Dipsacaceae, formed by the united bracteoles. involucral Forming an involucre. involucre Bracts forming a more or less calyx-like structure round or just below the base of a usually condensed inflorescence, e.g. Anthyllis, Asteraceae. involute With the margins rolled upwards. isodiametric More or less the same distance across in any plane. isomerous Having the same number of parts in two or more different floral whorls. jaculator The indurated funicle in Acanthaceae which acts as an ejector of its seeds. keel A longitudinal ridge on an organ, like the keel of a boat; the lower petal or petals when shaped like the keel of a boat as in Fumaria and Fabaceae. labellum The central inner perianth segment which appears to be the lower one, but actually the uppermost due to the flower twisting through 180◦ and usually different from all the other perianth segments. lacerate Deeply and irregularly cut and appearing as if torn. laciniate Irregularly and deeply toothed. laciniations The narrow divisions when something is laciniate. lamina The blade of a leaf. laminar In the form of a flat leaf. lammas growth Extra, usually abnormal growth, put on in summer by some trees. lanceolate Very narrowly ovate. lanuginose With woolly indumentum.

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Glossary

latex Milky juice. latiseptate Fruit with the septum across the widest diameter. lax Loose or diffuse, not dense. leaflet A division of a compound leaf. leaf-opposed A lateral organ borne on the stem on the opposite side from a leaf, not in a leaf axil as usual. leaf rosette A radiating cluster of leaves often at the base of the stem at soil level. leaf teeth Pointing outwards (dentate), pointing towards the apex (serrate), breast-like with a small nipplelike apex (mammiform), similar to last but narrower and curved upwards (aquiline-mammiform), deeply and jaggedly cut (laciniate). legume A usually dry, mostly many-seeded fruit dehiscent along two sides, formed from one carpel. lemma A bract in the Poaceae borne on the abaxial side of the flower. lenticular Convex on both faces and more or less circular in outline. ligulate Strap-shaped. ligule A minute membranous flap at the base of the leaves of Isoetes and Selaginella; the strap-shaped part of a ligulate flower in the Asteraceae; the short projection in the axil of a leaf in the Cyperaceae and Poaceae. limb The flattened, expanded part of a calyx or corolla the base of which is tubular. linear Long and narrow with more or less parallel margins. lip Part of the distal region of a calyx or corolla sharply differentiated from the rest due to fusion or close association of its parts. lobe Divided substantially, but not into separate leaflets. loculicidal Splitting down the middle of each cell of the ovary. lodicule Two minute scales at the base of the ovary in the Poaceae. long-shoot Stem of potentially unlimited growth, especially in trees or shrubs. lower side The under surface of a flat organ. lunate Crescent moon-shaped. lyrate More or less lyre-shaped. mamillate With small nipple-like projections. marsh A community on wet or periodically wet, but not peaty soil. meadow A grassy field cut for hay. mealy With a floury texture. megasporangium In a heterosporous plant, the sporangia bearing megaspores. megaspore In a heterosporous plant, the female spores which give rise to female gametophytes. meiosis Special form of cell division (in sporangia, pollensacs or ovules) in which the chromosome number is halved, producing haploid spores. membranous Like a membrane in consistency. mericarp A one-seeded portion split off from a syncarpus ovary at maturity. -merous Denoting the number of parts in a structure, e.g. 5-merous, having the parts in fives. microsporangium In a heterosporous plant, the sporangia bearing microspores.

microspore The male spores in a heterosporous plant which give rise to the male gametophytes. midrib The central main vein. moder A top layer of partly decomposed plant remains with associated mineral particles; looser, less acid and less nutrient poor than mor. monadelphous (of stamens) United into a single bundle by the fusion of the filaments. monocarpic Living for one year, flowering, fruiting and then dying. monochasium A cyme with one lateral branch at each node. monochlamydeous Having only one series of perianth segments. monoecious Having male and female organs on the same plant. monomorphic Occurring in one form, not dimorphic or trimorphic. monopodial Of a stem, growing continuously from year to year by the same apical growing point. moor Upland communities, often dominated by heather, on dry or damp but not wet peat. mucilage Mucus. mucilate With mucilage. mucro The tip of a mucronate object. mucronate Having a very short, bristle-like tip. mucronulate The diminutive of mucronate. mull A fertile woodland soil with no raw humus layer. muricate Rough with short firm projections. muticous Without an awn or mucro. mycorrhiza An association of roots with a fungus which may form a layer outside the roots (ectomycorrhizal) or within the outer tissues (endomycorrhizal). naked Devoid of hair or scales, or not enclosed. native A plant growing in an area where it was not put by the hand of Man. naturalised An alien plant which has become selfperpetuating in the British Isles, or a native plant which is transferred to a new locality by Man and is selfperpetuating. nec Nor, nor of. nectariferous Nectar-bearing. nectar-pit A nectariferous pit. nectary Any nectariferous organ, usually a small knob or a modified petal or stamen. nerve See vein. nodding Bent over and pendulous at the tip. node The position of a stem where leaves, flowers or lateral stems arise. nodule A small, more or less globose swelling. non Not, not of. nonaploid See polyploid. nothomorph One of more than two variants of a particular hybrid. nucellus The tissue between the embryo and the integument in an ovule. nut A dry, indehiscent, one-seeded fruit with a hard, woody wall. nutlet A small nut, or a woody-walled mericarp.

Glossary ob- The other way up from normal, usually flattened or widened at the distal rather than the proximal end. obdiplostemonous Having the stamens in two whorls, the outer opposite the petals, the inner opposite the sepals. oblong A flat shape with the middle part parallel-sided, 1.2–3.0 times as long as wide; if less, broadly so, if more, narrowly so. obtuse Blunt. ochreae Sheathing stipules in the Polygonaceae. octoploid See polyploid. opposite Of two organs, arising laterally at one node on opposite sides of the stem. orbicular Like an orb. orthotropous Of an ovule, straight and with the axis of the ovule in the same line as that of the funicle. oval Elliptical. ovary The basal part of the gynoecium containing the ovule. ovate A flat shape widest nearer the base 1.2–3.0 times as long as wide; if less, broadly so, if more, narrowly so. ovoid A solid shape, ovate in side view. ovule An organ inside the ovary of the Magnoliophyta, naked in the Pinophyta, that contains the embryo-sac, which in turn contains the egg developed into a seed after fertilisation. palate A projecting part of the lip closing the mouth of the corolla in Antirrhinum. palea A bract on the adaxial side of a flower in the Poaceae. palmate Consisting of more than three leaflets arising from the same point. panduriform Lanceolate in outline with a dip in each side below the middle and with an obtuse apex. panicle A compound or much-branched inflorescence, either racemose or cymose. papilla A small nipple-like projection. papillose Covered with small warts. pappus The hairs, scales or scarious margin at the top of a fruit (achene or cypsela) in the Asteraceae. parasite A plant that gets all or part of its nourishment by attachment (often under the ground) to another plant. parietal Of a placenta, formed by a central axis of an ovary that is connected by septa to the wall. paripinnate Pinnate without an unpaired terminal leaflet. partial septum A septum that is incomplete. pasture A grassy field grazed during summer. patent Projecting at more or less right angles, spreading. pectinate Lobed with the lobes resembling and arranged like the teeth of a comb. pedate With five leaflets of a leaf arising from the same point, the leaflets obovate. pedicel Flower stalk. peduncle The stalk of a group of flowers. peltate Of a flat organ, with its stalk inserted in the lower surface. pendant Pendulous. pentaploid See polyploid. perennial Living for more than two years and usually flowering each year. perianth The floral leaves including petals and sepals.

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perianth segments, lobes or tube Petals and sepals, usually used when they are not or little differentiated. pericarp The wall of a fruit, originally the ovary wall. perigynous A flower with a superior ovary, but with the calyx, corolla and stamens inserted above the base of the ovary on an extension of the receptacle (hypanthium) that is not fused with the ovary. perigynous zone The annular region between the gynoecium and the other floral parts in perigynous or epigynous flowers. perisperm The nutritive tissue derived from the nucellus in some seeds. perispore A membrane surrounding a spore. persistent Remaining attached longer than normal. petal One of the segments of the inner whorl(s) of the perianth. petaloid Brightly coloured and resembling petals. petiole The stalk of a leaf. petiolate With a petiole. phyllary One of the involucral bracts surrounding the capitulum in the Asteraceae, which in this account are called involucral bracts. phyllode A green, flattened petiole resembling a leaf. pilose Hairy. pinna The primary division of a more than two-pinnate leaf. pinnate A compound leaf, with more than three leaflets arising in opposite pairs along the rhachis; two-pinnate with the pinnae pinnate again. pinnatifid Pinnately cut, but not into separate portions, the lobes being connected by the lamina as well as the midrib or stalk. pinnatipartite Divided pinnately. pinnatisect Like pinnatifid but with some of the lower divisions reaching very nearly or quite to the midrib. pinnule The ultimate division of a more than two-pinnate leaf, usually applied only in ferns. placenta The part of the ovary to which the ovules are attached. placentation The position of the placentae in the ovary. The chief types of placentation are: apical, at the apex of the ovary; axile, in the axils formed by the meeting of the septa in the middle of the ovary; basal, at the base of the ovary; free-central, on a column or projection arising from the base in the middle of the ovary, not connected with the wall by septa; parietal, on the wall of the ovary or on an intrusion from it; superficial, when the ovules are scattered uniformly all over the inner surface of the wall of the ovary. plano-convex Having one surface plane and the other convex. plastic Varying in form according to environmental conditions, not according to genetic characteristics. pollen The microspores of Pinophyta and Magnoliophyta. pollen-sac The microsporangium of a species of Pinophyta or Magnoliophyta; one of the chambers in an anther in which the pollen is formed. pollinia Regularly shaped masses of pollen formed by a large number of pollen grains cohering, as in Orchidaceae.

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Glossary

polygamous Having male, female and bisexual flowers on the same or different plants. polyploid Having more than two sets of chromosomes, e.g. 3, triploid; 4, tetraploid; 5, pentaploid; 6, hexaploid; 7, heptaploid; 8, octoploid; 9, nonaploid; 10, decaploid. pome A fruit in which the seeds are surrounded by tough but not woody or stony layers, derived from the inner part of the fruit wall, and the whole fused with the deeply cupshaped, fleshy receptacle, e.g. apple. porrect Directed outwards and forwards. premorse Ending abruptly and appearing as if bitten off at the lower end. prickle Spiny outgrowth with a broadened base. pricklet A small spiny outgrowth without a broadened base. procumbent Trailing along or loosely lying on the ground. proliferating Inflorescences bearing plantlets instead of flowers and fruits. pro parte Partly; in part. prostrate Lying closely along the surface of the ground. protandrous Having stamens maturing before the ovary. prothallus The small gametophyte generation of a plant bearing the true sex organs, mostly applied to the freeliving gametophytes of Pteridophyta. protogynous Having the ovary maturing before the stamens. proximal At the end near the point of attachment. pruinose With a bloom. pseudogamous See apomictic. pteridophytes Ferns and fern allies, i.e. Lycopodiophyta and Pteridophyta. puberulous With very short hairs. pubescent With short, soft hairs, but sometimes used as a word for generally hairy. punctate Marked with dots or transparent spots. punctiform A small more or less circular dot. punctulate Minutely dotted. pungent Sharply and stiffly pointed so as to prick. raceme An unbranched racemose inflorescence in which the flowers are borne on pedicels. racemose Having an inflorescence, usually conical in outline, whose growing points commonly continue to add to the inflorescence and in which there is usually no terminal flower. A consequence of this mode of growth is that the youngest and smallest branches or flowers are normally nearest the apex. radical Of leaves, arising from the base of the stem or a rhizome. radiate A central region of tubular flowers and an outer region of ligulate flowers in the Asteraceae. rank A vertical file of lateral organs; e.g. 2-ranked etc. with two ranks of lateral organs. raphe The united portions of the funicle and outer integument in a conotropous ovule. ray Anything that radiates outwards, e.g. branches of an umbel; stigma ridges in Papaver or Nuphar. ray flowers The outer ligulate flowers in an Asteraceae capitulum. receptacle The flat, concave or convex part of the stem or peduncle from which the parts of the flower arise;

often used to include the perigynous zone, especially in Asteraceae. receptacular pits The pits in the receptacle of the capitulum in the Asteraceae in which the flowers are seated. receptacular scales Scales or hairs on the receptacle of the capitulum in the Asteraceae adjacent to each flower. recurved Curved down or back. reflexed Bent down or back. regular Actinomorphic. reniform Kidney-shaped. repand Having a slightly uneven waved margin. replum The adjacent wall tissue to the placentae. resilient Springing sharply back when bent out of position. resiniferous Producing resin. reticulate Marked with a network, usually of veins. reticulum A network. retrorse Turned backwards. retuse Notched at the apex. revolute Rolled downwards. rhachis The axis of an inflorescence or pinnate leaf. rhacilla The short, slender axis of the flower in the Poaceae. rhizomatous Bearing a rhizome. rhizome An underground stem lasting more than one growing season. rhombic Having the shape of a diamond in a pack of playing cards. rigid Stiff. rostellum A beak-like process formed by the sterile stigma in the flower of the Orchidaceae. rounded Without a point or angle. ruderal Growing on waste ground. rugose With a wrinkled surface. rugulose Finely rugose. ruminate Looking as though chewed. runcinate Pinnately lobed with the lobes directed backwards towards the base of the leaf. saccate Pouched. sagittate Condition in which the base of a leaf is cut by straight lines upwards on either side from the margin to the petiole to leave an inverted V. salt-marsh The series of communities growing on intertidal mud or sandy mud in sheltered places on coasts and in estuaries. samara A dry indehiscent fruit, part of the wall of which forms a flattened wing. saprophyte A plant deriving its nourishment from decaying organisms. scaberulous Rough with minute projections. scabrid Rough to the touch. scabridulous The diminutive of scabrid. scale-leaf A leaf reduced to a small scale. scape A flowering stem of a plant in which all the leaves are basal with none on the stem. scarious Of thin papery texture and not green. schizocarp A fruit which breaks into one-seeded portions or mericarps. sclerenchyma Woody tissue in a partly or mostly nonwoody organ. scorpioid A monochasial cyme that is coiled up like a scorpion’s tail when young.

Glossary scrambler A plant sprawling over other plants, fences, etc. scrub A community dominated by shrubs. secund All directed towards one side. seed A fertilised ovule. self-compatible Self-fertile; able to self-fertilise. self-incompatible Self-sterile; not able to self-fertilise. semi-inferior Of an ovary, having the lower part inferior but the upper part free and projecting above the sepals, etc. sensu lato In the broad sense. sensu stricto In the narrow sense. sepal One of the segments of the outer whorls of the perianth. sepaloid Resembling sepals. septicical Dehiscing along the septa of the ovary. septum A wall of membrane dividing the ovary into cells. sericeous With silky, appressed, straight hairs. serrate Toothed with the teeth pointing towards the apex. serration With serrate teeth. sessile Not stalked. setaceous Shaped like a bristle, but not necessarily rigid. sheath A long stem, sheathing and often cylindrical round the lower part of a leaf in the Poaceae. short-shoot A short stem of strictly limited growth usually lateral on a long-shoot, especially on trees and shrubs. shrub A woody plant branching abundantly from the base and not reaching a very large size. silicula A dehiscent, two-valved, two-celled capsule less than three times as long as wide in the Brassicaceae. siliqua A dehiscent, two-valved, two-celled capsule more than three times as long as wide. simple Not compound. sinuate Having a wavy outline. sinus The space of indentation between a lobe or teeth; the space at the base of a leaf both sides of the petiole. solitary Borne singly. sorus A group of sporangia in the Pteridophyta. spadix Sterile axis on which the flowers of an araceous inflorescence are packed, which often extends distinctly as a succulent appendix. spathe An ensheathing bract in the Lemnaceae, Araceae and Hydrocharitaceae. spathulate Paddle- or spoon-shaped. spermatophyte A seed plant belonging to the Pinophyta or Magnoliophyta. spermatozoid A male reproductive cell capable of moving by means of cilia. spike A racemose inflorescence in which the flowers (or spikelets in Poaceae) have no stalks. spikelet One to many flowers in a discrete group in Limonium, Cyperaceae and Poaceae. spine A sharp, stiff, straight, woody outgrowth, usually not greatly widened at the base (see hairs). spinose Spine-like. spiny With spines. spiral The arrangement of lateral organs on the axis, one per node, successive ones not at 180◦ to each other. sporangiophore A structure, not leaf-like, bearing sporangia. sporangium A structure containing spores.

571

spore The haploid product of meiotic division produced on the sporophyte and developing into the gametophyte. sporophyll A leaf-like structure or one regarded as homologous with a leaf, bearing sporangia. spreading Growing out divergently, not straight or erect. spur A protrusion or tubular or pouch-like outgrowth of any part of the flower. squarrose Rough with scales or tips of bracts projecting outwards. stamen The basic male reproductive unit of the Magnoliophyta, one to many per flower, sometimes fused. staminode A sterile stamen, sometimes modified to perform some other function. standard The large, often erect adaxial petal of the zygomorphic flowers of the Fabaceae. stellate Star-shaped with radiating arms. stem-leaves Leaves borne on the stem as opposed to basally. sterile Not producing seed capable of germination, or anthers with viable pollen. stigma The receptive surface of the gynoecium to which the pollen grains adhere. stipel A structure similar to a stipule but at the base of the leaflets of a compound leaf. stipitate Having a short stalk or stalk-like base. stipule A scale-like or leaf-like appendage usually at the base of a petiole, sometimes adnate to it. stipulate With stipules. stolon An aerial or procumbent stem, usually not swollen. stoloniferous With stolons. stoma (pl. stomata) A pore in the epidermis which can be closed by changes in shape of the surrounding cells. stomium The part of the sporangium wall in the ferns, which ruptures during dehiscence. striate Marked with long narrow depressions or ridges. strict Growing up at a small angle to the vertical. strigose With stiff, appressed hairs. strophiole A small, hard appendage outside the testa of a seed. style The part of the gynoecium connecting the ovary with the stigma. stylopodium The enlarged base of the style in the Apiaceae. sub- Almost, as in subacute, subglabrous, subglobose, subentire, subequal. subsetiform With slightly stiff hairs. subshrub A perennial with a short woody surface stem producing aerial herbaceous stems. subtended Of a lateral organ, having another organ in its axil. subulate Awl-shaped, narrow, pointed and more or less flattened. succulent Fleshy and juicy or pulpy. sucker A shoot arising adventitiously from the root of a tree or shrub often at some distance from the main stem. suffruticose A dwarf shrub or undershrub. superior Any ovary that is borne above the calyx, corolla and stamens, or if below or partly below them then not fused laterally to the receptacle. suture The line of junction of two carpals.

572

Glossary

sympatric Having overlapping ranges. sympodial Of a stem, having the growing point either terminating in an inflorescence or dying each year, growth being continued by a new lateral growing point. syncarpous Of an ovary, having the carpels united to one another. tap-root A main descending root bearing laterals. taxon Any taxonomic grouping such as family, genus or species. tendril A spirally coiled, thread-like outgrowth from a stem or leaf, used by the plant to climb and support itself. tepal One of the segments of the perianth; term sometimes used when sepals and petals are not differentiated. terete Round, not ridged, grooved or angled. terminal Borne at the end of a stem and limiting its growth. ternate A compound leaf with three leaflets, which may be similarly divided again, e.g. 2-ternate, etc. testa The outer coat of a seed. tetrad A group of four spores cohering in a tetrahedral shape or as a flat plate and originating from a single spore mother cell. tetragonous Four-angled. tetraploid See polyploid. tetraquetrous Square in section. thallus The plant body when not differentiated into a stem, leaf, etc. thorn A woody, sharp-pointed structure formed from a modified branch. throat The opening where the tube joins the limb of the corolla or calyx. thyrsoid Like a thyrse, an ovoid or ellipsoid panicle with cymose branches. tiller Leafy shoot. tomentose Having a dense covering of short cottony hairs. tooth A shallow division of a leaf, calyx or corolla, or the apex of a capsule. transverse Lying crossways. tree A woody plant usually more than 5 m with a single trunk. triangular Having a flat shape with three sides, widest at the base and 1.2–3.0 times as long as wide with the two sides gradually narrowing to a point. trifid Split into three, but not to the base. trifoliolate A term used in the Fabaceae for ternate. trigonous Triangular in section, with obtuse to rounded angles. trimorphic Occurring in three forms. tripartite Divided into three parts. triploid See polyploid. triquetrous Having a solid body triangular in section and acutely angled. trullate Having a flat shape, widest nearer the base and more or less angled (not rounded) there, 1.2–3.0 times as long as wide; if less, broadly so, if more, narrowly so.

truncate Of the base or apex of a flat organ, straight or flat. tube The fused part of a corolla or calyx, or a hollow, cylindrical, empty prolongation of an anther. tuber Swollen roots or subterraneous stems. tuberous Tubercle-like. tubercle A small more or less spherical or elliptical swelling. tuberculate With a surface texture covered in minute tubercles. tubular In the form of a hollow cylinder. tufted Of elongated organs or stems, clustered together. tunic A dry, usually brown and more or less papery covering round a bulb or corm. turbinate Top-shaped. turion A detachable winter-bud, by means of which many water-plants perennate. twig Ultimate branch of a woody stem. umbel An inflorescence in which all the pedicels arise from one point. unarmed Devoid of thorns, spines or pricklets. undulate Wavy at the edge in the plane at right angles to the surface. unguiculate Clawed. unifacial With only one surface, not with a lowerside and underside. unilocular Having a single cavity. unisexual Of a flower, bearing only one sex. urceolate More or less globular to cylindrical but strongly contracted at the mouth. upperside The upper surface of a flat organ. vallecular Furrowed between two ridges. valve A deep division or lobe or a lobe of a capsule apex. valvate Of perianth segments, having their edges in contact with each other but not overlapping in bud. vein A strand of vascular tissue consisting of more than one vascular bundle. verrucose Covered in small wart-like outgrowths. villous Shaggy. viscid Sticky. viscidium Two viscid bodies to which the pollinia are attached in the Orchidaceae. vitta (pl. vittae) The resin canals on the fruits of Apiaceae. viviparous With flowers proliferating vegetatively and not forming seed. waste place Uncultivated more or less open habitat much influenced by Man. whorl More than two organs of the same kind arising at the same level. wing Extension of an organ. woody Hard and wood-like. wool alien An alien introduced as a contaminant of raw wool imports. woolly Clothed with shaggy hairs. zygomorphic Having only one plane of symmetry.

Index

Accepted latin names and the page numbers on which the account of them occurs are in bold type. Synonym latin names are in italic. Vernacular names are in roman. When a vernacular name is the same as the latin genus, the latin genus and species are placed first. Absinthium Hill vulgare Lam., 467 Achillea L., 470 ageratum L., 473 × decolorans Schrad., 471 distans Waldst. & Kit. ex Willd., 471 subsp. tanacetifolia Janch., 472 eupatorium M. Bieb., 473 filicifolia M. Bieb., 473 filipendulina Lam., 473 ligustica All., 472 millefolium L., 472, 556 var. alpestris auct., 472 var. compacta Br´eb., 472 var. conspicua Druce, 472 var. densiloba P. D. Sell, 472, 556 var. lanata auct., 472 var. millefolium, 472 var. villosa Hartm., 472 nobilis L., 472 subsp. nobilis, 472 ptarmica L., 471 forma ligulosa Vilm., 471 × macrophylla L., 471 forma ptarmica, 471 serrata Sm., 471 sylvestris Gray, 471 tanacetifolia All., 472 tenuifolia Salisb., 472 tomentosa L., 473 Achyrophorus Adans., 98 maculatus (L.) Scop., 100 radicatus (L.) Scop., 99 Acosta Adans., 86 Acrolophus Cass., 86 paniculatus (L.) Cass., 86 Acroptilon Cass., 84 picris (Pall. ex Willd.) C. A. Mey., 84 repens (L.) DC., 84 ADOXACEAE, 48 Adoxa L., 48 moschatellina L., 48 tuberosa Gray, 48 Adventina Raf., 520 ciliata Raf., 521 parviflora (Cav.) Raf., 520 Aetheorhiza Cass., 111 bulbosa (L.) Cass., 111 African Thistle, Lobed, 423 Rigid, 424 Variable, 423

Agathysus D. Don tataricus (L.) D. Don, 118 Ageratum L., 528 conyzoides L. var. mexicanum (Sims) DC., 528 houstonianum Mill., 528 mexicanum Sims, 528 Ake-ake, 461 Akiraho, 460 Akylopsis Lehm. suaveolens (Pursh) Lehm., 484 Amberboa (Pers.) Less. moschata (L.) DC., 84 Ambrosia L., 513 aptera DC., 514 artemisiifolia L., 513 coronopifolia Torr. & A. Gray, 513 elatior L., 513 integrifolia Muhlenb., ex Willd., 513 longistylis Nutt., 513 media Rydb., 513 psilostachya DC., 513 striata Rydb., 514 trifida L., 514 variabilis Rydb., 514 Amellus Adans. microglossus DC., 440 strigosus (Thunb.) Less., 440 Ammobium R. Br. ex Sims alatum R. Br., 432 Anacylus L. australis Sieber ex Spreng., 487 Anaphalis DC., 428 margaritacea (L.) Benth., 428 var. angustior (Miq.) Nakai, 428 var. intercedens Hara, 428 var. margaritacea, 428 var. occidentalis Green, 428 var. revoluta Suksd., 428 var. subalpina A. Gray, 428 triplinervis (Sims) C. B. Clarke, 428 Andryala L., 208 integrifolia L., 209 Andryala, Common, 209 Anisanthus Willd. ex Roem. & Schult., 41 microphyllus (Kunth) Willd. ex Roem. & Schult., 42 Anisoderis Cass., 205 foetida (L.) Fisch. & C. A. Mey., 205 Antennaria Gaertn., 427 dioica (L.) Gaertn., 427 var. dioica, 427

574 Antennaria dioica (L.) Gaertn. (cont.) var. hyperborea (D. Don) DC., 427 var. pedicellata F. B. White, 427 hibernica Braun-Blanq., 427 hyperborea D. Don. 427 margaritacea (L.) Gray, 428 montana Gray, 427 triplinervis Sims, 428 × Anthemimatricaria P. Fourn. celakovskyi Geisenh. ex Domin, 478 maleolens P. Fourn., 478 gruettierana Asch., 478, 556 Anthemis L., 474 altissima L., 477 anglica Spreng., 476 arvensis L., 475 var. anglica (Spreng.) Syme, 476 subsp. arvensis, 476 var. arvensis, 476 × Tripleurospermum maritimus subsp. inodorum, 478 austriaca Jacq., 477 cota L., 477 cotiformis Velen., 477 cotula L., 476, 556 var. cotula, 476 var. maritima auct., 476 × Tripleurospermum maritimus subsp. inodorum, 478 var. vectensis (F. N. Williams) P. D. Sell, 476, 556 cupaniana Tod. ex Nyman, 475 discoidea (All.) With., 477 foetida Lam., 478 var. vectensis F. N. Williams, 478, 556 grandiflora Ramat., 482 hyrcana Sosn. ex Grossh., 477 mixta L., 474 montana L. subsp. cupaniana (Tod. ex Nyman) Arcang., 475 neilreichii Ortmann, 476 nobilis L., 474 pontica auct., 476 punctata Vahl., 475 subsp. cupaniana (Tod. ex Nyman) R. Fevn., 475 var. incana Guss., 475 var. sicula Guss., 475 ruthenica M. Bieb., 476 subtinctoria Dobrocz., 477 tinctoria L., 477 subsp. australis R. Fern., 477 var. discoidea (All.) DC., 477 subsp. fussii (Griseb.) Beldie, 477 var. fussii Griseb., 477 subsp. subtinctoria (Dobrocz.) So´o, 477 subsp. tinctoria, 477 var. tinctoria, 477 zangelana Sosn. ex Grossh., 477 Anthemis × Tripleurospermum, 478 Apargia Scop. autumnalis (L.) Hoffm., 101 var. alpina Gaudin, 102, 531 var. salina Aspegren, 531

Index Apargia Scop. (cont.) hirta auct., 103 hispida (L.) Hoffm., 103 pratensis Hornem., 102, 531 taraxaci (L.) Willd., 102 Aparine Guettard boreale (L.) Hill, 22 latifolia Moench, 28 vulgaris Hill, 26 Apuleje Gaertn. heterophylla (Thunb.) Less., 423 zeyheri Less., 424 Aracium Monnier attenuatum Opiz ex Domin, 203 paludosum (L.) Monnier, 202 Arctium L., 70 austriacum Teyber, 71 bardana Willd., 72 batavum Ar`enes, 71 crispum Cretz., 72 debrayi Senay, 71 intermedium Lange, 71 lappa L., 70, 530 subsp. minus (Hill) Hook. fil., 71 subsp. nemorosum (Lej.) P. D. Sell, 71, 530 subsp. pubens (Bab.) P. D. Sell, 71, 530 majus Bernh., 71 subsp. intermedium (Lange) Nyman, 71 subsp. nemorosum (Lej.) Rouy, 71 minus (Hill) Bernh., 71 subsp. nemorosum (Lej.) Syme, 71, 530 subsp. pubens (Bab.) Ar`enes, 71, 530 nemorosum Lej., 71, 530 newbouldii A. Benn., 71 notha (Ruhmer) J. Wiess, 71 personata Mill., 71 pubens Bab., 71, 530 ruderale Salisb., 71 subracemosum (Simonk.) Nyman, 71 tomentosum Mill., 72 var. glabrum (K¨orn.) Ar`enes, 72 vulgare (Hill.) Druce, 71 Arctotheca J. C. Wendl., 422 calendula (L.) Levyns, 422 calendulacea (Hill) Lewin, 422 Arctotis L., 421 breviscapa Thunb., 422 calendula L., 422 calendulacea Hill, 422 stoechadifolia Bergius, 422 var. grandis (Thunb.) Less., 422 Arctotis, White, 422 Arnoseris Gaertn., 96 minima (L.) Schweigg. & K¨orte, 96 pusilla Gaertn., 96 Artemisia L., 465 abrotanum L., 468 absinthium L., 467 afra Jacq., 465 anethifolia Weber, 465

Index Artemisia L. (cont.) annua L., 468 biennis Willd., 468 brittonii Rydb., 467 campestris L., 469 candicans Rydb., 467 diversifolia Rydb., 467 dracunculoides Pursh, 469 dracunculus L., 468 falcata Rydb., 467 glabescens Stokes, 469 glauca Pall. ex Willd., 468 gnaphalodes Nutt., 467 herba-alba Asso, 465 herbacea Willd., 468 herriotii Rydb., 467 incompta Nutt., 467 inodora Willd., 468 lindheimeriana Scheele, 467 ludoviciana Nutt., 467 var. candicans (Rydb.) D. D. Keck, 467 maritima L., 469 norvegica Fr., 468 paniculatum Lam., 468 papularis (A. Nelson) Rydb., 467 platyphylla Rydb., 467 pontica L., 465 procera Willd., 468 purshiana Besser, 467 redowskyi Ledeb., 468 scoparia Waldst. & Kit., 465 serrata Nutt., 467 stelleriana Besser, 467 verlotiorum Lamotte, 466 × vulgaris, 467 vulgaris L., 466 var. coarctata Forselles, 466 subsp. ludoviciana (Nutt.) H. M. Hall & Clem., 467 var. vulgaris, 466 × wurzellii C. M. James & Stace, 467 Ascalea Hill, 73 lanceolata (L.) Hill, 77 nutans (L.) Hill, 75 Asperula L., 16 arvensis L., 18 azurea Jaub. & Spach, 18 caucasica Pobed., 17 ciliata auct., 16 cynanchica L., 17 var. densiflora Gren. & Godr., 17 var. maritima Lange, 17 subsp. occidentalis (Rouy) Stace, 17 minor Gray, 17 occidentalis Rouy, 17 odorata L., 19 orientalis Boiss. & Hohen., 18 propinqua Pobed., 17 sherardii H¨ock ex Hallier, 16 taurina L., 17 tinctoria L., 17

Aster L., 443, 555 amelloides Besser, 444 amellus L., 444 annuus L., 455 bellidiflorus Willd., 448 bonariensis Spreng., 453 brevibracteatus Rydb., 447 braumalis Nees, 447 californicus Less., 453 canadensis (L.) Krause, 458 canadensis (L.) Kuntze, 441 chinensis L, 459 concinnus Willd., 446 conyzae Griess., 435 cordifolius L., 445 discoideus Sond., 452 dracunculoides Willd., 448 dumosus L., 446 var. dumosus, 446 var. strictior Torr. & A. Gray, 446 durus Lunell, 448 eminens Willd., 448 ericoides auct., 445 ericoides L., 450 var. ericoides, 450 var. pansus (Blake) Boivin, 450 firmus Nees, 449 floribundus auct, 447 fluvialis Osterh., 448 foliaceus Lindl., 449 foliosus Pers, 446 geyeri (A. Gray) Howell, 447 hesperius A. Gray, 448 var. wootonii Greene, 448 hispidus Lam., 449 interior Wiegand, 448 laevigatus Lam., 447 laevis L., 446 var. concinnus (Willd.) House, 446 var. geyeri A. Gray, 447 var. laevis, 447 × novi-belgii, 447 lanceolatus Willd., 448 subsp. hesperius (A. Gray) Semple & Chmiel., 448 subsp. interior (Wiegand) A. G. Jones, 448 var. hirsuticaulis Semple & Chmiel., 448 var. interior (Wiegand) Semple & Chmiel., 448 subsp. lanceolatus, 448 var. lanceolatus, 448 var. latifolius Semple & Chmiel., 448 × novi-belgii, 448 latissimifolius Kuntze var. serotina Kuntze 442 lautus Lunell, 448 laxus Willd., 448 linosyris (L.) Bernh., 451 var. minor (Wallr.) Rouy 451 longifolius auct., 447 macrophyllus auct., 445 maritimus Salisb., 451

575

576 Aster L. (cont.) multiflorus Aiton, 450 novae-angliae L., 450 novi-belgii L., 447 subsp. laevigatus Lam., 447 osterhoutii Rydb., 448 paniculatus Lam., 448 paniculatus Willd., 445 pansus Blake, 450 pickettiana Suksd., 447 pilosus Willd., 445 var. demotus Blake, 446 var. pilosus , 446 var. pringlei (A. Gray) Blake, 446 pringlei A. Gray, 446 pubentior Cronquist, 449 punctatus Waldst. & Kit., 450 puniceus L., 449 recurvatus Willd., 448 salicifolius Aiton, 448 × salignus Willd., 448 schreberi Nees, 445 sedifolius L., 450 simplex Willd., 448 var. interior (Wiegand) Cronquist, 448 solandri (Hook. fil.) F. Muell., 461 stenophyllus Lindl., 448 tradescantii auct., 448 tripolium L., 451, 555 var. condensatus Druce, 451 var. discoideus Rchb., 451 var. flosculosus (Gray) P. D. Sell, 451, 555 subsp. tripolium, 452 var. tripolium, 451 umbellatus Mill., 449 var. umbellatus, 449 var. pubens A. Gray, 449 × versicolor Willd., 447 wootonii (Greene) Greene, 448 Aster, Blue Wood, 445 Bushy, 446 Chinese, 459 Frost, 445 Goldilocks, 451 Heath, 450 Leafy, 449 Mexican, 525 Narrow-leaved Smooth, 446 Sea, 451 Stonecrop, 450 Tall White, 449 ASTERACEAE, 62 Asterocephalus Zinn columbarius (L.) Rchb., 59 Athanasia L. maritima (L.) L., 470 Baccharis L., 462 halimifolia L., 462 Bachelor Buttons, 464, 471

Index Balsamita Mill., 465 major Desf., 465 Bardana Hill, 70 arctium (Hill) Hill, 72 lappa (L.) Hill, 71 minor (Hill) Hill, 71 Barkhousia Moench, 205 byzantina DC., 205 candollei Spreng., 205 foetida (L.) F. W. Schmidt, 205 var. subdivisa Schur, 205 glandulosa (Guss.) C. Presl, 205 gracilis Freyn & Sint., 206 gracilis Lej. ex Rouy, 205 graveolens (P. Gaertn., B. Mey. & Scherb.) Link, 205 haenseleri Boiss., 207 heterocarpa Boiss., 207 hirta W. D. J. Koch, 205 hispida (Waldst. & Kit.) Link, 208 laciniata Lowe, 207 nicaeensis (Balb.) Link, 206 pinguis Rchb., 203 prostrata Dumort., 205 rhoeadifolia (M. Bieb.) Fisch. & C. A. Mey., 205 rodigioides Sch. Bip. ex W. D. J. Koch, 205 setosa (Haller fil.) DC., 208 supina Rouy, 205 taraxacifolia (Thuill.) DC., 207 triangularis K. Koch, 205 zacynthia Margot & Reut. ex DC., 205 Baziasa Steud. urticaefolia (Kunth) Steud., 521 Beadplant, 15 Beauty Bush, 47 Bedstraw, Common Marsh, 23 Fen, 20 Heath, 25 Hedge, 24 Lady’s, 23 Limestone, 25 Northern, 20 Slender, 25 Slender Marsh, 23 Tree, 15 Wall, 27 Beggar’s-ticks, 523 Eastern, 523 Bellflower, Adria, 6 Chimney, 5 Clustered, 6 Cornish, 5 Creeping, 5 Giant, 4 Italian, 7 Ivy-leaved, 9 Milky, 3 Nettle-leaved, 4 Peach-leaved, 3 Rampion, 3 Spreading, 2

Index Bellflower (cont.) Trailing, 6 Tussock, 4 Bellis L., 462 monstrosa auct., 463 perennis L., 462, 556 forma disciformis P. D. Sell, 464, 556 forma discoidea D. C. McClintock, 463, 555 var. discoidea Hus, 463, 555 forma perennis, 463 × sylvestris, 463 sylvestris Cyr., 463 Bellium L. bellidioides L., 440 Bennettia Gray, 72 alpina (L.) Gray, 72 Berinia Brign., 203 nicaeensis (Balb.) Sch. Bip., 206 Berkheya Ehrh., 423 heterophylla (Thunb.) O. Hoffm., 423 pinnatifida (Thunb.) Thell., 423 rigida (Thunb.) Ewart, J. White & Rees, 424 Bidens L., 521 aristosa (Michx) Britton, 521 bipinnata L., 524 biternata (Lour.) Merr. & Sherff, 521 californica DC., 523 camporum (Hutch.) Mesfin, 521 cernua L., 522 var. cernua, 522 forma discoidea DC., 522 forma minima (Hill) F. N. Williams, 522 var. radiata DC., 522 forma rugosa Druce, 522 cicutaefolia Tausch, 524 connata Muhlenb. ex Willd., 522 decomposita Wall. ex DC., 524 elongata Tausch, 524 fervida Hort. ex Colla, 524 frondosa L., 523 kotschyana Sch. Bip., 524 melanocarpa Wieg, 523 minima Hill, 522 myrrhidifolia Tausch, 524 peacockii Druce, 522 pilosa L., 523 var. bipinnata (L.) Hook. fil., 524 var. pilosa, 524 var. radiata DC., 524 radiata Thuill., 522 subalternans DC., 521 tenuifolia Tausch, 524 tenuisecta A. Gray., 521 tripartita L., 522 vulgata Greene, 523 Black-eyed Susan, 517 Black-jack, 523 Blanket Flower, 527 Bolelia Raf., 14 Brachyderea Cass., 203 nicaeensis (Balb.) Sch. Bip., 206

Brachyglottis J. R. Forster & G. Forster, 505, 556 × jubar P. D. Sell, 505, 556 laxifolia x compacta, 505, 556 monroi (Hook. fil.) B. Nord., 506 rangina Buchanan, 506 repanda J. R. & G. Forst., 506 cv. Sunshine, 556 Breea Less., 80 dioica Less., 80 Buphthalmum L., 437 cordifolium Waldst. & Kit., 438 helianthoides L., 520 salicifolium L., 438 speciosum Schreb., 438 Bur-daisy, Hairy, 439 Purple, 439 Yellow, 439 Burdock, 70 Greater, 71 Hairy, 72 Lesser, 71 Northern, 71 Woolly, 72 Bur-marigold, London, 522 Nodding, 522 Trifid, 522 Butterbur, 510 Great, 510 White, 511 Buttonweed, 486 Annual, 487 Cacalia L. tangutica (Maxim.) Hand.-Mazz., 506 Calcitrapa Heist. ex Fabr., 87 hippophaestum Gaertn., 87 stellalis Hill, 87 Calea L. leptophylla G. Forst. 432 Calendula L., 512 arvensis L., 512 officinalis L., 512 Calliopsis Rchb. bicolor Rchb., 524 tinctoria (Nutt.) DC., 524 Callistemma (Mertens & W. D. J. Koch) Boiss. chinensis (L.) Druce, 459 Callistephus Cass., 459 chinensis (L.) Nees, 459 hortensis Cass., 459 Calocephalus R. Br. knappii (F. Muell.) Ewart & J. White, 432 Calotis R. Br., 439 cuneifolia R. Br., 439 dentex R. Br., 439 glabrescens C. T. White, 439 hispidula (F. Muell.) F. Muell., 439 lappulacea Benth., 439 microphylla Benth., 439 suffruticosa Domin, 439

577

578 Calysphyrum Bunge florida Bunge, 43 Campanula L., 1 alliariifolia Willd., 5 biserrata K. Koch, 3 calydina Boeber ex Roem. & Schult., 3 carpatica Jacq., 4 cavolinii Ten., 7 celtidifolia Boiss. & A. Huet, 3 diffusa Vahl, 7 esculenta Salisb., 3 fragilis Cirillo, 7 subsp. cavolinii (Ten.) Dambolt, 7 giesekiana auct., 8 glomerata L., 6 var. dahurica Fisch. & Ker-Gawl., 6 var. nana C. Bailey, 6 hederacea L., 9 heterophylla Gray, 7 hybrida L., 8 lactiflora M. Bieb., 3 lambertiana DC., 3 lamifolia Adans., 5 latifolia L., 4 linifolia L., 7 macrophylla Sims, 5 medium L., 4 minor Lam., 7 patula L., 2 persicifolia L., 3 portenschlagiana Schult., 6 poscharskyana Degen, 6 pyramidalis L., 5 rapunculoides L., 5 rapunculus L., 3 rhomboidalis L., 7 rigida Stokes, 5 rotundifolia L., 7, 530 subsp. montana (Syme) P. D. Sell, 8, 530 var. montana Syme, 8, 530 subsp. rhomboidalis (L.) Bonnier, 7 var. speciosa A. G. More, 8 var. uniflora auct., 8 speculum-veneris L., 8 trachelium L., 4 urticaefolia Salisb., 4 variifolia Salisb., 7 virgata DC., 3 CAMPANULACEAE, 1 Canterbury Bells, 4 CAPRIFOLIACEAE, 29 Caprifolium Mill., 46 dumetorum Lam., 45 germanicum Delarbre, 47 henryi (Hemsl.) Kuntze, 45 hortense Lam., 47 involucratum (Richardson) Kuntze, 44 italicum (Mill.) Medik., 47 japonicum (Thunb. ex Murray) Dum.-Cours., 46 pallidum (Host) Schur, 47

Index Caprifolium Mill. (cont.) perfoliatum (R¨ohl.) Gray, 47 pileatum (Oliv.) Kuntze, 44 rotundifolium Moench, 47 tataricum (L.) Kuntze, 45 vulgare Medik., 47 xylosteum (L.) Gaertn., 45 Carbenia Adans., 92 benedicta (L.) Arcang., 92 Cardoon, 83 Carduncellus Adans. coeruleus (L.) C. Presl, 93 Carduus L., 73 acanthoides auct. et Dandy (1958) et CTW (1987), 74 acanthoides L., 74 subsp. multiflorus (Gaudin) Nyman, 75 acaulos L., 79 var. dubius auct., 79 anglicus Lam., 77 argentatus L., 73 arvensis (L.) Hill, 80 var. latifolius Bab., 81 subsp. setosus (Willd.) Hook. fil., 81 carolorum Howie & Jenner, 78 crispus auct. et CTW (1987), 74 crispus L., 74, 530 var. glareicola P. D. Sell, 75, 530 subsp. multiflorus (Gaudin) Franco, 75 var. multiflorus (Gaudin) DC., 75 × nutans 75 subsp. occidentalis Chass. & Ar`enes, 75 dissectus L., 77 × dubius Balb., 75 eriophorus L., 76 erisithales Jacq., 78 forsteri (Sm.) Bab., 77 heterophyllus L., 78 lactifolius Stokes, 83 lanceolatus L., 77 macrocephalus Desf., 73 marianus L., 83 multiflorus Gaudin., 75 nutans L., 75, 530 × Cirsium arvensis (L.) Scop., 75 × Cirsium vulgare, 75 var. litoralis P. D. Sell, 75, 530 × orthocephalus auct., 75 palustris L., 80 polyacanthus Schreb., 74 polyacanthus Schleich., 75 pratensis Huds., 77 pycnocephalus L., 73 tenuiflorus Curtis, 73 thoermeri Weinm., 73 tinctorius (L.) Scop., 83 tuberosus L., 78 vulgaris Savi, 77 woodwardii H. C. Wats., 79 Carlina L., 69 vulgaris L., 69

Index Carlina vulgaris L. (cont.) forma maritima Rouy, 70 var. maritima (Rouy) P. D. Sell, 70 var. vulgaris, 70 Carthamus L., 92 lanatus L. 93 subsp. baeticus (Boiss. & Reut.) Nyman, 93 subsp. creticus (L.) Holmboe, 93 tinctorius L., 92 Cassinia R. Br., 432 fulvida Hook. fil., 432 leptophylla (G. Forst.) R. Br., 432 Catananche L., 96 caerulea L., 96 Cat’s-ear, 98 Smooth, 98 Spotted, 100 Celtuce, 117 Centaurea L., 84 aspera L., 88 benedicta (L.) L., 92 calcitrapa L., 87 cineraria L., 86 cyanus L., 91 dealbata Willd., 84 debeauxii Gren. & Godr., 90, 531 × jacea, 90 var. minima (C. E. Briitton) P. D. Sell, 90, 531 subsp. nemoralis (Jord.) Dost´al, 90, 531 var. nemoralis (Jord.) P. D. Sell, 90, 531 × nigra, 90 subsp. thuillieri Dost´al, 90, 531 var. pumila P. D. Sell, 90, 531 var. thuillieri (Dost´al) P. D. Sell, 90, 531 diffusa Lam., 87 diluta Aiton, 89 eriophora L., 88 heterophylla Willd., 88 hyalolepis Boiss., 87 isnardii (L.) L., 88 jacea L., 89 macrocephala Muss. Puschk. ex Willd., 89 melitensis L., 88 × moncktonii C. E. Britton, 90 montana L., 91 nemoralis Jord., 90, 531 var. minima C. E. Briitton, 90, 531 nigra L., 90 var. nemoralis (Jord.) Gremli, 531 subsp. rivularis (Brot.) Cout., 91 obscura Jord., 91 pallescens Delile subsp. hyalolepis (Boiss.) Holmboe, 87 var. hyalolepis (Boiss.) Boiss., 87 paniculata L., 86 parviflora Sibth. & Sm., 87 picris Pall. ex Willd., 84 pratensis Thuill., 90, 531 repens L., 84 rivularis Brot., 91

Centaurea L. (cont.) scabiosa L., 85, 531 var. nana P. D. Sell, 86, 531 var. succisiifolia E. S. Marshall ex C. E. Britton, 86 solstitialis L., 88 Centranthus DC., 53 marinus Gray, 54 ruber (L.) DC., 54 Cephalanoplos (Necker ex DC.) Fourr. arvensis (L.) Fourr., 80 Cephalaria Schrad., 57 elata (Hornem.) Schrad., 57 gigantea (Ledeb.) Bobrov, 57 syriaca (L.) Roem. & Schult., 57 tatarica Roem. & Schult., 57 Cervicina Delile, 8 hederacea (L.) Druce, 9 Chamaecerasus Duham. tartaricus (L.) Billiard, 45 xylosteum (L.) Billiard, 45 Chamaemelum Mill., 473 arvensis (L.) Schreb., 475 cotula (L.) All., 476 discoidea All., 477 foetidum Baumg., 476 mixtum (L.) All., 474 nobile (L.) All., 474, 556 forma duplicatum (Gray) P. D. Sell, 474, 556 var. duplicatum Gray, 474, 556 forma nobile, 474 forma nudum (Gray) P. D. Sell, 474, 556 var. nudum Gray, 474, 556 Chamomile, 474 Austrian, 477 Corn, 475 Eastern, 476 Mediterranean, 474 Sicilian, 475 Southern, 477 Stinking, 476 Yellow, 477 Chamomilla Gray, 483 discoidea (DC.) J. Gay, 484 inodora K. Koch, 486 nobilis (L.) Gren. & Godr., 474 occidentalis (Greene) Rydb., 484 recutita (L.) Rauschert, 484 suaveolens (Pursh) Rydb., 484 vulgaris Gray, 484 Chartolepis Cass., 89 Cheiroloma F. Muell., 439 hispidulum F. Muell., 439 Chevreulia Cass. sarmentosa (Pers.) Blake, 427 Chicory, 94 Chondrilla L. muralis (L.) Lam., 120 Chrysanthemum auct., 478 Chrysanthemum L., 481 balsamita (L.) Baill., 465

579

580 Chrysanthemum L. (cont.) coronarium L., 479 subsp. discolor (d’Urv.) Rech. fil., 479 var. discolor d’Urv., 479, 556 crassifolium (Lange) Guinea, 481 erubescens auct., 482 hybridum Guss., 479 indicum L., 482 inodorum L., 486 ircutianum (DC.) Turcz., 481 japonense Nakai, 482 leucanthemum L., 480 var. cantabricum (Sennen) Guinea, 481 subsp. praecox Horvatic, 480 macrophyllum Waldst. & Kit., 464 makinoi Matsum., 482 maximum auct., 481 maximum Ramond, 481 × morifolium Ramat., 482 myconis L., 479 ornatum Hemsl., 482 parthenium (L.) Bernh., 464 × rubellum Sealy, 482 segetum L., 478 serotinum L., 479 sibiricum Turcz., 483 sinense Sabine, 482 × superbum Bergmans ex J. Ingram, 481 tanacetum Karsch., 464 uliginosum (Waldst. & Kit. ex Willd.) Pers., 479 vestitum Hemsl., 482 vulgare (L.) Bernh., 464 zawadskii Herbich, 483 Chrysanthemum, Florist’s, 482 Indian, 482 Korean, 482 Zawadski’s, 483 Chrysocoma L., 452 aurea Salisb., 452 coma-aurea L., 452 var. patula (P. J. Bergius) DC., 452 graminifolia L., 442 linosyris L., 451 microcephala DC., 452 nupera Gray, 451 patula P. J. Bergius, 452 tenuifolia P. J. Bergius, 452 Cicerbita Wallr., 118 alpina (L.) Wallr., 118 bourgaei (Boiss.) Beauverd, 119 gmelinii Beauverd, 119 macrophylla (Willd.) Wallr., 119 subsp. uralensis (Rouy) P. D. Sell, 119 muralis (L.) Wallr., 120 plumieri (L.) Kirschl., 119 tatarica (L.) Beauverd ex Sosn., 118 uralensis (Rouy) Beauverd, 119 Cichorium L., 94 divaricatum Schousb., 95

Index Cichorium L. (cont.) endivia L., 95 var. crispum Hegi, 95 subsp. divaricatum (Schousb.) Bonnier & Layens, 95 intybus L., 94 subsp. foliosum (Hegi) Janch., 95 var. foliosum Hegi, 95 convar. radicosum (Alef.) Holub, 95 convar. radicosum Alef., 95 subsp. sativum Janch., 95 var. sativum (DC.) Janch., 95 taxon sativum Bisch., 95 var. sativum DC., 95 subsp. silvestre (Bisch.) Janch., 95 taxon silvestre Bisch., 95 perenne Stokes, 94 sylvestre Lam., 94 Cineraria L. alpina L. var. integrifolia L., 504 ambigua (Biv.) DC., 490 aspera Thunb., 504 bicolor Willd., 490 burkei Burtt Davy, 504 campestris auct., 504 congesta R. Br., 505 geifolia L., 504 gigantea Sm., 494 integrifolia (L.) Murr., 504 lobata L’Her., 504 lyrata DC., 504 nebrodensis Guss., 490 palustris (L.) L., 505 repanda (J. R. & G. Forst.) Forst. fil., 506 serratifolia Gray, 493 Cineraria, 503 Cirsium Mill., 75 acaule Scop., 79 × arvense, 79 forma caulescens Rchb., 79 × dissectum, 79 × palustre, 79 tuberosum, 80 × vulgare, 80 acaulon (L.) Scop., 79 anglicum (Lam.) DC., 77 arvense (L.) Scop., 80 var. horridum Wimm. & Grab., 81 subsp. incanum (S. G. Gmel.) Iljin, 81 var. incanum (S. G. Gmel.) Lebeb., 81 var. integrifolium Wimm. & Grab., 81 var. maritimum Fr., 81 var. mite Wimm. & Grab,, 81 × palustre, 81 subsp. setosum (Willd.) Iljin, 81 var. vestitum Wimm. & Grab., 81 × babingtonii Rouy, 79 × boulayi E. G. Camus, 79 carolorum (Howie & Jenner) Nyman, 78 × celakovskianum Knaf., 81

Index Cirsium Mill. (cont.) dissectum (L.) Hill, 77 × palustre, 77 erisithales (Jacq.) Scop., 78 eriophorum (L.) Scop., 76 subsp. britannicum Petr., 76 × vulgare, 77 esculentum (Stev.) C. A. Mey., 79 × forsteri (Sm.) Loudon, 77 fraserianum Druce, 80 × gerhardii Sch. Bip., 77 × gibsonii H. C. Wats., 79 glutinosum Lam., 78 × grandiflorum Kitt., 77 helenioides auct., 78 heterophyllum (L.) Hill, 78 × palustre, 78 incanum (S. G. Gmel.) Fisch. ex M. Bieb., 81 × kirschlegeri Sch. Bip., 79 lanceolatum Hill, 77 lanceolatum (L.) Scop., 77 × medium All., 80 × mixtum Druce, 81 montanum Hill, 72 ochroleucum DC., 78 oleraceum (L.) Scop., 79 palustre (L.) Scop., 80 var. ferox Druce, 80 × tuberosum, 80 × vulgare, 80 pratense (Huds.) Druce, 77 rhizocephalum C. A. Mey., 79 rivulare (Jacq.) All., 75 × sabaudum M. Loehr, 80 × semidecurrens Richt., 80 setosum (Willd.) M. Bieb., 81 × subspinuligerum Peterm., 80 tuberosum (L.) All., 78 vulgare (Savi) Ten., 77, 531 var. litorale P. D. Sell, 77, 531 × wankelii Reichardt., 78 × woodwardii (H. C. Wats.) Nyman, 79 × zizianum W. D. J. Koch, 80 Cleavers, 26 Corn, 27 False, 26 Southern, 27 Clintonia Douglas ex Lindl. elegans Douglas ex Lindl., 14 Cnicus L., 92 acaulos (L.) Willd., 79 anglicus (Lam.) C. C. Gmel., 77 arvensis (L.) Roth., 80 subsp. setosus (Willd.) Hook. fil., 81 benedictus L., 92 britannicus Druce, 77 carolorum (Howie & Jenner) Hook. fil., 78 eriophorus (L.) Roth., 76 erisithales (Jacq.) L., 78 forsteri Sm., 77

Cnicus L. (cont.) heterophyllus (L.) Roth., 78 lanceolatus (L.) Willd., 77 oleraceus L., 79 palustris (L.) Willd., 80 pratensis (Huds.) Willd., 77 tuberosus (L.) Roth., 78 × wankelii (Reichardt.) F. Hanb., 78 woodwardii (H. C. Wats.) Hook. fil., 79 Cocklebur, Argentine, 516 Rough, 515 Spiny, 515 Coleostephus Cass., 479 myconis (L.) Rchb. fil., 479 Colobium Roth taraxacoides (Vill.) Holub, 103 Colt’sfoot, 509 Purple, 512 Columbaria J. & C. Presl, 59 minor Gray, 59 Colymbada Hill, 85 COMPOSITAE, 62 Coneflower, 518 Conyza L., 433 squarrosa L., 435 vulgaris Lam., 435 Conyza Less, 456 albida Willd. ex Spreng., 457 ambigua DC., 458 bilbaoana J. R´emy, 458 blakei (Cabrera) Cabrera, 456 bonariensis (L.) Cronquist, 458 canadensis (L.) Cronquist, 458, 555 var. canadensis, 459 × Erigeron acer, 459 var. incisa P. D. Sell, 459, 555 var. obovoidea P. D. Sell, 459, 555 var. robusta P. D. Sell, 459, 556 var. simplex P. D. Sell, 459, 555 daveauiana Sennen, 457 erigeroides DC., 457 floribunda Humb., Bonpl. & Kunth, 458, 555 var. floribunda, 458 var. linearifolia P. D. Sell, 458, 555 japonica (Thunb. ex Murray) Less., 456 sumatrensis (Retz.) E. Walker, 457 Conyza x Erigeron, 459 Conyzella Fabr., 456 canadensis (L.) Rupr., 458 linifolia (Willd.) Greene, 458 × Conyzigeron Rauschert, 459 huelsenii (Vatke) Rauschert, 459 Coprosma J. R. Forst. & G. Forst., 15 baueria auct., 15 bauriana Hook. fil., 15 repens A.Rich., 15 retusa Hook. fil., 15 stockii B. S. Williams, 15 Coralberry, 41

581

582 Cornflower, 91 Perennial, 91 Cornsalad, Broad-fruited, 51 Common, 49 Hairy-fruited, 51 Keel-fruited, 49 Narrow-fruited, 51 Corvisartia M´erat caucasica (Pers.) G. Don ex Loudon, 438 helenium (L.) M´erat, 433 Coryopsis L., 524 bidens L., 522 boykiniana Nutt., 524 corymbifolia Wall. ex DC., 524 elegans hort., 524 grandiflora Hogg ex Sweet, 524 heterophylla Nutt., 524 tinctoria Nutt., 524 Cosmos Cav., 525 bipinnatus Cav., 525 Costmary, 465 Cota J. Gay ex Guss., 477 altissima (L.) J. Gay ex Guss., 477 austriaca (Jacq.) Sch. Bip., 477 tinctoria (L.) J. Gay ex Guss., 477 Cottonweed, 470 Cotula L., 486 australis (Sieber ex Spreng.) Hook. fil., 487 bipinnata Thunb., 486 coronopifolia L., 486 var. integrifolia (Hook. fil.) T. Kirk, 486 dioica (Hook. fil.) Hook. fil., 487 integrifolia Hook. fil., 486 pusilla Thunb., 486 sororia DC., 486 squalida (Hook. fil.) Hook. fil., 487 turbinata L., 486 venosa Colenso, 487 zeyheri Fenzl ex Harvey, 486 Cotula Rupr., 476 foetida S. G. Gmel., 476 Crepis L., 201 adenantha Vis., 206 agrestis Fr. ex Bisch., 206 agrestis Waldst. & Kit., 207 angustifolia d’Urv., 204 banatica Willd., 208 barbata L., 208 barbata Mill., 208 biennis L., 203 brachypappa Borm., 206 bulbosus (L.) Tausch, 111 capillaris (L.) Wallr., 206 var. agrestis (Waldst. & Kit.) Dalla Torre & Sarnth., 207 var. anglica Druce & Thell., 207 var. glandulosa Druce, 207 caucasica C. A. Mey., 202 charbonnelii L´evl., 118 cinerea Desf., 207 cooperi A. Gray, 206 croatica (Waldst. & Kit.) Froel., 203

Index Crepis L. (cont.) diffusa DC., 206 dioscoridis Pollich, 204 druceana Murr, 207 echioides Ledeb., 205 fallax Boiss., 205 fistula Fisch. ex Besser, 204 foetens Link ex Buchanon, 205 foetida L., 205 subsp. commutata (Spreng.) Babc., 206 subsp. glandulosa (Guss.) Hayek, 205 var. hispida Bisch., 205 subsp. maritima (Boiss.) Hayek, 205 ˇ subsp. rhoeadifolia (M. Bieb.) Celak., 205 subsp. zacintha (Margot & Reut. ex DC.) Hayek, 205 gaditana Boiss., 206 glabra Boiss., 202 glabrata Porcius, 208 glandulosa Bastard ex M¨ossler, 203 glandulosa Guss., 205 gmelinii Schult., 203 graveolens Schrad. ex Steud., 205 hackelii Lange, 207 haenseleri (Boiss.) F. W. Schultz, 207 hieracioides (L.) Lam., 107 hieracioides Waldst. & Kit., 203 hispida Waldst. & Kit., 208 humilis Gaudin, 206 insularis Moris & De Not., 205 integrifolia Vest, 204 interrupta Sibth., 205 intybacea Brot., 207 lachenalii Gochnat, 204 laciniata (Lowe) F. W. Schultz, 207 lodomeriensis Bess., 203 maritima Boucher, 203 mollis (Jacq.) Asch., 203 subsp. hieracioides (Waldst. & Kit.) Domin, 203 subsp. velenovskyi (Domin.) Domin, 203 muricata Gilib., 203 murorum S. G. Gmel., 204 nestmeiri F. Herm & Degen, 205 nicaeensis Balb., 206 numidica Pomel, 207 paludosa (L.) Moench, 202 polymorpha Pourr., 207 polymorpha Willd., 206 var. diffusa Wallr., 207 praecox Balb., 207 praemorsa (L.) Walther, 204 pulchra L., 202 racemiformis Cariot & St. Lag., 204 racemosa Cariot & St-Lag., 204 recognita Hall. fil., 207 rhoeadifolia M. Bieb., 205 rubra L., 202 rubricaulis Pers, 207 ruderalis Bouch., 207 rumicifolia Boiss., 202 ruprechtii Boiss., 202 rutilans Lacaita, 207, 532

Index Crepis L. (cont.) sabauda Balb. ex DC., 204 sancta (L.) Babc. subsp. sancta (Gouan) Babc., 202 scabra DC., 206 scabra Willd., 207 setosa Haller fil., 208 strybrnyi Velen., 205 suberostris Coss. & Durieu ex Batt., 202 succisifolia (All.) Tausch, 203 taraxacifolia Thuill., 207 subsp. stellata Ball, 207 taurinensis Willd., 207 tectorum L., 204 subsp. barckhausioides Rouy, 204 umbellata auct., 206 umbellata Thuill., 207 uniflora Thuill., 207 varia Moench, 204 velenovskyi Domin, 203 vesicaria L., 207 subsp. haenseleri (Boiss.) P. D. Sell, 207 var. rutilans (Lacaita) P. D. Sell, 207, 532 subsp. stellata (Ball) Babc., 207 subsp. taraxacifolia (Thuill.) Thuill., 207 var. taraxacifolia (Thuill.) B. Boivin, 207 virens L., 206 var. agrestis (Waldst. & Kit.) Bisch., 207 var. elatior Cariot & St. Lag., 207 Crinita Moench, 451 Crinitaria Cass., 451 linosyris (L.) Less., 451 Crocodilodes Adans., 423 bilobum (DC.) Kuntze, 423 epitrachys (DC.) Kuntze, 423 heterophyllum (Thunb.) Kuntze, 423 pinnatifidum (Thunb.) Kuntze, 423 rigidum (Thunb.) Kuntze, 423 Crosswort, 28 Caucasian, 16 Crucianella L. stylosa Trin., 16 Cruciata Mill., 28 chersonensis auct., 28 laevipes Opiz, 28 Cryptostemma R. Br., 422 calendula (L.) Druce, 422 calendulacea (Hill) R. Br., 422 Crysogonum L. virginianum L., 517 Cudweed, America, 429 Broad-leaved, 425 Cape, 431 Common, 425 Dwarf, 429 Field, 426 Heath, 429 Highland, 429 Jersey, 431 Marsh, 430 Narrow-leaved 426

Cudweed (cont.) Red-tipped, 425 Small, 426 Cupidone, Blue, 96 Cupularia Gren. & Godr., 436 graveolens (L.) Gren. & Godr., 436 viscosa (L.) Gren. & Godr., 436 Cyanus Mill., 91 cyanus (L.) Hill, 91 montanus (L.) Hill, 91 segetum Baumg., 91 vulgaris Delarb., 91 Cyclachaena Fresen. xanthifolia (Nutt.) Fresen., 514 Cynara L., 82 carduncellus L., 82 var. sativa Moris, 83 var. scolymus (L.) Fiori, 83 diversifolia Stokes, 78 eriocephala Stokes, 76 horridus Aiton, 82 lanceata Stokes, 77 palustris (L.) Stokes, 80 repens Stokes, 80 scolymus L., 83 Dahlia Cav., 525 coccinea Cav., 525 × hortensis Guillaumot, 525 pinnata Cav., 525 variabilis (Willd.) Desf., 525 Dahlia, Garden 525 Daisy, 462 Autumn Oxeye, 479 Crown, 479 Garden, 463 Oxeye, 480 Seaside, 453 Shasta, 481 Daisybush, 461 Coastal, 461 Mangrove-leaved, 460 Sticky, 460 Dandelion, Acute-leaved, 179 Alderney, 144 Angular-lobed, 180 Apiculate-lobed, 190 Asymmetrical Hook-lobed, 166 Beautiful-leaved, 196 Beeftink’s, 158 Belgium, 143 Ben Lawes, 149 Berthe’s, 159 Black-toothed, 172 Blood-red, 158 Bluish-bracted, 178 Bluish-leaved, 201 Blunt-lobed, 177 B¨okman’s, 165 Brecon, 161 Bright, 157

583

584 Dandelion (cont.) Bright Green, 148 Brilliant-stalked, 196 British, 154 Broad-bracted, 184 Broad-leaved, 183 Broad-lobed, 194 Broad-stalked, 174 Broddeson’s, 176 Brown-fruited, 161 Brown-ribbed, 186 Brown-striped, 164 Bulbous-lobed, 192 Caledonian, 161 Celtic, 155 Cenabum, 139 Channel Island, 146 Cherwell, 176 Cinnamon-fruited, 142 City, 181 Claw-lobed, 162 Close-lobed, 170 Clova, 148 Common, 197 Complex-leaved, 195 Cornish, 152 Crowded-lobed, 187 Cut-leaved, 136 Cut-lobed, 184 Dahlstedt’s, 193 Dark-bracted, 154 Dark-green, 153 Dark Hook-lobed 164 Dark-leaved, 173 Decumbent, 172 Degelius’s 143 Deltoid-lobed, 199 Dense-bracted, 200 Dense-lobed, 169 De Retz’s, 142 Dirty-leaved, 192 Diverse-leaved, 197 Divided-leaved, 171 Double-toothed, 155 Druce’s, 152 Dull-leaved, 177 Dune, 139 Edmondson’s, 184 Ekman’s, 189 Elongate-bracted, 199 English, 147 Entire-lobed, 188 Equal-lobed, 181 Erect-bracted, 185 Essex, 141 Faeroes, 147 Fagerstr¨om’s, 195 Falcate-lobed, 188 False Fen, 160 False Hook-lobed, 164 Fen, 146

Index Dandelion (cont.) Few-lobed, 180 Fleshy-lobed, 164 Flintshire, 154 Gelert’s, 153 Golden-fruited, 141 Gotland, 140 Grassland, 192 Green, 170 Green-bracted, 141 Green-stalked, 168 Grey-bracted, 174 Hairy, 151 Hairy-ribbed, 183 Hairy-stalked, 182 Hairy-stemmed, 193 Hampshire, 158 Haworth’s, 139 Hexham, 189 Hook-lobed, 162 H¨ulphers’s, 194 Hump-lobed, 176 Imbricate-bracted, 185 Incise-leaved, 198 Incise-lobed, 168 Irish, 146 Irregular-bracted, 190 Jagged-leaved, 175 Kern’s, 167 Lacerate-leaved, 170 Lancashire, 160 Landmark, 158 Large-bracted, 200 Large-flowered, 187 Large-headed, 187 Large Hook-lobed, 163 Large-lobed, 153 Leaden-bracted, 178 Light-green, 173 Linear-lobed, 175 Long-bracted, 186 Lunar-lobed, 190 Lustrous-leaved, 167 Many-coloured, 197 Many-lobed, 145 Many-toothed, 143 Margetts’s, 173 Marklund’s, 163 Massive, 177 Multilobed, 181 Narrow-bracted, 165 Narrow-lobed, 142 Nordstedt’s, 159 Oblong-leaved, 191 Obtuse-leaved, 186 Ochre-styled, 198 Olga’s, 160 Øllgaard’s, 155 Orange-flowered, 175 Orange-tipped, 194 Orange-toothed, 179

Index Dandelion (cont.) Orkney, 144 Ostenfeld’s, 161 Oxford, 141 Pale-bracted, 151 Pale-green, 184 Pale-leaved, 149 Pale-stalked, 168 Parallel-leaved, 148 Parnassus, 139 Pectinate-leaved, 196 Pink-stalked, 172 Pointed-lobed, 200 Pollenless, 156 Porter’s 155 Pretty-leaved, 182 Prickly-leaved, 170 Pruinose, 166 Pruinose-bracted, 174 Pure Yellow, 162 Purple-blotched, 156 Purple-bracted, 137 Purple-stalked, 200 Recurved-bracted, 145 Recurved-lobed, 171 Reddish-bracted, 174 Red-fruited, 150 Red-tipped, 179 Reflexed-bracted, 185 Reflexed-lobed, 182 Regular-lobed, 195 Remarkable, 171 Richards’s, 152 Roadside, 199 Robust, 187 Rounded-lobed, 191 Ruddy, 138 Sagittate-lobed, 166 Sahlin’s, 166 Sand, 138 Saw-leaved, 165 Scottish, 142 Selland’s, 180 Serpentine, 148 Sharp-lobed, 182 Sharp-toothed, 137 Shetland, 147 Shining, 198 Shiny-leaved, 169 Sinuate-lobed, 193 Sk˚ane, 137 Slender Hook-lobed, 163 Small-headed, 173 Small-spotted, 149 Smooth, 188 Sociable, 138 Spear-toothed, 167 Spiky-leaved, 177 Spotted, 151 Spreading-bracted, 151 Spreading-lobed, 193

Dandelion (cont.) Spur-lobed, 191 Squat, 150 Stiff-leaved, 152 Straw-fruited, 140 Sundeberg’s, 201 Swollen-lobed, 179 Tailed, 190 Tall, 181 Tar-blotched, 171 Teesdale, 159 Texel, 157 Thames, 157 Thick-lobed, 194 Three-lobed, 196 Tongue-leaved, 145 Tongue-lobed, 168 Toothed, 189 Twisted-bracted, 183 Twisted-lobed, 144 Umber-fruited, 140 Unexpected, 136 Variable-leaved, 178 Violet-striped, 144 Water Meadow, 157 Wavy-leaved, 169 Webb’s, 146 Welsh, 160 Western, 156 Wetland, 150 White-stalked, 176 Wide-stalked, 150 Winged, 189 Yellowish-green, 199 Yellow-styled, 186 Delairea Lem., 505 odorata Lem., 505 Dendranthemum (DC.) Des Moul., 481 × grandiflorum (Ramat.) Kitam., 482 indicum (L.) Tzvelev, 482 morifolium (Ramat.) Tzvelev 482 rubellum (Sealy) Philp, 482 sinense (Sabine) Des Moul., 482 zawadskii (Herbich) Tzvelev 483 Dibothrospermum Knaf. agreste Knaf., 486 Diervilla Mill. florida (Bunge) Siebold & Zucc., 43 Diotis Desf., 470 candidissima Desf., 470 maritima (L.) Desf. ex Cass., 470 Diplemium Raf. Diplosastera Tausch tinctoria (Nutt.) Tausch, 524 DIPSACACEAE, 54 Dipsacus L., 54 × fallax Simonk., 55 ferox Loisel., 25 fullonum L., 55 subsp. fullonum auct., 55 × laciniatus, 55

585

586 Dipsacus fullonum L. (cont.) var. sativus L., 55 subsp. sativus (L.) Thell., 55 × sativus, 55 subsp. sylvestris (Huds.) P. Fourn., 55 laciniatus L., 56 pilosus L., 56 sativus (L.) Honck., 55 strigosus Willd., 56 sylvestris Huds., 55 Distegia Raf. nutens Raf., 44 Dittrichia Greuter, 436 graveolens (L.) Greuter, 436 viscosa (L.) Greuter, 436 Doellingeria Nees, 449 umbellata (Mill.) Nees, 449 pubens (A. Gray) Rydb., 449 Doria Fabr., 492 Doria Thunb. canadensis (L.) Lunell, 441 Doronicum L., 507 columnae Ten., 509 × pardelianches x plantagineum, 508 cordatum auct., 509 cordatum Lam., 508 cordifolium Stokes, 508 × draytonense hort., 508 × excelsum (N. E. Br.) Stace, 508 × hybridum hort., 508 pardalianches L., 508 × plantagineum, 508 plantagineum L., 508 var. excelsum N. E. Br., 508 forme wildenowii Rouy, 508 var. wildenowii (Rouy) A. B. Jacks., 508 × wildenowii (Rouy) A. W. Hill, 508 Dortmanna Hill, 12 erinus (L.) Kuntze, 12 lacustris Kuntze, 13 siphilitica (L.) Kuntze, 13 urens (L.) Kuntze, 12 Downingia Torrey, 14 elegans (Douglas ex Lindl.) Torr., 14 Dracunculus Ledeb. marschallianus Ledeb., 469 Ebulus Garcke, 31 humilis Garcke, 31 Echinops L., 68 banaticus Rochel ex Schrad., 69 commutatus Jur., 69 exaltatus Schrad., 69 mollis Schur, 69 multiflorus Lam., 68 paniculatus Jacq., 68 ritro auct., 69 rochelianus Griseb., 69 ruthenicus Rochel, 69

Index Echinops L. (cont.) sphaerocephalus L., 68 strictus Fisch., 69 viscosus Wierzb., 68 Elder, 30 American, 31 Dwarf, 31 Golden, 31 Marsh, 514 Parsley-leaved, 31 Red-berried, 30 Elecampane, 433 Endive, 95 Epaltes Cass. australis Less., 424 Erechtites Raf. glossanthus Sond., 488 minima (Poir.) DC., 503 prenanthoides DC., 503 pumila DC., 503 quadridentata (Labill.) DC., 503 Erigeron L. 452 acer L., 455 alpinus auct., 454 alpinus L., 454 ambiguus (DC.) Schultz Bip., 458 amplexicaulis Desf., 454 annuus (L.) Pers., 455 subsp. strigosus (Muhl. ex Willd.) Wagenitz, 455 blakei Cabrera, 456 bonariensis (L.) Cronquist, 458 borealis (Vierh.) Simmons, 454 canadensis L., 458 crispus Pourr., 458 eucephaloides Greene, 453 formosissimus Greene, 453 glaucus Ker Gawl., 453 grandiflorus Nutt., 453 graveolens L., 436 heterophyllus Muhl. ex Willd., 455 hispida Nutt., 453 × huelsenii Vatke, 459 hyemalis Raf., 454 japonica Thunb. ex Murray, 456 karvinskianus DC, 454 var. mucronatus (DC.) Aschers., 454 leiophyllus Greene, 453 linifolius Willd., 458 linosyris (L.) Clairv, 451 macranthus Nutt., 453 maritimus Nutt, 453 mucronatus DC, 454 neglectus A. Kerner, 454 philadelphicus L, 454 forma angustatus Vict. & Rousseau, 454 var. glaber Henry, 454 forma scatericola (Fern.) Cronquist, 454 provancheri Vict. & Rousseau, 454 pulchellus DC.

Index Erigeron L. (cont.) var. townsendii Peattie, 454 purpureus Aiton, 454 var. attenuatus Nutt., 454 scatericola Fern, 454 speciosus (Lindl.) DC., 453 var. macranthus (Nutt.) Cronquist, 453 var. speciosus, 453 strigosus Muhl. ex Willd., 455 sumatrensis Retz., 457 undulatus Moench, 458 uniflorus auct., 454 viscosum L., 436 vreelandii Rydb., 453 Eriolepis Cass., 76 Eritheis Gray, 433 maritima Gray, 435 Erythrochaete Sieb. & Zucc. dentata A. Gray, 507 Escarole, 96 Euchylia Dulac villosa Dulac, 45 Eupatorium L., 528 cannabinum L., 528 subsp. cannabinum, 528 Euphrosyne DC. xanthifolia (Nutt.) A. Gray, 514 Eurybia (Cass.) Cass., 460 avicenniifolia (Raoul) Hook. fil., 460 dentata Hook. fil. var. linearifolia Hook. fil., 461 var. oblongifolia Hook. fil., 461 forsteri Hook. fil., 460 maritima Gray, 451 var. flosculosa Gray, 451, 555 nummulariifolia Hook. fil., 460 solandri Hook. fil., 461 traversii F. Muell., 461 Euthamia Nutt., 442 graminifolia (L.) Elliott, 442 Everlasting-flower, Annual, 432 New Zealand, 432 Silver-bush, 432 Facelis Cass. retusa (Lam.) Sch. Bip., 427 Fedia Gaertn., auricula (DC.) Mert. & W. D. J. Koch, 51 carinata (Loisel.) Steven, 49 dentata (L.) Mirb., 51 ecalyculata Stokes, 49 eriocarpa (Desv.) Rchb., 51 morisonii Spreng., 51 olitoria (L.) Gaertn., 49 Feverfew, 464 Filaginella Opiz, 430 uliginosa (L.) Opiz, 430 Filago L., 424, 555 apiculata G. E. Sm. ex Bab., 425

Filago L. (cont.) arvensis L., 426 subsp. lagopus (Stephan ex Willd.) Nyman, 426 var. lagopus (Stephan ex Willd.) DC., 426 canescens Jord., 425 eriocephala auct., 425 filiformis Lam., 425 gallica L., 426 germanica auct., 425 germanica Huds., 425 germanica L., 425 lagopus (Stephan ex Willd.) Parl., 425 lutescens Jord., 425 maritima L., 470 minima (Sm.) Pers., 426 pyramidata L., 425 spathulata auct., 425 vulgaris Lam., 425 Fireweed, Cotton, 503 Toothed, 503 Flaveria Juss. bidentis (L.) Kunze, 521 Fleabane, Alpine, 454 Argentine, 458 Bilbao’s, 458 Blue, 455 Canadian, 458 Common, 437 Eastern, 434 Garden, 453 Guernsey, 457 Hooker’s, 434 Irish, 434 Many-flowered, 458 Meadow, 434 Mexican, 454 Small, 437 Small-headed, 457 Stinking, 436 Strigose, 455 Tall, 455 Woody, 436 Fleawort, Field, 504 Marsh, 505 Flossflower, 528 Fox-and-cubs, 211 Dense-flowered, 212 Glaucous, 213 Hybrid, 211 Small-flowered, 212 Yellow, 212 Franseria Cav., 513 artemisioides Willd., 513 Gaertneria Medik., 513 artemisioides (Willd.) Kuntze, 513 Gaillardia Foug., 527 aristata Pursh × pulchella Foug., 527 × grandiflora Van Houtte, 527

587

588 Galactites Moench tomentosa Moench, 81 Galatella Cass., 450 linosyris (L.) Rchb. fil., 451 punctata (Waldst. & Kit.) Nees, 450 Galedragon Gray, 54 pilosus (L.) Gray, 56 Galinsoga Ruiz & Pavon, 520 pilosus (L.) Gray, 56 aristulata E. P. Bicknell, 521 ciliata (Rafn.) S. F. Blake, 521 hirsuta Baker, 520 parviflora Cav., 520 quadriradiata Ruiz & Pavon, 521 quinqueradiata Ruiz & Pavon, 520 urticaefolia (Kunth.) Benth, 521 Galium L., 18 album Mill., 24 subsp. pycnotrichum (Heinr. Braun) Krendl, 24 anglicum Huds., 27 var. leiocarpum Tausch, 28 var. trichocarpum Tausch, 28 aparine L., 26, 530 subsp. agreste P. D. Sell, 26, 530 var. agreste P. D. Sell, 26, 530 forma condensata Druce, 26 var. marinum Fr., 26 var. spurium (L.) W. D. J. Koch, 27 subsp. vaillantii (DC.) Hook. fil., 27 aristatum auct., 24 asperum Schreb., 24 austriacum auct., 25 boreale L., 22 var. stenophyllum Druce, 22 cinereum auct., 24 constrictum Chaub., 23 cruciata (L.) Scop., 28 debile Desv., 23 diffusum D. Don ex Hook., 24 divaricatum Pour. ex Lam., 18 elatum Thuill., 24 elongatum C. Presl, 23 erectum Huds. 1778, non 1762, 24 fleurotii Jord., 25, 530 floridum Salisb., 23 harcynicum Weigel, 25 var. alpestre auct., 25 var. transiens auct., 25 hillardiae Druce 25, 530 infestum Waldst. & Kit., 27 insubricum Gaudin., 24 laeve Thuill., 25 mollugo L., 24 var. aristatum auct., 24 var. bakeri Syme, 24 subsp. elatum (Thuill.) Syme, 24 subsp. erectum Syme, 24 var. insubricum (Gaudin) Druce, 24 subsp. pycnotrichum (Heinr. Braun) O. Schwarz, 24 subsp. scabrum Hook. fil., 24

Index Galium mollugo L. (cont.) var. scabrum (Hook. fil.) Druce, 24 × verum, 24 montanum auct., 25 montanum Huds., 25 murale (L.) All., 18 nitidulum Thuill., 25 × ochroleucum Wolf ex Schweiff., 24 odoratum (L.) Scop., 19 palustre L., 23 forma angustifolium Druce, 23 var. angustifolium (Druce) Druce, 23 subsp. elongatum (C. Presl) Arcangeli, 23 var. elongatum (C. Presl) Syme, 23 var. gracile auct., 23 var. lanceolatum Uechtr., 23 var. microphyllum Lange, 23 var. serrulatum Druce, 23 subsp. tetraploideum A. R. Clapham ex Franco, 23 var. witheringii (Sm.) Syme, 23 parisiense L., 27 subsp. anglicum (Huds.) A. R. Clapham, 27 var. anglicum (Huds.) Hook. & Arn., 27 × pomeranicum Retz., 24, 530 nothosubsp. hillardiae (Druce) P. D. Sell, 25, 530 procumbens Stokes, 25 pulchellum Salisb., 25 pumilum Murray, 25, 530 subsp. fleurotii (Jord.) P. D. Sell, 25, 530 var. glabrum auct., 25 var. nitidum (Thuill.) Druce, 25 subsp. septentrionale Sterner ex Hyl., 25 pusillum auct., 25 pycnotrichum Heinr. Braun, 24 × rothschildii Druce, 24 rupicola Bertol., 25 saccharatum All., 27 saxatile L., 25 var. vivianum (Kliphuis) Erend., 26 × sterneri, 26 setaceum Lam., 18 spurium L., 26 subsp. infestum (Waldst. & Kit.) Sch¨ubler & Martens, 27 var. vaillantii (DC.) Gren., 27 sterneri Ehrend., 25 sylvestre Pollich, 25 var. nitidulum Thuill., 25 tricorne auct., 27 tricornutum Dandy, 27 uliginosum L., 22 umbellatum Lam., 25 uncinatum Gray, 26 vaillantii DC., 27 valantia Gaertn., C.A. Mey. & Scherb., 28 valantia G. Weber, 27 verrucosum Huds., 27 verum L., 23 var. littorale Br´eb., 24 subsp. maritima (DC.) Adema., 24

Index Galium verum L. (cont.) var. maritimum DC., 24 subsp. wirtgenii (F. W. Schultz) Oborny, 24 virgatum Nutt. ex Torr. & A. Gray, 18 vulgare Gray, 24 witheringii Sm., 23 Gallant Soldier, 520 Gamochaeta Weddell, 430 purpurea (L.) Cabrera, 430 Gazania Gaertn., 422 leucolaena DC., 423 rigens (L.) Gaertn., 423 var. leucolaena (DC.) Harv., 423 var. rigens, 423 var. uniflora (L. fil.) Roessler, 423 uniflora (L. fil.) Sims, 423 var. leucolaena (DC.) Harv., 423 Gentiana L. collina With., 6 Geracium Rchb., 202 paludosum (L.) Rchb., 202 praemorsum (L.) Rchb., 204 German Ivy, 505 Geropogon L. glaber L., 111 hirsutus L., 111 hybridus (L.) Sch. Bip., 111 Gifola Cass., 425 apiculata Chrtek & Holub, 425 germanica Dumort, 425 Glebionis Cass. roxburghii Cass., 479 Globe Artichoke, 83 Globe Thistle, 69 Blue, 69 Glandular, 68 Glossogyne Cass. hennedyi R. Br., 439 Gnaphalium L., 428, 555 arvense L., 426 bellidioides (G. Forst.) Hook. fil., 432 dichotomum Salisb., 425 dioicum L., 427 gallicum (L.) Hill, 426 lagopus Stephan ex Willd., 426 lanatum auct., 432 luteoalbum L., 431 margaritaceum L., 428 maritimum (L.) Hill, 470 minimum Sm., 426 norvegicum Gunnerus, 429 pensylvanicum Willd., 430 purpureum L., 430 rectum Sm., 429 supinum L., 430 sylvaticum L., 429 var. alpestre Br¨ugger, 429 var. nigrescens auct., 429 subsp. norvegicum (Gunnerus) Hook. fil., 429

589

Gnaphalium sylvaticum L. (cont.) var. norvegicum (Gunnerus) Hook. & Arn., 429 var. rectum (Sm.) Gaudin, 429 var. sylvaticum, 429 uliginosum L., 429 undulatum L., 431 viscosum Kunth, 428 Goat’s-beard, 110 Golden bush, 432 Goldenrod, 440 Canadian, 441 Early, 442 Grass-leaved, 442 Lime, 441 Rough-stemmed, 441 Golden Thistle, 94 Spotted, 93 Goldilocks, Fine-leaved, 452 Shrub, 452 Gomozia Lam. granadensis Mutis ex L. fil., 15 Gorteria L. rigens L., 423 uniflora L. fil., 423 var. pinnate Harv., 423 Grindelia Willd., 438 andersonii Piper, 439 hendersonii Greene, 439 lanata Greene, 439 macrophylla Greene, 439 oregana A. Gray, 439 stricta DC., 439 Grossheimia Sosn. & Takht. macrocephala (Muss. Puschk. ex Willd.) Sosn. & Takht., 89 Groundsel, 501 Eastern, 502 Heath, 502 Purple, 503 Sticky, 502 Tree, 462 Welsh, 500 Guelder Rose, 37 American, 40 Asian, 40 Guizotia Cass., 516 abyssinica (L. fil.) Cass., 516 oleifera (DC.) DC., 516 Gum-plant, Coastal, 439 Gymnocline Cass., 463 leucocephala Cass., 464 macrophylla (Waldst. & Kit.) Bluff & Fingerh., 464 Haplostephium D. Don, 202 Harebell, 7 Broad-leaved, 7 Hawkbit, Autumn, 101 Lesser, 103 Rough, 103 Scaly, 97 Star, 98

590 Hawk’s-beard, Beaked, 207 Bristly, 208 French, 206 Leafless, 204 Marsh, 202 Narrow-leaved, 240 Northern, 203 Rough, 203 Smooth, 206 Stinking, 205 Tuberous, 111 Hawkweed, Aberfeldy, 380 Aggregate-headed, 378 Alpine, 294 Ample-toothed, 401 Anglian, 371 Apiculate-toothed, 346 Aristate-toothed, 307 Armadale, 379 Arran, 311 Autumn, 241 Backhouse’s, 288 Baker’s, 245 Bald-leaved, 290 Baldock, 367 Balloon-leaved, 368 Bank, 418 Barras, 419 Basalt, 346 Beacons, 322 Beautiful, 375 Beautiful-leaved, 298 Beeby’s, 325 Beinn Dearg, 279 Ben Hope, 391 Ben More, 310 Berriedale, 331 Bettyhill, 343 Black-bracted, 412 Black-glandular, 303 Black-headed, 299 Bloody, 388 Bluish-leaved, 239 Blunt-leaved, 339 Blunt-toothed, 378 Boat-leaved, 386 Borrer’s, 259 Borrowdale, 276 Boswell’s, 330 Braemar, 258 Brecon 347 Breac, 301 Breadalbane, 384 Breast-toothed, 368 Brindled, 392 Bristol Channel, 355 British, 350 Broad-headed, 254 Broad-leaved, 252 Burnside, 388

Index Hawkweed (cont.) Caernarvon, 338 Caesius, 360 Cairngorm, 282 Caithness, 323 Calcareous, 392 Caledonian, 329 Candelabra, 403 Carboniferous, 389 Carnedd, 335 Cat’s-ear, 343 Cave, 268 Chapel-le-dale, 385 Chedder, 355 Cher, 377 Cille, 341 Claret-leaved, 361 Clasping-leaved, 314 Cliff, 297 Cliva Hill, 264 Close-toothed, 385 Clova, 306 Coldbackie, 329 Common, 357 Complete, 272 Confused, 351 Crag, 308 Craig Gleisiad, 381 Crass-leaved, 380 Cravon, 357 Cumbrian, 309 Cup-shaped, 337 Cusped-toothed, 303 Dales, 345 Dappled, 395 Dark-bracted, 380 Dark-flowered, 384 Dark-headed, 247 Dark-leaved, 369 Dark-styled, 248 Dartmoor, 420 Deganwy, 324 Dense-branched, 401 Dense-haired, 307 Dense-leaved, 244 Derby, 354 Dewar’s, 260 Diaphanous, 371 Disc-leaved, 395 Distinguished, 363 Donegal, 328 Double-toothed, 364 Dovre, 265 Doward, 408 Drummond’s, 247 Dumfries, 386 Dusky-headed, 302 Dyfed, 377 Ebudes, 344 Elegant, 288

Index Hawkweed (cont.) Elongate-leaved, 387 English, 316 Entire-leaved, 312 Equal-toothed, 360 Erin, 314 Excellant, 273 Exmoor, 241 Fang-toothed, 370 Farr, 323 Fat-leaved, 330 Fimbriate-pitted, 313 Fine-bracted, 371 Fionn Glen, 335 Fisher Ghyll, 310 Flat-leaved, 346 Fleck-leaved, 391 Fleshy-leaved, 347 Floccose, 319 Floccose-bracted, 311 Foreign, 419 Forked, 347 Four-toothed, 402 Garden, 409 Giant-toothed, 402 Gigantic-toothed, 402 Glabrous-headed, 238 Glamorganshire, 401 Glandular-toothed, 309 Glen Einich, 283 Glenridding, 256 Gloucester, 408 Golden-flowered, 307 Goring Gap, 366 Gouget’s, 396 Grampian, 340 Grand-toothed, 400 Granite, 281 Grassland, 374 Green-flowered, 326 Green-leaved, 374 Grey-bracted, 412 Grey-haired, 358 Grey-headed, 356 Griffith’s, 325 Groves’s 277 Hairless-bracted, 255 Hairy-bracted, 311 Hairy-headed, 381 Hairy-leaved, 376 Hairy-stemmed, 251 Hanbury’s, 270 Handsome, 266 Hard-headed, 393 Harris, 325 Hart’s, 315 Hastate-leaved, 298 Heart-leaved, 412 Hebridean, 318 Hepste, 321

Hawkweed (cont.) Highland, 284 Hill, 379 Hirsute, 292 Hjelt’s, 419 Humphrey Head, 337 Irish, 258 Irregular-toothed, 364 Isabel’s, 298 Island, 397 Jagged-toothed, 300 Javeline-leaved, 317 Jordan’s, 415 Keld Head, 366 Kenneth’s, 272 Kent, 250 Kent’s, 407 Kerry, 257 Killarney, 332 Kingshouse, 302 Kintyre, 360 Klingrahool, 261 Koehler’s, 406 Lacerate-leaved, 367 Laggan, 372 Lairig, 278 Lakeland, 307 Lance-leaved, 343 Langwell, 316 Large-flowered, 249 Large-fruited, 291 Large-headed, 315 Large-leaved, 327 Large-toothed, 239 Laxo Burn, 263 Leek-coloured, 394 Lesser-leaved, 331 Ley’s, 339 Liljeholm’s 397 Limestone, 354 Linton Falls, 359 Linton’s, 382 Llanfairfechan, 255 Llanwrtyd, 370 Long-bracted, 403 Long-haired, 254 Long-leaved, 240 Long-stalked, 378 Long-toothed, 363 Lortet’s, 376 Mainland, 262 Many-toothed, 375 Marbled, 312 Marigold, 274 Maritime, 244 Marshall’s, 300 McCallum Webster’s, 388 Meall Garbh, 394 Memorable, 286 Miles’s, 285

591

592 Hawkweed (cont.) Minute-toothed, 414 Moffat, 304 Mountain, 299 Narrow-bracted, 326 Narrow-leaved, 294 Narrow-toothed, 290 Naver. 420 Neat, 270 Nidd, 257 Nipple-toothed, 409 Noble, 287 North England, 389 Northern, 249 North Roe, 261 Oblong-leaved, 296 Okraquoy, 266 Orange-flowered, 361 Orkney, 379 Ormes Head, 351 Oval-leaved, 311 Oxford, 396 Painted-leaved, 334 Pale, 325 Pale-headed, 358 Pale-leaved, 340 Pale Spotted, 334 Patent-toothed, 406 Peak, 359 Pellucid-leaved, 415 Pentaploid, 293 Penwyllt, 321 Perth, 260 Petite-leaved, 369 Pettycur, 363 Picts, 387 Pointed-leaved, 316 Porrect-bracted, 336 Pruinose, 418 Pugsley’s, 263 Purple-flushed, 335 Purplish-leaved, 253 Purple-tinted, 262 Radyr, 373 Rare, 267 Ravens’, 305 Reay, 246 Red-flushed, 365 Red-tinted, 323 Remarkable, 275 Remote, 260 Repand-leaved, 344 Reticulate-leaved, 246 Rhombic-leaved, 362 Ribbleshead, 391 Riddelsdell’s, 344 Rigid, 238 Riverside, 362 Roberts’s, 360 Rock, 345

Index Hawkweed (cont.) Roffey’s, 359 Ronas Voe, 400 Rosette, 322 Rough, 367 Rough-leaved, 258 Round-headed, 280 Round-leaved, 317 Rubicund, 329 Rusty-red, 361 Sagittate-leaved, 382 Sand, 322 Sannox, 310 Scabrous, 253 Scandinavian, 303 Scarp, 314 Schmidt’s, 342 Scottish, 320 Scott’s, 322 Scully’s, 252 Semblister, 266 Shade, 336 Shaggy, 295 Shaggy-stalked, 317 Shale, 350 Sharp-toothed, 240 Shetland, 267 Shining-leaved, 340 Shoolbred’s, 319 Short-flowered, 420 Short-stemmed, 382 Silurian, 331 Silvery, 332 Sinuate-toothed, 383 Skerry, 390 Skipton, 370 Slender, 245 Slender-bracted, 338 Slender-leaved, 302 Slender-toothed, 254 Small-flowered, 318 Small-headed, 394 Small-leaved, 330 Small-spotted, 383 Small-toothed, 318 Snake-headed, 390 Snowdonia, 421 Sociable, 373 Solitary, 267 Sombre-headed, 305 Sommerfelt’s, 334 Southern, 373 Sowa Dee, 321 Sparse-leaved, 256 Spear-leaved, 289 Spence’s, 263 Splendid, 277 Spotted, 367 Spreading-toothed, 345 Stained-leaf, 366

Index Hawkweed (cont.) Star-haired, 421 Stellate-haired, 319 Stenstr¨om’s, 407 Stewart’s, 251 Sticky, 269 Stiff-haired, 341 Streamside, 305 Strict, 246 Strict-headed, 413 Subglobose, 279 Suffolk, 396 Summit, 327 Surrey, 365 Sutherland, 332 Taf-fechan, 408 Tain, 324 Tall, 372 Tapered-leaved, 320 Tavistock, 409 Tawe, 249 Tenuous-leaved, 283 Thick-leaved, 328 Three-coloured, 336 Thule, 264 Tongue-leaved, 257 Toothed, 251 Toothless, 414 Trackway, 403 Traligill, 312 Triangular-leaved, 385 Tufted-hair, 299 Tutshill, 351 Uig, 255 Uist, 383 Umbellate, 242 Undivided-leaved, 328 Uneven-toothed, 337 Unst, 265 Upland, 308 Valuable, 274 Variable, 369 Variable-leaved, 390 Variable-toothed, 306 Velvet-headed, 301 Vennicontes’s, 304 Violet-leaved, 364 Vorlick, 301 Wall, 269 Wandering, 413 Waterfall, 362 Wathamstow, 375 Wavy-leaved, 297 Wedge-leaved, 397 Weedy, 414 Welsh, 333 Western, 355 Westmorland, 339 West’s, 304 Willow-leaved, 240

Hawkweed (cont.) Wine-leaved, 308 Wine-stemmed, 261 Witley, 250 Wood, 384 Woolly, 313 Wye, 333 Yellow-glandular, 242 Yellow-styled, 248 Yorkshire, 342 Hedypnois Mill., 97 autumnalis (L.) Huds., 101 biennis (L.) Huds., 203 cretica (L.) Dum. Cours., 97 hieracioides (L.) Huds., 107 hirta auct., 103 hispida (L.) Huds., 103 monspeliensis Willd., 97 polymorpha DC., 97 rhagadioloides (L.) F. W. Schmidt, 97 taraxaci (L.) Vill., 102 tubiformis Ten., 97 Helenium L., 527 autumnale L., 527 Helianthus L., 518 annuus L., 518 × decapetalus, 519 aridus Rydb., 519 decapetalus auct., 519 integrifolius Nutt., 519 × laetiflorus Pers., 519 laevis L., 520 × multiflorus L., 519 patens Nutt., 519 pauciflorus Nutt., 519 petiolaris Nutt., 519 rigidus (Desf.) Cass. × tuberosus L., 518 tuberosus L., 519 Helichrysum Mill., 431 bellidioides (G. Forst.) Willd., 432 bracteatum (Vent.) Andrews, 432 capillaceum (Thunb.) Less., 431 expansum (Thunb.) Less., 431 italicum (Roth) Don, 431 odoratissimum (L.) Less., 431 petiolare Hilliard & B. L. Burtt, 432 petiolatum auct., 432 Heliopsis Pers., 520 helianthoides (L.) Sweet, 520 var. helianthoides, 520 var. scabra (Dunal) Fern., 520 laevis (L.) Pers., 520 platyglossa Cass., 516 scabra Dunal, 520 Heliotrope, Winter, 511 Helminthia Juss., 108 echioides (L.) Gaertn., 108 subsp. humifusa (Willd.) Arcangeli, 108

593

594 Helminthia echioides (L.) Gaertn. (cont.) var. mollis Duby, 108, 532 var. pratensis (Chevall.) DC., 108, 532 humifusa (Willd.) Trevis, 108 pratensis Chevall., 108 Helminthotheca Zinn, 108 echioides (L.) Holub, 108 var. incisa P. D. Sell, 108, 532 var. mollis (Duby) P. D. Sell, 108, 532 var. pratensis (Chevall.) P. D. Sell, 108, 532 Hemizonia DC. pungens Torrey & A. Gray, 521 Hemp Agrimony, 528 Heterochaeta DC., 454 Hieraciodes Kuntze, 201 caucasicum (C. A. Mey.) Kuntze, 202 foetidum (L.) Kuntze, 205 glabrum (Boiss.) Kuntze, 203 molle (Jacq.) Kuntze, 203 paludosum (L.) Kuntze, 203 praemorsum (L.) Kuntze, 204 rumicifolium (Boiss.) Kuntze, 202 ruprechtii (Boiss.) Kuntze, 202 setosum (Haller fil.) Kuntze, 208 taraxacifolium (Thuill.) Kuntze, 207 tectorum (L.) Kuntze, 204 vesicarium (L.) Kuntze, 207 Hieracioides Moench, 242 Hieracium L., 218 acamptum P. D. Sell & C. West, 250 acroleucum auct., 357 acroleucum (Stenstr.) Dahlst., 358 var. daedalolepium auct., 369, 357, 358 var. mutabile (Ley) W. R. Linton, 369 acuminatum Jord., 372 acutifolium Vill., 316, 542 adelphicum P. D. Sell, 401, 554 adlerzii auct., 369, 370, 371 adlerzii F. Hanb., 356 aequiserratum P. D. Sell, 360, 550 aestivum Fr. subsp. angustum auct., 245 subsp. hemitrachys Druce & Zahn, 248 subvar. hethlandiae (F. Hanb.) Zahn, 264 subsp. subcrocatum (E. F. Linton) Zahn, 248 subsp. subumbellatiforme Zahn, 245 aggregatifolium P. D. Sell, 322, 543 aggregatum Backh. fil., 378 var. prolongatum F. Hanb., 394, 554 alfvengrenii Dahlst. var. plurifolium Zahn, 373 alpinum L., 294 subsp. alpinum, 295 var. genuinum, 295 forma grande sensu F. N. Williams, 287 forma holosericeum (Backh. fil.) Zahn, 295 var. insigne auct., 294 var. insigne (Backh.fil.) Bab., 287 var. melanocephalum auct., 294 subsp. nigrosetosum auct., 287

Index Hieracium alpinum L. (cont.) subvar. tenellum (Backh. fil.) Zahn, 293 subvar. villosissimum, 295 amaurostictum Walter Scott & R. C. Palmer, 266, 533 amnicola P. D. Sell, 362, 551 amplexicaule L., 269 subsp. pulmonarioides (Vill.) Zahn, 269 var. pulmonarioides (Vill.) Gaudin, 269 subsp. speluncarum (Arv.-Touv.) Zahn, 268 var. subhirsutum Arv.-Touv., 268 ampliatiforme P. D. Sell., 318, 542 ampliatum (W. R. Linton) Ley, 317 var. gracilius Pugsley, 318, 542 var. jaculifolium (F. Hanb.) Pugsley, 317 amydrostictum P. D. Sell, 334, 545 anfractiforme E. S. Marshall, 300 anglicum Fr., 316 var. acutifolium (Backh. fil.) Backh. fil., 316 var. amplexicaule Backh. fil., 314 var. brevifurcatum F. N. Williams, 339 var. brigantum (F. Hanb.) W. R. Lint., 342 var. calcaratum E. F. Linton, 317 var. cerinthiforme (F. Hanb.) F. Hanb., 314 var. decipiens Syme, 314 forma hartii (F. Hanb.) W. R. Linton, 315 var. hartii (F. Hanb.) F. N. Williams, 315 var. jaculifolium F. Hanb., 317 × lancifolium auct., 342 var. longibracteatum F. Hanb., 319 anglorum (Ley) Pugsley, 371 anguinum (W. R. Linton) Roffey, 390 angustatiforme P. D. Sell & C. West, 330 angustatum auct., 330, 393 angustatum (Lindeb.) Lindeb., 331 var. amphiboloides auct., 253 subsp. cacuminum (Ley) Zahn, 327 angustifolium Hoppe, 246, 532, 542 angustisquamum (Pugsley) Pugsley, 323 angustum auct., 245 apheles P. D. Sell, 321, 543 aphyllopodioides F. N. Williams, 252 apiculatidens P. D. Sell, 346, 548 apiculatum (E. F. Linton) Druce, 371 ardaricum Pugsley, 251 ardisodon Dahlst., 379 argentatum (Pugsley) P. D. Sell, 332, 545 argenteum Fr., 332 var. septentrionale F. Hanb., 332, 545 var. subglabratum (F. Hanb.) Pugsley, 331 argentiforme P. D. Sell, 332, 545 argillaceum Jord., 373 argutidens Uechtr., 363 argutifolium Pugsley, 240 aristidens P. D. Sell, 307, 539 armadalense P. D. Sell, 379, 552 arranense P. D. Sell, 311, 541 arrectarium auct., 373 arrostocephalum Omang, 268 arvonense P. D. Sell, 338, 546 ascendentidens P. D. Sell, 325, 544

Index Hieracium L. (cont.) asperatum Jord. ex Boreau, 367 aspernatum Jord. var. anglicum Zahn, 373 asteridophyllum P. D. Sell & C. West, 421 aterrimum Hyl., 406 atraticeps (Pugsley) P. D. Sell & C. West, 272 atratum auct., 303 subsp. centripetale (F. Hanb.) Zahn, 301 subsp. cumbriense auct., 419 subsp. cumbriense (F. Hanb.) Zahn, 309 var. gracilifolium (F. Hanb.) Zahn, 302 subsp. molybdochroum Zahn, 299 subsp. sinuans (F. Hanb.) Zahn, 297 subsp. subnigrescens auct., 302 atriglandulosum P. D. Sell, 303, 538 attenuatifolium P. D. Sell & C. West, 263 aurantiacum L., 211 subsp. carpathicola N¨ageli & Peter, 211 auratiflorum Pugsley, 384 auratum auct., 248 var. thulense F. Hanb., 264 auricula auct., 213 australius (Beeby) Pugsley, 265 aviicola Jord. ex Boreau, 375 backhouseanum auct., 251 backhouseanum (Zahn) Roffey, 252 var. radnoricum Pugsley, 253 backhousei auct., 286 backhousei F. Hanb., 288 bakeranum Pugsley, 245 baliophyllum Dahlst. ex Hyl., 396 barbareifolium auct., 372 bartonii Pugsley, 248 basalticola Pugsley, 346 basicrinum (Zahn) Roffey, 335 bauhinii Besser subsp. arvorum (N¨ageli & Peter) Zahn, 213 beebyanum Pugsley, 325 bettyhillense P. D. Sell, 343, 547 bichlorophyllum (Druce & Zahn) Pugsley, 243 var. curtum (W. R. Linton) Pugsley, 243 bienne (L.) Karsch., 204 bifidum Kit. subsp. aggregatum (Backh. fil.) Zahn, 378 subsp. anguinum (W. R. Linton) Zahn, 390 var. anguinum (W. R. Linton) F. N. Williams, 390 subsp. caesionigrescens (Fr. ex Stenstr.) Zahn, 359 subsp. crebridens auct., 385 subsp. crebridens (F. N. Williams) Zahn, 385 subsp. cymbifolium (Pursh) Zahn, 386 var. delicatulum (W. R. Linton) Zahn, 394 subsp. duriceps (F. Hanb.) Zahn, 393 subsp. maculosum (Dahlst. ex Stenstr¨om) Zahn, 391 subsp. prolixum (Norrl.) Zahn, 391 subsp. scandinavicum auct., 384 subsp. subcyaneum (W. R. Linton) Zahn, 336 subsp. subtenue (W. R. Linton) Zahn, 380 subsp. tricolorans Zahn, 336

Hieracium L. (cont.) bladonii Pugsley, 241, 532 var. brunkeri Pugsley, 241 boreale auct, 238 var. calvatum F. Hanb., 238 var. dumosum (Jord.) Gren. & Godr., 242 var. hervieri auct., 241 var. lacerum Wimm. & Grab., 239 subsp. obliquum (Jord.) Sudre, 241 microg. propinquum Sudre, 241 var. propinquum (Sudre) Rouy, 241 var. rigens (Jord.) Sudre, 238 subsp. vagum (Jord.) Rouy, 238 var. vagum (Jord.) Gren. & Godr., 238 subsp. virgultorum (Jord.) Sudre, 240 boreoanglicum P. D. Sell, 389, 554 boreophilum (Zahn) Roffey, 253 borreri Syme, 259 boswellii E. F. Linton, 330 breacense P. D. Sell, 301, 537 breadalbanense F. Hanb., 384 breconense P. D. Sell, 347, 549 breconicola P. D. Sell, 322, 543 breve Beeby, 267 brevidentatum Jord. var. multiflorum Zahn, 373 brigantum (F. Hanb.) Roffey, 342 britanniciforme auct., 351 britanniciforme Pugsley, 351 britannicoides P. D. Sell, 351, 549 britannicum auct., 346, 347, 351 britannicum F. Hanb., 350 var. glaucinum Pugsley, 347 var. ovale A. Ley, 347, 549 var. stenolepiforme Pugsley, 355 var. subbritannicum auct., 351 var. subbritannicum (Ley) Pugsley, 354 var. vagense F. Hanb., 333 brunneocroceum Pugsley, 211 buglossoides auct., 324, 326 var. subrude (Arv.-Touv.) Arv.-Touv., 324 cacuminatum auct., 373 var. barbareifolium auct., 372 cacuminum (Ley) Ley, 327 caesioides Arv.-Touv. subsp. rionii (Gremli) Zahn, 312 caesiomurorum Lindeb., 387 var. umbraticum K. Johans., 378 caesionigrescens (Fr. ex Stenstr.) Omang, 359 caesiopilosum Pugsley, 360 caesitium Norrl., 391 caesium auct., 351 subsp. adlerzii auct., 370 var. alpestre auct., 306 var. angustatum Lindeb., 331 subsp. caesiomurorum (Lindeb.) Zahn, 378 var. cambricum Baker, 333 var. coracinum Ley, 381 subsp. cravoniense (F. Hanb.) Zahn, 357 var. decolor auct., 335

595

596 Hieracium caesium auct. (cont.) var. decolor W. R. Linton, 336 subsp. dissimile (Lindeb.) Zahn, 303 subsp. eustales (E. F. Linton) Zahn, 311 subsp. farrense (F. Hanb.) Zahn, 323 var. hypochaeroides (S. Gibson) Bab., 343 var. insulare auct., 306 var. insulare F. Hanb., 310 subsp. neomarshallianum Zahn, 300 subsp. petrocharis (E. F. Linton) Zahn, 308 var. petrocharis E. F. Linton, 308 var. rhomboides (Stenstr.) W. R. Linton, 362 var. smithii auct., 336 var. smithii Baker, 351 var. subminutidens Zahn, 370 caespitosum Dumort., 212 subsp. colliniforme (Peter) P. D. Sell, 212 calcaricola (F. Hanb.) Roffey, 251 caledonicum F. Hanb., 329 var. boswellii (E. F. Linton) Zahn, 330 var. platyphyllum (Ley) Ley, 330 var. pseudozetlandicum (Roffey) Pugsley, 322 subsp. rubicundiforme Zahn, 329 subsp. veterascens (Dahlst.) Zahn, 329 calenduliflorum Backh. fil., 274 caliginosum auct., 420 callistophyllum F. Hanb., 298 var. cremnanthes auct., 298 var. cremnanthes (F. Hanb.) F. Hanb., 297 calvatum (F. Hanb.) Pugsley, 238 calviceps Pugsley, 255 var. robustum Pugsley, 255 calvum P. D. Sell & D. J. Tennant, 290 cambricogothicum Pugsley, 255 var. glandulosum Pugsley, 253 cambricum (Baker) F. Hanb., 333 camptopetalum (F. Hanb.) P. D. Sell & C. West, 420 candelabrae auct., 392 candelabrae W. R. Linton, 403 caniceps F. Hanb., 388 cantianum F. Hanb., 250 var. subrigidum E. F. Linton ex F. Hanb., 250 cardiophyllum (Jord. ex Sudre) Juxip, 412 carenorum F. Hanb., 340 carneddorum Pugsley, 335 carpathicum Besser, 260 subsp. dewari (Syme) Zahn, 260 subsp. dovrense (Fr.) Zahn, 265 subsp. euchrysostylum auct., 322 subsp. perthense (F. N. Williams) Zahn, 260 subsp. scullyi (E. F. Linton) Zahn, 252 subsp. truncatum auct., 262 centripetale F. Hanb., 301 cerinthiforme F. Hanb., 314 var. hartii F. Hanb., 315 cerinthoides auct., 314 var. acutifolium Backh. fil., 316, 542 var. anglicum (Fr.) Backh. fil., 316 charitodon P. D. Sell, 254, 532 cheriense Jord. ex Boreau, 377

Index Hieracium L. (cont.) chloranthum Pugsley, 326 chloranthum Pugsley ex P. D. Sell, 326, 545 chlorophyllum auct., 373 chlorophyllum Jord. ex Boreau, 374 chrysanthum Backh. fil., 270 var. gracilentiforme F. Hanb., 276 var. marshallii (E. F. Linton) F. N. Williams, 300 var. microcephalum Backh. fil., 271 chrysolorum P. D. Sell & C. West, 307 ciliatiflorum auct., 385, 388 ciliatum auct., 365, 388 var. repandum Ley, 344 var. venosum Ley, 344 cillense Pugsley, 341 cinderella (Ley) Ley, 403 cinerascens auct., 341, 342 clivicola (F. Hanb.) Pugsley, 379 clovense auct., 307 clovense E. F. Linton, 306 var. griffithii F. Hanb., 325 var. uistense Pugsley, 397 colliniforme (Peter) Roffey, 212 collinum Tausch subsp. colliniforme Peter, 212 commixtum Jord., 366 commutatum (Lindeb.) Elfstr., 299 completum P. D. Sell & C. West, 272 concinnatum (F. Hanb.) Roffey, 215 congestum (Beeby) Roffey, 261 forma vinaceum (Beeby) Roffey, 261 coniops Norrl., 358 var. hebescens Brenner, 358 consociatum Jord. ex Boreau, 373 conspurcans Norrl. subsp. callistophyllum (F. Hanb.) Zahn, 298 subsp. cremnanthes (F. Hanb.) Zahn, 297 cordigerum Norrl. var. asymmetricum (Ley) Roffey, 337 var. subcyaneum (W. R. Linton) Roffey, 336 var. tricolorans (Zahn) Roffey, 336 corymbosum auct., 248 var. melanoglochin E. F. Linton, 247 var. pycnotrichum W. R. Linton, 244 var. salicifolium auct., 245, 248 forma. subumbellata F. N. Williams, 243 var. umbellatiforme W. R. Linton, 245 crassum Almq., 380, 552 cravoniense (F. Hanb.) Roffey, 357 var. pseudoduriceps Pugsley, 357 crebridens auct., 384 crebridens F. N. Williams, 385 crebidentiforme Pugsley, 385 cremnanthes (F. Hanb.) Pugsley, 297 crinellum Omang, 339 croaticum Waldst. & Kit., 203 crocatum auct., 245, 246 var. congestum Beeby, 261 forma. hethlandiae (F. Hanb.) Roffey, 264 var. hethlandiae (F. Hanb.) Druce, 264

Index Hieracium crocatum auct. (cont.) var. maritimum F. Hanb., 244 var. thulense (F. Hanb.) Druce, 264 var. vinaceum Beeby, 261 croceostylum Pugsley, 238, 532 cumbriense auct., 419 cumbriense F. Hanb., 309 cuneifrons (Ley) Pugsley, 397 var. decipiens Pugsley, 407 curvatum auct., 272 cuspidens P. D. Sell & C. West, 303 cyathis (Ley) W. R. Linton, 337 cyclicum P. D. Sell, 317, 542 cymbifolium Purchas, 386 daedalolepioides (Zahn) Roffey, 369 dalense P. D. Sell, 345, 548 dasypodum Dahlst., 347 dasythrix (E. F. Linton) Pugsley, 307 deargicola P. D. Sell & D. J. Tennant, 279, 533 decolor auct., 335 decolor (W. R. Linton) Ley, 336 deductum auct., 373 deductum Sudre, 375 deganwyense Pugsley, 324 demissum Str¨omfelt var. australius (Beeby) Druce, 265 var. cacuminum (Ley) W. R. Linton, 327 var. pulchelliforme (W. R. Linton) Zahn, 266 subsp. zetlandicum (Beeby) Zahn, 267 denticulatum auct., 259 dentifex E. F. Linton, 342 dentulum (E. F. Linton) P. D. Sell, 328, 545 devoniense (F. Hanb.) Roffey, 355 dewari Syme, 260 diaphanoides auct., 371 diaphanoides Lindeb., 371 var. apiculatum E. F. Linton, 371 var. ornatum auct., 369 diaphanum Fr., 369 subsp. acroleucum Stensr., 358 var. cacuminum Ley, 327 var. glaucovirens auct., 372 var. praestans W. R. Linton, 363 dicella auct., 347 dicella P. D. Sell & C. West, 347 difficile P. D. Sell & C. West, 266 dilectum P. D. Sell & C. West, 262 dipteroides Dahlst., 380 discophyllum P. D. Sell & C. West, 395 dispalatum Jord. var. macrodon Sudre, 239, 532 dissimile (Lindeb.) T. Durande & B. D. Jackson, 303 var. majus Pugsley, 303 var. poliaenum auct., 378 divaricatum G. Don, 296 diversidens P. D. Sell & C. West, 306 donegalense Pugsley, 251 dovrense Fr., 265 var. australius Beeby, 265 var. hethlandiae F. Hanb., 264

Hieracium dovrense Fr. (cont.) var. pulchelliforme (W. R. Linton) Beeby, 266 var. spectabile E. S. Marsh., 260 var. vulgare Lindeb., 265 dowardense P. D. Sell, 408, 555 drummondii Pugsley, 247 dumosum Jord., 242 duplicatum auct., 304 var. stenophytes (W. R. Linton) W. R. Linton, 362 duriceps F. Hanb., 393 var. camptopetalum (F. Hanb.) Pugsley, 420 var. cravoniense F. Hanb., 357 var. delicatulum (W. R. Linton) Pugsley 394 eboracense Pugsley, 249 ebudicum Pugsley, 344 einichense P. D. Sell & D. J. Tennant, 283 elatius (Ley) Druce, 408, 555 elatius Rehmann, 408, 555 elegantiforme Dahlst., 267 elevatum P. D. Sell, 408, 555 elongatifolium P. D. Sell, 387, 553 eminens auct., 238, 241 eminentiforme Pugsley, 241 epileucoides Dahlst., 256 erubescens Jord. ex Boreau, 365 erythraeum E. F. Linton ex Pugsley, 323 eucallum P. D. Sell & C. West 345 euprepes F. Hanb., 379 var. clivicola F. Hanb., 379 var. glabratum E. F. Linton, 304 var. pruiniferum W. R. Linton, 418 var. stenophyes (W. R. Linton) F. N. Williams, 362 eustales auct., 319 eustales E. F. Linton, 311 eustomon (E. F. Linton) Roffey, 355 var. devoniense (F. Hanb.) Pugsley, 355 eximium Backh. fil., 292 forma tenellum (Backh. fil.) P. D. Sell & C. West, 293 var. tenellum Backh. fil., 293 exotericum Jord. ex Boreau, 415 var. candelabrae (W. R. Linton) Pugsley, 403 var. cinderella (Ley) Pugsley, 403 var. glevense Pugsley, 408 forma grandidens (Dahlst.) Pugsley, 408 var. sublepistoides (Zahn) Pugsley, 412 faeroense Dahlst., 256 farrense F. Hanb., 323 fastigiatum Fr., 373 festinum Jord. ex Boreau, 376 subsp. deductum (Sudre) Sudre, 375 subvar. latebrosum (Jord. ex Boreau) Sudre, 375 fictum Jord. ex Boreau, 367 filisquamum P. D. Sell, 326, 544 firmiramum Hyl., 401 fissuricola P. D. Sell, 310, 540 flagellare Willd., 214 subsp. bicapitatum (P. D. Sell & C. West) P. D. Sell, 214 flocculipubens P. D. Sell, 319, 542 flocculosiforme P. D. Sell, 318, 542

597

598 Hieracium L. (cont.) flocculosum Backh. fil. ex Bab., 319 var. angustifolium Pugsley, 319, 542 var. cambricum (Baker) F. N. Williams, 333 var. denticulatum Pugsley, 318, 542 var. insulare (F. Hanb.) F. N. Williams, 318 florentinum All. subsp. praealtum (Vill. ex Gochnat) N¨ageli & Peter, 213 floribundum Wimm. & Grab., 212 subsp. helveolum Dahlst., 212 fragilicaule Pugsley, 253 forma subhirsutum Pugsley, 253 fratrum Pugsley, 350 friesii auct., 262 var. hirsutum F. Hanb., 254 var. latifolium (Backh.) E. F. & W. R. Linton, 252 var. stewartii F. Hanb., 251 fucatifolium P. D. Sell, 334, 546 fulvocaesium Pugsley, 361 furcilliferum Dahlst., 347 furculatum Elfstr. subsp. dissimile (Lindeb.) Elfstrand, 303 gentile Jord. ex Boreau, 419 glabrescens Murr, 360, 550 gibsonii Backh. fil., 343 glanduliceps P. D. Sell & C. West, 366 glandulidens P. D. Sell & C. West, 309 glevense (Pugsley) P. D. Sell & C. West, 408 globosiflorum Pugsley, 280 var. lancifolium Pugsley, 289 var. larigense Pugsley, 278 globosum Backh. fil., 280 gothiciforme auct., 254 gothicoides Pugsley, 254 gothicum auct., 254, 255 var. basifolium auct., 254 forma latifolium (Backh.) W. R. Linton, 252 var. latifolium Backh., 252 var. stewartii (F. Hanb.) F. Hanb., 251 gougetanum Gren. & Godr., 396 gracilentum auct., 283 gracilentum Backh. fil., 285 var. graniticola (W. R. Linton) W. R. Linton, 281 gracilifolium auct., 302, 303 gracilifolium (F. Hanb.) Pugsley, 302 grampianum P. D. Sell, 340, 547 grandescens auct., 253 grandidens Dahlst., 400 graniticola W. R. Linton, 281 gratum P. D. Sell & C. West, 266 gravestellum Dahlst. var. rhomboides Stenstr., 362 griffithii (F. Hanb.) F. Hanb., 325 grovesii Pugsley, 277 halfdanii Osk., 265 halleri auct., 292 var. calenduliflorum (Backh. fil.) F. N. Williams, 274 hanburyanum Zahn, 323 hanburyi Pugsley, 270 forma atraticeps (Pugsley) P. D. Sell & D. J. Tennant, 277

Index Hieracium hanburyi Pugsley (cont.) var. atraticeps Pugsley, 272 var. humile Pugsley, 271 var. microcephalum (Backh. fil.) Pugsley, 271 forma pusillum (Pugsley) P. D. Sell & D. J. Tennant, 271 hartianum Pugsley, 251 hartii (F. Hanb.) P. D. Sell & C. West, 315 hartzianum Dahlst., 268 hastiforme P. D. Sell & C. West, 298 hebridense Pugsley, 318 helveolum (Dahlst.) Pugsley, 212 hesperium P. D. Sell, 328, 545 heterophyllum Bladon, 241 hethlandiae (F. Hanb.) Pugsley, 264 hibernicum F. Hanb., 258 var. vennicniorum Pugsley, 258 hjeltii Norrl., 419 holophyllum W. R. Linton, 328 var. angustisquamum Pugsley, 323 var. dentulum E. F. Linton, 328, 545 holopleurum auct., 379 holosericeum Backh. fil., 295 hortense Hyl., 409 humidorum Almq., 264 hyparcticoides Pugsley ex P. D. Sell, 312, 542 hyparcticum auct., 312 hypochaeroides S. Gibson, 343 var. cyathis Ley, 337 var. griseum Ley, 341 var. lancifolium W. R. Linton, 343, 548 var. saxorum F. Hanb., 345 hypophalacrum P. D. Sell, 331 inaequilaterum P. D. Sell, 337, 546 incisifolium Jord. ex Boreau, 367 inquinatum Jord. ex Boreau, 336 insigne Backh. fil., 287 forma celsum P. D. Sell & D. J. Tennant, 287 forma insigne, 287 inspissatum P. D. Sell, 380, 552 insulare (F. Hanb.) F. Hanb., 310 var. petrocharis (E. F. Linton) F. Hanb., 308 integratum auct., 392 integratum (Dahlst. ex Stenstr¨om) Dahlst., 414 integrilaterum Dahlst. var. stenopholidium Dahlst., 355 inuloides Arv.-Touv. var. amplidentatum (F. Hanb.) Zahn, 245 forma angustifolium Dahlst., 244 subsp. latobrigorum Zahn, 248 subsp. opsianthum (Dahlst. ex F. Hanb.) Zahn, 246 var. pseudauratiforme Zahn, 248 subsp. pycnotrichum (W. R. Linton) Zahn, 244 subsp. strictiforme Zahn, 246 iricum Fr., 314 irregularidens P. D. Sell, 378, 551 irriguum auct., 371 var. haematophyllum Dahlst., 365 subsp. lepidiceps Dahlst., 370 subsp. lepidulum (Stenstr.) Dahlst., 364 subsp. violascens Almq. ex Dahlst., 364, 551

Index Hieracium L. (cont.) isabellae E. S. Marshall, 298 itunense Pugsley, 419 jaculifolium (F. Hanb.) Roffey, 317 var. calcaratum (E. F. Linton) Roffey, 317 johnstonii Dahlst., 248 jovimontis auct., 340 juranum Fr. subsp. pseudelatum auct., 259 kalsoense Dahlst. subsp. burnense Druce & Zahn, 379 kennethii P. D. Sell & D. J. Tennant, 272 kentii P. D. Sell, 407 killinense auct., 393 killinense (Zahn) Roffey, 394 kingshousense P. D. Sell, 302, 537 kintyricum P. D. Sell, 360, 550 klingrahoolense Walter Scott & R. C. Palmer, 261, 533 koehleri Dahlst., 406 lacerum Reut. ex Fr., 239 lachenalii auct., 373 forma nemorale Pugsley, 377 var. pseudoporrigens Pugsley, 377 var. radyrense Pugsley, 373 var. transiens Ley, 373 lactucaceum Schrank ex DC., 204 lactucella Wallr., 213 laetificum P. D. Sell & C. West, 299 laevicaule Jord. subsp. acroleucum (Stenstr.) Zahn, 358 subsp. sejunctum (W. R. Linton) Zahn, 357 subsp. triviale (Norrl.) Zahn, 356 var. baltorum Zahn, 357 laevigatum Willd. subsp. backhouseanum Zahn, 252 subsp. bichlorophyllum Druce & Zahn, 243 subsp. boreophilum Zahn, 253 subsp. calcaricola (F. Hanb.) Zahn, 251 subsp. cantianum (F. Hanb.) Zahn, 250 forma centonata F. N. Williams, 256 subsp. friesii auct., 262 subvar. glandulosiceps Zahn, 251 subsp. grandescens auct., 253 subsp. linguans Zahn, 257 subsp. lissolepium Zahn, 255 subsp. longiciliatum (F. Hanb.) Zahn, 254 subsp. nidense (F. Hanb.) Zahn, 257 subsp. placerophyllum auct., 253 var. polyphyllum Zahn, 251 subsp. scabrisetum Zahn, 253 var. serpentinum (F. Hanb.) Zahn, 256 subsp. sparsifolium (Lindeb.) Zahn, 256 subsp. stewartii (F. Hanb.) Zahn, 251 subsp. stictophyllum (Dahlst.) Zahn, 256 subsp. subgracilipes auct., 255 subsp. substrigosum Zahn, 254 subsp. tavense (Ley) Zahn, 249 subsp. trichocaulon (Dahlst.) Zahn, 251 lagganense P. D. Sell, 372, 551 lakelandicum P. D. Sell, 307, 539

Hieracium L. (cont.) lanatum Vill., 313 lanceolatum Vill., 258 lancifolium Vuk., 343, 548 langwellense F. Hanb., 316 lapeyrousii auct., 314 lapponicum auct., 254 larigense (Pugsley) P. D. Sell & C. West, 278 lasiophyllum auct., 342 lasiophyllum W. D. J. Koch, 341 var. euryodon auct., 342 var. planifolium auct., 345 var. planifolium F. Hanb., 345 var. semiglabrum auct. 338, 339 latebrosum Jord. ex Boreau, 375 latiusculum (N¨ageli & Peter) Roffey, 215 latobrigorum (Zahn) Roffey, 248 forma angustifolium Pugsley, 248 lawsonii auct., 314 lepidiceps (Dahlst.) Prain, 370 lepidulum (Stenstr.) Omang, 364 leptodon P. D. Sell & D. J. Tennant, 290 leucograptum auct., 306 leyanum (Zahn) Roffey, 330 leyi auct., 340 leyi F. Hanb., 339 var. vestitum Ley ex E. F. Linton, 338, 547 liljeholmii Dahlst., 397 lima F. Hanb., 342 var. brigantum F. Hanb., 342 linguans (Zahn) Roffey, 257 lineatum Almq. ex Stenstr¨om, 255 lingulatum Backh. fil. ex Hook. & Arn., 296 lintonianum Druce, 382 lintonii Ley., 382 lissolepium Roffey, 255 listerae Pugsley, 246 longiciliatum (F. Hanb.) Roffey, 254 longilobum (Dahlst. ex Zahn) Roffey, 304 longiramosum Pugsley, 255 × longisquamum Peter, 218 lortetiae Balbis, 376 lucidulum (Ley) Roffey, 415 lycopifolium Froel. var. norvegicum Lindb., 246 macrocarpum Pugsley, 291 macrodon N¨ageli & Peter, 239, 532 macrodon Sudre, 400 maculatum auct., 366 maculatum Sm., 367 subsp. asperatum (Jord. ex Boreau) Zahn, 367 subsp. commixtum (Jord.) Zahn, 366 var. genuinum auct., 371 subsp. fictum (Jord. ex Boreau) Zahn, 367 forma immaculatum Zahn, 368 forma incisifolium (Jord. ex Boreau) Zahn, 367 var. inquinatum (Jord. ex Boreau) Zahn, 366 subsp. naevuliferum (Jord. ex Boreau) Zahn, 367 subsp. pollichiae (Sch. Bip.) Zahn, 359 var. spilophaeum (Jord. ex Boreau) Zahn, 367

599

600 Hieracium L. (cont.) maculatum Teesdale ex Turner & Dillwyn, 367 maculoides P. D. Sell & C. West, 383 maculosum auct., 396 maculosum (Dahlst. ex Stenstr¨om) Omang, 391 magniceps P. D. Sell & C. West, 315 magyaricum N¨ageli & Peter subsp. arvorum N¨ageli & Peter, 213 subsp. thoumasium Peter, 213 mammidens P. D. Sell, 368, 551 marginatum P. D. Sell & C. West, 289 forma chaetocephalum P. D. Sell & C. West, 290 forma marginatum, 290 mariae P. D. Sell, 388, 553 maritimum F. J. Hanb., 244 var. pauciceps Pugsley, 244 marshallii E. F. Linton, 300 var. cremnanthes F. Hanb., 297 medium Jord., 409 medium Lindeb., 371 megapodium Dahlst., 371 melanocephalum auct., 294 melanocephalum Lindeb., 253 var. insigne (Backh. fil.) Bab., 287 melanochloricephalum Pugsley, 305 melanoglochin (E. F. Linton) P. D. Sell, 247, 532 melanolepis Almq. ex Norrl., 415 memorabile P. D. Sell & C. West, 286 micracladium (F. N. Williams) Ley, 394 micracladium (Zahn) Omang, 393, 554 microdon Dahlst., 332 microphyllum Hyl., 321, 543 microspilum (Jord. ex Sudre) A. W. Hill, 414 milesii P. D. Sell & C. West, 285 mirandum P. D. Sell & C. West, 260 molle Jacq., 203 molybdochroum Omang, 299 monstrosum Hyl., 409 mougeotii Froel. forma acutifolium (Backh.) Zahn, 316 subsp. anglicum (Fr.) Zahn, 316 var. cerinthiforme (F. Hanb.) Zahn, 314 forma hartii (F. Hanb.) Zahn 315 subsp. iricum (Fr.) Zahn, 314 subsp. skyense Zahn, 319 mucronellum P. D Sell & C. West, 319 mundum P. D Sell & C. West, 288 murorum L. subsp. caliginosum auct., 420 var. camptopetalum F. Hanb., 420 subsp. candelabrae (W. R. Linton) Zahn, 403 microg. cardiophyllum Jord. ex Sudre, 412 subsp. cardiophyllum (Jord. ex Sudre) Zahn, 412 var. cardiophyllum (Jord. ex Sudre) Rouy, 413 subsp. crassiceps auct., 387 var. crassiusculum auct., 392 var. cuneifrons (Ley) Zahn, 397 var. dissimile Lindeb., 303 subsp. exotericum (Jord. ex Boreau) Sudre, 415 var.. exotericum (Jord. ex Boreau) Rouy, 415 microgen. gentile (Jord. ex Boreau) Sudre, 419

Index Hieracium murorum L. (cont.) subsp. gentile (Jord. ex Boreau) Zahn, 419 subsp. grandidens (Dahlst.) Zahn, 400 subsp. hjeltii (Norrl.) Zahn, 419 subsp. integratum (Dahlst. ex Stenstr.) Zahn, 414 subsp. killinense Zahn, 394 subsp. lepidulum Stenstr., 364 subsp. longilobum Zahn, 304 subsp. lucidulum (Ley) Zahn, 415 var. lucidulum Ley, 415 microg. macrodon Sudre, 400 var. medium (Jord.) Gren. & Godr., 409 subsp. micracladium Zahn, 554 subsp. microspilum (Jord. ex Sudre) Zahn, 414 var. microspilum Jord. ex Sudre, 414 subsp. neosparsum Zahn, 418, 555 subsp. oblongum (Jord.) Zahn, 414 var. oblongum (Jord.) Gren. & Godr., 414 var. pachyllum auct., 384 subsp. pachyphylloides Zahn, 389 var. pachyphyllum auct., 346 var. pachyphyllum Purchas., 389 subsp. patale (Norrl.) Zahn, 420 subsp. pellucidum (Laest.) Zahn, 415 var. pellucidum (Laest.) F. Hanb., 415 var. phaeotrichum auct., 384 subsp. pictorum (E. F. Linton.) Zahn, 387 subsp. pollinarium (F. Hanb.) Zahn, 420 subsp. pruinale Zahn, 418 subsp. pulcherrimum (F. Hanb.) Zahn, 393 var. pulcherrimum auct., 421 var. pulcherrimum F. Hanb., 393 var. sagittatum Lindeb., 382 var. sanguineum Ley, 388 subsp. stenstroemii (Dahlst.) Zahn, 407 subsp. subcrassum (Almq. ex Dahlst.) Zahn, 403 var. sublepistoides Zahn, 412 subsp. subsanguineum Zahn, 388 var. subulatidens auct., 418 var. sylvivagum (Jord. ex Boreau) Zahn, 413 subsp. sylvularum (Jord. ex Boreau) Zahn, 401 var. sylvularum (Jord. ex Boreau) Sudre, 401 subsp. variicolor auct., 392 mutabile (Ley) Ley, 369 naevuliferum Jord. ex Boreau, 367 naviense J. N. Mills, 354 nemophilum Jord. ex Boreau, 374 neocoracinum Pugsley, 381 neocorymbosum Pugsley, 248 var. crocatiforme Pugsley, 248 neomarshallianum (Zahn) Roffey, 300 neomicrocladium P. D. Sell 393, 554 neopinnatifidum Pugsley, 363 neosparsum (Zahn) P. D. Sell, 418, 555 nidense (F. Hanb.) Roffey, 257 nigrescens auct., 272, 302 subsp. backhousei (F. Hanb.) Zahn, 288 subsp. calenduliflorum (Backh. fil.) Zahn, 274 subsp. chrysanthum Zahn, 270 subsp. commutatum (Lindeb.) Lindeb., 299 var. commutatum Lindeb., 299

Index Hieracium nigrescens auct. (cont.) var. curvatum auct., 272 subsp. eximium (Backh. fil.) Zahn, 292 subsp. globosum Zahn, 280 var. globosum Hook. fil., 280 var. gracilifolium F. Hanb., 302 subsp. graniticola (W. R. Linton) Zahn, 281 subsp. lingulatum (Backh. fil. ex Hook. & Arn.) Zahn, 296 var. lingulatum (Backh. fil. ex Hook. & Arn.) Hook. fil., 296 subsp. pseudocurvatum Zahn, 282 subsp. pseudopetiolatum Zahn, 294 var. senescens (Backh.) Hook. fil., 299 subsp. subgracilentipes Zahn, 276 nigrifactum P. D. Sell, 302, 538 nigrisquamum P. D. Sell & C. West, 299 nigroglandulosum L¨onnr., 415 nitidum Backh. fil., 340 var. carenorum (F. Hanb.) Pugsley, 340 var. siluriense F. Hanb., 331, 545 northroense Pugsley, 261 norvegicum Fr. var. confertum auct., 256 subsp. hibernicum (F. Hanb.) Zahn, 258 notabile P. D. Sell & C. West, 275 obesifolium Pugsley, 244 obliquum Jord., 241 oblongum Jord., 414 occitanum auct., 241 ochthophilum P. D. Sell, 305, 538 oenophyllum P. D. Sell, 308, 540 ogwenii E. F. Linton, 243 oistophyllum Pugsley, 382 var. abrasum (Dahlst.) Pugsley, 379 oligodon (E. F. Linton) Pugsley, 257 olivaceum Gren. & Godr. subvar. calcaratum (E. F. Linton) Zahn, 317 subsp. flocculosum (Backh. fil. ex Bab.) Zahn, 319 subsp. jaculifolium (F. Hanb.) Zahn, 317 subsp. langwellense (F. Hanb.) Zahn, 316 subsp. llynense Zahn, 345 subvar. longibracteatum (F. Hanb.) Zahn, 319 subsp. pseudolangwellense Zahn, 297 subsp. subrubicundum (Dahlst.) Zahn, 327 onosmoides auct., 324, 326 var. buglossoides auct., 324 subsp. subrude (Arv.-Touv.) Zahn, 324 onychodontum Hyl., 402 opsianthum (Dahlst. ex F. Hanb.) Roffey, 246 optimum P. D. Sell & C. West, 277 orarium Lindb. var. angustatum (Lindeb.) F. N. Williams, 331 var. erythraeum E. F. Linton, 323 var. fulvum auct., 323 var. fulvum F. Hanb., 361 var. ravusculum auct., 357 orbicans auct., 392 orcadense W. R. Linton, 379 oreades auct., 331 var. subglabratum F. Hanb., 331, 545 orimeles F. Hanb. ex W. R. Linton, 325

Hieracium L. (cont.) orimeles F. Hanb., 325 var. argentatum Pugsley, 332, 545 var. obscurum Pugsley, 326 orithales E. F. Linton, 382 ornatilorum P. D. Sell & C. West, 249 ostenfeldii Dahlst., 268 ovaliforme P. D. Sell, 311, 541 oxybeles P. D. Sell, 316, 542 oxyodus (W. R. Linton) W. R. Linton, 304 var. delicatulum W. R. Linton, 394 pachylodes N¨ageli & Peter, 218 pachyphylloides (Zahn) Roffey, 389 pachyphyllum (Purchas) F. N. Williams, 389 pallescens Waldst. & Kit. var. pollichiae (Sch. Bip.) Arv.-Touv., 359 pallidum auct., 347 subsp. crinellum (Omang) Zahn, 339 subsp. devoniense (F. Hanb.) Zahn, 355 subsp. eustomon (E. F. Linton) Zahn, 355 subsp. jovimontis auct., 340 subsp. lasiophyllum (W. D. J. Koch) Zahn, 341 subsp. leyi (F. Hanb.) Zahn, 339 var. planifolium (F. Hanb.) Zahn, 346 subsp. lima (F. Hanb.) Zahn, 342 var. persicifolium auct., 332 subsp. schmidtii (Tausch) Zahn, 342 paludosum L., 202 paniculatum Gilib., 202 paraliaeforme Dahlst., 379 patale Norrl., 420 patens Dahlst., 314 pauculidens P. D. Sell & C. West., 390 peccense (W. R. Linton) P. D. Sell, 359, 550 peleteranum M´erat, 218 var. peleteranum Pugsley, 218 subsp. subpeleteranum N¨ageli & Peter, 218 subsp. tenuiscapum (Pugsley) P. D. Sell, 218 pellucidum Laest., 415 var. lucidulum auct., 395, 421 var. lucidulum (Ley) W. R. Linton, 415 var. pulcherrimum (F. Hanb.) W. R. Linton, 393 pensum P. D. Sell & C. West, 274 pentaploideum P. D. Sell & D. J. Tennant, 293, 536 perampliforme auct., 316 perampliforme Dahlst., 314 peramplum Dahlst., 314 percissum Jord. ex Boreau, 363 pernigrescens (Zahn.) Roffey, 215 peroblongum P. D. Sell, 339, 547 perpropinquum (Zahn.) Druce, 241 perscitum P. D. Sell & C. West, 270 perthense F. N. Williams, 260 petiolatum auct., 294 petrocharis (E. F. Linton) W. R. Linton, 308 philanthrax auct., 382 phrixoclonum Omang, 265 pictorum E. F. Linton, 387 var. breadalbanense (F. Hanb.) F. N. Williams, 384 var. dasythrix E. F. Linton, 307 piligerum (Pugsley) P. D. Sell & C. West, 392

601

602 Hieracium L. (cont.) pilosella L., 214, 215 var. angustifolium Tausch, 215 subsp. angustius N¨ageli & Peter, 215 var. concinnatum F. Hanb., 215 var. concolor Tausch, 215 subsp. euronota N¨ageli & Peter, 215 subsp. latiusculum N¨ageli & Peter, 215 subsp. melanops Peter, 215 subsp. micradenium N¨ageli & Peter, 215 subsp. nigrescens (Fr.) P. D. Sell & C. West, 215 var. nigrescens Fr., 215 subsp. pernigrescens Zahn, 215 var. pseudopilosella auct., 215 var. stoloniflorum auct., 214 subsp. subvirescens N¨ageli & Peter, 215 subsp. tricholepium N¨ageli & Peter, 215 var. tricholepium (N¨ageli & Peter) Pugsley, 215 subsp. trichoscapum N¨ageli & Peter, 215 subsp. trichosomum N¨ageli & Peter, 215 var. viride F. Hanb., 214 pilosum Schleich. ex Froel, 313 pinnatifidum auct., 356 pinnatifidum L¨onnr. ex Dahlst., 363 subsp. scanicum Dahlst., 370 placerophylloides Pugsley, 253 placerophyllum auct., 253 platylepium auct., 261 platypyllum (Ley) W. R. Linton, 330 pollichiae Sch. Bip., 359 pollinarioides Pugsley, 392 pollinarium F. Hanb., 420 var. platyphyllum Ley, 330 polycomatum auct., 265 polycomum auct., 265 polyphyllum Dahlst., 248 porrigens auct., 377 subsp. floccifrons Elfstr., 303 praealtum Vill. ex Gochnat, 213 subsp. thaumasium (Peter) P. D. Sell, 213 praecox auct., 395 subsp. basicrinum Zahn., 335 subsp. ciliatum auct., 388 subsp. clovense (E. F. Linton) Zahn, 306 microg. fraternum Sudre, 401, 554 subsp. fraternum (Sudre) Zahn, 401 subsp. gougetanum (Gren. & Godr.) Zahn, 396 subsp. medium (Jord.) Zahn, 409 var. medium (Jord.) Sudre, 409 var. oxyodontoides (Sch. Bip.) Sch. Bip., 359 praemorsum L., 204 praesigne (Zahn) Roffey, 363 var. subdiaphanum Pugsley, 363 praetenerum auct., 384 praetermissum P. D. Sell & C. West, 345 praethulense Pugsley, 264 prasiophaeum Arv.-Touv. & Gaut., 396 pratense Tausch., 212 prenanthoides Vill., 258 subsp. lanceolatum (Vill.) Zahn., 258 subsp. strictissimum auct., 258

Index Hieracium L. (cont.) probum P. D. Sell & C. West, 273 prolixum Norrl., 391 prominentidens P. D. Sell, 239, 532 protentum P. D. Sell, 394, 554 promonteriale P. D. Sell, 337, 546 protractum auct., 249, 256 var. shetlandicum Dahlst., 256 proximum F. Hanb., 323 pruinale (Zahn) P. D. Sell & C. West, 418 pseudacrifolium Pugsley, 253 pseudamplidentatum Pugsley, 245 var. stenophyllum Pugsley, 245 pseudanglicoides J. E. Raven, P. D. Sell & C. West, 312 pseudanglicum Pugsley, 308 pseudocurvatum auct., 272 pseudocurvatum (Zahn) Pugsley, 282 pseudoleyi (Zahn) Roffey, 335 pseudomicrodon Dahlst., 332 pseudonosmoides auct., 326 pseudopetiolatum (Zahn) Roffey, 294 pseudopilosella auct., 215 pseudoprotractum Pugsley, 256 pseudorepandum Pugsley, 344 pseudosarcophyllum Pugsley, 386 pseudostenstroemii Pugsley (1941), 407 pseudostenstroemii Pugsley (1948), 389 pseudozetlandicum Roffey, 322 pugsleyi P. D. Sell & C. West, 263 pulchelliforme (W. R. Linton) Pugsley, 266 pulchellum auct., 266 pulcherrimum (F. Hanb.) Roffey, 393 pulchrius (Ley) W. R. Linton, 375 pulmonarioides Vill., 269 pulmonarium auct., 297 pulmonarium Sm., 269 purpurascens Dahlst., 365 pusillifolium P. D. Sell, 321, 543 pycnodon Dahlst. var. acutidens Dahlst., 347 pycnotrichum (W. R. Linton) Roffey, 244 quadridentatum Hyl., 402 radyrense (Pugsley) P. D. Sell & C. West, 373 raveniorum P. D. Sell, 305, 539 reayense (Pugsley) P. D. Sell, 246, 532 reclinatum auct., 369 rectulum Ley, 377 repandulare Druce, 344 repandum (Ley) Ley, 344 var. venosum (Ley) Ley, 344 repens Gray, 214 reticulatiforme P. D. Sell, 246 reticulatum auct., 246, 247 reticulatum Lindeb., 246 var. angustifolium Pugsley, 246, 532 var. melanoglochin (E. F. Linton) Pugsley, 247 var. reayense Pugsley, 246, 532 rhayaderense Pugsley, 253 rhomboides (Stenstr.) Johanss., 362 riddelsdellii Pugsley, 344 rigens Jord., 238

Index Hieracium L. (cont.) rigidum auct., 248 var. calcaricola F. Hanb., 251 var. friesii auct., 249, 262 subsp. lineatum Dahlst., 255 var. longiciliatum auct., 254 var. longiciliatum F. Hanb., 254 var. nidense F. Hanb., 257 var. pullatum auct., 253 var. rubefactum (E. F. Linton) Roffey, 253 var. serpentinum F. Hanb., 256 var. stewartii (F. Hanb.) W. R. Linton, 251 var. strigosum Ley, 254 var. tavense Ley, 249 subsp. trichocaulon Dahlst., 251 var. trichocaulon (Dahlst.) W. R. Linton, 251 rionii Gremli., 312 rivale F. Hanb., 388 var. dasythrix (E. F. Linton) W. R. Linton, 307 var. subhirtum F. Hanb., 381 rivulare Krocker, 362, 551 robertsii P. D. Sell, 360, 550 roffeyanum Pugsley, 359 ronasii P. D. Sell, 400, 554 rosulare Brenner, 322, 543 rotundatum auct., 392 rubefactum (E. F. Linton) Roffey, 253 rubicundiforme (Zahn) Roffey, 329 rubicundum F. Hanb., 329 var. boswellii (E. F. Linton) F. N. Williams, 330 var. glabrescens Pugsley, 360, 550 rubiginosum F. Hanb., 361 var. peccense W. R. Linton, 359, 550 var. rivulare Pugsley, 362, 551 sabaudum L., 241 forma bladonii (Pugsley) P. D. Sell, 241, 532 forma calvatum (F. Hanb.) Zahn, 238 subsp. dumosum (Jord.) Sudre, 242 var. latifolium Gaudin, 238 subsp. obliquum (Jord.) Schinz & Keller, 241 var. perpropinquum Zahn, 241 subsp. rigens (Jord.) Zahn, 238 var. rigens (Jord.) Schinz & Keller, 238 var. salicetorum (Sudre) Zahn, 240 subsp. scabiosum (Sudre) Zahn, 240 subsp. vagum (Jord.) Schinz & Keller, 238 subsp. virgultorum (Jord.) Schinz & Keller, 240 sagittaticeps Dahlst., 379 sagittatum (Lindeb.) Dahlst., 382 var. abrasum Dahlst., 379 var. ampliatum (W. R. Linton) Zahn, 317 subsp. breadalbanense (F. Hanb.) Zahn, 384 var. clivicola (F. Hanb.) Zahn, 379 var. dasythrix (E. F. Linton) Zahn, 307 subsp. euprepes Zahn, 379 var. glabratum (E. F. Linton) Zahn, 304 var. lanuginosum auct., 385 subsp. leyanum Zahn, 330 var. maculigerum (W. R. Linton) Zahn, 382 var. peccense (E. F. Linton) Zahn, 359 var. philanthrax auct., 382

Hieracium sagittatum (Lindeb.) Dahlst. (cont.) subsp. rivale (F. Hanb.) Zahn, 388 subsp. rubiginosum (F. Hanb.) Zahn, 361 var. subhirtum (F. Hanb.) W. R. Linton, 381 salicetorum auct., 248 salicetorum Sudre, 240 salticola (Sudre) P. D. Sell & C. West, 239 sanguineum (Ley) W. R. Linton, 388 sannoxense P. D. Sell, 310 sarcophylloides Dahlst., 347 sarcophyllum auct., 386 var. ampliatum W. R. Linton, 317 var. expallidiforme auct., 390 var. expallidiforme auct., 347 sarmentosum Salisb., 214 saxifragum auct., 296 subsp. argenteum (Fr.) Zahn, 332 subsp. carenorum (F. Hanb.) Zahn, 340 subsp. microdon (Dahlst.) Dahlst., 332 subsp. nitidum (Backh. fil.) Zahn, 340 subsp. nitidum var. silurense (F. Hanb.) Zahn, 331 var. oreades auct., 331 subsp. orimeles (F. Hanb. ex W. R. Linton) Zahn, 326 var. orimeles (F. Hanb. ex W. R. Linton) F. Hanb., 325 subsp. pseudoleyi Zahn, 335 subsp. pseudonosmoides auct., 326 var. pseudonosmoides auct., 326 var. septentrionale (F. Hanb.) Zahn, 332 var. siluriense (F. Hanb.) Zahn, 331 var. subglabratum (F. Hanb.) Zahn, 331 var. vimineum auct., 340 saxorum (F. Hanb.) P. D. Sell & C. West, 345 scabiosum Sudre var. salicetorum (Sudre) Sudre, 240 scabrisetum (Zahn) Roffey, 253 scandinavicum auct., 384 scanicum auct., 373 scanicum (Dahlst.) Ley, 370 var. anglorum Ley, 371 scarpicum Pugsley, 314 schmidtii auct., 343 schmidtii Tausch., 342 var. crinigerum auct., 325 var. devoniense F. Hanb., 355 var. eustomon E. F. Linton, 355 forma lasiophyllum (W. D. J. Koch) F. N. Williams, 341 var. superbum auct., 340 sciaphilum (Uechtr.) F. Hanb., 373 var. amplifolium (Ley) W. R. Linton, 368 forma paucifolia Ley ex Riddelsd., 373 var. pulchrius Ley, 375 var. strumosum Ley, 375 scoticiforme Dahlst., 320 scoticum F. Hanb., 320 var. microphyllum Pugsley, 321, 543 var. occidentale F. Hanb., 328, 545 var. rosulare Pugsley, 322, 543 var. submaculatum Dahlst., 322 scotostictum Hyl., 395 scottii P. D. Sell, 322, 544 scullyi E. F. Linton, 252

603

604 Hieracium L. (cont.) scytophyllum auct., 371 sejunctum (W. R. Linton) Roffey, 357 semicrassiceps Pugsley, 387 senescens Backh. fil., 299 var. brevifurcatum Pugsley, 299 subsp. marshallii (E. F. Linton) Zahn, 300 septentrionale Arv.-Touv., 253, 545 var. amphibolum auct., 253 var. simplex Ley, 321, 543 seriflorum Hyl., 402 serratifrons Almq. subsp. caliginosum auct., 420 var. caliginosum auct., 390, 420 var. cinderella Ley, 403 var. crassiceps auct., 387, 408 var. lepistoides auct., 407, 408 subsp. stenstroemii auct., 389 subsp. stenstroemii Dahlst., 407 var. stenstroemii auct., 389 var. stenstroemii (Dahlst.) W. R. Linton, 407 subsp. subcrassum Almq. ex Dahlst., 403 var. torticeps auct., 407, 408 var. triangulare auct., 407 severiceps Wiinst., 413 shoolbredii E. S. Marshall, 319 shoolbredii Zahn, 322 siluriense (F. Hanb.) P. D. Sell, 331, 545 silvaticoides Pugsley, 384 silvaticum auct., 384 var. asymmetricum auct., 390, 391 var. asymmetricum Ley, 337, 546 subsp. caesionigrescens Fr. ex Stenstr., 359 subsp. cordigerum Fr. ex Stenstr., 359 var. crassum Ley, 380, 552 var. lucidulum (Ley) F. N. Williams, 415 var. maculosum (Dahlst. ex Stenstr.) F. N. Williams, 391 subsp. maculosum (Dahlst. ex Stenstr.) Omang, 391 var. micracladium F. N. Williams, 394 var. pellucidum (Laest.) Almq., 415 subsp. prolixum (Norrl.) Dahlst., 391 var. prolixum (Norrl.) F. N. Williams, 391 var. prolixum auct., 307 var. rivale (F. Hanb.) F. N. Williams, 388 forma subhirtum (F. Hanb.) F. N. Williams, 381 subsp. sagittatum (Lindeb.) Stenstr., 382 var. sanguineum (Ley) F. N. Williams, 388 var. subcyaneum W. R. Linton, 336 var. subtenue W. R. Linton, 380 var. tricolor W. R. Linton, 336 simplex Viv., 321, 543 sinclairii Dahlst., 347 sinuans F. Hanb., 297 sinuolatum P. D. Sell, 383, 552 skyense (Zahn) Roffey, 319 snowdoniense P. D. Sell & C. West, 421 solum P. D. Sell & C. West, 267 sommerfeltii Lindeb., 334 var. griffithii (F. Hanb.) Zahn, 325 subsp. sommerfeltii, 334

Index Hieracium sommerfeltii Lindeb. (cont.) subvar. splendens (F. Hanb.) Zahn, 335 var. splendens F. Hanb., 335 var. subtruncatum Pugsley, 345 var. tactum F. Hanb., 335 sordidum W. R. Linton ex E. F. Linton, 350 sowadeense P. D. Sell, 321, 543 sparsidens Dahlst. var. elatius Ley, 408, 555 sparsifolium Lindeb., 256 var. grandescens auct., 249, 253 var. lingua Ley, 257 var. longiciliatum (F. Hanb.) W. R. Linton, 254 var. oligodon E. F. Linton, 257 var. placerophyllum auct., 253 var. strigosum (Ley) W. R. Linton, 254 sparsifrons P. D. Sell & C. West, 257 sparsum Friv., 418, 555 sparsum Jord. ex Boreau, 418 speluncarum Arv.-Touv., 268 var. lanceolatum Arv.-Touv., 269 spenceanum Walter Scott & R.C. Palmer, 263 spicatum Gilib., 204 spilophaeum Jord. ex Boreau, 367 spraguei Pugsley, 213 spurcatum Jord. ex Boreau, 367 stenolepiforme (Pugsley) P. D. Sell & C. West, 355 stenolepis auct., 346, 351, 354, 355 var. anguinum W. R. Linton, 390 var. subbritannicum A. Ley, 354 stenophyes W. R. Linton, 362 var. oxyodus W. R. Linton, 304 stenopholidium (Dahlst.) Omang, 355 stenotum auct., 332 stenstroemii (Dahlst.) Johanss., 407 var. subcordatum Pugsley, 408 sternbergii Hoppe ex Hornem., 203 stewartii (F. Hanb.) Roffey, 251 var. praegeri Pugsley, 251 stictophyllum Dahlst., 256 var. concolor Pugsley, 256 var. serpentinum (F. Hanb.) W. R. Linton, 256 stictum P. D. Sell, 343, 548 × stoloniflorum Waldst. & Kit., 211 subsp. schuriana N¨ageli & Peter, 212, 532 striatum Tausch. var. pseudauratum auct., 248 strictiforme (Zahn) Roffey, 246 strictum auct., 246 var. amplidentatum F. Hanb., 245 var. angustum auct., 245 var. humilius Beeby, 261 var. opsianthum Dahlst. ex F. Hanb., 246 var. subcrocatum E. F. Linton, 248 strumosum (Ley) Ley, 375 subaequialtum Hyl., 412 subalpestrifrons Dahlst., 379 subamplifolium (Zahn) Roffey, 368 subanfractum E. S. Marshall, 300 subbritannicum auct., 351

Index Hieracium L. (cont.) subbritannicum (Ley) P. D. Sell & C. West, 354 subcrassum (Almq. ex Dahlst.) Johanss., 403 subcrocatum (E. F. Linton.) Roffey, 248 subcyaneum (W. R. Linton) Pugsley, 336 subelatum auct., 258 suberectum auct., 238 subexpallescens Dahlst., 379 subfasciculare (W. R. Linton) Roffey, 357 subglobosum P. D. Sell & C. West, 279 subgracilentipes auct., 286 subgracilentipes (Zahn) Roffey, 276 subhirtum (F. Hanb.) Pugsley, 381 subimpressum Dahlst., 316 subincisum Freyn var. rionii (Gremli) Arv.-Touv., 312 subintegrifolium Pugsley, 356 subintegrum auct., 257 subirriguum Dahlst. ex Porat, 364 sublactucaceum auct., 238, 239 var. radnoricum Druce & Zahn, 238 sublasiophyllum P. D. Sell, 338, 547 sublepistoides (Zahn) Druce, 412 subminutidens (Zahn) Pugsley, 370 submurorum auct., 306 submutabile (Zahn) Pugsley, 369 subnigrescens auct., 302 var. gracilifolium (F. Hanb.) Pugsley, 302 subplanifolium auct., 308, 345 subplanifolium Pugsley, 346 subpolyphyllum Pugsley, 248 subprasinifolium Pugsley, 394 subquercetorum Pugsley, 238 subramosum auct., 357 subramosum L¨onnr., 363 subsp. coniops (Norrl.) Zahn, 358 subsp. holophyllum (W. R. Linton) Zahn, 328 subsp. insulare (F. Hanb.) Zahn, 310 var. megalodon (F. Hanb.) Zahn, 365 subsp. orcadense (W. R. Linton) Zahn, 379 var. rhomboides (Stenstr.) Zahn, 362 subsp. surrejanum (F. Hanb.) Zahn, 365 subravusculum (W. R. Linton) Roffey, 357 subrectum auct., 238 subrubicundum Dahlst., 327 subrude Arv.-Touv., 324 subsanguineum (Zahn) Roffey, 388 subscoticum P. D. Sell, 320, 543 substrigosum (Zahn) Roffey, 254 subtenue (W. R. Linton) Roffey, 380 var. canispense Pugsley, 380 subtenuifrons P. D. Sell & D. J. Tennant, 284, 535 subtruncatum Beeby, 262 var. glussburnense Pugsley, 262 subulatidens auct., 407, 418 var. cuneifrons Ley, 397 subumbellatiforme (Zahn) Roffey, 245 subviolascens P. D. Sell, 264, 551 subvirescens (N¨ageli & Peter) Roffey, 215 succisifolium All., 203

Hieracium L. (cont.) surrejanum F. Hanb., 365 var. megalodon E. F. Linton, 365 sylvaticum Lam. subsp. integratum Dahlst. ex Stenstr., 414 sylvivagum Jord. ex Boreau, 413 sylvularum Jord. ex Boreau, 401 syngenes Jord. ex Boreau, 415 taraxaci L., 102 tavense (Ley) Ley, 249 tectorum (L.) Hornem., 204 tenuifrons auct., 284 tenuifrons P. D. Sell & C. West, 283 thalassinum P. D. Sell, 311, 541 thuringorum Sag., 367 triangulifolium P. D. Sell, 385, 553 trichocaulon (Dahlst.) Johanss., 251 tricholepium (N¨ageli & Peter) Roffey, 215 tricolorans (Zahn) Pugsley, 336 tridentatum auct., 249, 251 tridentatum Fr., 251 var. acrifolium auct., 253 var. decipiens Ley, 249 forma glandulosiceps (Zahn) Pugsley, 251 var. polyphyllum (Zahn) Pugsley, 251 var. setigerum Ley, 253 trinitatis Pugsley, 255 triviale (Norrl.) Norrl., 356 truncatum auct., 262 tumescens Norrl. subsp. molybdochroum Dahlst., 299 tunbridgense Pugsley, 377 uiginskyense Pugsley, 255 uistense (Pugsley) P. D. Sell & C. West, 397 uisticola Pugsley, 383 umbellatum L., 242 subsp. bichlorophyllum (Druce & Zahn) P. D. Sell & C. West, 243 var. commune Fr., 243 var. coronopifolium Bernh. ex Hornem., 243 var. curtum E. F. Linton, 243 var. dunense Reyn, 243 var. filifolium Fr., 243 forma latifolium W. R. Linton, 243, 532 var. latifolium (W. R. Linton) P. D. Sell, 243, 532 var. linariifolium Wallr., 243 var. litorale Lindb., 243 var. littoreum auct., 243 subsp. maritimum (F. Hanb.) Zahn, 244 var. monticola auct., 243 subsp. ogwenii (E. F. Linton) W. R. Linton, 243 var. ogwenii (E. F. Linton) F. Hanb., 243 var. paniculatum auct., 243 var. paniculatum Cariot, 243 var. pauciflorum auct., 244 var. sarniense P. D. Sell, 243, 532 var. serum Sudre, 243 var. taylori Bab., 243 vagense (F. Hanb.) Ley, 333 vagicola P. D. Sell, 351, 549

605

606 Hieracium L. (cont.) vagum Jord., 238, 532 forma croceostylum (Pugsley) P. D. Sell, 238, 532 forma vagum, 238 variicolor auct., 392 var. piligerum Pugsley, 392 variifolium P. D. Sell & C. West, 390 vennicontium Pugsley, 304 vestitum Gren. & Godr., 338, 547 veterascens Dahlst., 329 vinaceum (Beeby) Pugsley, 261 vinicaule P. D. Sell & C. West, 261 vinifolium P. D. Sell, 361, 550 violascens (Almq. ex Dahlst.) Dahlst., 364 violascens Borbas, 364, 551 virgultorum Jord., 240 var. salticola Sudre, 239 vorlichense P. D. Sell, 301, 536 vulgatum Fr., 357, 549 subsp. acuminatum (Jord.) Sudre, 372 var. acuminatum (Jord.) Arv.-Touv., 372 var. amplifolium Ley, 368 subsp. argillaceum (Jord.) Sudre, 373 var. argillaceum (Jord.) Arv-Touv., 373 var. asperatum Sudre, 374 subsp. aviicola (Jord. ex Boreau) Zahn, 375 var. aviicola (Jord. ex Boreau) Sudre, 375 var. cacuminum (Ley) F. Hanb., 327 var. cheriense (Jord. ex Boreau) Zahn, 377 subsp. chlorophyllum (Jord. ex Boreau) Zahn, 373 var. chlorophyllum (Jord. ex Boreau) Sudre, 374 var. cinereum Backh., 328 subsp. consociatum (Jord. ex Boreau) Zahn, 373 var. consociatum (Jord. ex Boreau) Sudre, 373 subsp. daedalolepium Zahn., 369 var. daedalolepium auct., 369 subsp. deductum auct., 369 subsp. diaphanum (Fr.) Zahn., 369 var. fastigiatum Zahn, 373 microg. festinum (Jord. ex Boreau) Sudre, 376 subsp. festinum (Jord. ex Boreau) Sudre, 376 var. genuinum (Jord. ex Boreau) Sudre, 363 var. genuinum Syme, 373 subsp. lachenalii auct., 373 forma latebrosum (Jord. ex Boreau) Zahn, 375 subvar. latebrosum (Jord. ex Boreau) Sudre, 375 subsp. lepidiceps (Dahlst.) Zahn, 370 subsp. lepidulum (Stenstr.) Zahn, 364 var. lortetiae (Balbis) Sudre, 376 forma maculatum Backh. fil., 367 var. maculatum F. Hanb., 367 var. medium auct., 373 var. mutabile Ley, 369 var. nemorosum auct., 356 subsp. percissum (Jord. ex Boreau) Zahn, 363 var. percissum (Jord. ex Boreau) Sudre, 363 subsp. pinnatifidum Zahn, 363 subsp. praesigne Zahn, 363 forma pseudosubramosum (Pugsley) P. D. Sell, 357, 549 var. pseudosubramosum Pugsley, 357, 550 subsp. pulchrius (Ley) Zahn, 375

Index Hieracium vulgatum Fr. (cont.) var. rosulatum Syme, 357 var. rubescens Backh. fil., 361 subsp. scanicum (Dahlst.) Zahn, 370 var. sciaphilum Uechtr., 373 forma sejunctum (W. R. Linton) P. D. Sell, 357, 550 var. sejunctum W. R. Linton, 357, 550 subsp. subamplifolium Zahn, 368 forma subfasciculare (W. R. Linton) P. D. Sell, 357, 549 var. subfasciculare W. R. Linton, 357, 549 subsp. subirriguum (Dahlst.) Zahn, 364 var. submutabile Zahn, 369 var. sublaeve Sudre, 377 forma subravusculum (W. R. Linton) P. D. Sell, 357, 550 var. subravusculum W. R. Linton, 357, 550 subsp. triviale Norrl., 356 var. triviale (Norrl.) Pugsley, 356 subsp. violascens (Almq. ex Dahlst.) Zahn, 364 westii P. D. Sell, 304 wiesbaurianum Uechtr. subsp. britannicum (F. Hanb.) Zahn, 350 subsp. cambricum (Baker) Zahn, 333 subsp. cyathis (Ley) Zahn, 337 var. griseum (Ley) Zahn, 341 subsp. hypochaeroides (S. Gibson) Zahn, 343 var. saxorum (F. Hanb.) Zahn, 345 subsp. vagense (F. Hanb.) Zahn, 333 zetlandicum Beeby, 267 zygophorum Hyl., 396 Hippophaestum Gray, 84 vulgare Gray, 87 Homogyne Cass., 511 alpina (L.) Cass., 512 Honeysuckle, 46 Box-leaved, 44 Californian, 44 Fly, 45 Garden, 47 Henry’s, 45 Himalayan, 43 Perfoliate, 47 Tartarian, 45 Wilson’s, 44 Hostia Moench, 205 foetida (L.) Moench, 205 Hyoseris L. cretica L., 97 hedypnois L., 97 minima L., 96 rhagadioloides L., 97 taraxacoides Vill., 103 Hypochaeris L., 98 ascendens Brot., 98 dimorpha Brot., 95 glabra L., 99 var. erostris Boiss., 99 var. loiseleuriana Godr., 99 × radicata, 99 hispida Roth., 99 infesta Salisb., 99 × intermedia Richt., 99

Index Hypochaeris L. (cont.) maculata (L.) Scop., 100 radicata L., 99 var. salina (Gren.) Rouy, 100 subsp. ericetorum Van Soest, 100, 106 salina Gren., 100 Hypochaeris auct., 98 Ifloga Cass. verticillata (L. fil.) Fenzl., 427 Intybellia Monnier, 204 praemorsa (L.) Monnier, 204 Intybus Fr., 204 bulbosus (L.) Fr., 111 praemorsus (L.) Fr., 204 Inula L., 433 aspera Poir., 434 britannica L., 434 caucasica Pers., 438 conyza DC., 435 conyzae (Griess.) Meikle, 435 cordata Boiss., 434 crithmifolia L., 435 crithmoides L., 435 cylindrica With., 437 dysenterica L., 437 glandulosa Willd., 434 grandiflora Gray, 433 grandiflora Willd., 434 graveolens (L.) Desf., 436 helenium L., 433 hookeri C. B. Clarke, 434 oculus-christi L., 435 orientalis Lam., 434 pulicaria L., 437 pulicaris Stokes, 437 salicina L., 434 subsp. aspera (Poir.) Hayek, 434 squarrosa (L.) Bernh., 435 suaveolens Pall., 435 uliginosa Sibth., 437 viscosa (L.) Aiton, 436 vulgaris Trevis, 435 Iva L., 514 xanthifolia Nutt., 514 Ixine Hill, 75 arvensis (L.) Hill, 80 Jacea Mill., 89 alata Lam., 91 cineraria (L.) Delarbre, 86 communis Delarbre, 89 nigra Gray, 89 nigra (L.) Hill, 90 paniculata (L.) Lam., 86 pratensis Lam., 89 scabiosa (L.) Lam., 85 segetum Lam., 91 vulgaris Bernh., 90 Jacobaea Mill., 494 aquatica P. Gaern., B. Mey. & Scherb., 495

Jacobaea Mill. (cont.) carnosa C. Presl, 497 doria (L.) P. Gaertn., B. Mey. & Scherb., 492 elegans (L.) Moench, 503 erucifolia (L.) P. Gaertn., B. Mey. & Scherb., 496 incisa C. Presl, 497 nemorosa Fourr., 494 ovata P. Gaertn., B. Mey. & Scherb., 493 tomentosa Moench, 490 vulgaris Gaertn., 494 Jaegeria Humb., Bonpl. & Kunth abyssinica (L. fil.) Spreng., 516 urticaefolia (Kunth) Spreng., 521 Jasione L., 10 appressifolia Pau, 11 espadanae Pau, 11 mediterranea Rouy 11 montana L., 11 var. boraei Rouy, 11 var. glabra Peterm., 11 var. gracilis Lange, 11 var. hirsuta Duby, 11 var. imbricans J. Parn., 11 var. laevis Duby, 11 var. latifolia Pugsley, 11 var. litoralis Fr., 11 var. major Mert. & W. D. J. Koch, 11 var. maritima Br´eb., 11 var. megaphylla Vicioso, 11 var. nana Boreau, 11 var. nana Gren. & Godr., 11 var. prolifera A. DC., 11 var. sebularia Cout., 11 var. stolonifera DC., 11 var. tenella Peterm., 11 var. timbali Rouy, 11 undulata Lam., 11 vulgaris Gaterau, 11 Jerusalem Artichoke, 519 Kentranthus auct., 53 Kentrophyllum Necker ex A. DC., 93 baeticum Boiss. & Reut., 93 lanatum (L.) DC., 93 Kerneria Moench bipinnata (L.) Gren. & Godr., 524 Knapweed, Brown, 89 Chalk, 90 Common, 90 Greater, 85 Jersey, 86 Ragwort, 86 Russian, 84 Knautia L., 57 arvensis (L.) Coult., 57 var. integrifolia (Roth) DC., 58 Kolkwitzia Graebn., 47 amabilis Graebn., 47 Lacinaria Hill ligulistylis A. Nelson, 529

607

608 Lactuca L., 115 adulterina Gren. & Godr., 117 alpina (L.) A. Gray, 118 ambigua Schrad., 117 angustifolia Gilib., 117 bourgaei (Boiss.) Irish & N. Taylor, 119 caucasica K. Koch, 117 cyanea K. Koch, 117 dubia Jord., 116 flavida Jord., 117 integrata (Gren. & Godr.) A. Nelson, 116 kochiana Beauverd, 118 lactucarii Lamotte, 117, 532 macrophylla (Willd.) A. Gray, 119 minima Gray, 117 muralis (L.) Gaertn., 120 officinarum Crantz, 115 plumieri (L.) Gren. & Godr., 119 salicifolia (K. Koch) Grossh., 118 salicifolia Salisb., 117 saligna L., 117 var. cracoviensis auct., 117 var. runcinata Gren. & Godr., 117 sativa L., 116 var. angustana Irish ex Bremer, 117 subsp. asparagina Janch., 117 subsp. capitata (L.) Sch¨ubl. & G. Martens, 116 var. capitata L., 116 subsp. crispa (L.) Sch¨ubl. & G. Martens, 116 var. crispa L., 116 var. longifolia Lam., 116 subsp. romana Sch¨ubl. & G. Martens, 116 scariola L., 115 var. altissima Lecoq & Lamotte, 117 var. integrata Gren. & Godr., 116 var. integrifolia (Gray) Bogenhard, 116 var. lactucarii (Lamotte) Rouy, 117, 532 var. typica Rouy, 116 var. vulgaris Bisch., 116 schimperi Jord., 115 serratifolia Sennen, 117 serriola L., 115 var. dubia (Jord.) Rouy, 116 forma integrifolia (Gray) S. D. Prince & R. N. Carter, 116 silvestris Garsult, 115 sonchifolia DC., 118 sylvestris Lam., 115 tatarica (L.) C. A. Mey., 117 viminea (L.) J. & C. Presl, 115 virosa L., 117, 532 var. integrifolia Gray, 116 forma lactucarii (Lamotte) P. D. Sell, 117, 532 Lagedium Soj´ak tataricum (L.) Soj´ak, 118 Lappa Scop., 70 arctium Hill, 72 glabra Lam., 71 intermedium (Lange) Reichb. fil., 71 macrosperma Wallr., 71

Index Lappa Scop. (cont.) major Gaertn., 71 var. racemosa (Lej.) G. Mey., 71 minor Hill, 71 mixta E. G. Camus, 71 nemorosa (Lej.) K¨orn., 71 notha Ruhmer, 71 pubens (Bab.) Boreau, 71 ruhmerio Gu´etrot, 71 subracemosa Simonk., 71 tomentosa (Mill.) Lam., 72 var. glabra K¨orn., 72 vulgaris Hill, 71 Lapsana L., 96 capillaris L., 206 communis L., 97 subsp. communis, 97 subsp. intermedia (M. Bieb.) Hayek, 97 intermedia M. Bieb., 97 pusilla Willd., 96 stellata L., 98 Lasallea Greene, 450 ericoides (L.) Semple & Brouillet, 450 novae-angliae (L.) Semple & Brouillet, 450 Laurustinus, 37 Lavender-cotton, 470 Legousia Durande, 8 durandii Delarbre, 8 hybrida (L.) Delarbre, 8 parviflora Gray, 8 speculum-veneris (L.) Chaix, 8 Leontodon L., 102 autumnalis L., 101, 531 var. alpinus (Gaudin) Gren. & Godr., 102 subsp. borealis Ball, 102 var. cinerascens Briq., 102, 531 var. coronopifolius Lange, 102, 531 var. dentatus Holuby ex Schur, 102, 531 var. latifolius Schur, 102, 531 var. minimus auct., 101 var. nigrolantus Fr., 101, 531 var. pinnatifidus Schur, 102, 531 var. pratensis (Hornem.) W. D. J. Koch, 102 var. salinus (Aspegren) Lange, 101 var. simplex Duby, 101, 531 var. taraxaci (L.) Hartm., 102 var. vulgaris Neilr., 102 bulbosus L., 111 danubialis Jacq., 103 hastilis L., 103 var. arenarius Duby, 106, 531 var. glabratus W. D. J. Koch, 103 hirtus auct., 103 hispidus L., 103 subsp. danubialis (Jacq.) Simonk., 103 var. glabratus (W. D. J. Koch) Bisch., 103 × saxatilis, 103, 531 leysseri Beck, 103 nudicaulis auct., 103 var. pristis Druce, 106, 531

Index Leontodon L. (cont.) obliquus Fr., 145 palustris Lyons, 146 pratensis (Hornem.) Rchb., 102 psilocalyx (Rchb.) M´erat, 106 saxatilis Lam., 103, 531 var. arenarius (Duby) P. D. Sell, 106, 531 var. pristis (Druce) P. D. Sell, 106, 531 taraxacoides (Vill.) M´erat, 103 taraxacum L. × vegetus Finch & P. D. Sell, 103, 531 Leontopodium (Pers.) R. Br. ex Cass. alpinum Cass., 433 Leopard-plant, 507 Przewalski’s, 507 Leopard’s-bane, 508 Eastern, 509 Harper-Crewe’s, 508 Plantain-leaved, 508 Willdenow’s, 508 Lepidanthus Nutt. suaveolens (Pursh) Nutt., 484 Lepidoseris Rchb., 207 Leptilon Raf., 456 bonariense (L.) Small, 458 canadensis (L.) Britton, 458 linifolium (Willd.) Small, 458 Leptinella Cass., 486 dioica Hook. fil., 487 squalida Hook. fil., 487 Leptinella, 487 Hairless, 487 Lettuce, Blue, 118 Cabbage, 116 Cos, 116 Curled, 116 Garden, 116 Great, 117 Least, 117 Prickly, 115 Wall, 120 Leucantha Gray cyanifolia Gray, 88 Leucanthemella Tzvelev, 479 serotina (L.) Tzvelev, 479 Leucanthemum Mill., 480 cantabricum Sennen, 481, 556 crassifolium (Lange) Lange, 481 ircutianum DC., 481, 556 cantabricum (Sennen) Vogt, 481 crassifolium (Lange) Vogt, 481 lacustre (Brot.) Samp. × maximum, 418 maximum auct., 481 maximum (Ramond) DC., 481 pallens DC. var. crassifolium Lange, 481 praecox (Horvatic) Horvatic, 480 × superbum (Bergmans ex J. Ingram) Kent, 481

Leucanthemum Mill. (cont.) vulgare Lam., 480, 556 subsp. cantabricum (Sennen) P. D. Sell, 481, 556 subsp. crassifolium (Lange) Rouy, 481 var. crassifolium (Lange) Merino, 481 subsp. ircutianum (DC.) P. D. Sell, 481, 556 var. nanum P´erard, 481 subsp. vulgare, 480 var. vulgare, 481 Leucojum L. aestivum L. subsp. aestivum, xviii subsp. pulchellum (Salisb.) Briq., xviii Leysera L. capillifolia (Desf.) Spreng., 433 tenella DC., 433 Leycesteria Wall., 43 formosa Wall., 43 Liatris Gaertn. ex Schreb. ligulistylis (A. Nelson) A. Nelson, 529 Ligularia Cass., 507 clivorum Maxim., 507 dentata (A. Gray) H. Hara, 507 przewalskii (Maxim.) Diels, 507 tangutica (Maxim.) Mattf., 506 Limbardia Adans., 433 tricuspis Cass., 435 crithmifolia Raf., 435 crithmoides (L.) Dumort., 435 Linnaea L., 42 americana Forbes, 43 borealis L., 42 subsp. americana (Forbes) Hult´en, 43 var. americana (Forbes) Rehder, 43 Linosyris Cass., 451 vulgaris Less., 451 var. minor Wallr. 451 Liparis Rich loeselii (L.) Rich var. loeselii, xviii var. ovata Ridd. ex Godfrey, xviii Lobelia L., 12 angulata G. Forst., 14 dortmanna L., 13 erinus L., 12 lacustris Salisb., 13 siphilitica L., 13 urens L., 12 Lobelia, California, 14 Garden, 12 Great, 13 Heath, 12 Lawn, 14 Water, 13 Logfia Cass., 426 arvensis (L.) Holub, 426 gallica (L.) Coss. & Germ., 426 minima (Sm.) Dumort., 426

609

610 Lonicera L., 43 americana auct., 47 caprifolium L., 47 × etrusca Sant., 47 cochinchinensis G. Don, 46 dumetorum Moench, 45 henryi Hemsl., 44 hispanica Boiss. & Reut., 45 involucrata (Richardson) Banks ex Spreng., 44 italica (Mill.) Wood, 47 × italica Schmidt ex Tausch, 47 japonica Thunb. ex Murray, 46 ledebourii Eschsch., 44 ligustrina Wall. var. pileata (Oliv.) Franch., 44 var. yunnanens Franch., 44 mocinianan DC., 44 nitida E. H. Wilson, 44 ochroleuca St-Lag., 45 pallida Host, 47 perfoliata (R¨ohl.) Edwards, 47 periclymenum L., 46, 530 var. clarkei Hesl.-Harr., 46 var. glaucohirta Kuntze, 47 var. hirsuta (Rouy) P. D. Sell, 45, 530 subvar. hirsuta Rouy, 45, 530 subsp. hispanica (Boiss. & Reut.) Nyman, 47 var. quercifolia Aiton, 46 pileata Oliv., 44 forma yunnanensis (Franch.) Rehd., 44 pubescens Stokes, 45 racemosa (Michx) Fern., 41 repens Miq., 45 rotundifolia (Moench) Medik., 47 suavis Salisb., 47 symphoricarpos L., 41 tatarica L., 45 villosa (Dulac) K. Koch, 45 vulgaris (Borkh.) Rehd., 45 xylosteum L., 45 Lopholoma Cass., 85 scabiosa (L.) Cass., 85 Lyonnettia Cass., 475 Madder, 29 Field, 16 Wild, 28 Magaris DC., 41 Malacothrix DC. crepoides A. Gray ex J. G. Cooper, 206 Mantisalca Cass. salmantica (L.) Briq. & Cavill., 84 Maori Holly, 461 Margaria DC. nudiflora DC., 42 Mariana Hill, 83 lactea Hill, 83 mariana (L.) Hill, 83 Marianthemum Schrank, 1 medium (L.) Schur, 4

Index Marigold, African, 526 Corn, 478 Dwarf, 527 Field, 512 French, 526 Mediterranean, 479 Pot, 512 Southern, 526 Marsea Adans., 456 canadensis (L.) Badillo, 458 Maruta Cass., 478 cotula (L.) DC., 476 foetida Gray, 476 vulgaris Bluff & Fingerh., 476 Matricaria L., 483 aurea (Loefl.) Sch. Bip., 483 chamomilla L., 485, 486 chamomilla L. (1763), non L. (1753), 484 decipiens (Fisch. & C. A. Mey.) K. Koch, 483 discoidea DC., 484, 556 subsp. discoidea, 484 subsp. occidentalis (Greene) P. D. Sell, 484, 556 grandiflora (Thunb.) Fenzl ex Harv., 483 indica (L.) Desr., 482 inodora L., 485, 486 var. maritima (L.) Wahlenb., 485 var. phaeocephala auct., 485 var. salina auct., 486 maritima L., 485 subsp. inodora (K. Koch) A. R. Clapham, 486 subsp. phaeocephala auct., 485 var. phaeocephala auct., 485 matricarioides auct., 484 occidentalis Greene 484, 556 parthenium L., 464 perforata M´erat, 486 recutita L., 484 sauveolens L., 484 sauveolens (Pursh) Buchenau, 484 vulgaris Gray, 464 Matricaria sensu Flora Europaea, 484 Mayweed, Scented, 484 Scentless, 485 Swollen-flowered, 483 Mecoschistum Dulac, 12 urens (L.) Dulac, 12 Meridiana Hill uniflora (L. fil.) Kuntze, 423 Michaelmas Daisy, Common, 448 Confused, 447 European, 444 Glaucous, 446 Hairy, 450 Late, 447 Narrow-leaved, 448 Nettle-leaved, 445 Red-stalked, 449 Micropsis DC. spathulata (Pers.) Cabrera, 427

Index Milfoil, Yellow, 473 Millefolium Adans., 471 nobile (L.) Fourr. 472 officinale Ledeb., 472 tomentosum (L.) Fourr., 473 vulgare Ledeb., 472 Millotia Cass. perpusilla (Turcz.) P. S. Short, 432 Molpadia (Cass.) Cass., 438 sauveolens Cass., 438 Moschatel, 48 Mountain Everlasting, 427 Mouse-ear Hawkweed, 214 Few-headed, 214 Shaggy, 218 Mugwort, 486 Annual, 468 Chinese, 466 Hoary, 467 Norwegian, 468 Slender, 468 Western, 467 Mulgedium Cass., 118 alpinum (L.) Less., 118 bourgaei Boiss., 119 hispidum Korsh., 119 macrophyllum (Willd.) DC., 119 plumieri (L.) DC., 119 salicifolium K. Koch, 118 tataricum (L.) DC., 118 uralensis Rouy, 119 Mycelis Cass., 120 muralis (L.) Dumort., 120 Myconia Sch. Bip., non Lapeyr. chrysanthemoides Sch. Bip., 479 Myriocephalus Benth. rhizocephalus (DC.) Benth., 432 Nardosmia Cass., 511 denticulata Cass., 511 fragrans (Vill.) Rchb., 511 japonica Siebold & Zucc., 510 Nardostachys DC. jatamansii DC., 435 Nemauchenes Cass., 208 Nertera Banks & Sol. ex Gaertn., 15 depressa Banks & Sol. ex Gaertn., 15 granadensis (Mutis ex L. fil.) Druce, 15 montana Colenso, 15 New Zealand Holly, 461 Niger, 516 Nintoa Sweet japonica (Thunb. ex Murray) Sweet, 46 Nipplewort, 95 Oglifa (Cass.) Cass. minima (Sm.) Rchb. fil., 426 Olearia Moench, 460 avicenniifolia (Raoul) Hook. fil., 460 × moschata, 461

Olearia Moench (cont.) dentata Hook. fil., non Moench, 461 forsteri (Hook. fil.) Hook. fil., 460 × haastii Hook. fil., 461 hillii Colenso, 460 ilicifolia Hook. fil., 461 macrodonta Baker, 461 multibracteolata Colenso, 461 nummulariifolia (Hook. fil.) Hook. fil., 460 paniculata (J. R. & G. Forst.) Druce, 460 solandri (Hook. fil.) Hook. fil, 461 traversii (F. Muell.) Hook. fil., 461 Oligosporus Cass. condimentarius Cass., 468 Omalotheca Cass., 429 norvegica (Gunnerus) Sch. Bip. & F. W. Schultz, 429 supina (L.) DC., 430 sylvatica (L.) Sch. Bip. & F. W. Schultz, 429 Oncosiphon K¨allersjo, 483 grandiflorum (Thunb.) K¨allersjo, 483 piligerum (L. fil.) K¨allersjo, 483 suffruticosum (L.) K¨allersjo, 483 Onopordon L., 81 acanthium L., 82 macranthum Schousboe, 82 nervosum Boiss., 82 vulgare Gray, 82 Oporinia D. Don, 100 autumnalis (L.) D. Don, 101 var. alpina (Gaudin) DC., 102 var. minima auct., 101 var. sordidus Bab., 102 pratensis (Hornem.) Less., 102 Opulus Mill., 37 palustris Gray, 37 Orchis L. militaris L. var. militaris, xviii var. tenuifrons P. D. Sell, xviii Ormenis (Cass.) Cass. mixta (L.) All., 474 nobilis (L.) J. Gay, 474 Otanthus Hoffmanns. & Link, 470 maritimus (L.) Hoffmanns. & Link, 470 Othonna L. integrifolia L., 504 maritima L., 490 palustris L., 505 Oxeye, Rough, 520 Oxtongue, Bristly, 108 Hawkweed, 107 Pearly Everlasting, 428 Three-nerved, 428 Pectis L. pinnata Lam., 527 Pentzia Thunb. grandiflora (Thunb.) Hutch., 483 Pericallis D. Don, 503 hybrida R. Nordestam, 503

611

612 Periclymenum Mill., 46 italicum Mill., 47 perfoliatum Gray, 47 vulgare Mill., 46 Petasites Mill., 510 albus (L.) Gaertn., 511 amplus Kitam., 511 elatior Hill, 510 fragrans (Vill.) C. Presl, 511 hybridus (L.) P. Gaertn., B. Mey. & Scherb., 510 subsp. hybridus, 510 japonicus (Siebold & Zucc.) Maxim., 510 subsp. giganteus Kitam., 511 officinalis Moench, 510 ovatus Hill, 510 petasites (L.) Karst., 510 vulgaris Desf., 510 Phaeopappus (A. DC.) Boiss. macrocephalus (Muss. Puschk. ex Willd.) Boiss., 89 Phalacroderis DC., 205 coa DC., 206 Phalacroloma Cass. 455 Phrygia (Pers.) Gray, 89 major Gray, 85 nigra (L.) Gray, 90 Phuopsis (Griseb.) Hook. fil., 16 stylosa (Trin.) Benth. & Hook. fil. ex B. D. Jacks., 16 Phyteuma L., 9 orbiculare L., 10, 530 subsp. tenerum (R. Schulz) P. D. Sell, 10, 530 scheuchzeri All., 10 spicatum L., 10 subsp. coeruleum R. Schulz, 10 tenerum R. Schulz, 10, 530 Picris L., 106 auriculata Sch. Bip., 107 autumnalis (L.) All., 101 crepoides Saut., 107 echioides L., 108 foetida (L.) Lam., 205 grandiflora Ten., 107 hieracioides L., 107, 531 var. arenaria (Ny´ar.) P. D. Sell, 107, 531 subsp. auriculata (Sch. Bip.) Hayek, 107 subsp. crepoides (Saut.) Nyman, 107 var. gracilis auct., 107 subsp. grandiflora (Ten.) Arcangeli, 107 var. incana Druce, 107 var. leteae (Ny´ar.) P. D. Sell, 107, 531 subsp. paleacea (Vest) Domin & Podp., 107 subsp. setulosa (Guss. ex Ces., Pass. & Gibelli) Arcangeli, 107 subsp. sonchoides (Vest) Thell., 107 subsp. spinulosa (Bertol. ex Guss.) Arcangeli, 107 var. stricta (Jord.) Druce, 107 subsp. tatrae Domin & Podp., 107 var. umbellata auct., 107 subsp. villarsii (Jord.) Nyman, 107 var. villarsii (Jord.) P. D. Sell, 107, 531 humifusa Willd., 108

Index Picris L. (cont.) paleacea Vest, 107 pyrenaica auct., 107 ruderalis Salisb., 108 setulosa Guss. ex Ces., Pass. & Gibelli, 107 sonchoides Vest, 107 var. arenaria Ny´ar., 107 var. leteae Ny´ar., 107, 531 spinulosa auct., 107 spinulosa Bertol. ex Guss., 107 stricta Jord., 107 tatreae Borb´as, 107 villarsii Jord., 107, 531 Pilosella Hill, 209 aurantiaca (L.) F. W. Schultz & Sch. Bip., 211 subsp. brunneocrocea (Pugsley) P. D. Sell & C. West, 211 subsp. carpathicola (N¨ageli & Peter) Soj´ak, 211 × officinarum, 211, 532 caespitosa (Dumort.) P. D. Sell & C. West, 212 subsp. colliniformis (Peter) P. D. Sell & C. West, 212 × lactucella, 212 euronota (N¨ageli & Peter) Dost´al, 215 flagellaris (Willd.) P. D. Sell & C. West, 214 subsp. bicapitata P. D. Sell & C. West, 214 × floribunda (Wimm. & Grab.) Arv.-Touv., 212 lactucella (Wallr.) P. D. Sell & C. West, 213 subsp. helveola (Dahlst.) P. D. Sell & C. West, 212 × longisquama (Peter) Holub, 218 melanops (Peter) Dost´al, 215 officinarum F. W. Schultz & Sch. Bip., 214 subsp. concinnata (F. Hanb.) P. D. Sell & C. West, 215 subsp. euronota (N¨ageli & Peter) P. D. Sell & C. West, 215 subsp. melanops (Peter) P. D. Sell & C. West, 215 subsp. micrandenia (N¨ageli & Peter) P. D. Sell & C. West, 215 × peleteranum, 218 subsp. tricholepia (N¨ageli & Peter) P. D. Sell & C. West, 215 subsp. trichoscapa (N¨ageli & Peter) P. D. Sell & C. West, 215 subsp. trichosoma (Peter) P. D. Sell & C. West, 215 pachylodes Soj´ak, 218 peleterana (M´erat) F. W. Schultz & Sch. Bip., 218 subsp. subpeleterana (N¨ageli & Peter) P. D. Sell, 218 subsp. tenuiscapa (Pugsley) P. D. Sell & C. West, 218 praealta (Vill. ex Gochnat) F. W. Schultz & Sch. Bip., 213 subsp. arvorum (N¨ageli & Peter) P. D. Sell & C. West, 213 subsp. spraguei (Pugsley) P. D. Sell & C. West 213 subsp. thaumasia (Peter) P. D. Sell, 213 × stoloniflora (Waldst. & Kit.) F. W. Schultz & Sch. Bip., 211, 532 nothosubsp. schuriana (N¨ageli & Peter) P. D. Sell, 212, 532 thaumasia (Peter) Dostˆal, 213 tricholepia (N¨ageli & Peter) Dostˆal, 215 trichoscapa (N¨ageli & Peter) Dostˆal, 215 trichosoma (N¨ageli & Peter) Dostˆal, 215 Pinardia Cass., 478 coronaria (L.) Less., 479 Pineapple Weed, 484

Index Ploughman’s Spikenard, 435 Podolepis Labill. gracilis (Lehm.) Graham, 433 longipedata A. Cunn. ex DC., 433 Polycantha Hill, 73 acanthoides (L.) Hill, 74 palustris (L.) Hill, 80 procumbens Gray, 88 Polymnia L. abyssinica L. fil., 516 frondosa Bruce ex Murray, 516 Porcellites Cass. maculata (L.) Cass., 100 radicata (L.) Cass., 99 Pratia Gaudich., 13 angulata (G. Forst.) Hook. fil., 14 var. arenaria (Hook. fil.) Hook. fil., 14 arenaria Hook. fil., 14 treadwellii auct., 14 Prenanthes L. bulbosus (L.) DC., 111 muralis L., 120 Prismatocarpus L’H´er., 8 hybridus (L.) L’H´er., 8 speculum-veneris (L.) L’H´er., 8 Pseudognaphalium Kirp. luteoalbum (L.) Hilliard & B. L. Burtt, 431 undulatum (L.) Hilliard & B. L. Burtt, 431 Pseudolinosyris Novopokr., 451 Ptarmica Mill., 471 acuminata Ledeb., 471 vulgaris DC., 471 Pulicaria Gaertn., 437 dysenterica (L.) Bernh., 437 var. discoidea Druce, 437 subvar. flosculosa Corb., 437 var. longiradiata Druce, 437 graveolens (L.) Nyman, 436 prostrata Aschers., 437 pulicaria (L.) Karst., 437 uliginosa (Sibth.) Gray, 437 viscosa (L.) W. D. J. Koch, 436 vulgaris Gaertn., 437 Pyrethrum Zinn, 463 balsamita (L.) Willd., 465 indicum (L.) Cass., 482 inodorum Moench, 486 macrophyllum (Waldst. & Kit.) Willd., 464 major (Desf.) Tzvelev, 465 maritimum (L.) Sm., 485 myconis (L.) Moench, 479 parthenium (L.) Sm., 464 sinense (Sabine) DC., 482 tanacetum (Karsch) Grecescu, 464 uliginosum Waldst. & Kit., ex Willd., 479 zawadskii (Herbich) Nyman, 483 Ragweed, 513 Giant, 514 Perennial, 513

Ragwort Broad-leaved, 492 Chamois, 493 Chinese, 506 Common, 494 Fen, 493 Golden, 492 Hedge, 506 Hoary, 496 Magellan, 494 Marsh, 495 Monro’s, 506 Narrow-leaved, 492 Narrow-lobed, 491 Oxford, 496 Purple, 491 Shoddy, 491 Shrub, 505 Silver, 490 Welsh, 500 Wood, 493 Woad-leaved, 491 York, 500 Rampion, Oxford, 10 Round-headed, 10 Spiked, 10 Ramtilia DC. oleifera DC., 516 Rapuntium Mill., 12 dormanna (L.) C. Presl, 13 erinus (L.) Mill., 12 siphiliticum (L.) Mill., 13 urens (L.) Mill., 12 Ratibida Raf. columnifera (Nutt.) Wooton & Standley, 518 Rhagadiolus Juss., 98 creticus (L.) All., 97 edulis Gaertn., 98 intermedius Ten., 98 lyratus Lam., 98 stellatus (L.) Gaertn., 98 var. eriocarpus Faure & Maire, 98 var. hebelaenus DC., 98 var. indivisus Viv., 98 var. leiocarpus DC., 98 var. leiolaenus Boiss., 98 Rhodanthe Lindl. charsleyae (F. Muell.) Paul G. Wilson, 432 corymbiflora (Schtdl.) Paul G. Wilson, 432 floribunda (DC.) Paul G. Wilson, 432 manglesii Lindl., 432 moschata (Cunn. ex DC.) Paul G. Wilson, 432 Robin’s Plantain, 454 Rodigia Spreng., 205 bulgarica Velen., 205 commutata Spreng., 206 Rubia L., 28 anglica Huds., 28 peregrina L., 28 tinctorum L., 29

613

614 RUBIACEAE, 14 Rudbeckia L., 517 amplexicaulis Vahl, 517 fulgida Aiton, 517 hirta L., 517 laciniata L., 518 serotina Nutt., 517 trilobata L., 517 Sabazia Cass. urticaefolia (Kunth) DC., 521 Safflower, 92 Downy, 93 Sagmen Hill, 85 jacoidea Hill, 85 scabiosa (L.) Hill, 85 Salsify, 110 Black, 109 Slender, 111 Sambucus L., 29 aurea (Sweet) Carri`ere, 31 canadensis l., 31 ebulus L., 31 herbacea Stokes, 31 humilis Mill., 31 laciniata (L.) Mill., 31 melanocarpa A. Gray, 30 nigra L., 30 forma aurea (Sweet) Schwer., 31 var. aurea Sweet, 31 var. chlorocarpa Hayne, 31 var. heterophylla Endl., 31 forma laciniata (L.) Zabel, 31 var. laciniata L., 31 var. leucocarpa Sm., 31 forma luteovariegata (Weston) Schwer., 31 var. luteovariegata Weston, 31 forma pendula Dippel, 31 var. rotundifolia DC., 31 var. virescens Sweet, 31 forma viridis (Weston) Schwer., 31 var. viridis Weston, 31 pubens Michx, 30 racemosa L., 30 var. melanocarpa (A. Gray) McMinn, 30 subsp. pubens (Michx) House, 30 var. pubens (Michx) Koehne, 30 subsp. sieboldiana (Miq.) Hara, 30 var. sieboldiana Miq., 30 sieboldiana (Miq.) Schwer., 30 Samphire, Golden, 435 Santolina L., 469 chamaecyparissus L., 470 subsp. chamaecyparissus, 470 subsp. tomentosa (Pers.) Arcangeli, 470 maritima (L.) Crantz, 470 suaveolens Pursh, 484 tomentosa Pers., 470 Saussurea DC., 72 alpina (L.) DC., 72

Index Saw-wort, 83 Alpine, 72 Scabiosa L., 59 ambigua Ten., 62 arvensis L., 57 var. integrifolia (Roth) Peterm., 58 var. pinnatisecta auct., 57 ateridoi Pau, 62 atropurpurea L., 59 subsp. maritima (L.) Arcangeli, 62 borealis Salisb., 58 columbaria L., 59 var. patens (Jord.) Car. & St-Lag., 59 cupanii Guss., 62 elata Hornem., 57 fuchsii Gray, 58 gigantea Ledeb., 57 grandiflora Scop., 62 integrifolia Roth., 58 maritima L., 62 paui Sennen, 62 subacaulis Bernardin, 58 succisa L., 58 var. arenaria Rouy, 58, 530 var. glabrata auct., 58 var. grandifolia Rouy, 59, 530 var. hispidula auct., 58 var. ovalis Rouy, 58, 530 var. subacaulis (Bernardin) Rouy, 58, 530 tatarica M. Bieb., 57 Scabious, Devil’s-bit, 58 Field, 57 Giant, 57 Small, 59 Sweet, 59 Scarole, 96 Schkuhria Roth, 526 abrotanoides Roth, 527 advena Thell., 527 bonariensis Hook. & Arn., 527 isopappa Benth., 527 pedata A. Gray, 527 pinnata (Lam.) Thell., 527 Scolymus L., 93 hispanicus L., 94 maculatus L., 93 Scorzonera L., 109 autumnalis (L.) Lam., 101 hispanica L., 109 humilis L., 109 Scorzoneroides Moench, 100 autumnalis (L.) Moench, 101, 531 var. alpina (Gaudin) P. D. Sell, 102, 531 var. cinerascens (Briq.) P. D. Sell, 102, 531 var. coronopifolia (Lange) P. D. Sell, 102, 531 var. dentata (Holuby ex Schur) P. D. Sell, 102, 531 var. latifolia (Schur) P. D. Sell, 102, 531 var. nigrolanata (Fr.) P. D. Sell, 101, 531 var. pinnatifida (Schur) P. D. Sell, 102, 531

Index Scorzoneroides autumnalis Moench (cont.) var. pratensis (Hornem.) P. D. Sell, 102, 531 var. salina (Aspegren) P. D. Sell, 101, 531 var. simplex (Duby) P. D. Sell, 101, 106, 531 Senecilis Gaertn. tangutica (Maxim.) Kitam., 506 Senecio L., 488 aethnensis Jan ex DC., 497 × albescens Burb. & Colgan, 491 alpestris Gaudin, 493 ambiguus (Biv.) DC., 490 × erucifolius, 491 × jacobaea, 491 aquaticus Hill, 495, 556 subsp. aquaticus, 495 var. aquaticus, 495 var. aquaticus auct., 495 forma aurantiacus Druce, 495 var. barbareifolius auct., 495 subsp. barbareifolius (Wimm. & Grab.) Walters, 495 var. barbareifolius Wimm. & Grab., 495 forma discoideus Druce, 495 var. dubius Druce, 495 subsp. erraticus (Bertol.) V. A. Mathews, 495 var. erucoides (Wimm. & Grab.) V. A. Mathews, 495 var. flosculosus Gray, 495 var. intermedius Druce, 495 × jacobaea, 496 var. lanatus Gray, 495 var. major Bab., 495 subsp. ornatus (Druce.) P. D. Sell, 495, 556 var. ornatus Druce, 495, 556 var. pennatifidus Gren. & Godr., 495 arcticus Rupr., 505 × baxteri Druce, 500 bicolor (Willd.) Tod., 490 subsp. cineraria (DC.) Chater, 490 subsp. nebrodensis (Guss.) Chater, 490 brasiliensis Less., 488 burchellii auct., 492 cambrensis Rosser, 500 campestris auct., 504 var. maritima Syme, 504 carnosus (C. Presl) C. Presl, 497 chrysanthemifolius Poir., 497 subsp. aethnensis (Jan ex DC.) Lambinon, 497 cineraria DC., 490 clivorum (Maxim.) Maxim., 507 congesta (R. Br.) DC., 505 crithmifolius Scop., 435 cruentus auct., 503 denticulatus O. F. M¨ull., 501 doria L., 492 subsp. transylvanicus (Boiss.) Nyman, 493 doronicum (L.) L., 493 dunensis Dumort., 494 eboracensis R. J. Abbott & A. J. Lowe, 500 elegans L., 503 erectus Trow, 501

Senecio L. (cont.) erraticus Bertol., 495 subsp. barbareifolius auct., 495 subsp. barbareifolius (Wimm. & Grab.) Beger, 495 erucifolius L., 496 var. communis Rouy, 496 var. erucifolius, 496 forma glabra Druce, 496 × jacobaea, 496 var. latilobus Boiss., 496 var. linnaeanus Rouy, 496 var. viridulus (Matrin-Donos) Rouy, 496 flosculosus Jord., 494 fluviatilis Wallr., 492 forsteri Hook. fil., 506 fuchsii auct., 493 georgii Endl., 506 glaber Ucria., 497 glastifolius L. fil., 491 glossanthus (Sond.) Belcher, 488 grandidissimus Schur, 492 grandiflorus P. J. Bergius, 491 greyi auct., 505 × helwingii Begar ex Hegi, 502 henryi Hemsl., 506 hybrida Hyl., 503 × hybridus Burb., 501 inaequidens DC., 492 incisus (C. Presl) C. Presl, 497 integrifolius (L.) Clairv., 504 subsp. maritimus (Syme) Chater, 504 var. maritimus (Syme) A. R. Clapham, 504 jacobaea L., 494 var. abrotanoides Druce, 494 subsp. aquaticus (Hill) Bonnier & Layens, 495 var. condensatus Druce, 494 var. discoideus Wimm. & Grab., 494 subsp. dunensis (Dumort.) Kadereit & P. D. Sell, 494 subsp. erraticus (Bertol.) Sudre, 495 var. flosculosus Lam. & DC., 494 subsp. jacobaea, 494 var. jacobaea, 494 var. nemorosus (Jord.) Loret & Barrandon, 494 var. nudus Weston, 494 var. stenoglossus Brenan & N. D. Simpson, 494 juniperinus L. fil., 488 laciniatus Gray, 494 var. maritimus Gray, 494 lanuginosus Trow, 501 latifolius Trow, 501 lautus auct, 492 × leichtensteinensis Murray, 496 leucanthemifolius Poir. subsp. vernalis (Waldst. & Kit.) C. Alexander, 502 lividus L. var. denticulatus (O. F. M¨ull.) DC., 501 × londinensis Lousley, 500 mikanioides Otto ex Walp., 505 minimus Poir., 503 monroi Hook. fil., 506

615

616 Senecio L. (cont.) montanus L., 502 multicaulis Trow, 501 nebrodensis auct., 497 var. bipinnatifidus (Vis.) Hayek, 497, 556 var. calabricus (Fiori) Fiori, 497 var. laciniatus Vis., 499, 556 nemorosus Jord., 494 nudiusculus DC., 488 oederiifolius DC., 488 × ostenfeldii Druce, 496 ovatus (P. Gaertn., B. Mey. & Scherb.) Willd., 493 subsp. alpestris (Gaudin) Herborg, 493 paludosus L., 493 palustris (L.) Hook., 505 × patersonianus R. M. Burton, 496 pinnulatus Thunb., 491 polyanthemus L. var. subserratus DC., 491 praecox Trow, 501 przewalskii Maxim., 507 × pseudovernalis Zabel ex Nyman, 502 pterophorus DC., 491 var. apterus Harv., 491 quadridentatus Labill., 503 rapistoides DC., 502 repens Stokes, 492 rupestris Waldst. & Kit., 497, 556 var. bipinnatifida Vis., 497 var. calabricus (Fiori) Pign., 497 var. pinnatifidus Evers ex Hegi, 497, 556 forma sinuatodentatus Evers ex Hegi, 497 var. sinuatodentatus (Evers ex Hegi) E. I. Ny´arˆady, 497 salicetorum Godr., 492 sarracenicus auct., 492 scandens DC., 505 semperflorens Stokes, 501 smithii DC., 494 spathulifolius auct., 504 squalidus L., 496, 556 subsp. aethnensis (Jan ex DC.) Greuter, 497 var. aethnensis (Jan ex DC.) Fiori, 497 var. bipinnatifidus (Vis.) P. D. Sell, 497, 556 var. calabricus Fiori, 497 subsp. chrysanthemifolium (Poir.) Greuter, 497 subsp. glaber (Ucria) Nyman, 497 var. glaucescens Spruce, 497 var. laciniatus (Vis.) P. D. Sell, 497, 556 var. pinnatifidus (Evers ex Hegi) P. D. Sell, 497, 556 subsp. rupestris (Waldst & Kit.) P. D. Sell, 497, 556 var. rupestris (Waldst & Kit.) P. D. Sell, 497, 556 subsp. squalidus, 497 forma subinteger Druce, 497 var. subinteger (Druce) Druce, 497 × viscosus, 500 × vulgaris, 500 subnebrodensis Simonk., 500

Index Senecio L. (cont.) sylvaticus L., 502 var. denticulatus (O. F. M¨ull.) Wahlenb., 501 var. nanus Rouy, 502 var. sylvaticus, 502 × viscosus, 502 tangutica Maxim., 506 tenuifolius Jacq., 496 × thuretii Briq. & Cavill., 491 transsilvanicus Schur, 492 venustus Aiton, 491 verbascifolius Hombr. & Jacq., 494 vernalis Waldst. & Kit., 502 × vulgaris, 502 viridulus Matrin-Donos, 496 × viscidulus Scheele, 502 viscosus L., 502 vulgaris L., 501, 556 subvar. crassifolius Rouy, 501, 556 var. crassifolius (Rouy) P. D. Sell, 501, 556 subsp. denticulatus (O. F. M¨ull.) P. D. Sell, 501 var. dunensis Knuth, 501 var. erectus (Trow) Druce, 501 var. hibernicus Syme, 501 var. lanifer Wilmott, 501 var. lanuginosus Druce, 501 var. latifolius (Trow) Druce, 501 var. multicaulis (Trow) Druce, 501 var. praecox (Trow) Druce, 501 forma radiatus Hegi, 501 var. radiatus auct., 501 var. radiatus W. D. J. Koch, 501 var. rubricaulis Trow ex C. E. Salmon, 501 subsp. vulgaris, 501 var. vulgaris, 501 × whitwellianus Lees ex Cheetham, 496 Seridia Juss., 88 asper (L.) Sweet, 88 melitensis (L.) Sweet, 88 solstitialis (L.) Sweet, 88 Seriola L. commutata (Spreng.) Less., 206 Seriphidium (Besser ex Hook.) Fourr. maritimum (L.) Poljakov, 469 Serratula L., 83 alpina L., 72 arvensis L., 80 incana C. G. Gmel., 81 inermis Gilib., 83 monticola Boreau, 84 picris (Willd.) M. Bieb., 84 setosa Willd., 81 tinctoria L., 83 var. monticola (Boreau) Syme, 84 var. reducta Rouy, 84 Shaggy Soldier, 521 Shawia J. R. & G. Forst., 460 avicenniifolia Raoul, 460 paniculata (J. R. & G. Forst.) Druce, 460 Sheep’s-bit,

Index Sherardia L., 16 arvensis L., 16 var. hirsuta Baguet, 16 var. maritima Griseb., 16 var. mutica Wirtg., 16 var. ovata Fisch.-Benz., 16 var. walravenii Wirt., 16 Sigesbeckia L., 516 caspica Fisch. & C. A. Mey., 517 cordifolia auct., 517 iberica Willd., 517 jorullensis auct., 517 microcephala DC., 516 orientalis L., 517 serrata DC., 517 Silybum Adans. 83 mariae Gray, 83 marianum (L.) Gaertn., 83 Sincalia H. Rob. & Brettell, 506 henryi (Hemsl.), H. Rob. & Brettell, 506 tangutica (Maxim.) B. Nord., 506 Sixalis Raf. atropurpurea (L.) Greuter & Burdet, 59 subsp. maritima (L.) Greuter & Burdet, 62 Sneezeweed, 527 Sneezewort, 471 Snowball Tree, 40 Snowberry, 41 Doorenbos, 42 Pink, 41 Small-leaved, 42 Solidago L., 440 altissima Aiton, 241 altissima L., 442 angustifolia Mill., 441 calcicola (Fernald) Fernald, 441 cambrica Huds., 441 canadensis L., 441 subsp. altissima (L.) O. Bol`os & Vigo, 442 subsp. canadensis, 442 var. scabra Torr. & A. Gray, 442 × virgaurea, 442 doronicum L., 493 gigantea Aiton, 442 subsp. gigantea, 442 var. leiophylla Fernald, 442 subsp. serotina (Kuntze) McNeill, 442 var. serotina (Kuntze) Cronquist, 442 graminifolia (L.) Salisb., 442 graveolens (L.) Lam., 436 jailarum Juz., 441 lanceolata L., 442 lapponica With., 441 latifolia Mill., 441 minor Mill., 441 minuta L., 441 minuta Mill., 441 × niederederi Khek, 442 rugosa Mill., 441 serotina Aiton, 442

Solidago L. (cont.) virgaurea L., 440 var. alpestris Rchb., 441 var. angustifolia (Mill.) Sm., 441 var. arctica Hartm., 441 var. calcicola Fernald, 441 var. cambrica (Huds.) Sm., 441 var. corymbosa Druce, 441 var. dentatifolia Druce, 441 var. interrupta Druce, 441 var. lanceolata Druce, 441 var. latifolia (Mill.) Sm., 441 var. minor (Mill.) Sm., 441 subsp. minuta (L.) Arcangeli, 441 var. minuta (L.) Simmons, 441 var. plukenetiana Druce, 441 var. pumila (Willd.) Gaudin, 441 subsp. virgaurea, 441 var. vulgaris Mill., 441 viscosa (L.) Lam., 436 vulgaris Mill., 441 Solstitiaria Hill, 88 flava Hill, 88 melitensis (L.) Hill, 88 solstitialis (L.) Hill, 88 Sonchus L., 112 alpinus L., 118 arvensis L., 113 var. angustifolius auct., 113 subsp. glabrescens (Wimm. & Grab.) Guenth., 113 var. glandulosus Coss. & Germ., 113 var. integrifolius Bisch., 113 var. laevipes W. D. J. Koch, 113 var. maritimus G. F. W. Meyer, 113 forma uliginosus (M. Bieb.) Javorka, 113 subsp. uliginosus (M. Bieb.) Nyman, 113 asper (L.) Hill, 114, 532 subsp. glaucescens (Jord.) Ball, 114 var. inermis Bisch., 114 var. integrifolius Lej., 114 var. laevis Wallr., 114 subsp. nymanii (Tineo & Guss.) Hegi, 114 × oleraceus, 115 var. sabulosus P. D. Sell, 114, 532 caeruleus Sm., 118 canadensis Froel., 119 glaucescens Jord., 114 hispidus Ledeb., 119 macrophyllus Willd., 119 nymanii Tineo & Guss., 114 oleraceus L., 113, 532 var. albescens Neuman, 114 subsp. asper (L.) Hook. fil, 114 var. asper L., 114 var. glandulosus auct., 114 var. integrifolius Wallr., 114 var. lacerus Wallr., 114 var. litoralis P. D. Sell, 114, 532 var. purpurascens Druce, 114 var. triangularis Wallr., 114

617

618 Sonchus L. (cont.) palustris L., 112 piquetii Druce, 113 plumieri L., 119 tataricus L., 118 uliginosus M. Bieb., 113 vulgaris Rouy, 113 var. marinus Rouy, 113 Southernwood, 468 Sow-thistle, Alpine Blue, 118 Common Blue, 119 Hairless Blue, 119 Marsh, 112 Perennial, 113 Pontic Blue, 119 Prickly, 114 Smooth, 113 Soyeria Monnier, 201 paludosa (L.) Godr., 202 Spanish Needles, 524 Specularia Heist. ex A. DC., 8 hybrida (L.) A. DC., 8 speculum-veneris (L.) A. DC., 8 Squinancywort, 17 Star-thistle, Eastern, 87 Large-headed, 89 Lesser, 89 Maltese, 88 Red, 87 Rough, 88 Small-flowered, 87 Woolly, 88 Yellow, 88 Stenactis Cass. annuus (L.) Nees, 455 glauca (Ker Gawl.) Nees 453 speciosa Lindl., 453 Stephanocoma Less. rubricaulis (DC.) Sch. Bip., 424 Stobaea Thunb., 423 apitrachys DC., 423 biloba DC., 423 heterophylla Thunb., 423 pinnatifida Thunb., 423 rigida Thunb., 424 rubricaulis DC., 424 zeyheri (Less.) DC., 424 St Paul’s-wort, Eastern, 517 Western, 517 Strongylosperma Less. australe (Sieber ex Spreng.) Less., 487 Stuartina Sonder hamata Philipson, 427 Succisa Haller, 58 praemorsa Asch., 58 pratensis Moench, 58, 530 var. arenaria (Rouy) P. D. Sell, 58, 530 var. grandifolia (Rouy) P. D. Sell, 59, 530 var. ovalis (Rouy) P. D. Sell, 58, 530 var. subacaulis (Bernardin) P. D. Sell, 58, 530 subacaulis Bernadin, 58, 530

Index Sunflower, 518 Lesser, 519 Perennial, 519 Thin-leaved, 519 Symphoria Pers., 41 glaucescens (Kunth) Spreng., 42 glomerata Pursh, 41 microphylla (Kunth) Spreng., 42 montana (Kunth) Spreng., 42 racemosa (Michx) Pursh, 41 rubra Raf., 41 Symphoricarpos Duhamel, 41 albus (L.) S. F. Blake, 41, 530 subsp. laevigatus (Fernald) P. D. Sell, 41, 530 subsp. laevigatus × microphyllus × orbiculatus, 42 var. laevigatus (Fernald) S. F. Blake, 41 × chenaultii Rehder, 41 × doorenbosii Kr¨ussm., 42 glaucescens Kunth, 42 imberbis Tausch, 41 mexicanus K. Koch, 42 microphyllus Kunth, 42 × orbiculatus, 41 montanus Kunth, 42 orbiculatus Moench, 41 racemosus Michx, 41 var. laevigatus Fernald, 41, 530 rivularis Suksd., 41 spicatus Engelm., 41 vulgaris Michx, 41 Symphyotrichum Nees hesperius (A. Gray) Å. L¨ove, 448 laeve (L.) Å. L¨ove, 446 lanceolatum (Willd.) G. L. Nesom, 448 novi-belgii (L.) G. L. Nesom, 447 × salignum (Willd.) G. L. Nesom, 448 simplex (Willd.) Å. L¨ove, 448 × versicolor (Willd.) G. L. Nesom, 447 Tagetes L., 525 erecta L., 526 erythrocephala Rusby, 526 gracilis DC., 526 minuta L., 526 patula L., 526 tenuifolia Cav., 526 Tanacetum L., 463 angulatum Willd., 473 balsamita L., 465, 556 var. balsamita, 465 subsp. balsamitoides (Sch. Bip.) Grierson, 465 var. balsamitoides (Sch. Bip.) P. D. Sell, 465, 556 var. tanacetoides Boiss. 465 balsamitoides Sch. Bip., 465, 556 breviradiatum Sch. Bip., 464 coccineum (Willd.) Grierson, 463 grandiflorum Thunb., 483 indicum (L.) Sch. Bip., 482 leucanthemum (L.) Sch. Bip., 480 leucophyllum Regel, 463

Index Tanacetum L. (cont.) macrophyllum (Waldst. & Kit.) Sch. Bip., 464 morifolium (Ramat.) Kitam., 482 parthenium (L.) Sch. Bip., 464 forma discoideum Sch. Bip., 464 forma flosculosum (DC.) Beck, 464 forma hortense (Schur) Beck, 464 forma parthenium, 464 sinense (Sabine) Sch. Bip., 482 serotinum (L.) Sch. Bip., 479 suaveolens (Pursh) Hook., 484 vulgare L., 464 zawadskii (Herbich) Pawl., 483 Tansy, 464 Rayed, 464 Taraxacum Wigg, 120 aberrans Hagend., Soest & Zevenb., 190 acidodontum Dahlst., 150 acidotum M. P. Christ., 148 acroglossum Dahlst., 184 acromaurum Dahlst., 149 acutifidum M. P. Christ., 200 acutifrons Markl., 179 acutum A. J. Richards, 144 adamii auct., 153 adiantifrons Ekman ex Dahlst., 182 adsimile Dahlst., 199 aequatum Dahlst, 186 aequilobum Dahlst., 183 aequisectum M. P. Christ., 181 affine G. E. Haglund, 145 akteum Hagend., Soest & Zevenb., 158 alatum H. Lindb., 170 altissimum H. Lindb., 181 amarellum J. Kirschner & Stepanek, 146 amphiodon Dahlst. ex G. E. Haglund, 195 amplum Markl., 189 ancistrolobum Dahlst., 180 angliciforme Dahlst., 160 anglicum Dahlst., 147 angulare Hagend., Soest & Zevenb., 180 angustisquamum Dahlst. ex H. Lindb., 181 ardisodon Dahlst., 197 ardlense A. J. Richards, 156 arenastrum A. J. Richards, 138 argutum Dahlst., 137 atactum Sahlin & Soest, 165 atonolobum Hagend., Soest & Zevenb., 171 atrovirens Dahlst., 164 aurosulum H. Lindb., 190 austrinum auct., 146 beeftinkii Hagend., Soest & Zevenb., 158 bercheriense Dahlst., 195 berthae C. C. Haw., 159 biforme auct., 161 boekmanii Borgv., 165 brachycranum (Dahlst.) A. W. Hill, 142 brachyglossum (Dahlst.) Raunk., 137 bracteatum Dahlst., 153 breconense C. C. Haw., 161 britannicum Dahlst., 154

Taraxacum Wigg (cont.) broddesonii G. E. Haglund, 176 caledonicum A. J. Richards, 161 calophyllum Dahlst., 152 caloschistum Dahlst., 196 cambricum A. J. Richards, 160 cambriense A. J. Richards, 159 campylodes G. E. Haglund, 148 canulum G. E. Haglund, 142 caudatulum auct., 190 celticum A. J. Richards, 155 cenabense Sahlin, 139 ceratolobum Dahlst., 148 cherwellense A. J. Richards, 176 chloroleucophyllum Dahlst., 162 chloroticum Dahlst., 177 chloroticum Dahlst. ex Florstr¨om, 177 chrysophaenum Rail., 179 ciliare Soest, 146 cimbricum Wiinst., 147 clovense A. J. Richards, 148 coartatum G. E. Haglund, 190 commixtum G. E. Haglund, 138 commutatum Dahlst., 138 connexum Dahlst., 189 cophocentrum Dahlst., 191 cordatum Palmgr., 188 cornubiense A. J. Richards, 152 corynodes G. E. Haglund, 169 craspedotum Dahlst., 149 crispifolium auct., 154 croceiflorum Dahlst., 175 croceum auct., 148 curtifrons H. Øllg., 178 cyanolepis Dahlst., 178 cymbifolium H. Lindb. ex Dahlst., 149 dahlstedtii H. Lindb., 193 degelii G. E. Haglund, 143 densiflorum M. P. Christ., 193 densilobum Dahlst., 170 diastematicum Markl., 192 dilaceratum M. P. Christ., 170 dilatatum H. Lindb., 192 discretum auct., 139 disseminatum G. E. Haglund, 141 drucei Dahlst., 152 dunense Soest, 139 duplidens H. Lindb., 161 duplidentiforme auct., 155 duplidentifrons Dahlst., 155 edmondsonianum H. Øllg., 184 edmondsonii A. J. Richards, 151 ekmanii Dahlst., 189 erythrospermum Andrz. ex Bess. subsp. brachyglossum Dahlst., 137 subsp. gotlandicum Dahlst., 140 subsp. lacistophyllum Dahlst., 136 subsp. proximum Dahlst., 140 subsp. rubicundum Dahlst., 138 euryphyllum (Dahlst.) Hjelt, 150 exacutum Markl., 185

619

620 Taraxacum Wigg (cont.) excellens Dahlst., 156 eximium auct., 147 expallidiforme Dahlst., 174 exsertiforme Hagend., Soest & Zevenb., 185 exsertum Hagend., Soest & Zevenb., 184 faeroense (Dahlst.) Dahlst., 147 fagerstroemii S˚altin, 195 falcatum Brenner, 142 fasciatum Dahlst., 200 firmuliforme A. J. Richards, 158 fulgidum G. E. Haglund, 157 fulvicarpum Dahlst., 161 fulviforme Dahlst., 141 fulvum Raunk., 142 subsp. brachycranum Dahlst., 142 fusciflorum H. Øllg., 164 geirhildae (Beeby) R. Palmer & Walter Scott, 147 gelertii Raunk., 153 glauciniforme Dahlst., 143 gotlandicum (Dahlst.) Dahlst., 140 grandifolium G. E. Haglund, 165 gravinense Dahlst., 180 haematicum G. E. Haglund ex H. Øllg. & Wiezell, 158 hamatiforme Dahlst., 166 hamatifrons auct., 161 hamatifrons Dahlst., 150 hamatulum Hagend., Soest & Zevenb., 163 hamatum Raunk., 162 hamifarum Dahlst., 164 haworthianum Dudman & A. J. Richards, 139 helvicarpum Dahlst., 141 hemicyclum G. E. Haglund, 182 hemipolyodon Dahlst., 195 hepaticum Rail., 195 hesperium C. C. Haw., 156 hexhamense A. J. Richards, 189 hibernicum G. E. Haglund, 154 hirsutissimum C. C. Haw., 151 hispanicum auct., 137 hoedicense Soest, 146 horridifrons Rail., 170 huelphersianum Dahlst. ex G. E. Haglund, 194 hygrophilum Soest, 157 hypochaeris auct., 148 inane A. J. Richards, 156 incisum H. Øllg., 198 inopinatum C. C. Haw., 136 insigne Ekman ex M. P. Christ. & Wiinst., 171 interveniens G. E. Haglund, 181 intumescens G. E. Haglund, 179 johnstonii Dahlst., 162 kernianum Soest, Hagend. & Zevenb., 167 lacerabile Dahlst., 160 lacerifolium G. E. Haglund, 175 laciniosifrons Wiinst., 193 laciniosum Dahlst., 171 lacinulatum Markl., 175 lacistophyllum (Dahlst.) Raunk., 136 laeticolor Dahlst., 168

Index Taraxacum Wigg (cont.) laetiforme auct., 139 laetifrons Dahlst., 152 laetum auct., 139 lainzii auct., 152 lamprophyllum M. P. Christ., 167 lancastriense A. J. Richards, 160 lanceolatum Poiret, 146 lancidens Hagend., Soest & Zevenb., 167 landmarkii Dahlst., 158 latens H. Øllg., 184 laticordatum Markl., 172 latisectum H. Lindb., 194 latispina Dahlst., 199 latissimum Palmgr., 183 lepidum M. P. Christ., 174 leptodon Markl., 186 leucopodum G. E. Haglund, 176 limnanthes G. E. Haglund subsp. limnanthoides Soest, 146 linguatum Dahlst. ex M. P. Christ. & Wiinst., 168 lingulatum Markl., 186 longisquameum H. Lindb., 199 lucidum Dahlst., 201 lunare M. P. Christ., 190 luteum C. C. Haw. & A. J. Richards, 162 macranthoides G. E. Haglund, 187 macrolobum Dahlst., 168 maculatum Jord., 199 maculigerum auct., 151, 152 maculigerum H. Lindb. subsp. europhyllum Dahlst., 150 maculosum A. J. Richards, 151 magnihamatum auct., 156 margettsii C. C. Haw., 173 maritimum Hagend., Soest & Zevenb., 146 marklundii Palmgr., 163 melanthoides Dahlst. ex M. P. Christ. & Wiinst., 201 mimulum Dahlst. ex H. Lindb., 182 monspeliense (Dahlst.) Dahlst., 138 mucronatum auct., 199 multicolorans Hagend., Soest & Zevenb., 197 naeviferum auct., 197 naeviferum Dahlst., 171 naevosiforme Dahlst., 150 naevosum Dahlst., 150 necessarium H. Øllg., 173 nietoi A. J. Richards, 154 nigridentatum T. Edmondson, 172 nitidum Hagend., Soest & Zevenb., 198 nordstedtii Dahlst., 159 obliquilobum Dahlst., 194 obliquum (Fr.) Dahlst., 145 subsp. platyglossum (Raunk.) Nordh., 145 oblongatum Dahlst., 191 obscuratum G. E. Haglund, 199 obtusifrons Markl., 186 obtusilobum Dahlst. ex G. E. Haglund, 177 ochrochlorum G. E. Haglund ex Rail., 189 oellgardii C. C. Haw., 155

Index Taraxacum Wigg (cont.) officinale Wigg., 148 officinale auct. subsp. gelertii (Raunk.) Dahlst., 153 subsp. ostenfeldii (Raunk.) Dahlst., 161 subsp. speciosum (Raunk.) Dahlst., 177 subsp. tenebricans Dahlst., 169 olgae A. J. Richards, 160 oncolobum Dahlst., 174 orcadense Dahlst., 153 ordinatum Hagend., Soest & Zevenb., 171 ornatum auct., 147 ostenfeldii Raunk., 161 oxoniense Dahlst., 141 pachylobum Dahlst., 194 pachymerum G. E. Haglund, 192 pallescens Dahlst., 172 pallidipes Markl., 174 pallidipes Markl. ex G. E. Haglund, 174 palustre (Lyons) Symonds, 146 palustrisquamum A. J. Richards, 160 pannucium Dahlst., 168 pannulatiforme Dahlst., 171 pannulatum Dahlst., 186 parnassicum Dahlst., 139 parvuliceps auct., 161 pectinatiforme H. Lindb., 196 pectinosum G. E. Haglund, 142 perhamatum Dahlst., 191 pericrispum M. P. Christ., 170 perlaciniatum Dahlst., 153 piceatum Dahlst., 178 placidum A. J. Richards, 144 planum Raunk., 197 platyglossum Raunk., 145 plicatum Dahlst., 150 pollichii auct., 146 polyodon Dahlst., 197 porrigens Markl. ex Puol., 176 porteri C. C. Haw., 155 praeradians auct., 184 prionum Hagend., Soest & Zevenb., 165 proceriquameum H. Øllg., 187 procerum auct., 187 protractifrons Dahlst. ex M. P. Christ. & Wiinst., 168 proximiforme Soest ex Lambinon & Soest, 140 proximum (Dahlst.) Raunk., 140 pruinatum M. P. Christ., 166 pseudohamatum auct., 162 pseudohamatum Dahlst., 164 pseudolacistophyllum Soest, 145 pseudolarssonii A. J. Richards, 151 pseudonordstedtii A. J. Richards, 159 pseudoproximum Soest, 141 pseudoretroflexum M. P. Christ., 191 pulchrifolium Markl., 196 pycnostictum M. P. Christ., 149 quadrans H. Øllg., 164 raii auct., 146 raunkiaeri Wiinst., 155

Taraxacum Wigg (cont.) reclinatum auct., 147 remanentilobum Soest, 188 retroflexum H. Lindb., 182 retzii Soest, 142 rhamphodes G. E. Haglund, 187 richardsianum C. C. Haw., 152 rubellum M. P. Christ., 150 rubicundum (Dahlst.) Dahlst., 138 subsp. monspeliense Dahlst., 138 subsp. pulvigerum H. Lindb., 138 sagittipotens Dahlst. & R. Ohlsen ex G. E. Haglund, 188 sahlinianum Dudman & A. J. Richards, 166 sarniense A. J. Richards, 146 scanicum Dahlst., 137 scoticum A. J. Richards, 142 scotiniforme Dahlst. ex G. E. Haglund, 199 sellandii Dahlst., 180 semiglobosum H. Lindb., 183 semiprivum Dahlst., 170 serpenticola A. J. Richards, 148 serratilobum Dahlst., 162 severum M. P. Christ., 191 silesiacum Dahlst. ex G. E. Haglund, 139 similatum Dahlst., 195 simile auct., 141 sinuatum Dahlst., 193 speciosum Raunk., 177 spectabile auct., 147 var. faeroense Dahlst., 147 var. geirhildae Beeby, 147 spiculatum M. P. Christ., 166 spilophyllum auct., 185 stenacrum Dahlst., 175 stenoglossum Dahlst., 176 stereodes Ekman ex G. E. Haglund, 182 stictophoreum M. P. Christ., 149 stictophyllum Dahlst., 152 subbracteatum A. J. Richards, 154 subcyanolepis M. P. Christ., 174 subdilatatum Dahlst., 194 subditivum Hagend., Soest & Zevenb., 166 subexpallidum Dahlst., 168 subhamatum M. P. Christ., 163 subhuelphersianum M. P. Christ., 173 sublaciniosum auct., 168 sublaearifolium Hagend., Soest & Zevenb., 195 sublaeticolor Dahlst., 173 sublaetum auct., 139 sublatissimum Dahlst., 200 sublongisquameum M. P. Christ., 199 subnaevosum A. J. Richards, 151 subpallescens Dahlst., 186 subpraticola auct., 174 subundulatum Dahlst., 195 subxanthostigma M. P. Christ. ex H. Øllg., 200 sundebergii Dahlst., 201 tamesense A. J. Richards, 157 tanylepioides Dahlst., 153 tanylepis Dahlst., 144

621

622 Taraxacum Wigg (cont.) tanyphyllum Dahlst., 193 tarachodum Hagend., Soest & Zevenb., 186 tenebricans (Dahlst.) Dahlst., 169 texelense Hagend., Soest & Zevenb., 157 tortilobum Florstr¨om, 144 trilobatum Palmgr., 196 tumentilobum Markl. ex Puol., 179 uncosum auct., 172 undulatiflorum M. P. Christ., 177 undulatum H. Lindb. & Markl., 169 unguilobiforme Dahlst., 162 unguilobum Dahlst., 162 vachelliae Dahlst., 137 valens Markl., 185 vastisectum Markl. ex Paol., 187 wallonicum Soest, 143 webbii A. J. Richards, 146 westhoffii Hagend., Soest & Zevenb., 146 xanthostigma H. Lindb., 198 xiphoideum G. E. Haglund, 148 Tarragon, 468 Russian, 469 Teasel, Cut-leaved, 56 Fuller’s, 55 Small, 56 Wild, 55 Yellow-flowered, 56 Telekia Baumg., 438 cordifolia (Waldst. & Kit.) DC., 438 ovata K. Koch, 438 speciosa (Schreb.) Baumg., 438 Tephroseris (Rchb.) Rchb., 504 helenitis (L.) B. Nord., 504 integrifolia (L.) Holub, 504 subsp. integrifolia, 504 subsp. maritima (Syme) B. Nord., 504 palustris (L.) Fourr., 505 subsp. congesta (R. Br.) Holub, 505 Tessenia Bub. philadephica (L.) Lunell, 454 var. attenuata (Nutt.) Lunell, 454 Tetragonotheca L. abyssinica (L. fil.) Ledeb., 516 Tetralix Hill, 75 eriophora (L.) Hill, 76 Thistle, Blessed, 92 Broad-winged, 74 Cabbage, 79 Carline, 69 Cotton, 82 Creeping, 80 Golden, 94 Marsh, 80 Meadow, 77 Melancholy, 78 Milk, 83 Musk, 75 Plymouth, 73 Reticulate, 82 Slender, 73

Index Thistle (cont.) Spear, 77 Spotted Golden, 93 Tuberous, 78 Welted, 74 Woolly, 76 Yellow, 78 Thrincia Roth, 102 hirta Roth, 103 var. arenaria (Duby) DC., 106 var. psilocalyx (Rchb.) DC., 106 hispida auct., 103 laevis Lag., 106 leysseri Wallr., 103 nudicalyx Lag., 106 nudicaulis auct., 103 psilocalyx Rchb., 103 taraxacoides (Vill.) Gaudin, 103 Throatwort, 9 Thyrsosma Raf., 32 Tickseed, Garden, 524 Large-flowered, 524 Tolpis Adans., 208 barbata (L.) Gaertn., 208 umbellata Bertol., 208 Tolpis, 208 Trachelium L., 9 caeruleum L., 9 Tragopogon L., 110 australis Jord., 111 coelesyriacus Boiss., 111 eriosperma Ten., 111 geropogon Rouy, 111 glaber (L.) Hoffm., 111 hirsutus (L.) Kotschy, 111 hybridus L., 111 longirostris Bisch. ex Sch. Bip., 111 minor Mill., 110 × mirabilis Rouy, 111 orientalis L., 111 porrifolius L., 110 subsp. australis (Jord.) Nyman, 111 subsp. longirostris (Bisch. ex Sch. Bip.) Holmboe, 111 × pratensis, 111 subsp. sativus (Gaterau.) Br.-Bl., 111 pratensis L., 110 subsp. minor (Mill.) Wahlenb., 110 subsp. orientalis (L.) Celak., 110 sinuatus Av´e-Lall., 111 Treasureflower, 423 Plain, 422 Trichera Schrad. ex Roem. & Schult., 57 arvensis (L.) Schrad. ex Roem. & Schult., 57 var. subintegerrima Lange, 58 Trimorpha Cass., 455 acris (L.) Gray, 455 alpina Gray, 454 borealis Vierh., 454 uniflora auct., 454 vulgaris Cass., 455

Index Tripleurospermum Sch. Bip., 484, 556 inodorum Sch. Bip., 486 maritimum (L.) W. D. J. Koch, 485 subsp. inodorum (K. Koch) Hyl. ex Vaar., 486 subsp. maritimum, 485 subsp. nigriceps P. D. Sell 485, 556 subsp. phaeocephalum auct., 485 var. salinum auct., 486 subsp. vinicaule P. D. Sell 486, 556 perforatum (M´erat) Lainz, 486 × Tripleurothemis Stace, 478, 556 gruetterana (Asch.) P. D. Sell, 478, 556 maleolens (P. Fourn.) Stace, 478 Tripolion Raf., 451 maritimum Raf., 451 Tripolium Nees, 451 vulgare Besler ex Nees, 451 Troglophyton Hilliard & B. L. Burtt parvulum (Harvey) Hilliard & B. L. Burtt, 431 Tussilago L., 509 alba L., 511 alpina L., 512 farfara L., 509 fragrans Vill., 511 hybrida L., 511 petasites L., 510 ruderalis Salisb., 509 vulgaris Lam., 509 Twinflower, 42 Urostemon B. Nord., 505 Vaccinium L. album L., 41 Valantia L. aparine L., 27 ciliata Opiz ex J. & C. Presl, 28 cruciata L., 28 Valerian, Common, 52 Marsh, 53 Pyrenean, 53 Red, 54 Turkey, 53 Valeriana L., 52 collina Wallr., 52 dentata (L.) All., 51 dioica L., 53 subsp. sylvatica (Rich.) F. G. Mey., 53 locusta L., 49 var. dentata L., 51 var. olitoria L., 49 officinalis L., 52, 530 var. latifolia auct., 52 subsp. collina (Wallr.) Nyman, 52 subsp. dunensis P. D. Sell, 52, 530 var. mikanii auct., 52 subsp. sambucifolia (J. C. Mikan ex Pohl) Hayw., 52 olitoria (L.) Willd., 49 phu L., 53 pyrenaica L., 53 rubra L., 54 sambucifolia J. C. Mikan ex Pohl, 52 sylvestris Gray, 53

VALERIANACEAE, 48 Valerianella Mill., 48 auricula DC., 51 carinata Loisel., 49 var. pubescens auct., 49 dentata (L.) Pollich, 51 var. mixta Dufour, 52 eriocarpa Desv., 51 var. glabrescens Cosson, 51 kotschyi Boiss. 49 locusta (L.) Laterr., 49 subsp. dunensis (D. E. Allen) P. D. Sell, 49 var. dunensis D. E. Allen, 49 var. oleracea (Schltr.) Breist., 49 morisonii (Spreng.) DC., 51 olitoria (L.) Pollich, 49 rimosa Bastard, 51 var. dasycarpa auct., 51 Vellereophyton Hilliard & B. L. Burtt dealbatum (Thunb.) Hilliard & B. L. Burtt, 431 Venus’s Looking-glass, 8 Large, 8 Verbesina L. encelioides (Cav.) Benth. & Hook. fil. ex A. Gray, 517 sativa Roxb. ex Sims, 516 Viburnum L., 32 americanum auct., 40 × bodnantense Aberc. ex Stearn, 33 davidii Franch., 37 edule Pursh, 40 farinosum Stokes, 33 farreri Stearn, 33 × grandiflorum, 33 fragrans Bunge, 33 lantana L., 33 var. glabratum Chab., 36 × rhytidophyllum, 36 var. rugosum Lange, 36 var. viride A. Kern., 36 opulus L., 37, 530 var. americanum Aiton, 40 forma flavum (Horwood) P. D. Sell, 40, 530 var. flavum Horwood, 40, 530 var. pimina Michx, 40 forma roseum (L.) Hegi, 40 var. roseum L., 40 var. sterilis DC., 40 oxycoccus Pursh, 40 × rhytidophylloides J. V. Suringar, 36 rhytidophyllum Hemsl. ex Forbes & Hemsl., 36 sargentii Koehne, 40 forma flava Rehd., 40 tinus L., 37 trilobum Marsh., 40 veitchii C. H. Wright, 36 Viburnum, Bodnant, 33 David’s, 37 Farrer’s, 33 Hybrid, 36 Wrinkled, 36 Viper’s Grass, 109

623

624 Virea Adans. autumnalis (L.) Gray, 101 hispida (L.) Gray, 103 pyreniaca Gray, 101 Virga Hill, 54 pilosa (L.) Hill, 56 Virgulus Raf., 450 ericoides (L.) Reveal & Keener, 450 var. pansus (Blake) Reveal & Keener, 450 novae-angliae (L.) Reveal & Keener, 450 Vittadinia Rich. triloba auct., 454 Wahlenbergia Schrad. ex Roth, 8 hederacea (L.) Rchb., 9 nutabunda (Guss.) A. DC., 9 trichogyna Stearn, 9 Wayfaring Tree, 33 Asian, 36 Weigela Thunb., 43 florida (Bunge) DC., 43 rosea Lindl., 43 Weigela, 43 Wibelia P. Gaertn., B. Mey. & Scherb., 205 foetida (L.) Sch. Bip., 205 graveolens P. Gaertn., B. Mey. & Scherb., 205 setosa (Haller fil.) R¨ohl., 208 Wiborgia Roth, 520 acmella Roth, 520 parviflora (Cav.) Kunth, 520 urticaefolia Kunth, 521 Wiestia Sch. Bip. virosa (L.) Sch. Bip., 117 Woodruff, Annual, 18 Blue, 18 Pink, 17 Sweet, 19 Woodvillea DC. calendulacea DC., 453 Wormwood, 467 Field, 469 Sea, 469 Xanthidium Delpino, 513 artemisioides (Willd.) Delpino, 513

Index Xanthium L., 514 albinum (Widder) H. Scholz, 515 ambrosioides Hook. & Arn., 516 basilicum Vell., 515 californicum Greene, 515 catharticum Coste & Sennen, 516 cavanillesii Schouw & Didr, 515 echinatum Murray, 515 eriocarpon Wallr., 516 italicum Moretti, 515 leucocarpon Wallr., 516 macrocarpum DC., 515 orientale L. 515 riparium Itzigs & Hertsch., 515 saccharatum Wallr., 515 spinosum L., 515 strumarium L., 515 subsp. italicum Moretti D. L¨ove, 515 subsp. strumarium, 515 Xanthophthalmum Sch. Bip., 478 coronarium (L.) P. D. Sell, 479, 446 var. discolor (d’Urv.) P. D. Sell, 479, 556 segetum (L.) Sch. Bip., 478 Xeranthemum L. bracteatum Vent., 432 bellidiodes G. Forst., 432 Ximenesia Cav. encelioides Cav., 517 Xylosteon Mill., 44 album (L.) Moldenke, 41 cordatum Moench, 45 dumetorum Moench, 45 involucratum Richardson, 44 tataricum (L.) Dum.-Cours., 45 vulgare Borkh., 45 Yarrow, 472 Fern-leaf, 473 Noble, 472 Southern, 472 Sweet, 473 Tall, 471 Yellow Ox-eye, 438 Willow-leaved, 438