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

Citation preview

This page intentionally left blank

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

978-0-511-16796-6 eBook (EBL) 0-511-16796-2 eBook (EBL)

isbn-13 isbn-10

978-0-521-55338-4 hardback 0-521-55338-5 hardback

Cambridge University Press has no responsibility for the persistence or accuracy of urls for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate.

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

xxiv

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

xxvi

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

3

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.

4

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

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

5

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

6

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

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

7

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.

8

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

(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

9

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

10

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

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

11

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

12

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

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.

13

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

14

1 4 3 . RU B I AC E A E

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,

15

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

16

1 4 3 . RU B I AC E A E

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

17

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.

18

1 4 3 . RU B I AC E A E

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

19

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

20

1 4 3 . RU B I AC E A E

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

21

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.

22

1 4 3 . RU B I AC E A E

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.

23

(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,

24

1 4 3 . RU B I AC E A E

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.

25

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

26

1 4 3 . RU B I AC E A E

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

28

1 4 3 . RU B I AC E A E

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.

29

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

30

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

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.

31

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

32

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

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

33

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.

34

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

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

35

36

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

(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

37

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

38

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)

39

40

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.

41

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,

42

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

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

43

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.

44

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

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.

45

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,

46

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

47

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

48

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

49

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

50

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

51

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,

52

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,

53

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

54

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.

55

× 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),

56

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

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,

57

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,

58

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

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

59

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

60

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

61

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.

62

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

64

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

65

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

66

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

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

67

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

68

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

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,

69

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,

70

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

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

71

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

72

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

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

73

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

74

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

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

75

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.

76

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

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-

77

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.

78

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

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,

79

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).

80

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

× 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

81

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.

82

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

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

83

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

84

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

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

85

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.

86

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

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-

87

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.

88

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

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.

89

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.

90

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

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,

91

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

92

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

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.

93

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

94

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

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

95

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

96

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

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

97

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.

98

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

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

99

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

100

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

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,

101

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.

102

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

(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

103

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,

104

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

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

105

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)

106

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

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

107

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

108

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

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

109

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

110

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

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

111

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

112

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

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.

113

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,

114

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

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.]

115

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

116

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

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

117

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

118

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

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

119

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

120

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

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

121

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

122

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

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

123

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

124

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

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.

125

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.

126

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

¨ 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

127

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

128

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

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

133

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

134

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

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

135

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

136

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

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.

137

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

138

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

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

139

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,

140

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

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;

141

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

142

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

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.

143

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

144

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

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

145

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

146

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

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

147

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

148

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

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,

34. Taraxacum

149

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.

150

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

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

151

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

152

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

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

153

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

154

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

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

155

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

156

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

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

157

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

158

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

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.

159

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

160

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

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-

161

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

162

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

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 ×

163

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

164

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

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

165

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

166

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

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

167

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

168

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

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.

169

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