Flora of Ethiopia and Eritrea. Volume 4, part 2 Asteraceae (Compositae) 9789197128599, 9197128597

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FLORA OF ETHIOPIA AND ERITREA VOLUME 4, FART 2 Editors: Inga Hedberg, lb Friis & Sue Edwards

ASTERACEAE (COMPOSITAE) by

Mesfin Tadesse

A ddis A b a b a , E thiopia U ppsala, Sw eden 2004

wiinrom the National nerDarium(EffiF U

... Department of Plant Biology Biodiversity "wanagement A d r i |ec eA ^ Nu ‘Ura' SC,en“ *

Addis Ababa U n i v , ^ .

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 2

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MAP OF THE FLORISTIC REGIONS OF ETHIOPIA AND ERITREA These are the regions used in Volume 3 and kept in all the volumes for continuity They do not bear any relation to the new administrative structures EE - Ertirea East, below and to the cast o f the 1000 m contour

TU - Tigray region, above and to the west o f the 1000 m contour

AR - Arsi region VVG - W elega region

EW - Eritrea west, above and to the west o f the 1000 m contour

GD - Gonder region GJ - Gojam region

IL - Ilubabor region KF - Kefa region

AF - Afar region, below and to the east o f the 1000 m contour to the Eritrean border in the east and the Harerge border in the south

WU - Welo region, above and to the west o f the 1000 m contour

GG -- Gamo Gofa region SD -- Sidamo region

KENYA

SU - Shewa region, above and to the west o f the 1000 m contour

BA - Bale region HA - Harerge region

MAP OF ETHIOPIA AND ERITREA SHOWING MAJOR PHYSIOGRAPHIC FEATURES

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 2 Editors: Inga Hedberg, ib Friis, Sue Edwards

ASTERACEAE (COMPOSITAE) by Mesfin Tadesse

Addis Ababa, Ethiopia Uppsala, Sweden 2004

Published by The National Herbarium, Biology Department, Science Faculty, Addis Ababa University, Ethiopia and The Department o f Systematic Botany Uppsala University, Sweden Prepared by the Ethiopian Flora Project, funded by Addis Ababa University and the Swedish International Development Cooperation Agency (SIDA) through the Ethiopian Science and Technology Commission. Project Leader - Dr. Sebsebe Demissew, and European Coordinator - Dr Inga Hedberg. Final pages prepared at the Department o f Systematic Botany, Uppsala University using a Personal Computer and a LaserJet 4L printer. Text formatted with Ventura Publisher 8. Printed in Times New Roman. Printed by the Educational Materials Production and Distribution Agency (EMPDA), Textbook Production Department, Addis Ababa, Ethiopia. © The National Herbarium, Addis Ababa Univeraity, 2004 ISBN 91-971285-9-7

D istributed by: In Ethiopia

Outside Ethiopia

The National Herbarium Addis Ababa University

The Swedish Science Press P O Box 118 SE-751 04 Uppsala SWEDEN Fax: +46-18-36 52 77 e-mail mfb ffjssp nu

P O Box 3434,

Addis Ababa. ETHIOPIA Tel. +251-1-11 43 23 Fax: +251-1-55 23 50 e-mail: nat.heb^telecom .net.et

Cover illustrations: Front: Aspilia mossambicensis drawn by Birhanu Mihretu Back: Kleiniapendula drawn by Pat Halliday. Spine: C ypselaof Bidens macroptera drawn by Mesfin Tadesse.

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 2 CONTENTS Map o f the floristic regions o f Ethiopia and Eritrea Map o f Ethiopia and Eritrea showing msyor physiographic features Editorial Board Editorial Team Foreword Acknowledgements List o f tribes and genera

ASTERACEAE Key to taxa Mutisieae 1. Tarchonanthus 2. Gerbera 3. Dicoma Cynareae 4. Echinops 5. Carduus 6 . Cirsium 7. Cynara 8 . Silybum 9. Ochrocephala 10. Plectocephalus 11. Volutana 12. Carthamus 13. Centaurca 14. Gomocaulon 15. Notobasis Cichorieae 16. Scolymus 17. Cichorium 18. Crepis 19. Dianthoseris 20. Taraxacum 21. Lactuca 22. Prenanthes 23. Launaea 24. Reichardia 25. Sonchus 26. Tolpis 27. Helminthotheca

28. Picris 29. Tragopogon Vernonieae 30. Vemonia 31. Distephanus 32. Gutenbergia 33. Erlangea 34. Bothriocline 35. Ethulia 36. Elephantopus Arctoteae 37. Arctotis 38. Haplocarpha 39. Berkheya 40. Gazania 41. Hirpicium Inuleae 42. Anisopappus 43. Geigeria 44. Inula 45. Pentanema 46. Blumea 47. Pegolettia 48. Pulicaria 49. Iphiona 50. Iphionopsis Plucheeae 51. Delamerea 52. Pseudoconyza 53. Laggera 54. Pluchea 55. Pseudoblepharispermum 56. Sphaeranthus 57. Litogyne Gnaphalieae 58. Athrixia 59. Macowania 60. Phagnalon 61. Helichrysum 62. Pseudognaphalium 63. Chiliocephalum 64. Gnaphalium 65. Filago 6 6 . Bracteantha Calenduleae 67. Osteospermum 6 8 . Calendula Astereae 69. Ceruana 70. Grangea 71. Dichrocephala 72. Psiadia

73. Microglossa 74. Felicia 75. Symphyotrichum 76. Erigeron 77. Conyza 78. Nidorella Anthemideae 79. Ursinia 80. Tanacetum 81. Artemisia 82. Dendranthema 83. Santolina 84. Matricaria 85. Cotula 8 6 . Anthemis 87. Argyranthemum 8 8 . Chrysanthemum 89. Leucanthemum Senecioneae 90. Senecio 91. Solanecio 92. Cineraria 93. Emilia 94. Crassocephalum 95. Gynura 96. Mikaniopsis 97. Euryops 98. Kleinia Helenieae 99. Athroisma 100. Belpharispermum 101. Schkuhria 102. Pectis 103. Tagetes 104. Flaveria Heliantheae 105. Eclipta 106. Acmella 107. Spilanthes 108. Zinnia 109. Aspilia 110. Verbesina 111. Melanthera 112. Blainvillea 113. Tithonia 114. Sclerocarpus 115. Helianthus 116. Galinsoga 117. Tridax 118. Acanthospermum 119. Guizotia 120. Enydra

1 2 1 . Sigesbeckia 122. Micractis 123. Bidens 124. Cosmos 125. Dahlia 126. Chrysanthellum 127. Parthenium 128. Ambrosia 129. Xanthium Eupatorieae 130. Mikania 131. Adenostemma 132.Ageratum 133. Stomathanthes 134 Barlettina

3 16 316 3

jg

335 336 33g 339 340 341

342 343 344 346 34g 3 ^q

A Glossary o f Botanical Terms Abbreviations

369

Index to Scientific Names

373

Vernacular Names for Volume 4 , Part

35

2

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393

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 2

Editorial Team Prof. Inga H edberg, D ep artm en t o f System atic B otany, U p p sala U n iv ersity , U ppsala, Sw eden. E m eritu s Prof. O lov H edberg, D ep artm en t o f S ystem atic B otany, U ppsala U n iv ersity , U ppsala, Sw eden. M s Eva Persson D epartm ent o f S ystem atic B otany, U ppsala U niversity, U ppsala, Sw eden. Prof. Ib Friis, B otanical M useum and L ibrary, U n iv ersity o f C openhagen, C openhagen, D enm ark. P rof. Sebsebe D em issew , T he N atio n al H erbarium , B iology D epartm ent F acu lty o f S cience, A d d is A b ab a U niversity, A ddis A baba, E thiopia. D r E nserm u K elbessa T he N a tio n a l H erbarium , B iology D epartm ent F aculty o f S cience, A ddis A baba U niversity, A ddis A baba, E thiopia. M s Sue E dw ards T he N atio n al H erbarium , B iology D epartm ent F aculty o f S cience, A ddis A baba U niversity, A ddis A baba, E thiopia. M r M elaku W ondafrash T he N atio n al H erbarium , B iology D epartm ent F aculty o f S cience, A d d is A baba U niversity, A ddis A baba, E thiopia. M r Solom on K assie T he N atio n al H erbarium , B iology D ep artm en t F aculty o f S cience, A d d is A baba U niversity, A ddis A baba, E thiopia.

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 2 Editorial Board D r T ew olde B erhan G ebre E gziabher, D .Sc. H onoris C ausa, D irector G en eral, E nvironm ental P rotection A uthority, A ddis A baba, E thiopia E m eritu s Prof. O lov H edberg, D ep artm en t o f S ystem atic B otany, U p p sala U niversity, U ppsala, Sw eden. Prof. Sebsebe D em issew , K eeper, T he N atio n al H erbarium , B iology D epartm ent F aculty o f Science, A ddis A b ab a U niversity, A ddis A baba, E thiopia. D r M esfln T adesse, (alternate m em ber) C ollege o f B iological Sciences, D ept, o f Plant B iology, T he O hio U niversity, C olum bus, U SA form erly: T he N atio n al H erbarium , B iology D epartm ent F acu lty o f S cience, A ddis A baba U niversity, A ddis A baba, E thiopia. D r E n serm u K elbessa, (alternate m em ber) C urator, T he N atio n al H erbarium , B iology D epartm ent F aculty o f Science, A d d is A b ab a U niversity, A ddis A baba, E thiopia. Prof. Ib Friis, B otanical M useum and L ibrary, U n iv ersity o f C o p enhagen, C openhagen, D enm ark. Prof. Inga H edberg, D ep artm en t o f S ystem atic B otany, U p p sala U n iv ersity , U ppsala, Sw eden. M s Sue E dw ards T he N atio n al H erbarium , B iology D ep artm en t F acu lty o f Science, A d d is A b ab a U niversity, A ddis A baba, E thiopia.

FOREWORD tie present volume is the second part of Volume 4 o f Flora o f Ethiopia and Eritrea, and treats the family A :teraceae (Compositae). This family has been the subject o f many collaborative studies. Chemical and molecular data have relatively recently been added to the morphological data gathered about members o f the family since 1816. Information on the biology and chemistry o f the family was compiled for the first time in 975 and published in 1977 [The Biology and Chemistry o f the Compositae, Heywood et al. (eds.)]. Advances . ; Compositae systematics was published in 1994 (Hind et al .) .The author o f the present volume has consulted d utilized the above and other recent publications. The taxonomic account reflects the current situation in steraceae Systematics, which involves changes in the circumscription o f many species, genera and even bes, triggered by new information obtained from studies on DNA and applications o f newer methods. 35 : neranot recorded in Cufodontis’ Enumeratio are here substantiated, 12 genera are reduced to synonymy, and genera are excluded since they occur only outside o f the Flora area. Many new records represent recent Elections o f naturalized and/or introduced weedy taxa, fewer represent newly discovered wild plants. The Flora Project was initiated in 1980 and the volumes so far published are numbers 2, 3, 4(1), 6 and 7. v olume 3 provides background information about the writing of a modem Flora, notes on the collecting and d entification o f plants, and information on the format used for presenting families, genera and species. The tader is referred to that volume for information dealing with these matters. Arrangements to make the Flora available on CD-ROM are under way, but for obvious reasons it is sential that the work also remains available in printed form.

I.H., I.F.,M .T.,S.E. October 2004

ACKNOWLEDGEMENTS The production of Flora o f Ethiopia and Eritrea enjoys the support o f many institutions and individuals, both* within and outside Ethiopia. The editors particularly wish to emphasize the contributions o f the following, all of whom are gratefully acknowledged for the support they have rendered to the project: The Ethiopian Government through the Ethiopian Science and Technology Commission, in particular the Commissioners, Ato Asrat Bulbula and Ato Mulugeta Amha, and the Swedish Government through the Swedish Internationa] Development Co-operation Agency (Sida/SAREC), which has provided the financial support for the preparation and publication o f all volumes o f the Flora including this one; the past and present officers of Addis Ababa University, especially the Office of the President and its current President, Prof Andreas Eshete, the current Vice Presidents, Ato Mohammed Habib (VP for Business and Development), Prof Endashaw Bekele, Associate VP for the Research and Graduate Programme, the Research Director, Dr Shiferaw Taye, the past and present Deans of the Faculty o f Science, especially the present Dean, Dr Gezahegn Yirgu, and the past and present Heads o f the Department o f Biology, especially the present Head, Dr Kifle Dagne. This volume would not have been prepared without the support and assistance of the curators, keepers and directors o f the following herbaria: Addis Abeba (ETH), Copenhagen (C), Florence (FT and FI), London (BM. K), Missouri (MO), Nairobi (EA), Paris (P), Stockholm (S), Stuttgart (STU), Uppsala (UPS), Wageningen (WAG), Vienna (W, WU) and Zurich (Z). Thanks are especially due to the Royal Botanic Gardens, Kew, England, particularly the past and present Directors, Prof. G. Prance and Prof. P. Crane, and the past and present Keepers of the Kew Herbarium, Prof. G. Lucas and Prof. S. Owens, and their scientific and technical staff Thanks are also due to the Botanical Museum and Library o f the University o f Copenhagen, to the Carlsberg Foundation for supporting extensive field work, which has contributed much to improving the coverage of species and their distributions in the present volume, and to the staff o f the Biology Library, Uppsala University, for their indefatigable assistance in tracing relevant literature and titles. Thanks to the courtesy o f various authorities, editors and editorial boards, it has been possible to use illustrations already published in a number of botanical publications. The following authorities and sources a it gratefully acknowledged: TTie Linnaean Society o f London (Botanical Journal o f the Linnaean Society) Botanical survey of India (Flora o f India), Botanischer Garten und Botanisches Museum Berlin-Dahlem (Englera)\ Botanische Jahrbiicher fu r Systematik; Mitteilungen der Botanischen Staatssammlung Mimchen. Danish Botanical Society (Botanisk Tidsskrift)', Field Museum o f Natural History, Chicago (Fieldiana); Flora de la Province de Jujuy, Flora o f West Tropical Africa; Flora Zambesiaca Managing Committee {Flora Zambesiaca): Laboratoire de Phanerogamie, Paris (Flore de M adagascar et des Comores, Flore des Mascareignes); The Missouri Botanical Garden Press (Annals o f the Missouri Botanical Garden)\ Mitteilungen aus dem Institut fu r allgemeine Botanik in Hamburg; National Botanic Garden o f Belgium (Bulletin du Jardin botanique a Bruxelles, Flore d'A fhque Centrale, Flore des Spermatophytes du Parc National Albert): The New York Botanical Garden Press (Bnttonia, Intermountain Flora), Nordic Journal o f Botany, Norwegian Botanical Society (Norwegian Journal o f Botany)-, Opera Botanica: Oklahoma State University Press (Families o f Flowering Plants in Ethiopia); Oxford University Press (Upland Kenya Wild Flowers)', Publications Scientifiques du Museum National d’Histoire Naturelle, Paris (Adansonia), Royal Botanic Gardens, Kew (Advances in Compositae Systematics, Flora o f East Tropical Africa, H ooker’s leones Plantarum, Journal o f Botany, Kew Bulletin)', Springer-Verlag (Plant Systematics and Evolution)', Strauss & Cramer (Phanerogamarum Monographie): Students' Flora o f Egypt; Symbolae Botanicae Upsalienses: The Israel Academy of Sciences and Humanities (Flora Palaestina), The Jepson Herbarium (The Jepson M anual Higher plants o f California)', The Weed Flora o f Egypt', University o f California Press (University o f California Publications in Botany)-, University o f Hawaii Press (Manual o f the Flowering Plants o f Hawaii)', University o f Michigan Press (Flora Novo Galiciana), University o f Natal Press (Compositae in Natat)\ University of Washington Press (Vascular plants o f the Pacific Northwest). Over the years many institutions and a number o f individuals have assisted in various ways. For discussions and valuable inputthe author would like to thank Mr. C. Jeffrey. Dr. N. Hind, Dr. H. Beentje, Dr. G. Pope, Dr. T. Stuessy and Dr. D. J Crawford. Efforts were also made to stud\ and understand the chemistry o f

I u ts o f the Asteraceae and the assistance of Prof. Berhanu Abegaz and Prof. Ermias Dagne (Addis Abeba diversity) is gratefully acknowledged. The contributions by Dr. S. Ortiz {Dicoma), Prof. I. Friis (Goniocaulon and Asteraceae nomenclature), 1r. D. Otieno (Senecio), and Dr. T. Eriksson (Athroisma and Blepharispermum) are much appreciated. Since moving to Ohio State University in October 1993, the author has enjoyed the support o f the >apartments o f Plant Biology and Evolution, Ecology and Organismal Biology, where the final pages o f the ■steraceae manuscript were completed. We would like to thank Prof. R. Boemer, Prof. D. J. Crawford and Dr. V. Freudenstein o f EEOB for this support.

I.H., IF ., M.T., S.E. October 2004

Tribes and genera of the Compositae in the Flora area The sequence o f genera follows the classification o f the Asteraceae/ Compositae by Bremer (1994) with the M utisieae appearing first and the Eupatorieae last (see under the Mutisieae for a more recent attempt at classification o f the Asteraceae); 134 genera with c 473 species are included. 36 o f the genera are introduced (* - below and in text), especially from the M editerranean area, Mexico/Central America or Peru, 12 are naturalized (•* - below and in text). A few genera are endemic (+) in the Flora area or in NE Africa. The known number o f species is given in parentheses.

Tribe 1 . Mutisieae 1. Tarchonanthus ( 1) 2. Gerbera (3) 3. Dicoma (4) Tribe 2 . Cynareae 4. Echinops (12) 5. Carduus (5) 6 . Cirsium (5) 7. Cynara* ( 1) 8 . Silybum* (2) 9. Ochrocephala ( 1) 1 0 . Plectocephalus ( 1 ) + 1 1 . Volutaria (2) 1 2 . Carthamus* (2 ) 13. Centaurea (4) 14. Goniocaulon ( 1) 15. Notobasis* ( 1) Tribe 3. Cichorieae 16. Scolymus* ( 1) 17. Cichorium* (2) 18. Crepis (7) 19. Dianthoseris ( 1) + 2 0 . Taraxacum* ( 1 ) 2 1 . Lactuca (4) 2 2 . Prenanthes (1) 23. Launaea (11) 24. Reichardia** ( 1 ) 25. Sonchus(9) 26. Tolpis (1) 27. Helminthotheca** ( 1 ) 28. Picris (2) 29. Tragopogon* ( 1) Tribe 4. Vernonieae 30. Vemonia (49) 31. Distephanus (2) 32. Gutenbergia (3) 33. Erlangea (2) 34. Bothriocline (1) 35. Ethulia (2) 36. Elephantopus* ( 1) Tribe 5. Arctoteae 37. Arctotis* (1) 38. Haplocarpha (3) 39. Berkheya (2) 40. Gazania* (?2) 41. Hirpicium (2) Tribe 6 . Inuleae 42. Anisopappus (2) 43. Geigeria (2) 44. Inula (4)

45. Pentanema* ( 1) 46. Blumea (3) 47. Pegolettia ( 1) 48. Pulicaria ( 11 ) 49. Iphiona ( 1) 50. Iphionopsis ( 1) Tribe 7. Plucheeae 51. Delamerea ( 1) + 52. Pseudoconyza** ( 1) 53. Laggera (6 ) 54. Pluchea (6 ) 55. Pseudoblepharispermum ( 1) + 56. Sphaeranthus (8 ) 57. Litogyne (?l)

Tribe 8 . Gnaphalieae 58. Athrixia(l) 59. Macowania (2) 60. Phagnalon (5) 61. Helichrysum (23 ) 62. Pseudognaphalium ( 5 ) 63. Chiliocephalum (2 ) + 64. Gnaphalium (4) 65. Filago (2) 6 6 . Bracteantha* ( 1) Tribe 9. Calenduleae 67. Osteospermum (2) 6 8 . Calendula* (2 ) Tribe 1 0 . Astereae 69. Ceruana* ( 1) 70. Grangea* ( 1) 71. Dichrocephala (3) 72. Psiadia (2). 73. Microglossa (2 ) 74. Felicia (2) 75. Symphyotrichum* ( 1) 76. Erigeron ( 1) 77. Conyza (25) 78. Midorella (2 ) Tribe 11. Anthem ideae 79. Ursinia ( 1) 80. Tanacetum* (2 ) 81. Artemisia (4) 82. Dendranthema * ( 1 ) 83. Santolina* ( 1) 84. Matricaria* ( 1) 85. Cotula (3) 8 6 . Anthemis ( 1 ) 87. Argyranthemum* (2) 8 8 . Chrysanthemum* (2 ) 89. Leucanthemum* ( 1 )

Tribe 12. Senecioneae 90. Senecio (25) 91. Solanecio (6 ) 92. Cineraria (3) 93. Emilia (12) 94. Crassocephalum (5) 95. Gynura ( 1 ) 96. Mikaniopsis (I) 97. Euryops (3) 98. Kleinia (16) Tribe 13. Hdenieae 99. Athroisma (3) 1 0 0 . Bleparispermum (4) 1 0 1 . Schkuhria* ( 1 ) 102. Pectis* (? 1) 103. Tagetes** (3) 104. Flaveria** (2) Tribe 14. Hetiantheae 105. Eclipta** ( 1) 106. Acmella** (1) 107. Spilanthes ( 1) 108. Zinnia* (2 ) 109. Aspilia (5) 110. Verbesina* ( 1 ) 1 1 1 . Melanthera (2 ) 112. Biainvillea** (I) 113. Tithonia* ( 1) 114. Sclerocarpus ( 1 ) 115. Helianthus* ( 1 ) 116. Galinsoga** (2) 117. Tridax* (1) 118. Acanthospermum ( 1) 119. Guizotia(6) + 120. Enydra{2) 1 2 1 . Sigesbeckia ( 1 ) 1 2 2 . Micractis (2 ) 123. Bidens (20) 124. Cosmos* (2 ) 125. Dahlia* (3) 126. Chrysanthellum* ( 1 ) 127. Parthenium** ( 1 ) 128. Ambrosia* ( 1 ) 129. Xanthium** (2 ) Tribe 15. Eupatorieae 130. Mikania (2 ) 131. Adenostemma (3) 132. Ageratum** (2) 133. Stomathanthes (2 ) 134. Bartlettina* ( 1)

ASTERACEAE (COMPOSITAE) by Mesfin Tadesse* (except as specified) '»alpers, G.G. (ed.), Repertorium botanices systematicae 2: 945-955, (1843) & 6: 88-366, (1846) [in which names roposed by C.H. Schultz Bipontinus are validated]; A. Richard: Tent. FI. Abyss. (1848); Schweinfurth, G. & Ascherson, P. Beitrage zur Flora von Aethiopiens, I, (1867) [in which names proposed by C.H. Schultz Bipontinus re validated]; Oliver, D. & Hiern, W.P. FI. Trop. A f r 3: 253-461 (1877); Cufodontis, Enum.: 1062-1192 (1967); effrey, C., Kew Bull. 21: 177-223 (1967), Kew Bull. 22: 107-140 (1968),t e w Bull. 41: 873-943 (1986), Kew Bull. 43: 195-277 (1988); Hind, D.J.N., Jeffrey, C. and Scott, A.J. (eds.), Composees, FI. Mascareignes 109: 1-113 1993); Pope, G. C om positae, FI. Zamb., 6(1) (1992); Bremer, K. A steraceae - Cladistics and Classification, 1994); Panero, J.-L. & Funk, V.A., Proc. Biol. Soc. Washington 115: 909-922 (2002). Herbs or shrubs, less com m only trees or clim bers; som etim es with milky latex. Leaves alternate, som etim es opposite or w horled, basal or cauline, sim ple to com pound, blade com m only 1-veined, rarely 3-5-veined rom base, sessile-auriculate or decurrent or petiolate, w ithout stipules. Flow ers small (often referred to placed Dicoma in a new tribe. Dicomeae Panero & V A. Funk, and Tarchonanthus in the Tarchonantheae. nit they did not indicate to which tribe Gerbera should be placed, although thc> rccogni/.cd a third tribe. Cardueae within the new subfamily they erected, the Carduoideae. Until this newly proposed classification of the Asteraceae provided fully indicating the placement of all genera ithin tribes, the Muiiseae is here applied in its traditional and broadest sense.

T. cam phoratus L. (1753) type: "Habitat in Aethiopia.” South Africa. Herb. Cliff. 398. 'Tarchonanthus no. 1' [BM lecto.. selected bv Andcrberg in Jarvis & al. (ed.). Regnum Veg. 127: 92(1993)1. T. abyssinicus Sclnvcinf. & Asch.; Beitr. FI. A e th io p 287 (1867). nom. nud.

Plants dioecious, unisexual; shrubs or trees: pappus absent. 1. Tarchonanthus Plants monoecious, with both unisexual and bisex­ ual florets, perennial or annual herbs from thick orwood\ rootstock: pappus present. 2 Plants stcmlcss. scapigcrous: phyllarics herba­ ceous. not spine-tipped. 2. G erbera Plants with evident stems, not scapigcrous; phyllarics chartaceous to coriaceous, often pun­ gent. 3. Dicoma

1 TARCHONANTHUS L. (1753) H. Bcentje in Kew Bull 54: 81 95 (1999).

Trees or shrubs, dioecious, often aromatic. Leaves simple, alternate, petiolate or subsessilc. entire, subcoriaceous to coriaccous. prominent beneath, tomentosc to densely vvoolK below, midrib and lateral ribs depressed above. Capitula in terminal or axillary pamclcs. 1- to fewflowered. densely woolly, unisexual. Receptacle convex or flat, epaleate. usually with long silky hairs. Phyllaries 1-3-scnate. imbricate, free or slightly connate below, broader in female capitula. Male florets up to 20 per capitulum; corolla tubular or funnel-shaped, much longer than those of the female florets. 5-lobcd. lobes recurved, villous or woolly externally, often glandular; anthers exsertcd. connective lanceolate, base sagittate and tailed, filaments glabrous; ovary abortive, style exserted. undi­ vided or slightly 2-dentate at apex: pappus absent. Female florets solitary or 2 or 3 per capitulum: corolla cylindric.

A genus with 2 species in tropical and southern Africa and Arabia. Only one species in the Flora area.

Strongly aromatic (with smell of camphor) shrub or tree up to 8 m high. Stem up to 40 cm wide at base; young branches grey or brown-tomcntellous. Leaves narrowly or broadly oblong-elliptic to narrowly obovate. (1-) 1.5-8 (-10) x (0.6—) 1—3(— 4) cm (larger in coppice shoots), coriaccous to subcoriaceous, subsessilc to shortly petiolate. tomentellous, shallowly canaliculate above, glabrcsccnt and green above with tessellate reticulation and minutely glandular dots, whitish tomentellous be­ neath. hairs interspersed with minute golden glands; base cuneatc: margins entire, rarely dentate (in young leaves), apex obtuse to sub-mucronulate; petiole 1-5 nun long, shallowh canaliculate above. Inflorescence terminal, 10-25 x 5-15 cm. florets cream-white, aromatic. Male capitula: 4-5 x 4-7 mm. campanulatc to urccolate. often 10-20-flowercd: phy llaries 1-2-seriate. free, linear, 1-3 x 1-2 mm. densely w oolly outside, glabrous inside; corolla tubular to funnel-shaped, woolly on outside. 2.5-3.8 mm long: anthers 1.7-2 111111 long; ovaiy abortive, c 0.4 mm long, style c 5 111111 long. Female capitula: 3-5 x 3-4 mm. elliptic or obovate. (3-)4-5-flowered; phyllaries 2seriatc. free or slightly connate at base. 4—4.5 x 1.5-3 min, linear-obovate to obovatc-oblong. densely woolly on out­ side orciliatc in the upper half only; corolla tubular, pilose externally. 1.2-2 111111 long: ovary 2-2.5 x 1 nun. woolly, style 1.7-3 111111 long, stigmatic branches glabrous. Cypsclas elliptic, with shiny and wavy liairs. (1.5—)3—4 x 1.2-1.5 nun. Fig. 145.1. Open woodland orbushland with Combretum. Acacia. ('ussonia and Pappaea. usually in derived vegetation re­ placing Juniper us Olea drv evergreen forest; 1300-1950 (-2500) 111. EW TU TU/GD BA SD; dirough E Africa south to S Africa (Natal. Cape), west to Angola: Saudi Arabia. B u rg e r 1871; F riis cl al. 7 7 9 ; M o o n e y 5547.

2. GERBERA L. (1753). nom. cons. H.V. Hansen in Opera Hot. 78: 1 36 (1985) & Nordic J. Bol. 9 469 485 (1990).

Perennial herbs from a spreading, slightly thickened, silky-pilose or velvety rootstock and with l-4(-7) scapes, liairs whitish or brownish. Leaves in basal rosettes, spreading or ascending, sessile or petiolate. alternate, sim­ ple to deeply pinnatifid. petiole terete to dorsiventrally

8

145. A SIERA C EA E: 1. Tarchonanthus, 2. Gerbcra

Figure 1.

TARCHONANTHUS CAM PHOR4TUS: 1 flowering branch x Vy, 2 female capitulum x 4; 3 - male capitulum x 4; 4 female floret x 8; 5 - male floret x 8. 1 ,2 & 4 from Hansen 3 141; 3 & 5 from Norrgrann 160. Drawn by Eleanor Catherine. (M odi­ fied and reproduced with permission from FI Zamb. vol. 6(1), Tab. 2.)

flattened. Scapes naked or bracteate. terete, striate, th in ly to densely toinentose, sometimes inflated near the recep­ tacle. Capitula radiate, heterogamous, solitary, erect. Re­ ceptacle flat, convex, epaleate. honey-combed. Phyllaries 2-3-seriate. gradually elongating inwards, lanceolate to narrowly elliptic, greenish or brownish, often purpletinged. pilose on outside, glabrous on inside, margins scarious. Ray florets female. 1-2-seriate, bilabiate, tube narrow , dorsiventrally compressed. Outer series radiate, lower limb lanceolate to oblong-elliptic. 3-toothed at apex, upper limb of two narrowly linear, often curled, lobes, style terete, branch short. Inner series not much protruding, slender, lower limb short, lanceolate to ob­ long-elliptic, upper limb of two small linear segments.

sometimes with stanunodes. style long. Disc florets bisex­ ual, lower limb of corolla broadly elliptic, 3-fid at apex, upper limb of 2, usually curled, linear to narrowly elliptic lobes Anthers glabrous, brown, connective narrowly ovate. Style terete, arms short oblong, papillose. Cypselas all fertile or those of the central disc florets sterile, brown­ ish. dorsiventrally compressed, elliptic, tapering or beaked at the apex, 5-10-ribbed, pilose, rarely subglabrous to gla­ brous Pappus of many-barbellate bristles in about two rows, wliitish, rarely brownish or purplish. A genus of about 13 species found mainly in South Af­ rica w'ith a few species also occurring in South-Central. East and mid-West Africa (one species), and Asia. Two indigenous and one cultivated species in the Flora area.

145. ASTERACEAE: 2. Gerbera

9

Figure 2. GERBERA JA M E SO N II 1 habit x 2/.v, 2 - outer phyllary x 514; 3 in­ ner phyllarv x 5'/2; 4 - pappus bristle x 11 Vi; 5 ray floret x 2/.v. 6 - ray floret without cypsela x 2 l/y. 7 disc floret, without ovary 5'-4; 8 - tailed anthers x Th\ 9 - style o f disc floret x 7 l/2'. 10 cypsela x 4; 11 - longitudinal section o f a capitulum x 7. All from Vaughan 10346. Drawn by Fat Halliday. (Reproduced from FI. M ascareignes. 109. C om posees, PI. 1.)

Scapes wider below the receptacle, cypselas with long beak, pilose, with inflated hairs; pappus silky, wliitish or brownish. 1. G. pilosclloides Scapes more or less uniform, not wider below the receptacle; cypselas shortly beaked, liairs. when present, slender, tapering; pappus not silky, whit­ ish. sometimes violet-purple at least at apex. 2 Lca\ es pinnatifid, up to 70 cm long; capitula large, up to 4-5 cm w ide; ray florets up to 5 cm long. 2. Leaves simple, dentate or undulate, only up to 45 cm long; capitula up to 3 cm wide; ray florets up to 2 cm long. 3. G. viridifolia G. pilosclloides (L.) ('ass. (1820); Arnica piloselloides L. (1760) - type; "Habitat [in Aethiopia.j” South Africa. Oldenland s.n. in Herb. Burman s.n. (G-Herb. Bunnan). selected bv Hansen in Opera Bot. 78: 19 (1985).

A. hirsuta Forssk. (1775); Piloselloides hirsuta (Forssk.) Cufod. (1967) - type: Yemen. Forsskal in Herb. Forssk. 958 (C holo.. BM iso ). Gerbera schimperi Hochsl. (1841), nom. nud.. G. schimperi Sell. Bip. (1844) - type: TU, Mt. Scholoda. PI. abyss. I, Schiniper 185 (?P holo., BM K iso.). Perennial herb with 1-3(—7), usually 20-60 cm long, tomentose scapes, from a slightly thickened rootstock cov­ ered with light brownish, silky-villose hairs. Leaves sim­ G.orjamesonii ple. oblong-elliptic ovate, alternate. 4-12 x 1.0-5.7 cm, sessile to petiolate, sometimes decurrent and forming nar­ row wings on scape, bright or pale green above with appressed, scattered or dense light or golden-brown liairs. lower surface pale grey, usually with denser cob-webby whitish liairs intermingled with golden brown liairs, mar­ gins entire or obscurely wavy; petiole up to 14 cm long, pilose to villose. with whitish or brownish hairs. Capitula solitary', 12—35(—45) x 12-25 mm at anthesis. scape densely tomentose and broadened (6-12 mm wide) near

—

10

145. ASTERACEAE: 2 Gerbera. 3. Dicoma

base of receptacle, hairs golden-brown. Receptacle flat or convex, ebracteate. Involucre hemispheric Phyllaries 4-25 x 1-3 mm. graded monomorphic. pale green, pur­ plish-red at apex. Ray florets 2-senate, numerous, or­ ange-red. white or pink tinged, rarely yellow tsh or pale purplish. 9-30 x 1-2 mm. female; outer series longer and broader, tube 3-10 mm long, lower limb 3-10 mm long, upper limb 2-5 mm long; inner scries few. slender, tube 4-10 mm long, limb 1-5 mm long. Disc florets yellow or white, rarely tinged pink, tube 5-10 mm long, lower limb 1-3.5 mm long, upper limb 1-3 mm long. Anthers 2-2.5 mm long. Style bifurcate. 10-12 mm long. Cypselas 6-20 x 1.2-1.8 111111 (incl. beak), body 4-9 mm long, beak 5-10 mm long, papillose to glabrous. Pappus white or pale brownish-white. 9-10 nun long, barbellate. Fig. 145.3.5 & 6. Upland or montane grassland with scattered Acacia trees or patches of evergreen bushiand w ith Rosa. Osyris, Euclea. etc.. often on steep, grassy hillsides with Aloe. Becium. etc.. among rocks in dry Juniperus- Hagenia for­ est; 1700-3300 m. EWTU GD GJ SU WG ILGG SD BA HA; Somalia. E Africa. Burundi. Zambia. Zimbabwe. South Afnca (Cape region). Afesfm T. et al. 2880; Moonev 7865; W. de Wilde 6441. 2 G. jamesonii Bolus ex Adlam (1888) type: South Afnca. Natal. Adlam. s.n (Hcib no 40170) (SAM ho Jo). Perennial licit? w ith one to several, crcct. 13-70 cm long. 2-5 111111 wide scape, from a fasciclcd rootstock crowned with brownish feltv orvillosc tomentum. Leaves simple to pinnatifid. alternate, spreading. 15— 45(—55) x (l-)4-14 cm. pctiolate: petiole 2-15(-40) cm long, flattened or grooved at base, blade 9-40 111111 long, upper surface dark-green, lower surface shinv-green. thinly tomentose to glabrous, terminal leaf segment tnangular. margins sinuate to runcinately pinnatifid. Capitula solitary, large, 20-50 x 10-20 111111 at anthesis. scape narrowed at base of rcccptaclc. Involucre campanulate. Phyllaries 4—17(—21) x 1-3 mm. apex long, acuminate. Ray florets severalscnate. yellow, orange, orange-red. red. white or pink on upper surface, low ersurfacc paler, female. Outcnnost flo­ rets: tube 4 - 9 111111 long, low cr limb 15-40 mm long, upper limb 2—t mm long. Inner florets: tube 3-7 mm long. lower limb 2-3 111111 long, upper limb 1-3 mm long. Disc florets: corolla tubular. 3-8 111111 long, lower limb 1-3 nun long, upper limb 0 8-2 111111 long. Cypselas 6—13 x 1.5-2.5 nun (incl beak), pilose. Pappus creamy-white. Fig. 145.2. Cultivated in gardens and as an outdoor pot plant; 2300-2440 111. EW SU; native of southern Afnca but in­ troduced as a garden ornamental elsew here. Afeworki & Me hrel 25. 3 G. viridifolia (DC.) Sch. Bip. (1844); Las1opus viridifolius DC. (1838) - type: S Africa. Keiskama & Buffelnver. Drege 5110 (G-DC lecto.). Gerbera abyssimca Sch. Bip (1844); Perdicium abvssinicum (Sch. Bip.) Hiem (1898)- type: TU. Mt.

Scholoda. PI. abyss. I, Schimper 193 (P holo.. BM BR K iso ). subsp viridifolia Perennial herb with 1^4, usually 10-70 cm long, 1-3 mm wide scapes from a fleshy or thickened, silky-pilose rootstock; liairs white, silky or dark, both types often found together on scapes, leaves and phyllaries. Leaves simple or lobed. alternate. 5-45 x 1.5-10 cm. sessile to petiolate. petiole 0 5-25 x 1-3 mm, up to 10 111m wide at base, blade elliptic, oblong-lanceolate to ovate-lanceo­ late. base often cuncate. sometimes cordate, rarely trun­ cate to decurrcnt. upper surface green, long silky-pilose (espccialh in young ones), strigose. sometimes velvety to totalh glabrous, low cr surface thinly tomentose. margins cihate. entire or shallow ly lobed to crenate. Capitula radi­ ate. vanable in size. 12-40 x 10-22 mm. Receptacle flat. Involucre obconic Ray florets pinkish w hite above and below , or pink. red. or purple below , outer series: tube 3-6 mm long, lower limb 8-24 x 2—3.5 111111. upper limb 1-4 mm long; style 1.5-4.5 mm long, inner series: tube 3-7 mm long, lower limb 1-4 mm long, upper limb 1-3 nun long, style 1-4 111111 long. Disc florets white, yellow or green-tipped, rarely violet, stigma yellow Corolla (3—)4—8 mm long. Cypselas beaked. (4-) 6-12 x 1.5-2.5 mm. pilose to subglabrous. beak 2-5 nun long. Pappus pale white to brownish. Fig. 145.3.1—4. In dry open woodland, rocky grassland, frequently in the ecotone between open grassland and dry forest or among scattered bushes; 1350-2400 m; EW TU GD SU WG KF SD BA; Sudan. Somalia. E Africa, Cameroun. Rep Deni du Congo. Malaw i, Zambia. Zimbabwe. Mo­ zambique. Swaziland Lesotho. South Africa. Mesfin T. 8270; Friis et al. 3751; Mooney 5508 G. viridifolia subsp. natalensis (Sch. Bip.) H.V. Hansen occurs in Swaziland and S Africa.

3. DICOMA Cass. (1817)> Hochstetteria DC. (1838); Schajfnera Sch. Bip. (1841). nom. nud. S Ortiz, .1 Rodrigucz-Dubina & Mesfin Tadesse 111 Ann. Miss. Hoi. Card 85: 440-459(1998). Annual or perennial herbs. Stem simple to branched, senccous to lanate Leaves simple, alternate, sessile, often w ith a pseudopetiolc. margins entire to serrulate. Capitula obconic to campanulate. discoid and homogamous (radi­ ate and heterogamous outside the Flora area). Phyllaries in many rows, acuminate to pungent, coriaceous. Receptacular paleae absent. Corollas white, cream, yel­ lowish or lilac, actinomorphic, deeply 5-lobed. with subsessile or long glandular liairs. Anthers with lanceolate, acute to acuminate apical appendages and long tapenng, retrorse-pilose tails. Style-branches not separating, with 1 bv Santiago Ortiz, Laboratorio de Botamca, Facultade de nmacia. Universidade de Santiago, 15706 Santiago de C umpostela. Galicia. Spain & Mesfin Tadesse, The Ohio Uni­ versity. USA.

145. ASTERACEAE: 2. Gerbera. 3. Dicoma

Figure 3.

GERBERA

IIR ID IE O LIA 1 habit x 2/3; 2 marginal floret, fe­ male. bilabiate with strap-shaped outer lip and 2-lobed inner lip \ 5: 3 ray floret, female, bilabiate \ 5: 4 centra) floret, functionally male, bilabiate, cvpscla rudimentary \ 5 G. PILO SELLOID ES 5 capitu­ lum. note thickened peduncle \ 2 .v 6 cypsela x 3. 1 from Robinson 1277. 2 4 from Phillips 1X7: 5 from IViehe N/238; 6 from Brumniit X577 Drawn by Eleanor Catherine. (R e­ produced with permission from /*'/ Zarnh., vol. 6(1). Tab. 9.)

m

obtuse apex and short, generally subapieal. acute, sweep­ ing hairs. Cypselas obovoid to turbinate. (5—)8—K). ob­ scurely to prominently ribbed, densely pilose, basal hairs shorter and spreading. Pappus of barbellate or plumose bristles in several row s, with or without one internal row of scales.

Ethiopia but no Ethiopian material of this species has been seen by the authors, and. therefore, it is not included here.

About 50 species in tropical and southern Africa and Madagascar; one species reaches the Arabian Peninsula and another one is found in India and Pakistan.

-

Cufodontis [in Bull Jcird. Bot. Not. Belg. 37, Suppl.: 1178-1179 (1967)] cited D. bangueolensis Buscal. from

1.

2.

Leaves conduplicatc; phyllaries long acuminatepungcnt; pappus dimorphic, the outer series of numerous barbellate bristles, the inner of c 10 scales. 1 D. tomento.sa Leaves not conduplicatc; phyllaries short acu­ minate-pungent; pappus isomorphic. Pappus 1-seriate, of (4 -)8 -10 ( - 11). rigid, flattened bristles. 2. D. schimperi Pappus multiseriate. of more than 10 bristles. 3

12

145. ASTERACEAE: 3. Dicoma

Figure 4. DICOMA TOM ESTO SA 1 - habit x c 14; 2 - capitulum x 2; 3 leaves, often folded partly or all along the mid-vein x c 2 ; 4 - floret excluding ovary x 7; 5 - cypsela with pappus x 4 All from Ash 2106. Drawn by Alfredo L6pez.

3.

-

Leaves 11-13 cm long; phyllaries without median vein, the innermost phyllaries shorter than the ad­ jacent outer series; pappus of 9-10 mm long plumose bristles. 3. D. sessiliflora Leaves up to 9 cm long; phyllaries with a conspicu­ ous median vein, the innermost phyllaries longer than the outer series; pappus of baibellate bristles up to 6 mm long. 4. D. aethiopica

1. D. tomentosa Cass. (1818) - type; from Senegal. Adanson s.n. (P-JU, holo.). D. lanuginosa DC. (1838). D. gnaphaloides Mattei (1908). Schaffnera carduoides Sch. Bip.. nom. nud. Erect annual herb. 15-60 cm tall. Stem ramified above, greyish white, woolly to glabrescent often with purplish bark. Leaves subsessile. 10-80 x 2-16 mm, linear-elliptic

to oblanceolate. discolorous, upper surface thinly cob­ webby to glabrescent lower surface greyish lanate, atten­ uating gradually to a pseudopetiole, margins serrulate, apex mucronate or cuspidate. Capitula obconic-campanulate, 10-20 x 14-20 mm, solitary to many, on peduncles of 1—5(—8) mm, with 0-4 subtending leaves. Phyllaries 30-50. arranged in 5-6 rows, stiff, straight 2-18 x 0.8-3 mm. usually purple-tinged with a darker purple stripe on either side of the midrib, margins sharply semilateciliolate. outermost puberulous, subulate; innermost nar­ rowly triangular, apex long acuminate, pungent. Corollas creamy-white to lilac. 4.5-7 mm long, with subsessile glandular twin hairs, deeply 5-Iobed, lobes linear, erect or recurved at the apex Anthers c 3.5 mm long, apical ap­ pendage c 0.3 mm long, anther tails c 1.2 mm long. Style c 8 mm long, branches 1-1.5 mm long. Cypselas 1 .6 -3 x 1-1.5 m m slightly compressed, with 10 prominent ribs.

145. ASTERACEAE: 3. Dicoma

13

irbinate. densely covered with long ascending hispid::ose liairs especially between the ribs. Pappus diitiorDhic: outermost series of 50-70 barbellate bristles, 5-7.5 nm long: the inner series of c 10 scales. 5-7.5 x 0.5-1 inn. margins broadly scarious, barbellate near the apex. :ig. 145.4. Open Acacia woodland. Acacia Commiphora bush,md. Acacia grassland, roadsides, on sandy, gravelly and Dcky granitic or calcareous soils and lava nows'. 00-2600 in. EE AF EW TU SU GG SD HA: widespread tropical and southern Africa, also in Socotra. Pakistan, ndia. Bally 6786; Gilbert et al. 295; \yslrom 77. 1 D. schimperi (DC) Baill. ex O. Hoffm. (1893); Hochstetteria schimperi DC. (1830) - type: Saudi Arabia, “in montibus ad Vallem Fatmensis”. Schimper 864 (G holo. not seen). ubsp schimperi Irect annual herb, up to 45 cm high; stem branched, iranchcs grevish-white tomentose with sessile to sub-essile glands. Leaves (8-) 15-20 x (2-) 5-10 mm. lanceoaie. elliptic to broadh ovate, pseudopetiolatc-aunculate. margins sermalte. greyish-white tomentose with sessile glands. Capitula several, in open corymbosc cymes. 5-12 . 15-20 nun. obconic-cainpanulate: peduncle 0.5-10 cm long, bracteatc. Phyllaries 4-6- seriate. 40-70 (-100). lin­ ear-deltoid. yellowish-white with a greenish stripe on ei­ ther side of the midrib, rigid with pungent-acuminate apex, glabrcsccnt to tomentose. with broad scarious mar­ gins. 2-10 x 0.3-1.5 mm. all reflcxed in fruit. Corolla greenish-vellow or greenish-white, 5-9 nun long, glandular-hairy. lobes 2.5—3.5 mm long Anthers 3 4-4.5 mm long, appendage 1-1.7 mm long: anther tails 1—1.3 mm long. Style 5-8 mm long, branches 1-15 nun long. Cypsela turbinate. 1.5—2.5 x 1-2.2 mm. crcci-luspid. 8-10 -12)-ribbcd. Pappus bristles. (4-) 8-11. isomorphic, rigid, barbellate. flattened toward the base No habitat oraltitudinal records available for the Flora irca. EW (Mt. Ghedein): Egypt. Sudan. Djibouti, and Saudi Arabia. Terraciano & Pappi 29 (FT!) Subsp. cinerea (S. Ortiz & Rodr.Oubina) S. Orti/. & Rodr.Oubina occurs in northern Somalia. It is distin­ guished by being densely tomentose. with leaves broadh ovate to sub-orbicular with pustullatc teeth and larger api­ cal mucro.

1 igure 5 DICOMA SF SS1U FLORA I habit x » x 2 capitu­ lum x c 1.3 middle phyllary x c 1.4 inner phyllary x c 1. 5 floret and detail o f plumose pappus bristle \ 2 I Irom Salubeni 59: 2 5 from Faulkner Kew 64. Drawn b\ Eleanor Catherine. (Rearranged and reproduced with permission from FI Zarnb. vol. 6(1). Tab 7 p.p.)

3. D. sessiliflora Ilarw (1865) in 10-12 rows, lanceolate-subulate. 6.5-23 x 2.3-5 nun. type: Malawi. Manganja Hills. Metier s.n. (K stramineous, chartaceous. glabrous, with inrollcd mar­ holo). gins in upper part, attenuate-pungent at the apex, outer series without scarious margins, median series with nar­ i.rect perennial herb up to 1 in tall. Stem simple, row scarious margins, innermost phyllaries shorter than iraneose-lanatc. Leaves sessile, narrowly elliptic to the adjacent outer scries, white scarious. apiculatc. Co­ broadly oblanceolate. 11-13 x 1.5-3.7 cm. attenuate at rolla yellowish. 9-10 nun long, with long, glandular tw in lie base, margins serrulate, upper surface green, gla­ hairs, lobes 5-6 mm long, erect, acute. Anther c 9 mm brous. lower surface white araneosc-lanatc. Capitula long, apical appendage 1.5-2 mm long, apex acute not obconic to campanulatc. few. more or less sessile, invo­ apiculate. anther tails c 4.5 mm long. Style c 9 nun long. lucres 30-35 x 30-35 nun. Phyllaries 100-120. arranged

14

145. ASTERACEAE: 3. Dicoma

Figure 6. DICOMA AETHIOPICA 1 habit x Vi; 2 - leaf x 4Vi; 3 - capitulum x 2'/a; 4 - outer phyllary; 5 - floret x 3 '/ 3 . 6 - cypsela with pappus x 3 Vi; 7 - apex o f a pappus bristle. All from Frits et al 3686. Drawn by M. Guitian. (Repro­ duced with permission from Nordic J Bol. 16, 1996, p. 280. fig. 2.)

swollen al the base, branches 2-3 mm long. Cypselas (onl> immature seen) 3 x 2-2.5 inm. obovoid. with 8-10 inconspicuous veins, densely covered w ith long ascend­ ing hispid-setose hairs. Pappus of 70—90 plumose bris­ tles in several rows, the longest bristles 9-10 mm long Fig 145.5.

subsp. stcnophylla Pope in Kew Bull. 46; 701 (1991) - type: Nigeria. Lokoja. Mt. Patti. Dalziel 33 (K

scanous margins, outermost phyllaries erecl-paieni lo erect, seldom reflexed. ■ Combre turn Temimaha woodland; 1500-1800 m. WG; Senegal. Guinea Bissau Ghana, Togo, Nigeria Central Af­ rican Republic W. de Wilde 8941, Friis et al. 9105. 9266. Subsp. sessiliflora with leaves broadly oblanceolate, pin llaries with broadly white scarious margins and outer­ most phyllaries reflexed occurs in Tanzania, Rep. Dem. du Congo, Malawi, Mozambique, Angola and Namibia.

liolo.). Leaves narrow ly elliptic-oblanceolate; outer and median phy Hanes more or less concolorous or with narrow

4. D. aethiopica S. Ortiz & Rodr. Oubiha (1996) type. BA. Sof Omar caves, Friis, Gilbert & Vollesen 3686 (K holo., C ETH iso.).

145. ASTERACEAE: 3. Dicoma, 4. Echinops

irect annual or perennial herb, up to 40 cm tall. Stemramfied, moderately araneose-lanate. Leaves elliptic to iblanceolate, 1.5-9 x 0.2-1.7 cm, margins serrulate, at­ tenuating gradually to a pseudopetiole, green con:olorous, often rugose and slightly to moderately tomenose on both faces. Capitula many, widely cylindrical to campanulate, 8-12 x 12-20 mm, on peduncles of 30-90 rim, with 3-8 small, subtending leaves. Phyllaries 70-100, 4-6-seriate, deltate to lanceolate, acuminatcmngent, 3.5-11 x 0.7-1.7 mm, straw-coloured to dark :urple with a darker stripe on each side of the midrib, gla­ brous to sparsely hairy especially on the outermost •hyllaries. erect to erect-patent. Corolla white, 5-8 mm to ng, covered with subsessile glandular twin liairs, lobes c 3 5 mm long, acute, becoming recurved. Anthers c 3.5 mm long, apical appendage c 1 mm, narrowly apiculate, mthertails 1.3 mm long. Style 6.5-9 mm long, swollen at the base, branches c 2.3 nun long. Cypselas turbinate. 1 5-2.5 x 0.7-1.2 mm, with 8-U> prominent ribs, densely .overed with long ascending hispid-setose liairs espe.ially between the ribs. Pappus of 40-80(-100) barbellate Dristles in 3-4 rows, widening slightly at the tip. the lon­ gest bristles 4-6 mm long. Fig. 145.6. Acacia-Commiphora bushland and open habitats, on imestone with red soil and granite outcrops; 1400-1500 m. SD BA; not known elsewhere. Gilbert et al. 7781. 7870.

Tribe 2. C Y N A R E A E Lam. (1806) C ardueae C ass. (1 8 1 9 ) ^ornual or perennial herbs, rarely small shrubs; stem sim­ ple or branched, often spiny and winged. Leaves alternate, pinnatifid or incised, rarely simple, surfaces densely tomentose or lanate, rarely glabrous; margins dentate, ;pinescent. Capitula solitary or corymbose, homogamous md discoid with linear, usually unequal corolla lobes, or lieterogamous and disciform with marginal florets female or neuter, radiate. Receptacle flat or obconic, usually pa­ leaceous or with long bristles or hairs. Phyllaries coriaceous or scarious, imbricate, multiseriate. Florets yellowish or purplish, rarely white, red or pink. Anthers sagittate or caudate, with entire, irregularly laciniate or marginally pinnatifid auricles. Style near point of bifurca­ tion with a ring of minute liairs or slightly thickened, branches separate or connate, short, obtuse. Cypselas more or less terete or compressed, obovate. not beaked, sometimes uinbonate (in the Carduineae), glabrous or sparsely hairy . Pappus a crown of basally connate multiieriate bnstles, scales or 1-seriate bristles or pales. A tribe with about 80 genera, mainly found in Europe, the Mediterranean region and in Central Asia; 11 genera in the Flora area.

2

Capitula subtended by 5 pinnatifid-spinescent leaves/bracts. 15. Notobasis Capitula not subtended by pinnatifid leaves. 2 Capitula one-flowered, aggregated into globose heads. 4. Echinops

15

Capitula many-flowered, distinct. 3 3. Phyllaries with distinct spiniferous, pectinate or scarious appendages. 4 - Phyllaries without terminal appendages, spiny at apex. 8 4. Phyllary appendage spoon-sliaped, scarioussilvery; leaves white-tomentose beneath; capitula homogamous. 9. Ochrocephala - Phyllary appendage lanceolate, spiniferous or pectinate; leaves green to arachnoid; capitula heterogamous. 5 5 Phyllary appendages pectinate; marginal florets greatly enlarged, conspicuous, pink, white or pur­ ple. 10. Plectocephalus Phyllary appendages spiniferous or pectinate; mar­ ginal florets not or only slightly enlarged 6 6. Leaf margin entire or serrulate. 13. Centau rea - Leaves lobed or pinnatifid. lobes ending in sharp spines. 7. Leaves pale green, mottled white along veins, large (c 10-50 cm long); florets pink to purple. 8. Silybum - Leaves green, concolorous. small (up to 10 cm long); florets yellow to orange. 12. C artham us 8. Pappus of unequal long scales, surrounded bv a low narrow rim; phyllaries not spinescent; annual herbs. 14. Goniocaulon - Pappus of barbellate or plumose hairs; phyllaries spinv at apices and sometimes also on margins; perennial herbs, rarely annual. 9 9. Pappus hairs plumose. 10 Pappus hairs glabrous or barbellate. or absent. 11 10. Capitula small, c 2-3 cm across or high; phyllaries coriaceous with pectinate-ciliate margins. 6. Cirsium - Capitula large, over 5 cm across or high; phyllaries fleshy with smooth entire margins. 7. Cynara 11. Leaves spiny at the margins; plants thistles or thistle-like. 5. C arduus - Leaves not spiny at the margins; plants not thistle-like. ll.V o lu ta ria

4. ECHINOPS L. (1737) C. Jeffrey in Kew Bull. 22: 107-140 (1968), Meslm Tadesse & Barhanu Abegaz in Mitt. hist. All. Bot. Hamburg 23b 605-629 (1990); Mesfin Tadesse in Kew Bull. 52: 889-901 (1997). Spiny perennial herbs, cliamaephytes or tall slrnibs with marcesccnt leaves. Leaves simple to twice-pinnatisect. petiolate or sessile. Capitula single-flowered, aggregated into globose heads (compound capitula or pseudocephalia); each capitulum sessile, w ith a ring of simple or branched extraphyllary bristles and 3^4-seriate phyllaries. Phyllaries imbricate; the outermost smallest, mi­ nutely spathulate above and often pinnately long-ciliate near base; median and inner ones gradually longer, bristh or produced into long spines; innermost phyllaries usually 5. connate or fused, differentiated from the outer scries in

16

145. A S T E R A C E A E 4. Echinops

r\

\

shape (usually linear-lanceolate or oblong) and in the api­ cal part (obtuse, fimbriate or lacimate. 3~fid. etc.). Florets bisexual, white to cream, pale to deep blue, pink to red or puiplish-red, rarely violet. Corolla tubular, actinomorphic. with a slender glandular or glabrous tube. 5-lobed. lobes 5. cut to base, linear. Anthers violet or purplish, nectary big to small, at junction of tube and lobes. Style bifurcate, stigma white or purple. Cypselas oblong or obovate, densely apprcssed-villous. bristles wliite or yel­ low. Pappus coroinform, persistent, of free or connate scaJe-like ciliate bristles. The one-seeded, deciduous capitulum is the unit of dispersal. A genus of about 120 species found in Europe, the Mediterranean region. Centra] Asia, and in North and Tropical Africa: 12 species in the Flora area.

ev

-

Peduncle white-woolly; corolla glabrous; cypselas 7.5-10 mm long. 4

4

Florets deep purplish-red. corolla 3—3.5 cm wide; head 5-10 cm wide; capitulum 4.5-5 cm long; in­ ner phyllaries 2-3 cm long. 2. E. amplexicaulis Florets white, corolla 1—1.3 cm long; head 3-4 cm wide; capitulum 1.5-2 cm long; inner phyllaries 1.2-1.5 cm long I E. hoehneiii

-

5.

Phy llanes (except sometimes the outennost) with hooked bnstles at apex and along margins near apex. I I E. hoehneiii Phyllanes with soft or stiff bristles at margins and apex, bnstles never hooked at apex, or absent.

6.

Corolla red. pink or reddish-brow n. rarely white or violet. 30-45 mm long; capitulum 45-70 nun long; inner phy llaries 20-55 mm long. Corolla wliite. blue lilac or mauve. 10-20 mm long; capitulum 15—30 mm long; inner phyllaries 8-18 mm long. 9

7.

Perennial herb. 1-1.5 (-2) m high; head 6-10 cm wide, sessile or on up to 4 cm long peduncle; phyllaries pectinate-ciliate. surface glabrous. 2. E. amplexicaulis Tall shrubs to small trees: head 10-25 cm wide. long-pedunculate, peduncle 10-45 cm long; phyllaries with 2-7 tufts of ciliate bristles in the upper part, surface hispid.

1 Leaves simple or shallowly lobed. margins coarsely and often irregularly toothed. - Leaves pinnatisect to bipinnatisect, segments lin­ ear or broad, or deeply lobed. 5 2. Straggling perennial herbs, up to 1 rn high; leaves linear-lanceolate, c 6-30 times as long as broad; inner phyllanes 8-10 mm long; corolla 1.5-2 cm long 12. E. longifolius - Erect heibs or shrubs; leaves ovate-elliptic to ovate-lanceolate, less than 6 times as long as broad; inner phyllaries 12-30 mm long; corolla (1.8—)2—3.5 cm long 3

-

8. 3. Peduncle densely hispid, liairs greenish-yellow; corolla pubescent; cy pselas 13-15 mm long. 5. E. giganteus

Figure 7. ECH INO PS spp.: phyllaries (arranged starting from the outermoft on the left hand side) and cypselas. All x c 3 . l - £ LONGIFOLIUS a-e outer, f - inner, g cypsela. a - f fron P u ff & Ensermu 820808 -2 ) 1, g from Mercier 2929. 2 - E. PAPPI. a d outer, e - inner, f - cypsela. From Yehenew M. s.n. 3 - E MACROCHAETUS : a-d: outer, e - inner, f cypsela. a-e from Mesfin 6866, f fron Mercier 2090. 4 E GIGANTEUS a-d: outer, e-f: inner; g - cypsela From Mesfm et al. 3582. Drawn b\ Damtew Teferra. (Reproduced with permission from Mitt Inst. Allg. Bot Hamburg, 23b, 1990. p. 628, fig. 4.)

Peduncle whate-woolly. exposing dark reddishbrown bark; leaf segment linear; cypselas 20-23 mm long; phyllaries 40-58 mm long. 4. E. ellenbeckii

145. ASTERACEAE: 4. Echinops

Peduncle glandular-hairy; leaf segment broadly tri­ angular, ovate or rounded; cypselas 16-18 mm long; phyllaries 20-35 mm long. 3. E. longisetus Peduncle and leaves densely hispid or hispidpilose; outer phyllaries with a tuft of glabrous or 10 ciliate bristles at apex. Peduncle and leaves sparsely or densely arachnoidpubescent, woolly-tomentose, glandular hairy or glabrous; outer pity llaries devoid of apical tufts of setae, merely with soft bristles along margins and ciliolate at margins toward apex. 11 10. Erect herbs, 50-100 cm long, leaf lobes 1 cm or more across; mature cypselas 13-15 mm long. 5. E. giganteus Plant often acaulescent, rarely stem up to 50 cm high; leaf segments about 0.5 cm wide; mature cypselas 9-10 mm long. 6. E. hispidus 11. Head with stiff pungent spines exserted up to 2 cm from surface; corolla 2.5-3 cm long; leaf segment triangular or narrowly so, tapering toward the apex. 12 Head without spines; corolla 2-2.5 cm long; leaf segment narrowly linear, almost parallel-sided. 14 .2. Tall bushy herbs, l-2(-3)m high, peduncle and stem glandular-hairy, not white-woolly; cypselas 5-8 mm long. 7. E. macrochaetus Herbs or shrubs, up to 1 m high; peduncle arachnoid-pubescent; stem sparsely or densely white-woolly with dispersed short purple hairs; cypselas 8-10 mm long. 3. Middle and innermost outer phyllaries dagger­ shaped with cuspidate apex and glabrous or ciliolate collar, some produced into stiff pungent spines, 8—15(—20) mm long; cypselas 8-10 mm long. 9. E. pappii Phyllaries narrowly ovate-lanceolate, with long ta­ pered apex, 15-40 nun long, and with soft mar­ ginal bristles; cypselas unknown. 10. E. buhaitensis 14. Leaves bi- to tri-pinnatisect with narrow (c 3-5 mm wide) segments; phyllaries monomorphic, all with dense, soft bristles marginally, corolla tube glandular-hairy. 11. E. angustilobus - Leaves lobed or pinnatisect with 5-15 mm wide segments; phyllaries dimorphic, inner ones fimbriate at apex; corolla tube glabrous except near base where glandular. 8. E. kebericho 1. E. hoehnelii Schweinf (1892) - type: Tanzania, Kilimanjaro, von H oehnel s.n. (B holo destroyed); Kilimanjaro, Greenway 3814 (K neo.. selected by Lisowski). Shrub, or densely branched herb, up to 1.5 m high; stem dark-brown or blackish, arachnoid-pubescent with sharp riangular ultimate segments, basal or lower cauline leaves up to 45 x 22 cm, middle up to 1 5 x 8 cm, amplexicaul, dark-green above and glabrous, woolly-tomentose beneath, segments with sharp pungent spines. Head 3-4 cm in diam­ eter. Peduncle absent or up to 3 cm long, 3-4 mm wide, white-tomentose exposing dark-purple bark. Capitulum '

17

1.5-2 cm long. Phyllaries in 3-4 series; outermost 4-5 x 0.3-0.5 mm, spathulate with a few marginal bristles; sec­ ond row boat-shaped, dark-puiple on upper half, with usu­ ally 3-5-hooked apical bristles and few marginal ones to­ ward the apex, c 8-10 x 2 mm, narrowly hy aline at margins, inner row similar to second row except in size (c 10-14 mm long); innermost row fused and enclosing cypsela, with many hooked bristles. Corolla white, 10-13 mm long; tube glabrous; anthers purple and stigma reddishpurple at anthesis. Cypselas densely hispid-pilose, c 8 x 3 mm, obovate, bristly hairs yellow, overtopping pappus. Pappus of numerous contiguous, about 1 mm long, ciliolate, narrow connate scales. Fig. 145.8.6. Erica arborea bushland, wet Hagenia-Schefflera, Juniperus-Olea forests, often on stream banks; 2050-3750 m. SU GG BA; E Africa, Rep. Dem. du Congo, Rwanda, Burundi. Ash 1532, 3531; Mesfin T. 5410. 2. E. amplexicaulis Oliv. (1873) -ty p e; Tanzania Usui, Grant 141 (K holo ). E. neumannii O. Hoffm. (1906) - type: GG, Fofa plateau, W of L. Abaya Neumann 201 (B holo. de­ stroyed). E. sidamensis Cufod. (1966) - type; SD, near Yirgalem, Kuls 25 (FR holo., K photo.). Erect unbranched herb, sometimes several stems from the rootstock, (0.5—)1—1.5(—2) m high; lower parts of stem smooth or very sparingly setulose, woolly, upper parts white-cottony; root tuberous. Leaves stiff, ovate or ellip­ tic in outline, sessile, amplexicaul and auriculate; blade with slightly to deeply pinnatifid segments, margins dou­ bly dentate, surface thinly tomentose, scabridulous and light-green above (turning black upon drying), densely white-tomentose beneath, lobes terminating in spines. Head 6-10 cm in diameter, often solitary. Peduncle white-woolly, up to 4 cm long and 3.6 mm wide. Capitulum 4.5-5 cm long. Outer phyllaries linearlanceolate, 8-32 mm long, up to 35 mm long in fruit, 3-4-seriate, reddish brown, pectinate-ciliate, smoothly ta­ pered at apex. Inner phyllaries glabrous, ciliate, spinulous, 20-30 mm long, up to 32 mm long in fruit. Unear-lanceolate or subulate, some similar to outer phyllaries, acuminate at apex. Corolla deep reddish brown or pink, 3-3.5 cm long, tube glabrous; anthers red or red­ dish brown. Cypselas appressed-pilose, hairs yellow, bris­ tly, 7.5-10 x 1.5-2 mm. Pappus of numerous contiguous, narrow, ciliolate, connate bracts. Fig. 145.8.3. Terminalia-Combretum-Cussonia woodland, along edge of forests, dry grassland, in roadside thickets and rarely in open wasteland, found on red clayey or sandy soil overlying granite; 1400—2400 m. GJ SU WG IL KF GG SD BA; Sudan, E Africa. Rwanda Burundi, Nigeria Cameroun. Central African Republic, Rep. Dem. du Congo. Mesfin T. & Kagnew 2081; Mooney 6918; Ash 1483. 3. E. longisetus A Rich. (1848) - type: SU, ‘Choa’, Petit s.n. (P holo.). E. steudneri O. Hoffm. (1898) - type; GD, Mt. Guna, Steudner 268 (B lecto., K isolecto.).

18

145 ASTERACEAE: 4. Echinops

Figure 8. EC H IN O P S spp . phyHanes (arranged starting from the outerm ost on the left hand side) and cypselas. 1 - E H ISPID VS. a-d: outer, e - inner, f - cypscla. From Mesfin T. 3327. 2 - E E LLE N BECKII : a-c: outer, d -e: inner, f cypsela. From Mesfin T. 6980A. 3 - E A M P LEXIC AU LIS : a-d: outer, e-f: in­ ner; g - cypsela. a - f from Mesfin —3000(—3600) m. EW TU GD GJ WU SU AR A HA; Sudan. Mooney 5306; Mesfin T. 729; Evans and •- lenly 135. 8 .E. kebericho Mesfin (1990) - type. SU. 15 km on Tikur Inchini-Shenen road, c 9° 52' N; 37° 42' E. Mesfin T. 6990 (ETH holo., C K UPS iso.). i:rect perennial herb or shrub, up to 1.2 m high, usually : om a massive (c 30-50 x 10-20 cm) rootstock; stem leafy throughout, sparsely arachnoid-pubescent, somei mes mixed with dark-purple 1-seriate hairs. Leaves el­ liptic. amplcxicaul. basal ones up to 25 x 15 cm. pinnati­ sect with segments divided into triangular lobes all ending in sharp pungent spines, or bipinnatisect with linearhmccolate segments; upper surface arachnoid pubescent mainly along median vein, lower surface densely tomen1lose, dull, with I -seriate hairs along midvein. Head 3-4 .m in diameter, spineless. Peduncle up to 5.5 cm long, iraclmoid-pubesccnt, 1 -2 mm wide. Capitulum 2-2.5 cm

21

long. Outer phyllaries 3-4-seriate, outermost row spathu­ late. 4-6 mm long, with a few white, soft bristles along margins in the lower half, middle row spathulate with apiculate apex, 6-10 mm long, with several white, soft bris­ tles on margins toward apex; innermost series lanceolate, tapered at base and at apex, 9-14 mm long. 1 nun wide, with bristles al margins in middle and toward tip; apex acute. Inner phyllaries oblong-linear, fused for up to 3/4 of their length, fimbriate at apex, c 10-13.5 mm long, 1 nun wide, glabrous, ciliate at margins near apex. Corolla white or bright blue, c 12-15 mm long, tube glabrous ex­ cept near base of limb where papillose; anthers light brown. Cypselas oblong-elliptic, 7.5-10.5 x 1.5-2.5 mm. covered with appressed silvery-white ciliolate bristles. Pappus of numerous, contiguous, narrow, ciliolate scales that are connate at base. Fig. 145.11. Montane Acacia woodland, disturbed Gnidia, Carissa, Rhamnus staddo bushland with dense, short grassy undergrowth; 2300-2600 m. TU GJ SU; not known elsewhere. Mesfin T. 886; Evans 350; Lawesson 5001. E. kebericho is variable in habit and in the dissection of the leaf blade. Populations from dry and stony lateritic soils are perennial herbs w ith narrowly dissected leaves arranged on nodes of wide interval along the stem arachnoid-pubescent stems and short (c 15-25 x 7 cm) tu­ berous roots, e.g. Mesfin T. 6995. Those found growing in deep vertisols arc usually low shrubs; their sterns are arachnoid-pubescent with mixed short erect hairs. The leaves are pinnatisect with broad segments, and are ar­ ranged on nodes at short intervals. On tlie roots are mas­ sive tubers (c 30-50 x 10-25 cm). Friis et al. 2264 from KF may represent such a broad-leaved form of this spe­ cies. The chemical compounds isolated from the roots of specimens of the populations from stony lateritic soils and deep vertisols respectively, were identical [cf. Berhanu et al. in Biochem. Syst. Ecol. 19: 323-328 (I9 9 l)|. 9. E. pappii Chiov. (1911)

- type: EW, Baza, Badum. on road to Mereb R., Pappi 6848. 6849, 6852 (FT syn.. K photo). E. boranensis Lanza (1939) - type: SD, Negelle Borenna, Cufodontis 191 (FT holo., W WU iso.). E. spinosus L. var. homoilepis Cliiov. in Ann. R. 1st. Bot. Roma 8: 193 (1904) - type: EW. Amba, Terracciano & Pappi 1324, 1980, 1988 (FT syn.). E. boranensis Lanza var. minor Cufod. in N. Giorn. Bot. Ital., n.s. 50: 115 (1943) - type: SD. Negelle. Corradi 1970 (FT holo. not seen, W WU iso.). E. spinosus auct., non L. (1767): A. Rich. (1848); Oliv. & Hiem (1877: 431); Cluov. (1904); Engler (1892). Densely branched low shrub, 40-75 cm high, rarely bushy herb; stem striate-sulcate, arachnoid-pubescent, often mixed w ith dark-purple liairs, c 5-8 mm wide, bark often dark-purple; rootstock soft and corky. Leaves obovate or elliptic, up to 25 x 12 cm, pinnatisect, segments lacerate with margins terminating in numerous pungent spines, about 2-5 cm long and up to 1 cm wide; upper surface

22

145 ASTERACEAE: 4. Echinops

Figure 11 ECHINOPS KEBER1CUO 1 - upper stem parts x c 1; 2 - leaf de­ picting narrow segm ents in populations from Mullo (SU) x c 1; 3 - capitulum x c 2; 4-8: outer phyllaries, from the out­ erm ost (4) to the innermost (8) x 3W. 9 & 10: - inner phyllaries x 3 '/i;11 cyp­ sela x 314. 1,4-11 from Mesfin T 6990, 2 -3 from Mesfm T. 6995. Drawn by Damtew Teferra. (Reproduced with per­ mission from Mitt. Inst. Allg. Bot.. Ham­ burg, 23b, 1990, p. 626, fig. 2.)

light green, hispid-pubescent or sparsely arachnoidpubescent, lower surface densely white-tomentose. Head 4-5 cm wide. Peduncle up to 4 cm long, arachnoidpubescent often mixed with dark-purple hairs. Capitulum 2.5-3 cm long. Outer phyllaries 3-seriate, all daggershaped (with acuminate apex and broad shoulders), gla­ brous; outermost narrow. 8-10 x 0.5—1 mm, middle series 12-14 mm long, innermost series 18-20 mm long. Inner phyllaries oblong-linear, blunt at apex, toothed, 14-16 mm long. Corolla wliite or pale blue, c 14-17 mm long, tube papillose; anthers pale purple. Cypselas densely appressed setose-pilose, c 8-10 x 1 mm. obovate, bristly

hairs of surface overtopping pappus. Pappus of numerous, contiguous, 1 mm long ciliolate narrow scales, connate at base Fig. 145.7.2. Acacia wooded grassland, low evergreen scrub with scattered Jumper us, Barbeya, Pistacia, Cussonia and Heteromorpha, usually on rocky, shallow soil on moun­ tain slopes or on brown clay soil over limestone; 1350-2200 m. EW TU WU SU SD HA; Ghana, Nigeria, Cameroun, Chad. Mali. Niger, Central African Republic, Sudan. Somalia, Kenya; Yemen, Saudi Arabia. Schimper 951; Friis et al. 3245.

145. ASTERACEAE: 4. Echinops. 5. Carduus

0. E. buhaitensis Mesfin (1990) - type: GD, Buhait, 3350-3660 m, Schimper 1070 (P holo ). Perennial herb, 30-40 cm high; stem sparsely arachnoidpubescent mixed with long purple hairs, densely leafy t uxmghout. Leaves pinnatisect with deeply-lobed seg­ ments, each ending in a pungent spine, up to 20 x 6 cm; upper surface glabrous or sparsely arachnoid-pubescent on veins; lower surface densely white-tomentose, up to 3 cm long. Phyllaries multiseriate, stalked; outermost nari ikvly spathulate with long, soft membranous bristles ng, 5-lobed, lobes c 10 mm long; tube glabrous. Anlers light-brown Cypselas (only immature seen) Dpressed-hispid, bristles white. Fig. 145.8.4. Mountain grassland; 3350-3800 m. GD; not known Isewhere O. Hedberg & G. Aw eke 5464; Schimper 1070. This species seems to be a rarely collected endemic ?m the Simen Mts., known so far from these collections :>nly.

11. E. angustilobus S. Moore (1902) - type: Kenya, Mau to Nolongoselli, Scott-Elliot 7005 (BM holo., K iso.). : rect perennial herb, up to c 1 m high; stem arachnoidubescent especially at upper parts and often mixed with • -own, c 1 mm long 1-seriate hairs. Leaves obovate, up to 25 x 14 cm, bipinnatisect (upper leaves pinnatisect), pri­ mary segment up to 9 cm long, secondary segment up to 4 5 cm long and c 5 mm wide, rachis narrowly winged, ul: imate segments narrowly linear-lanceolate ending in -12 x 0.1—0.3 mm, middle row 14-16 x 0.7-1 mm. Ini>er phyllaries oblong, c 12 x 1 mm, dark-grey on upper tialf, fimbriate at apex; innermost 18-20 x 1.5-2 mm. Co­ rolla white or faintly blue, c 15 mm long, tube glandular uiry, anthers light brown. Cypselas densely appressed uspid, c 8-10 x 1-1.5 mm, oblong. Pappus of numerous, iliolate scales, connate at base. Fig. 145.8.5. Mountain slopes with low shrubs and herbs; 1800-1900 m. SD; rare in the Flora area but common in '.enya with the mountains around Mega being the north­ ernmost limit of distribution. Gillett 14447; M esfn T. 3409 (ETH).

23

12. E. longifolius A Rich. (1848) -type: GD, Kouaieta, 1840, Quartin-Dillon s.n (P holo., K photo.). Echinops luckii R E. Fries (1923) - type: Kenya Lumbwa, Fries & Fries 2850 (UPS holo.). E. schweinfurthii Mattf. (1924) - type: Border Su­ dan/Ethiopia |GD1, Gallabat, Meteinma. 1865. Schweinfurth 480 (B syn. destroyed, K isosyn.), 481 (B syn destroyed, BM P WU isosyn.) & 482 (B syn. destroyed, BM P WU isosyn.). E. serratifolius Sch. Bip. (1861), nom. nud., quoad PI. abyss. II, Schimper 941. Straggling perennial herb, up to 1 m high; stem striate-sulcate, arachnoid-pubescent exposing dark-purple bark, slender, c 5 mm wide, solitary or several from a woody rootstock. Leaves simple, linear-lanceolate, up to 25 cm long, and 1—2.5 cm wide, auriculate and amplexi­ caul at base, acute at apex; margins regularly doubly-serrate or sometimes with long triangular spinous lobes in the lower half; upper surface glabrous, dark green, lower white-tomentose, midrib prominent below. Head 3-5 cm in diameter, solitary or up to 6 per plant. Peduncle up to 4 cm long, arachnoid. Capitulum 2-2 .5 cm long. Outer phyllaries green 3-seriate, surface glabrous except at margins, which are ciliolate in upper parts, out­ ermost spathulate with narrow lower parts. 4-8 mm long, middle linear-rhomboid. 8-12 x 1-2 mm. innermost subulate or oblong-ovate, c 1 5 x 2 nun. Inner phyllaries oblong-linear. 8-10 x 1 mm. glabrous except the ciliolate margins; apex fimbriate or 3-cleft. Corolla pale to dark-blue, c 15-20 mm long, tube glabrous. Cypselas densely setose-pilose especially toward the apex, c 10-12 mm long. Pappus of many contiguous, ciliate. narrow scales, connate at base. Figs 145.7.1; 145.10.8-10. Wooded grassland and deciduous woodland on black clay soil or on sandy soil of volcanic origin; 500-3050 m. EW TU GD GJ SU WG IL; Sudan, E Africa, Togo. Guinea, Mali. Ghana. Cote d ’Ivoire. Nigeria. Cameroun. Chad, Central African Republic, Zaire. Hillier 945; Smeds 740; Friis et al. 2407.

5. CARDUUS L. (1753) S.M.A. Kazmi in Mitt. Bot. Staatssconl. MiXnchen 5. 139-198 (1963) & 279-550 (1964). Perennial or annual herbs; stem winged, spinulose. L e av es alternate, lobed to twice p in n a tise c t. sc a b rid -

pubescent to glabrous, margins spiny. Capitula homo­ gamous, solitary' or several aggregated together, sessile or shortly pedunculate, globose or campanulate. Phyllaries multiseriate, imbricate, patent or deflex innermost phyllaries usually membranous. Receptacle flat or con­ vex, paleaceous. Florets purple, pink or white, tubular; co­ rolla actinomorphic or zygomorphic, deeply 5-lobed; fila­ ments free, pilose, anthers tailed; style bifid. Cypscla obovate to ovate-oblong. 2-4.5 mm long. 1.5-1.8 mm w ide, compressed, glabrous. 6-10-striate-sulcate. annular and umbonate at apex. Pappus multiseriate. white; bristles

---

24

145. ASTERACEAE 5. Carduus

Annual heibs: peduncle and phyllaries densely arachnoid; cypselas shiny, white, 6-10 sulcate 5. Perennial lierbs. peduncle and phyllaries hispid; cypselas sluny, black or brown streaked with black, without evident striae. 4. C. leptacanthus I C. schimperi Sch Hip. ex A. Rich. (1847)

type GD. Simen Mts., Mt. Selki. PI. abyss. II, S chtm per 1301 (P holo . BM K L M P S UPS iso.). ( nicus cham aecephalus Vatke (1875); C arduus cham aecephalus (VaUie) Oliv & Hiem (1877) - type:

GD. Debre Tabor. 1863. Schimper 1412 (B holo. de­ stroyed. BM K P iso ). C. erem ocephalus Chiov. (1912) - type: SU. Ciafedenza. Negri 51 (FT syn.). SU, Entoto, Negri 309 (FT syn). Carduus ellenbeckii R E. Fries (1923) - type: SD. Jam-Jam. Ellenbeck 1754 (B holo.. destroyed) Ethio­ pia jcxact region not known], between Harer and Addis Abeba. X. 1898. IVel/bvs.n. (K neo ).

Figure 12. CAROL I S SC H IM P E R I Habit x •/?. From Thuiin et al 1393. Drawn by Daintcw Teferra

scabrous or baibcllatc to glabrous, connate at base and de­ ciduous as a unit. A genus with about 90 species distributed in Eurasia. North and East Africa and widely introduced as weed elsewhere. A world-wide revision of the genus was made b\ Kaznii (loc. cit.) who. like Cufodontis (1967). recorded 9 species from the Flora area; the present study reduces the number to 5 species only. The citation of Ethiopia as pari of the distributional area for C. afromontanus R E. Fries in FI. Trop. E. Afr. Compositae (Part I. 2000) appears to be an error. No specimen from the Flora area matching the said species has been found. 1 2.

-

3.

Plants steinless; capitula sessile in centre of leaf ro­ sette I. C. schimperi Plants with evident stems; capitula al apex of stem or branches. 2

subsp schimperi Steinless perennial herb. with a long, white or brow n tap­ root Leaves flat on ground or slightly raised from liall-wa> up. 3-20 cm long, 1.5-6.5 cm wide, rigid, ellip­ tic or oblanccolatc. pinnatisect. shortly petiolate. seg­ ments rounded with triangular lobules, margins w ith up to Xmm long spines; upper surface hispid-pilose to glabrous, olive or greyish-green, red-violet tinged at base, lower surface arachnoid-pubescent, or, if glabrescent, with long luspid liairs only along veins, pale or bluish-green Capitula sessile, at the centre of leaf rosette, usually soli­ tary oblong-campanulate. (l-)2 -4 .5 cm high. ( J—)2—2.5 cm wide at base. Phyllarics inultiseriate, imbricate, elon­ gate-lanceolate. adpressed. rigid, sparsely arachnoidpubescent to glabrous, exccpt sometimes for the hispid margins; apex acute w ith up to 2 nun long spine: outer se­ ries about 3- 15 mm long. 2-5 mm w idc at base; innermost ph> llaries oblong-linear, much longer. Florets white to pale-lilac, sweet-scented; corolla 18-35 mm long, lobes 4 -1 1 nun long; anthers 8-10 mm long. Cypsela obovate, light-brown. 9-10 mm long. 3.5 mm wide, annular and umbonate at apex. Pappus multisenate, barbellate or gla­ brous. light brown. (15—)3()—40 111111 long. Figs 145.12;

Outer phyllaries 2.5-3.5 mm wide at base, margins scarious. yellow, sharply laciniatc; plant densely white-woollv. 2. C. m acracanthus Outer phyllaries less than 2 mm wide at base, mar­ gins glabrous or hispid; plant hispid or thinly arachnoid. 3

Montane grassland, encaceous bushland. secondary scrub, open Juniper us forest; abundant in heavilv grazed area; 2000-4600 m. GD WU SU WG KF GG SD BA. Su­ dan E Africa, ( iillett 14707; Ash 1632; Afoonev 8516.

Phyllarics subequal, hispid, all ending, in long erect or spreading spines, about 0.5 nun wide at base, the outer series obscuring the lower parts of the median ones. 3. C. nyassanus Outer phyllaries short, phyllaries progressively longer towards centre, usually reflexed from mid­ dle, glabrous or thinly pubescent, the median se­ ries clearly visible. 1-2 mm wide at base. 4

The material from the Flora area is all subsp. schimperi. Oilier subspecies in East Africa: subsp. platyphyllus (R.E. F r ) C. Jeffrey is endemic to Mt. Kenya and subsp. nanus (R E Fr.) C. Jeffrey is found in Uganda, Kenya and Rep. Dein. du Congo. According to C. Jeffrey & H J Beentje Jin FI Trop. E. Afr. Compositae (Part 1): 46 (2000)), subsp. schimperi is distinguished mainly by its larger (26-40 mm) corolla.

145.13.1 —4.

145. ASTERACEAE: 5. Carduus

25

Figure 13. CARDUUS SC H IM P E R I 1 - leaf x V3; 2 - capitulum x c 1; 3 phyllaries x '/?, (longest - the innermost); 4 - capitulum x 'A. C. MACRACANTHUS. 5 - leaf x 1/ 3; 6 - capitulum x IV2 . C. N Y A SSA N U S : 7 - capitulum x I. C LEPTACANTHUS. 8 capitulum x c l.C. PYCNOCEPHALUS: 9 - upper part o f stem with capitula x >/4 ; 10 capitulum x 1. 1 from Wellby s.n; 2 & 3 from Schimper 1301; 4 from Negri 51; 5 & 6 from Schimper 51; 7 from Whyte s.n.; 8 from Schimper 910; 9 & 10 no specimen cited. 1-8 artist not given in orig.; 9 & 10 drawn by John H. Rumely. (1 -8 combined and reproduced with per­ mission from Milt. Bot. Staatssaml. M im chen,\o\. 5 ,1 9 6 3 , pp. 176-181, figs 4a, 6b & c, 9b, 12, 13, 15a & b, 9 & 10 from Vascular Plants o f the Pacific Northwest, part 5 ,1 9 5 5 , p. 117 p.p.)

2. C. m acracanthus Sch. Bip. ex Kazmi (1963) type: [TU/GD. probably from. Agau], 1853, Schimper 51 (G holo., P K iso.). C. macranthus Schweinf. & Aschers. (1867), nom. nud. Erect perennial herb. 0.7-1.3 m high, stem branched, con­ ical, up to 3 cm wide at base, hollow, sulcate, sparsely arachnoid-tomentose, liairs white, winged with orbicular or triangular, spinescent, 6-35 mm long lobes. Leaves ovate-lanceolate, pinnatisect, up to 35 x 10 cm; segments 3-4 cm long, 1-1.5 cm wide, incised-dentate with up to 4 cm long and erect yellow spines; upper surface scabrous, ower densely white-woolly. Capitula sessile, several clustered at apices of stem or branches, 25-32 mm high, about 25 mm wide. Phyllaries multiseriate, imbricate, adpressed. densely brown-tomentose; outer series 15-25 x 2.5-5 mm wide at base, linear-lanceolate to broadly oblong, coriaceous, arachnoid, margins scarious, ciliate. often cleft below apical spine on both sides, dark-purple toward apex; innermost phyllaries membranous, lanceo­ late, c 15 mm long, weakly spinescent at apex. Corolla li­ lac, 12-15 mm long, tubular, 5-lobed. lobes about 2.5 mm

long, anthers about 7 mm long. Cypselas black or daikbrown, streaked with black, ovate-oblong, about 2.5 x 1.2 mm; apex annular. Pappus white, connate at base and de­ ciduous in a ring, glabrous or baitoellate, about 10-12 mm long. Fig. 145.13.5 & 6. Open montane grassland, steep slopes in Erica scrub; 3950-4000 m. TU GD AR BA; not known elsewhere. W. de Wilde 9187; O. Hedberg 5623; Mesfin T. 7837. 3. C. nyassanus {S. Moore) R.E. Fries (1923); C. leptacanthus Fresen. var. nyassana S. Moore in. J. Linn. Soc. Bot. 37: 326 (1906) - type: Malawi, Mt. Malosa, Whyte s.n. (K lecto.). C. leptacanthus Fresen. var. steudneri Engl, in Abh. K. Preuss. Akad. Wiss. 1891: 450 (1892); C. steudneri (Engl.) R.E. Fries (1923) - type: GD, Simen Mts., Nori. Steudner 308 (B syn., destroyed); Tanza­ nia, Kilimanjaro, Johnston s.n. (B syn., destroyed, K lecto., BM isolecto.); ibid., 1900-2800 m., H. Meyer 237 & 231 (B syn. destroyed); v. Hdhnel 121 (B syn. destroyed). C. buchingeri Schweinf. & Aschers. (1867), nom. nud.; C. steudneri (Engl.) R.E. Fries subsp. buchingeri

26

145. ASTERACEAE: 5. Carduus

Figure 14.

CARDUUS N YA SSA N U S 1 habit x */s; 2 -4 - outer, middle and inner phyllary x 4; 5 - floret, most setae removed x 6; 6 - seta, detail x 16; 7 - pappus x 2; 8 cypscla x 6. 1-5 from Robson 365; 6 -8 from Fanshawe 7316. Drawn by Eleanor Catherine. (Reproduced with permission from/*7. Zamb. vol. 6(1), Tab. 10.)

Kazmi in Mitt. Bot. Staatssaml. Mtinchen 5: 163 (1963); C. nyassanus (S. Moore) R E Fries subsp. buchingeri (Kazmi) Cufod. in Bull. Jard. Bot. Etat Brux. 36, Suppl. : 1173 (1967) - type: Ethiopia [exact region not known), Schimper 44 (G holo., K iso.). C. afromontanus auct., non. R. E. Fries (1923); Kazmi (1963); Jeffrey (1968; 125); Cufodontis (1967: 1172), quoad Gillett 14951,15007 and Mooney 8298. subsp. nyassanus Erect perennial herb, 0.5-1.5 m high; stem green with white stripes, sulcate with irregularly incised spinescent

wings on ridges, wings broader in lower parts of stem, pu­ bescent toward apex, branched in the inflorescence. Basal leaves narrowly oblanceolate, 14-40 cm long, 4-17 cm wide including spine tips, decreasing in size toward apex, uppermost sessile, pinnatipartite; segments subrectangular or oblong, margins with many, 1-9 mm long spines; upper surface hispid, lower hirsute along veins. Capitula several at the apex of main steni and branches, sessile and congested rarely solitary, campanulate, 15-25 mm high including florets, c 10 mm wide at base. Phyllaries numer­ ous, imbricate, strigose or hispid, linear-subulate, 0.5-1 mm wide at base. 5-15 mm long, terminating in a long spreading spine, if reflexed, only at apex. Corolla white to

145. ASTERACEAE: 5. Carduus, 6. Cirsium

27

pale pink, 12-15 mm long, 5-lobed, lobes 6 mm long; anihers about 5 mm long. Cypselas obovate-oblong, 4-4.5 x 1-1.5 mm, smooth, glabrous, brown streaked with daikbrown, slightly curved toward apex. Pappus light-brown, 2—15 mm long, glabrous or barbellate, connate at base and deciduous in a ring. Figs 145.13.7; 145.14.

5. C. pycnocephalus L. (1763) type: “Habitat in Europa australi.” Linnaean Her­ barium No. 966.9 [LINN lecto., selected by Kazmi in Mitt. Bot. Staatssamml. Miinchen 5: 445 (1964)].

Degraded Juniperus-Podocarpus forest, open dis­ turbed or grassy areas along road in forest, damp valley '•ottom in Aningeria forest, open montane woodland; 1780-3510 m. TU GD GJ SU AR WG IL KF GG SD BA blA; Uganda. Rwanda. Burundi. Rep. Dem. du Congo. Tanzania. Mooney 5724; Mesfin T. & Kagnew 2508; Friis et al. 348.

style often with a hairy ring below the point of bifurcation. Cypselas oblong-obovate, compressed, or slighdy qua­ drangular, glabrous, smooth, with hemispherical umbo apically, 6-10-striate-sulcate. Pappus multiseriate, plu­ mose, connate at base into a ring, deciduous as a unit, or persistent; setae longer toward centre, scabrous or thick­ ened at apex.

Erect annual herb, c 75 cm high; stem slender, solitary, paniculately branched above, narrowly winged with trian­ Arundinaria alpina bamboo thicket, along streams gular, spinose or spinulose lobes. Leaves grey-green espe­ cially on lower side, pinnatilobed to pinnatisect, lobes and ravines in Juniperus forest, open Hagenia2-4-fid, lobules triangular, sparsely hairy above, Hypericum forest; (2000-)2300-3350 m. TU GD SU AR WG GG SD BA; Sudan. E Africa, Cameroun, Nigeria, canescent-tomentose beneath. Capitula solitary or 2-5 clustered together at apices of branches, or on up to 5 mm Rep. Dem. du Congo, Rwanda Burundi, Malawi, Zam­ bia. Gillett 15007; Mooney 8298; Friis et al. 5691. long, densely white tomentose peduncles, 17-25 mm long, 7-8 mm wide at base, arachnoid-pubescent to Subsp. kikuyorum (R. E. Fr.) C. Jeffrey occurs inE Af­ glabrescent. Phyllaries multiseriate, imbricate, adpressed, rica, Rep. Dem. du Congo and Rwanda. Subsp. nyassanus green toward apex, stramineous at base, lanceolate with differs by having subequal, strongly spine-tipped narrow spinous apices; imier series scarious, longer than phyllaries and the basal leaves being 7-21 cm wide and median ones. Corolla tubular, 14-16 mm long, slightly 2.5-5 (-8) times as long as wide [versus 4-6.5 cm wide. exserted, limb lilac-red. tube whitish-yellow. Cypselas 5.5-10 times as long as wide]. white, shiny, c 4.5 mm long, glabrous. Pappus glabrous or barbellate. connate at base and deciduous, falling as a 4. C. leptacanthus Fresen. (1840) type: EW/TU. between Halei and Tembien. ring. Fig. 145.13.9 & 10. Riippel s.n. (FR holo.. P iso., not seen). C. abyssinicus Sch. Bip. ex A. Rich. (1848) - type: subsp. pycnocephalus GD. Simen Mts.. Mt. Aber, near Genausa, PI. abyss. II, Schimper 910 (P holo., K S iso ). Median phyllaries strongly 1-veined in distal end, con­ C. semiensis Pic.Serin. (1951) - ty pe: GD. Simen tracted into a lanceolate prolongation about as long as Mts., Mt. Chiddis-Arit. Pichi-Sermolli 2625 (FT proximal part; phyllaries not purplish distally. holo ). Roadside weed; 1700-2100 m. SU; native of W Eu­ Erect perennial herb. 0.75-2 in high; stem usually un­ rope, Mediterranean region. NW Africa, SW and C Asia branched except in the inflorescence, flattish. sulcatc. to Afghanistan and Pakistan. W. de Wilde 8758; Mesfin T. Dilose-tomentose toward apex, winged throughout with 7057. sinuate or pinnatilobed. semiorbicular. and densely This appears to be a recent introduction into the Flora spinescent wings. Basal leaves pctiolate, oblanceolate. area, currently known only from SU (Ambo). 10-50 cm long. 3-25 cm wide, pinnatilobed to pinnati­ sect; segments subrectangular or ovatc-lanceolate, with 2-6 mm long pale, yellowish-brown spines at margins; upper surface dark-green hispid pilose, low er hirsute with 6. CIRSIUM Mill. (1754) ong crisp hairs along veins: cauline leaves similar to 1. Friis in N orw egianR ot. 22 201-207 (1975). basal, decreasing in size toward apex, segments lin­ ear-lanceolate. Capitula several, sessile and densely ag­ Erect perennial, rarely annual heibs, 0.5-4 m high; stem gregated at the apex of branches and main stem, obtuse simple to branched, winged, rarely acaulescent. Leaves .ampanulate. 15-20 mm high. 5-10 mm wide at base. alternate, sinuate-dentate to pinnatisect, rarely subentire, :>hyllanes multiseriate, imbricate, linear-lanceolate. glabrous to luspid-pubescent. spiny at margins. Capitula 1.5-2 mm wide at base, gradually narrowed to a homogamous. discoid, ovoid or globose, solitary or sev­ subuliform apex, markedly reflexed at middle, glabrous eral aggregated together. Phyllaries multiseriate, imbri­ or shortly strigose. Corolla purple-red or pink, tubular. cate. graded monomorphic. adpressed or patent in upper 15-18 mm long. 5-lobed. lobes 7-8 nun long; anthers parts, spinescent or membranous, margins and outer sur­ A’hite, 4 mm long; sty le purple. Cypselas oblong-obovure 21 P LE C T O C E P H A L U S VA R IA N S: upper part o f tem x '/2. From M elaku W. & Andargachew 292. Drawn by hrhanu Mihretu.

11. VOLUTARIA Cass. (1816) 1mberboi Adans.(1763), nom. illegit., non Amberboa Less. (1832), nom. cons. Locellia Viv. (1824). Volutarella Cass. (1826). j Wagemtz in Candollea 46(2): 407-410 (1991). innual herbs, pilose, unarmed; stem low-lying or up to 1 n high, often with m an y capitula. L e av es sim p le or often

innatifid or pinnatisect. Capitula medium sized, heteroamous. rarely homogamous, radiate. Receptacle setose, eiae smooth. PhyHanes multiseriate, imbricate, acute or rucronate at apex. Florets pink, purple, violet, blue or ellow. rarely the disc florets yellow; ray florets sterile, jbular, commonly funnel-shaped. Cypselas laterally impressed, sparsely villose, liairs long, longitudinally ibbed and transversely rugose; hilum lateral or subbasal. aith deeply hollowed-out pit and cartilaginous margin, 'appus persistent, multiseriate, paleaceous; pales narowly spathulate, scabrous.

Erect annual herb. 10-60 cm high; stem solitary or densely branched, scabrous with short white liairs. striate-sulcate. slightly tetragonal. Leaves oblanceolate in outline, pinnatifid to pinnatisect, scabrid-pilose. resin­ ous-dotted, lobes of basal and lower cauline leaves 5-10 cm wide with dentate or lobulate margins, the tenninal one usually the largest, those of upper leaves 1-4 mm wide: radical and lower cauline leaves petiolate. upper leaves sessile with shortly decumbent margins. Capitula campanulate. heterogamous, (5-) 10-20 mm high includ­ ing florets. Receptacle slightly cartilaginous, peduncle up to 10 cm long, white-woolly near base of capitulum. Phyllaries 5-6-seriate, imbricate, appressed, 3-5-veined. 2-15 nun long, gradually longer inwards, lanceolate to linear-lanceolate, tapering into a scarious. acute to spinescent apex, black-margined, surface thinly arach­ noid and scabrid. Florets white, pink, pale purple or pale blue; marginal florets 10-15 mm long, sterile; disc florets corolla 5-lobed. Cypselas oblong-elliptic, light brown, minutely but densely pitted. 3.5—4(—5) mm long. 1.2-2 nun wide, costate, sparsely to densely villose. hairs white, long; hilum lateral with deeply hollowed pit and yellow­ ish cartilaginous margin. Pappus of numerous multiseri­ ate pales, 0.2-5 mm long, outennost pales smallest; mar­ gins ciliate. Fig. 145.22. Grassy areas in degraded Juniperus forest, open Aca­ cia-Commiphora woodland, grassland; 1325-1850 m. SD. Djibouti, Socotra. Kenya. Tanzania. Friis et al. 866; Ash 1196; Puff & Ensermu K. 821224-4/12. 2. V. abyssinica (Sch. Bip. ex A.Rich.) C. Jeffrey ex Cufod. (1967); Amberboa abyssinica Sch. Bip. ex A. Rich. (1848) - ty pe: TU. Ouadgerate. Ouartin-Dillon et Petit 256 (P holo). Centaurea somalensis Oliv. & Hiem (1877): Amberboa somalensis (Oliv. & Hiem) Sojak (1962): Volutaria somalensis (Oliv. & Hiem) C. Jeffrey ex

36

145 ASTERACEAE II Volutaria

M

|

$7

Figure 22. VOLUTARIA BORANupper part o f stem x Vy. 2 capitulum x 2; 3 capitulum sec­ tion, some florets removed x 2. 4 receptacular hair x 3. 5 marginal floret x 4; 6 central floret, cypsela and pappus removed x 6; 7 sta­ mens x 9; 8 - style x 6; 9 immature cypsela with pappus x 6. 10 m a­ ture cypsela with pappus x 4 All from Gillett 20840 Drawn bv Pat Halliday (Reproduced with permis­ sion from FI. Trop. E. Afr. C o m ­ positae. fig. 14.)

E N SIS. 1

Cufod. (1967), syn. nov. - type: Somalia. Hildebrandt 1448 (BM holo ). Centaurea saltii Philipson (1939); Amberboa saltii (Philipson) Sojak (1962) - type: Ethiopia (exact region not known; probably EW/TU), Salt s.n. (BM holo.). Amberboa hochstetteri Buchinger ex Hochst., nom. nud., quoad Schimper 2183 (K). Centaurea hochstetteri auct., non Oliv. & Hiem: Lanza (1939), quoad Cufodontis 591. Erect annual herb, 30-100 cm high; stem repeatedly branched, striate-sulcate, puberulous below, dense.

\\ lute-cottony above, some branches slightly tetragonal, low er cauline leaves obovate pinnatifid to pinnatipartite, sometimes simple with lacerate margins, scabrid pilose, resinous-dotted, petiolate, up to 12 x 5 cm. lobes 5-12 mm wide, terminal lobe the longest and widest: upper leaves sessile. Capitulum campanulate or ureeolate, solitary heterogamous, 20-30 mm high, including florets, c 10-12 mm wide at base, receptacle cartilaginous tliickened. c 5-7 mm deep. Peduncle up to 9 cm long, arachnoidtomentose, striate-sulcate. Phyllaries 5-7-seriate, imbri­ cate. 2-18 mm long, gradually longer inwards, ovate to linear-lanceolate, veinsruot distinct, cartilaginous: surface

145. ASTERACEAE: 11. Volutaria, 12. Carthamus

s.abrid. thinly cottony, especially on the margins toward a-iex, black-margincd; apex acute, cartilaginous-spinesc:nt. Florets pink, violet, wliite or purple; marginal florets 1 >-22 mm long, sterile; disc floret corolla 5-lohed. Cyps .las oblong-elliptic. light brown, minutely but densely pitted. 4.5-5.5 mm long, 1.5-2 mm wide, costate, sparsely t 1densely villose. hairs white; liilmn lateral with deeply 1 allowed pit and yellowish cartilaginous margin. Pappus of numerous, multiseriate pales. 0.2-4 mm long, outerr lost pales the smallest; margins ciliate. Dry Acacia woodland, grassy mountain slopes on I roestone; 1520-2400 m. EW GD SU TU HA; Somalia. A looney 809; Tekle Hagos 62; Burger & A. Getahun 265. The two species, although they occur in different parts c the countiy and in different soil types, cannot be separ iied by discrete morphological characters. The capitula a id cypselas are generally larger in V. abyssinica than in \boranensis. In V boranensis these appear to be apical ly r uch more elongate than in V. abyssinica throughout the r .nge of its distribution. Also the receptacle is thick and 1 irdened and thus not easily flattened as in V. abyssinica. Wagemtz (1991) subdivided V abyssinica into 3 subspecies: subsp. abyssinica, subsp. inornata Wagenitz. and subsp. ayimeri (Bak.) Wagenitz.

12. CARTHAMUS* L. (1753)

P Hanelt in Feddes Repert. 67: 41 -180 (1963). I rect annual herbs; stems branched, glabrous or with \ lious-lanate indumentum. Cauline leaves alternate, 1m the Flora area.

Cufodontis \Bull. Jard. Bol. EtatBrux. 36. Suppl.: 177 ( 967)] cited Carthamus persicus Desf. ex Willd. (1803) giving Newbould 1001 as the voucher specimen. This soeciinen is from Somalia. Ghura Heis (Graheis), c 11° 01 N; 48° 53’ E and since the species could probably oc­ cur in eastern Ethiopia, it is included in the key. 1

Stem villosc-lanate. yellowish-brown, striatesulcate. 2. C. lanatus Stem glabrous, smooth white, terete.

2.

-

37

Florets orange-red or orange-yellow; leaves undi­ vided, merely spinulose-denticulate at margins. 1. C. tinctorius Florets yellow, leaves deeply pinnatifid with lobes ending in sharp pungent spines. ( ' persicus

1. C. tinctorius L. (1753) -type: “Habitat in Aegypto.,? Herb. Clifford: 394. “Carthamus 1" [BM lecto.. selected by Rcchinger in FI. Iranica 139b: 434 (1980)1. Erect stiff annual herb, up to 75 cm high; stem much branched, glabrous, smooth, white. Cauline leaves green, ovate to linear. 3-10 x 1-2 cm. sessile or slightly clasping the stem, glabrous, with 3-5 veins from base: margins en­ tire to finely spiny-serrate, lower leaves deciduous. Capitula lax. deeply 5-lobcd; anthers 7-8 mm long, brown. Cypselas smooth, glabrous, white. 8-10 x 4-5 nun. obpyramidal. tetragonal, truncate at apex, usually epappose but in some w ith short fcatheiy bnstles at centre and small scale-like pales externally; hiluin lateral. Fig. 145.23.7-1 1. Cultivated for the edible oil-producing fmits (cyp­ selas), sometimes an escape in wasteland; 1000-2400 m. EW TU GD WU SU AR WG KF HA: widely cultivated. Ash 1378; Mooney 5108; Mesfin T. 851. Not known in the wild state: widely cultivated in the E Mediterranean area. Afghanistan. India and California (U.S.A.) for its oil bearing fruits; sometimes also culti­ vated as an ornamental: it is reported that the flowers are used in cosmetics for making a rouge. 2. C. lanatus L. (1753); Kentrophyllum lanatum (L.) DC. & Duby (1806) type: “Habitat in Gallia. Italia. Crcta.” Loefling 610a in IAnnaean Herbarium No. 973.2 |LINN lecto.. se­ lected by Hanelt in Feddes Repert. 67: 136 (1963)J. K. lanatum (L.) DC. & Duby var. abyssinicum Sch. Bip. cx A. Rich. Tent. FI. Abyss, i: 454 (1848); ('arthamus lanatus L. var. abyssirticus (Sch. Bip. ex A. Rich.) Schweinf. & Asch.. Beitr. FI. Aethiop.: 143 (1867) - type: TU. Adwa. near Genniam in Memsach distr.. PI. abyss. I. Schimper 207 (P holo.). Annual herb. 15-100 cm high; stem grey-green to strawcoloured. sparsely to densely villous. Cauline leaves ovate to ovate-lanceolate, amplexicaul. glandular-villous, usu­ ally green, rarely greyish green. 2-6 x 0.5-1.5 cm exclud­ ing spines; v ein s 3-5. p ro m in en t below; margins spinvpinnatifid. Capitula campanulate, corymbose. 30-40 nun high, including fmits. 20-25 nun wide. Phyllaries multiseriate, imbricate, hispid-tomcntosc: outer ones leaf-like. 25-40 x 8-10 mm. erect to patent or somewhat recurved; median phy llaries gradually acuminate or appendaged. ap­ pendage leaf-like, c 15 x 5 nun. 3-veined; innermost c 25 x 5 nun long, scarious. nanowly lanceolate with acuminate apex. Florets lemon yellow, tubular; corolla 25-28 mm long, deeply 5-lcbcd: anthers 7-8 nun long. Cypselas slightly compressed-tetragonal, obpyramidal. 4-5 x 3.5-4 mm. brown. Pappus of multiseriate. scale-like, liispid

---------------------

38

145. ASTERACEAE: 12. Carthamus, 13. Centaurea

Figure 23. CARTHAM US LAN ATU S 1 - upper part o f plant with capitula x ‘/'a; 2 - outer phyllary x IV4; 3 - inner phyllary x 1V2\ 4 - corolla x 1W, 5 - cypsela with pappus x lVi; 6 - scale-like bristle x 3. C. TINCTORIUS 7 - upper part o f plant with capitula x V2; 8 - outer phyllary x 114; 9 - inner phyllary x IVi; 10 - floret excluding ovary x 3; 11 - flo­ ret with immature cypsela x 1V4. 1-6 from AmanuelB. A -7 5 ; 7-11 from AiooMey 5108. Drawn by Birhanu Mihretu.

bristles, 1-11 nun long, outermost ones the smallest, often streaked with black. Fig. 145.23.1-6. Weed of fallow-land and roadsides. 1000-2440 m EW TU GD GJ SU AR HA. W. C and S Europe. N Africa. S Russia. Iraq, W and C Iran, extending to W China. Moo­ ney 8245, Boulos 9548; Westphal 1923.

13. CENTAUREA L. (1753) M esfin T adesse in Kew Bull 47: 737-740 (1992).

Annual or perennial herbs; stem solitary or severalbranched, thinly pubescent to white-toinentose. Leaves alternate or basal, simple or pinnatisect, toothed or entire at margins, some^mes decurrent on stem. Capitula soli­ tary or cymose. pedunculate or sessile, heterogamous or rarely homogamous, disciform or radial, ovoid-globose or campanulate. Receptacle flat, bristly. Phyllaries multise­ riate, imbricate, appressed. coriaceous or scarious.

appendaged at apex with appendage having pectinate margins and rigid spines. Florets pink, purple or pale yellow, marginal florets neuter (sometimes with staminodes), funnel-formed with 5-8 lobes, or filiform with 4-5 linear segments; disc floret bisexual, corolla tubular, deeply 5-lobed, yellow, pink, purple or white; anthers sagittate, style bifurcate with a ring of short liairs below the point of bifurcation. Cypselas glabrous (when ripe) or puberulent, compressed or tetragonal, articulating later­ ally; apex rounded or truncate. Pappus persistent or rarely caducous, multiseriate, unequal, scabrous, barbellate or plumose. About 450 species chiefly in the Mediterranean region and SW Asia some in Tropical Africa and N and S Amer­ ica; 4 species in the Flora area. I.

Annual herbs; leaves lyrate-pinnatifid to sinuately lobed, decurrent, capitula 8-12 mm wide; cyp­ selas 2-3 mm long. 2

39

145. ASTERACEAE: 13. Centaurea

Figure 24 C E N T A U R E A M E L IT E N S IS 1 - upper stem part x Vi, 2 - floret x 3‘/2', 3 - basal leaf x Vi. 1 & 2 from M ac bride & Feather stone 1029; 3 from Sagastegui & Araujo 7785. Drawn by M. Werner. (Re­ produced with permission from Fieldiana Bot., N ew Series 10, 1982, fig. 1.)

Perennial herbs with woody rootstocks; leaves simple, decurrent or not; capitula 12-15 mm wide; cypselas 5-6 mm long. 3 Leaves narrowly-linear; disc corolla eglandular; terminal spine of phyllaries 7-16 nun long, stout. 2.

Leaves oblong-lanceolate; disc corolla glandular; terminal spine of phyllaries 3-7 nun long, slen­ der. 1. C. melitensis Leaves decurrent, margins coarsely serrate. 4. Leaves not decurrent. margins denticulate, revolute, disc florets pink, glandular. 3. C. praecox

1 C. melitensis L. (1753) No.

- type: “ H ab itat in M e h ta .” Linnaean Herbarium 1030.62 [L IN N lecto ., se le c te d by D illo n in

Fieldiana. Bot. n.s. 10: 2 (1982)]. Annual herb. 15-70 cm high; stem narrowly winged by solstitialis the C. decurrent leaf bases, little branched at middle, tliinly arachnoid when young, liairs coarser and crisp in older plants, purple-striate. Basal and lower cauline leaves oblanceolate. sinuately lobed to lyrate-pinnatifid, crispatc-pubcrulent to arachnoid, 2-15 cm long. 5-30 mm wide, usually deciduous, margins toothed, scabrid; mid­ C. jeffreyana dle and upper leaves smaller, becoming lanceolate, mar­ gins entire to irregularly undulate. Capitula homogamous.

40

145 ASTERACEAE: 13. Centaurea

Figure 25.

CENTAUREA PRAEC O X 1 habit, capitula with mature cyp­ selas, leaves fully developed x 14; 2 - habit, capitula in flower, young leaves x '/*; 3 - outer floret, sterile x 4; 4 - inner floret, fertile x 4; 5 - 7: outer, middle and inner phyllaries x 3; 8 - pappus scale x 20; 9 - cypsela with lateral at­ tachment scar x 5. 1 from Biegel 4 1 1 2 ,2 - 7 from Angus 3772; 8 & 9 from Greenway & Brenan 8111. Drawn by Eleanor Catherine. (Reproduced with per­ mission from FI. Zamb. vol. 6(1), Tab. 12.)

solitary or cymose, 12-15 x 8-12 mm. ovoid-globose, phyllaries glabrous to arachnoid, without evident veins; outer and middle phyllaries ovate, 4-8 x 2-3 m m indurate, stramineous, scarious w ith narrow, patent apical spines and 1-3 remote, short, lateral spines on each side, the central spine 5-9 mm long; innermost phyllaries weakly spinose or merely tapered, not enlarged apically, purple-tinged, 8-9 x 1.5-2 mm. glabrous, scarious. Co­ rolla yellow glandular, tubular, about 10 mm long. 5-lobed; lobes 1-2 mm long, linear. Cypselas oblong. 2.5-3 mm long, puberulent to glabrous. Pappus of numer­ ous, multiseriate bristles; outermost narrow, scale-like.

0.5-1.5 mm long; inner setaceous, about 3 mm long. Fig 145.24. Roadside ditches; 2000-2350 m. EW GJ SU; native of southern Europe, introduced elsewhere. Mesfin T. & kagnew 1615; Fichtl & Admassu 738; Pichi-Sermolli 2138 2 . C.

solstitialis L. (1753) type; "Habitat in Gallia, Anglia, Italia.” Linnaean Herbarium No. 1030.59 [LINN lecto., se­ lected by Jeffrey in Kew Bull. 22. 136 (1968)].

145. ASTERACEAE: 13. Centaurea

41

Annual herb, up to 50 cm high: stem thinly greyishUjmentose. much branched from base. Leaves decurrent, ^:;abrid and arachnoid or lanate, or greyish tomentose; basal and lower cauline leaves 5-10 cm long, 1-3 mm wide, lyrate to pinnatifid with triangular-oblong segi icnts; middle and upper leaves smaller, becoming linear ; nd entire, mucronate at apex. Capitula solitary, ovoidI lobose, 15-20 x 8-12 mm. Phyllaries multiseriate, imb ncate, broadly ovate, puberulent or thinly arachnoid; lat( d spines about 3 mm long; inner phyllaries with a small * aline appendage. Florets pale yellow, eglandular. Cypelas 2-3 mm long, dimorphic; marginal ones epappose, blick, central or disc cypselas greyish to brown, with . riite, 3-5 nun long pappus. Fallow fields; c 2380 in. EW SU; a species native of S urope, S Russia, the Middle East; introduced elsewhere. looney 7900; Terracciano & Pappi 1144 (FT). . C. praecox Oliv. & Hiern (1877)

- type; Nigeria. Bauchi, Barter 1223 (K holo.). C. rhizocephala Oliv. & Hiem (1877), non Trautvetter (1873), nom. illegit. - type: Sudan. Sabbi. Schweinfurth 2657 (K holo.). C. hamaseniana Cliiov., nom. nud.. quoad Fiori 1863, Pappi 4380. Terracciano & Pappi 1145 (FT). nect, perennial herb, up to 50 cm high, with woody r:>otstock; stems several, branched below, densely arach­ nid in upper parts. Cauline leaves simple, alternate. ■ blanceolate. 5-7 cm long. 5-10 mm wide, white-woolly .) glabrescent; margins denticulate, revolute. Capitula jnpanulate, in simple or compound cymes, 13-15 mm high including florets, 12-15 mm wide at base; peduncle l p to 2 cm long, lanate. Phyllaries multiseriate, imbricate, i i Itoid or oblong-deltoid, coriaceous, glabrous; outer and uddle series with patent spines, central spine longest, 5—10 mm long, lateral spines 3-4 on each side, 1-2 mm long; innermost phyllaries scarious with purplish, acute, oineless apex. 10-15 x 1.5-2.5 mm. Florets pink; disc corolla tubular, 12-15 mm long, deeply 5-lobed, lobes . landular. Cypselas light-brown, ovate, c 6 x 2 mm. thinly ubescent. terete, slightly arcuate. Pappus of numerous, mltisenate bristles; outer narrow, scale-like, 0.5-2 mm mg. ciliate at apex and margins; inner setaceous, spirally .oiled, glabrous or sparsely hairv, 10-12 mm long. Fig. 45.25. Combretum-Terminalia-Dombeya, etc. woodland, grassland; 1550-2400 m. EW WG; Sudan, E Africa, jhana. Benin, Niger, Nigeria, Central African Republic, i urundi. Rep. Dem du Congo. Malawi. Zambia, Zimba­ bwe. Mooney 7740; Firoi 1863 (FT). Pappi 4380 (FT). 4 C. jeffreyana Mesfin (1991) - tvpe: EW. Afdeyo, O. Rvding 2048 (ETH holo.. K UPS iso.). Centaurea spec, sensu Cufod. (1967: 1175). quoad Baldrati 4837, 1767. ubshrub or branched perennial herb up to c 50 cm high I 'om a thick rootstock; stem thinly grey-haiiy or arach-

Figure 26. CENTAUREA JEFFREYAN A. 1 - habit x 1; 2 outeT phyllary x 4; 3 - middle phyllary x 4 , 4 - profile o f tip o f middle phyllary x 4; 5 - inner phyllary x 4; 6 - floret x 3; 7 - an­ thers x 6, 8 -s ty le tip x 18. All from Ryding204$. Drawn by Pat Halliday. (Reproduced with permission from Kew Bull. 47, 1992, p. 738, fig. 1.)

noid. narrowly winged by the decurrent leaf bases, leafy especially toward apex, leaves deciduous below. Leaves simple, decurrent, oblong-oblanceolate, up to 6 x 1.5 cm; blade thinly arachnoid but denser on the veins, hairs coarser and crisp especially on older leaves, densely resin­ ous-punctate; margins coarsely serrate, recurved with apiculate, callose-thickened teeth, densely arachnoid. Capitula homogamous, solitary at the apex of branches, 12-16 x 15-18 mm. hemispherical, epedunculate or peduncle only up to 2 mm long, densely arachnoid. Phyllaries 5-6-seriate with numerous dark-purple spots except in area of overlap; outermost series arachnoid at base and tliinly so on median vein up to the apex, c 3.8-4 x 3.5-4 mm; middle and inner phyllaries glabrous, middle c 12-13.5 x 4.5-5 mm, innermost c 14.5-15.5 x 2.8-3 mm; apex of outer and middle phyllaries with 4-6 mm long central spine flanked on each side by 3-4 basally co­ alesced smaller spines; innermost phyllaries with one weak spine at apex, or sometimes tip merely apiculate. Corolla white, glandular. 2 .5-2.8 cm long, tube 1.9-2.2 cm long, lobes 5-6 mm long; style with a fringe of liairs at the point of bifurcation, branches 2. broad. Cypselas (only

42

145. ASTERACEAE: 13. Centaurea. 14. Goniocaulon. lS .N otobasis

immature seen) black, shiny, 5-6 x 1.8-2.2 mm. glabrous Pappus of 1-seriate bristles, c 8-9 mm long, pale white Fig. 145.26. Open bushland on shallow soils overlying rocks; 2300-2400 m. EW, not known elsewhere. Baldrati 4837, 1767; Ryding 2048.

14. GONIOCAULON Cass (1817)1 Annual herbs: stems acutely angled. Leaves alternate, ses­ sile, lanceolate, margins serrate. Capitula narrowly cylin­ drical or slightly widening towards the apex, fascicled in cory mbose inflorescences, homogamous. Phy llaries spi­ rally airanged. narrow, rigid, not spinescent. the inner ones gradually longer. Receptacle small, paleaceous. Flo­ rets relatively few, all bisexual and similar. Corolla 5-lobed. lobes of approximately the same length as tube or longer. Stamens with hairy filaments; anther bases sagittate, tails short Styles bifurcate, branches filiform Cypselas glabrous, subcy lindrical. with about 20 faint ribs forming a low rim around the pappus. Pappus multi­ seriate. scale-like, with narrowly lanceolate scabrid un­ equally long scales, surrounded bv a low, narrow pm. A monotypic genus found in India. Pakistan, Sudan and the Flora area. G. indicum (Klein ex Hi IId.) CB. Cl. (1876); Serratula indica Klein ex Willd (1803): A thanasia indica (Klein ex Willd.) Roxb (1832); A m berboa indica (Klein ex Willd.) DC. (1838) - type. India, probably Tranquebar. Klein s.n. in Heit> Willdenow (B-W holo ). Goniocaulon glabrum Cass (1818); A m berboa goniocaulon DC., nom. lllegit. (1838) - type: India. Tranquebar. unknown collector, probably Klein, sent to Jussieu from Copenliagen by A/. I'ahI in 1799 (P-Juss. no. 8303. holo ). Erect, annual herb up to c 1.7 m. from a taproot, little branched in lower parts, whole plant slightly glaucous Stem glabrcscent. with 5-7 very marked angles Leaves lincar-lanceolate or linear-oblong, sessile and partly ainplcxicaul. 7-13 x 1.4-2.3 cm. with 25-35 blunt to sharply pointed teeth or shallow lobes on both sides, glabrcscent. apex rounded to acuminate, base gradually narrowing Capitula clustered at the end of and along 3-5 paniculatclv arranged branches, sessile or on up to 1.3 cm long peduncles. 1.3-1.6 x 0.3-0.6 cm. with 4-8 flowers Phy llaries up to c 1.2 x 0.2 cm. long acuminate, pale green with darker apical parts and hyaline margins Paleae nar­ row. straw-coloured, almost bnstle-like. 0.4-0.6 cm. Co­ rolla pale purple. 1.2-1 4 cm long, 5-lobed. lobes c 0.4 cm long Cypselas finely ridged, glabrous, c 4 x 1.5 nun Pappus of flat, unequally long scales. 3-4 x c 0.7 mm. Balanites Combretum-Acacia woodland on black cotton soil, weed in Sorghum and Sesame cultivations on 1 by lb 1 rus. Botanical Museum, Gothersgade 130. Copenhagen K. Denmark

black cotton soil; 700-1150 m. TU GD; Sudan Pakistan. India. Friis et al. 6768 and 6844. Both in the Flora area and, according to literature and collector’s labels, in Sudan and India, G. indicum may form almost monospecific stands on areas with black cot­ ton soil, which have recently been cleared of vegetation, or it may invade recently harvested fallow fields. One Su ­ danese collector records it as having become a trouble­ some weed in Sesame cultivations, and this may also happen in the western Ethiopian lowlands.

15. NOTOBASIS* (L.) Cass. (1825) Annual herbs, spiny Leaves simple with lobed and spinescent margins, alternate, glabrous. Capitula discoid, subtended by 5 pinnatipartite and spinescent bracts. Phyllaries imbricate, held together by cob-webby hairs, pale green, outermost and median ending in spines. Flo­ rets pink or purple, tubular. Cypsela smooth, laterally flat­ tened Pappus muluseriate. plumose, deciduous as a unit A monotypic genus known from the Canary Islands, Madeira. N Africa. Spain, Corsica, southern Italy. Greece, Cyprus. Syria. Armenia, Turkey . Iran, and Iraq

N. syriaca (L.) Cass (1825); Carduus syriacus L. (1753); Cirsium syriacum (L.) Gaertn. ( 1791); Cnicus syriacus (L.) Roth ( 1794) 1 - type: “Habitat in Syria. Creta. Hispama.” Linnaean Herbarium No. 966.31 (LINN lecto., selected by Kupicha in Davis (ed.). FI. Turkey 5: 420 (1975)]. Annual heib. Leaves oblong or oblong-lanceolate, sessile, up to 15 x 5 cm, glabrous or sparsely hairy' beneath, base aunculate-amplexicaul, lobed, lobes irregularly triangu­ lar. ending in yellowish or whitish, 1-4 mm long spinules. margins minutely spinulose. Capitula solitary or in two’s, sessile or shortly pedunculate. globose or ovoid, 2.5-3 x 1 5-2.5 cm at anthesis, subtended by 5. up to 5 cm long pinnatipartite bracts Phyllaries glabrous or held together by cob~webby hairs, outer and median ovaie-oblong, sim­ ilar. all ending in 1—1.5 mm long spines: inner linear. gradually longer. Florets pink or pale purple. Corolla tu­ bular below. w ider above, 5-lobed, c 15-17 mm long. An­ thers c 5 nun long. Style c 17 mm long, shortly papillose w ith longer circular tuft of papillae c 3 mm below apex Cypsela brown, glabrous, obliquely obovoid-oblong. 4 7-5 8 x 2.8-3 9 nun Pappus 12-14 mm long, multi­ seriate. whitish brown, plumose, connate at base and de­ ciduous as a unit. Habitat unkown; c 2300 m; EW. Widespread in north Afnca. the Mediterranean area and in the Middle East. Pappi 74 (FT). It is not known whether this is native to Eritrea or a ca­ sual introduction. There is only a single record of this spe­ cies from Eritrea, Mai Ciuet (=Chewet), collected in October 1927 by Pappi.

43

145. ASTERACEAE: 16. Scolvmus

Tribe 3. C IC H O R IE A E Lam. & D C (1806) Lactuceae Cass. (1819) \nnual or perennial, caulescent or acaulescenl, steinless icrbs, rhizomatous or with deep-seated taproot or steins sometimes climbing; whole plant often producing milky atex. Leaves alternate or rosulate. simple to pinnatisect. inear to ovate. Capitula homogamous. ligulate. Recepta:le usually epaleate, rarely paleate with narrow pales Crepis, Scolymus), setose or fimbnllate (Cichorium. J;cris). Phyllaries multiseriate, imbricate, graded mononorphic Florets yellow, rarely white, blue, pink or purile. Cypselas grey to black, truncate to beaked, smooth to verrucose, glabrous to densely appressed-hispid. Pappus .)! erect bristles and/or soft feather-like liairs, white or yeliowish white, or absent. A tribe of about 98 genera with more than 1550 speJies found commonly in southern Europe. North Africa. Southwest and Central Asia, and western North America, :t few of the genera are tropical in origin (Dianthoseris, Launaea, Sonchus, Tolpis and Pier is)', 14 genera in the Flora area. 1 Stems with spiny wings; plants thistle-like. 16. Scolymus Stems not spiny-winged; plants not thistle-like. 2 2 Pappus of up to 2 mm long scales. 17. Cichorium Pappus of hairs or bristles, at least 4 mm long. 3 3

Hairs of at least the inner pappus feathery' (plumose). 4 Hairs of pappus not feathery, at most somewhat barbellate. 6

4 Plants glabrous at maturity, sometimes floccose when young; hairs simple; leaves grass-like. 29. Tragopogon - Plants coarsely hairy; hairs glocliidiate with 2-4 (-6) hooked barbs at apex; leaves not grass-like. 5 5. Cypselas distinctly beaked, beak yellow; outer phyllaries ovate-lanceolate, glochidiate at mar­ gins. 27. Helminthotheca - Cypselas not beaked, apex truncate or attenuate; outer phyllaries linear, margins glabrous. 28. Picris 6. Plants stemless; capitula solitary', rarely several, usually sessile amongst leaf rosette, or on long or short scapes, usually broader than long. 7 - Plants with evident stems or scapigerous; capitula usually many, rarely solitary, distinctly longer than broad. 8 7. Apex of cypselas beaked; phyllaries lengthening inwards; introduced weed. 20. Taraxacum - Apex of cypselas not beaked, truncate; phyllaries shorter inwards; plants restricted to high afroalpine meadows (alt. 3600 m and more).

19. Dianthoseris 8. Peduncle conspicuously expanded or inflated below capitula; annual herbs with broadly scarious-margined outer phyllaries; pappus of fine, down-like hairs. 24. Reichardia

Peduncle not expanded or inflated below' the capitula, annual or perennial herbs, outer phylla­ ries and pappus not with the above combined fea­ tures, the latter not or only partly of fine down-like hairs. 9 9. Cypselas tuberculate in upper parts, with a long slender beak; scapes onc-headed. hollow. 20. Taraxacum - Cypselas and scapes not w ith the above combined features, or plants not scapigerous. 10 10. Stem climbing, with numerous slender branches: leaves sub-peltate; florets white, pink or whitish yellow. 22. Prenanthes - Stem not climbing, erect; leaves not peltate; florets yellow, rarely blue or white. 11 11. Pappus dimorphic, of mixed bristles and fine down-like hairs. 12 - Pappus of bristles only, monomorphic, rarely with an outer ring of very short bristles. 13 12. Cypselas compressed, smooth or muricate, gla­ brous or with downwardly pointed liairs; ligules narrow. 25. Sonchus Cypselas columnar, patent-hairy or papillose, or glabrous and verrucose; ligules broad. 23. Launaea 13. Cypselas tnmcate. 14 - Cypselas beaked or at least somewhat attenuate above. 16 14. Florets pink or purple. 22. Prenanthes - Florets yellow, rarely blue. 15 15. Cypselas quadrangular, glabrous; pappus bnstles 4-17, separate to the base; inner florets usually much shorter, purplish-brown. 26. Tolpis - Cypselas compressed to cylindric with ascending hairs on ribs or margins; pappus bristles numer­ ous, fused at base and deciduous as a unit, florets ± uniform, yellow or rarely blue. 23. Launaea 16. Cypselas compressed and unequal-ribbed, with as­ cending, liairs on the ribs or at least on the mar­ gins (evident in microscope); involucre mostly cylindrical; plants sometimes stemless or scandent; florets yellow, blue, purple or white. 21. Lactuca - Cypselas terete and equal-ribbed, rough with as­ cending projections on the ribs; involucre usually obconic; plants never scandent, florets yellow. 18. Crepis 16. SCOLYMUS* L. (1753) Erect, thistle-like, annual or perennial herbs. Stems winged, wings spinous. Leaves alternate or radical, sim­ ple, variously lobed, spinous-dentate, thick. Capitula homogamous, ligulate, terminal or axillary, sessile or shortly pedunculate. Receptacle conical, paleaceous. In­ volucre ovoid or subglobose. Phyllaries multiseriate, im­ bricate. outermost large, exceeding the florets, stiff, mar­ gins spinous-pinnatiFid or pectinate. Paleae conduplicate. obovate. enclosing the ovaries or cypselas. Florets yel­ low; ligules truncate at apex, 5-fid. Anthers sagittate at

44

145 ASTERACEAE: 16. Scolymus. 17. Cichorium

c 10 x 3 mm. with long dark liairs externally Cypselas grev, glabrous. 2-4 nun long, obovate. Pappus absem Fig 145.27. Weed of cultivated fields, in wasteland and on roadsides. 2000-2400 nv EW TU SU; native of N Afnca S Europe and S Russia; introduced elsewhere; Sudan de Lotto 16; Afesfm T .1686; Albers 63042. Recently introduced in the Flora area.

17 CICHORIUM * L. (1753) Erect, annual or perennial herbs, stems glabrous or sparsely hispid, sap milky. Leaves alternate. Capitula homogamous. ligulate. terminal or axillary, shortly pedunculate. Receptacle flat. epaJeate or rarely sparsely fimbriate. Involucre cylindrical. Phyllaries 2-3-seriate outer few. short, inner subcoriaceous. conduplicatc and enclosing outer cypselas. Florets blue; ligules truncate. 5-dentate at apex Anthers sagittate at base. Style branches slender, apex obtuse. Cypselas glabrous, angu­ lar or compressed, obovoid Pappus of short scales or ab­ sent. A genus with 4 species, native of temperate Europe and Asia. Two species have been reported from the Flora area but only one of these is currently substantiated by vouchers. Other species may also be in cultivation. ligure 27 SCOLYM US M ACULATUS : 1 —upper branch with capitula in forks: 2 - cypsela x c 3l/i. (No specimens cited in orig.) Drawn by Ruth Koppel. (Reproduced from FI. Palaestina vol. 3, PI. 687.)

I

Plants annual; cypselas without pappus. 1 C. calvum Plants perennial, cypselas with pappus 2. C. intvbus

base, mucronate-acuminate at apex. Sty le bifurcate, arms 'penicillate at apex. Cypselas grey, compressed or angu­ lar. surrounded by, and adnate to. the obovate and winged paleae. Pappus of 2-3 bristles, or absent and represented bv an annular crown.

1. C. calvum Sch. Bip. ex Schweinf. & Asch. (1867) type: a cultivated plant raised from seeds col­ lected in Etliiopia by Schimper in the Berlin Botaiuc Garden in 1859 (?B destroyed. ?FR not seen).

Annual herb Stem erect, sparingly setose. Upper leaves oblanceolate. obtuse, rounded at the base, sessile, sparingly setose beneath especially on the midnb. unequally denti­ culate. ciliate-scabrous; die floral ones lanceolateacuminate from a cordate base. Capitula solitary, terminat­ S. maculatus L. (1753) ing stem, pedunculate, or 2-3 together, sessile. Phyllaries type: "Habitat in G. Narbonensi, Italia.” acuminate, setose-cilate. sparingly tuberculate-scabrous on Linnaean Herbarium No. 963.1 [LINN lecto.. selected the back, more or less callous at the base; the outer ones by Jeffrey in Jan is & al. (ed), Regnum Veg. 127 86 ovate-oblong, shorter, the inner ones oblong-lanceolate. (1993)). Cypselas straw-colored, obovate. angular, subcomprcsscd. altogether without pappus, c 1 nun long. Erect glabrous, stiff or rigid annual herb, up to 1 m high. Cultivated; altitude not known, but presumably from Stein branched near the apex, whitish, spinose-winged by the Ethiopian highlands. Region not known; not known the decurrence of the leaves. Radical leaves widely elsewhere. spathulate. spinose-dentate. shortly petiolate. 25-50 x 10 cm. Cauline leaves oblong, sinuous-pinnatifid. sessile. The above description is taken from Oliver & Hiern. 5-10 cm long, margins cartilaginous, apex spinous. FI. Trop. Afr. 3. 446-447 (1877). No specimen was Capitula solitary and terminal or axillary. Involucre c 15 x found to satisfy this description. The description seems 10 mm. Outer phyllaries green, margins spinousto combine the cliaractcrs o f C. endivia L. and ('. intvbus pinnatifid or pectinate, 10-15 x 3-5 mm. glandular pubes­ L.. the former is grown as a salad plant and the latter is cent. Inner phyllaries linear-lanceolate, c 15 x 4 mm, mar­ cultivated in many parts of the world for its edible roots. gins entire, apex acute and pointed. Ligules lemon yellow. The two species are very similar, with C. endivia differA genus with 3 species naturally distributed in the Mediterranean area It resembles Carthamus in its spiny habits. One species in the Flora area.

145. ASTERACEAE: 17. Cichorium

45

Figure 28 CICHORIUM INTYBU S 1 - habit x c 1/ 20; 2 - flowering axis x c V2: 3 - cypsela with pappus x 14. 4 phyllary, outer surface x 3V2. 1 from H ubbard 11476; 2 -4 from Whellan 2099. Drawn by Eleanor Catherine. (Re­ produced with permission from FI. Zamb. vol. 6(1). Tab. 35.)

tg in having usually less dissected leaves and longer • -ppus (\.e.. cypselas only 2-5 times longer than appus). Schweinf. & Asch. (1867) noted that C. intybus may probably be similar to C. endivia. Until more speci1 ;ns are obtained of the taxa from the Flora area, C. Ivum is retained with its original description as a taxon istinct from C. intybus. See also the notes on this by Uerxmiiller in Milt. Bot. Slaatssaml. Miinchen 1 7 -1 8 . 7 (1957).

lecto., selected by Lack in Rechinger (ed.), FI. lranica 122: 6(1977)).

Erect, perennial herb, up to 1 111 high, with a stout taproot. Stem and leaves hispid-pubescent to glabrescent. Basal leaves rosulate, shortly petiolate, oblanceolate. margins denticulate to runcinate, usually 8-15 cm long, rarely up to 35 x 10 cm; cauline leaves similar but sessiie. smaller, auriculate-amplexicaul. Capitula 2.5-3.5 cm wide at anthesis. axillary or terminal at the apex of thickened peduncles. Involucre cylindric. Outer phyllaries ovatelanceolate, often spreading. 7-8 x 2-2.5 mm at anthesis. 2 C. intybus L. (1753) type: “Habitat in Europa ad inargines agrorum sparsely glandular hairy , margins narrowly scarious. sometimes tinged purple. Corolla tube hairy, c 0.1 mm viarumque.” Linnaean Herbarium No. 962.1 {LINN.

46

145. ASTERACEAE: 17. Cichorium, 18. Crepis

long. Cypselas light brown. 1.5-2 x 0.5-0.8 mm, gla­ brous. c 10-ribbed, truncate at apex. Pappus of a series of 0.1-0.3 mm long, narrow, while, scales. Fig. 145.28.

-

Cypselas 6.5-12 mm long; peduncle sparsely white or brownish tomentose with black, inter­ mixed. long hairs. ligule 10-18 mm long.

Cultivated and sometimes remaining as a weed from previous cultivation in well drained farmland. 1800-2400 m. EW SU HA; cultivated and escaped as a weed in many parts of the world IECAMA $. rhizomatous or with deep-seated taproot: scapes pubescent or glabrous, erect or decumbent, branched Leaves caudical and/or cauline; caudical leaves numerous, usu­ ally rosulate, entire to pinnately divided, petiolate, persis­ tent or ephemeral, cauline leaves few or absent. Capitula homogamous. solitary orcymosely arranged, ligulate. cylindnc to campanulate or ureeolate. pedunculate. Recep­ tacle flat or convex, arcolate and epaleaceous. or alveolate with ciliate or narrowly membranous paleae Phyllaries 2to several-seriate, glabrous, tomentose, setose or hirsute, outer ones shorter than the inner, ovate-lanceolate, inner ones linear-lanceolate, in 2 or more series. Florets few to many, ligulate, yellow tinged reddish: ligule often tw o-third the length of the whole corolla teeth 4 or 5 An­ thers yellow, base sagittate. Style branches filiform, yel­ low. truncate or attenuate at the apex. Cypselas black or browa subterete or angular, usually narrowed above into a beak, often 10-20-ribbcd or striate, glabrous or spiculate. Pappus of numerous, white or light brown, barbellate or scabrous, persistent or deciduous bristles. A genus with c 200 species found mostly in temperate areas of the world; 7 species are known in the Flora area 1 Plant annual, tap root slender - Plant perennial; tap root stout, woody. 2

-

Stem and leaves densely hispid-setose: involucre 8-9 mm long at anthesis. ligule c 10 nun long: phyllaries callose-thickened in fruit. 1. C. foetida Stem glabrous or sparsely setose, leaves pubescent. liairs fine; involucre 9-13 mm long at anthesis: ligule 13-14 mm long, phyllaries not thickened in fruit. 2. C. tencrrim a

3. Stem leafy1throughout, luspid-scabrous. 5-10 mm wide, angled. 3. C. schultzii - Stem with few leaves, sparsely setose. 2-5 mm wide, terete; leaves mostly radical, stem leaves few. small 4

Cypselas 5-6 mm long; peduncle with dense brown woolly indumentum near summit; ligule 5-6 mm long. 4. C. carhonaria

7. -

Cypselas 7-9 mm long; ligule 10-14 mm long: in­ volucre 9-13 x 4-8 mm. Cypselas 11-12 mm long; ligule 16-18 mm long; involucre 13-15 x 9-11 mm. 6. C. achy rophoroides Stem leaves pinnatifid. ligule without hair. 2. C. tenerrima Stem leaves denticulate; ligule hairy at least in lower half. 7. C. xy lorrhiza

1. C. foetida L. (1753) type. "Habitat in Gallia." ? Gouan s.n. in Linnaean Herbarium. (This specimen seems to have been lost. Babcock (in Univ. Calif. Publ. Bot. 22: 688 691. pi. 17a, 1!M9) and Lamond (in Davis (ed.), ft. Turkey 5: 831 1975) designated Linnaean Herbarium No. 955.6 (LINN) as lectoty pe, while Jeffrey (in KeM Bull. 18. 462. 1966) designated Linnaean Herbarium No. 955.9 (LINN). However, both collections lack the relevant Species Plantarum number (i.e. “ 13”) and are ev idently post-1753 additions to the herbarium). Barkhausia schimperi Sch. Bip. ex A. Rich (1848); Crepis schimperi (Sch. Bip. ex A. Rich.) Schweinf. & Asch. (1867); Hieraciodes schimperi (Sch. Bip. ex A Rich.) O. Kuntze (1891). nom. illegit - type TU near Adwa. Memessah. Quart in Dillon s.n. (P sy n ); TU. near Adwa. PI. abyss. 1, Schimper 295 (P syn.. K isosyn ): SU. 'Choa .Petits.n (Psyn.) Crepis eritreensis Babe. (1938) - ty pe: EW. E of Mt Bizert Mogod. Schweinfurth & Riva 2031 (US holo., K P iso ) Erect, annual herb. 10-40 cm liigh. Stem and branches hispid throughout, with whitish setiform hairs, usualh much branched: branches ascending or spreading. Leaves simple. Jyrate-pmnatifid to deeply pinnatifid, oblanceo­ late. up to 12x3.5 cm. hispid with whitish setiform liairs; apex acute; margins of simple leaves dentate, base attenu­ ate into a narrowly winged petiole; upper leaves lacimate-subauriculatc. Capitula cylindric-turbinate. nod­ ding when young, erect at anthesis and in fruit, often 2-3 per branch or stem, arranged in cory mbose cymes; pe­ duncle 1—5(—10) cm long, pubescent with denser white hairs near the rcccptacle and 1-2 mm long setiform liairs all along ridges. Involucre 8-9 x 6-7 mm at anthesis. up to 10 x 8 mm in fruit Phyllaries 2-3-seriate. densely white pubescent with setiform liairs. outer 10-11, linear. 2-2 5 x 0 3-0.5 mm at anthesis. up to 3.5 nun long in fruit: inner 12-13, oblong-linear w ith acuminate purplish apex. 7-8 x 1-1.2 mm at anthesis. up to 11 x 2.5 nun in fruit, cartilaginous-thickened medially. Ligule c 15 x 1 mm, tube c 5 nun long, glabrous below', sparsely hairy above. Cy pselas reddish to dark brown, 8-11 x 0.5-0.8 mm. 15- ribbed, slightly tapered near the base, gradually attenuate into a

145. ASTERACliAE: 18 Crepis

47

Figure 29 CREPIS FOETID A: 1 habit \ ' v 2 capitula after fruit dis­ persal x c2 \ 3 inner phyllary \ 3 ‘ 2. 4 detail o f rcceptacle x 21; 5 & 6 cypselas x 7 'i; 7 pappus bristle x IVi. 8 - caudical leaf x c 1:9 4 cauline leaves x c I : 10 part o f in­ florescence x c 2; 11 2 nearly ma­ ture capitula x c 2. 12 floret without ovarv x 3'4: 13 detail o f ligule teeth x c 50; 14 stamens x 7’ 2; 15 anther base x 30. I 7 from Schweinfurth ith wide variation in nearly all features. Kilian (1997) who lias recently studied this as well as related species.

61

stated that it is “... well recognizable by its small, pale yel­ low (never golden yellow!) florets (with short anther tubes, 1.1-1.3 mm long) in medium-sized capitula. which have wiry (never capillaceous!) peduncles or are clustered and sessile.” For distinction from L. petitiana, see note under the latter species. 6. L. petitiana (A. Rich.) N. Kilian (1997); Lactuca petitiana A. Rich. (1848) - type: TU Djeladjeranne, Sept. 1840, Schimper 1448 (P lecto., BM K isolecto.). Microrhynchus hochstetteri Sch. Bip., nom. nud.; hi. octophyllus Hochst. var. (b) quoad Schimper 1448; Sonchus hochstetteri Sch. Bip. ex Schweinf. & Asch., nom. illeg. (1867) - type: SU, kChoa", Petit & Quartin-Dillon s.n. (P syn.); TU Djeladjeranne, Schimper 1448 (P svn.; selected as lectotype by Kilian (1997)1. Sonchus octophyllus Schweinf. & Asch. (1867), nom. nud., quoad Schimper 1448. Perennial herb, 10-75 cm high, with a slender trailing rhi­ zome. Stems one to several arising from a short caudex, decumbent or ascending to erect, openly branched, leaf­ less to leafy. Leaves crowded at base, or rosulate, (3-) 6—18(—27) x (0.5)2-6(-9) cm. obovate to narrowly spathulate or almost elliptical in outline, with wliitecartilaginously, sinuate-dentate to runcinate margins, soft and thin to somewhat leathery, tapered at base. Cauline leaves variable, narrowly lanceolate or rarely ribbon-like to broadly spathulate, sinuate-dentate to runcinate, with clasping, amplexicaul base. Capitula solitary or clustered at the nodes; peduncle absent or up to 15 mm long, densely bracteate. Involucre 11-16 x 2-3 mm at anthesis. Phyllaries 3-4-seriate; outer ones 8-12. broadly ovateacute. 1.5-2 nun long; with broad scarious margins, con­ tinuous with bracts on peduncle; inner ones 11-15 x 1-3 nun, linear-lanceolate, with broad, scarious margins. Lig­ ules golden yellow with tinges of violet on the veins, 7.6-10 x 1.5-2.5 mm; anthers heteromorphic, (3-) 3.4-5.8 x 0.5-1.0 mm, mature ones sometimes intermixed with thinner, pale, sterile ones; marginal ones compressed and curved, columnar or slender-cuneate. 5-ribbed; inner ones columnar, apically truncate to cuspidate. 4-angled with 4 ribs, transv ersely and sharply wrinkled. Pappus 6-9 mm long, persistent, white, dimorphic, with numerous dow ny outer bristles and a smaller number of glabrous in­ ner ones. Wooded grassland, open Acacia bushland on volcanic soil, grassland, dry' river banks, open busliland on dry limestone hills, rarely weed of cultivations; 1000-3000 m. EW TU WU SU AR SD HA; Sudan. Somalia. Kenya. NTanzania. (filbert ei al. 7412;A Anderberg 1606; Bur­ ger 2209. Launaea petitiana has. for a long time, been consid­ ered synonymous with either L. intybacea or L. taraxacifolia. The recent study by Kilian (1997) on both herbarium specimens, field collected material and plants raised in the Botanic Garden Berlin-Dalilcm lias con-

62

145 ASTERACEAE: 23. Launaea

finned that it is a distinct entity which can usually be readily distinguished from the annual L. intybacea by the different habit, the colour, size and shape of the ligules, as well as the length of the anther-tube...” and from L. taraxacifolia by the‘\.. number of inner involucral bracts" which is "constant in L. petitiana (being 8) and i taraxacifolia (being 5)”. Kalian found 3 cy to types in ma­ terial from S Somalia and also recognized 3 “aberrant forms due to putative hybridization and introgression of L petitiana with L. taraxacifolia Sonchus hochstetteri is an illegitimate name because it shares a type specimen with an earlier name, Lactuca petitiana, the epithet of which has to be adopted.

7. L. massauensis (Fresen.) Sch. Bip. ex Kuntze (1891); Heterachaena massauensis Fresen. (1839); Lactuca massauensis (Fresen.) Sch. Bip. ex A. Rich (1848); Sonchus massauensis (Fresen.) Schweinf & Asch. (1867) - type: EE, Massaua. 3.1832. Riippell s.n. (FR holo.). Erect, annual herb, up to 50 cm high. Stem slender, repeat­ edly dichotomously branched, glabrous, terete, glaucous, leaves mostly radical, simple with denticulate margins, to deeply pinnaiifid with rounded lobes, oblanceolate in out­ line. nanowed into a broad petiole, up to 12x5 cm; mar­ gins denticulate with white, callous-thickened teeth, cauline leaves sessile, cordate to auriculate-amplexicaul Capitula c 10-flow ered. solitary at apices of branches, ar­ ranged in lax paniculate cymes. Involucre cylindric. 8-9 x 1-1.5 mm at anthesis. Phyllaries 3—t-seriate, light green, tinged purple, glabrous or some with sessile glands, mar­ gins scarious; outer ones deltoid to linear. 2.8-3 x 0.8 nun. inner ones c 5, oblong-linear. 8-10 x 1 nun Ligules pale yellow, 4-5.6 x 0.8-1.1 mm; anthers without basal ap­ pendages. 0.8—1(—1.2) mm. Cypselas black, heteromorphic. transversely rugulose. compressed to subterete. attenuate more tow ard the apex, 4-5-ribbed on each face, smooth, 2.5-2.8 x 0.4-0.6 mm. Pappus w liite, dimorphic, of stiff, smooth, bristles and flexible (downy) hairs. 4-5 nun long. Grassland with red sandy soil or on gypsum, in wadis; 60-2400 m. EW EE TU SU AR GG SD BA HA: Sudan. Somalia, Socotra. N Kenya, Egy pt, Yemen. Saudi Arabia. W Pakistan. Ballv 6859, Gilbert & Thulin 993; Gillett 14399

8. L. pseudoabyssinica (Chiov.) M. Kilian (1997); Lactuca pseudoabyssinica Chiov. (1904) |as kpseudo-abyssinica] - type: EW, Mt. Bizen, 2000 m. Schweinfurth & Riva 2054 (FT syn.. K isosyn); EW. Mensa, Rora Ualicaue, 1900 m., Terracciano & Pappi 626 (FT syn., K photo). Lactuca boranensis Cufod. (1943); Launaea boranensis (Cufod.) Cufod. (1967) - types: SD, Mega Corradi 1906\ Corradi 1736, 1877, 1878, 1879, 1906, (FT syn, K photo). Launaea massauensis auct., non (Fresen.) Sch. Bip. ex Kuntze: C. Jeffrey (1966) p.p.

Perennial he/b, 15-50 cm high, from a short wood\ caudex. Stems one to several, each arising from the ape\ of the caudex, slender, erect, leafy or leaf-less, branchec from below- Leaves both rosulate and cauline. Rosulate leaves 3-9 x (0.5-) 1-3 cm, narrowly spathulate to nar rowly elliptic or linear-lanceolate in outline, with a nar row seini-amplexicaul base, sinuate-dentate to pinnatific segments nanouly triangular with acute to acuminate apex Cauline leaves entire, linear-lanceolate or sinuatedentate or pinnaiifid. margins denticulate, semi-amplexicaul to clasping at base. Capitula solitary and cymoseiy arranged, each with 7-12 flowers; peduncle 0.5-2(-5) cm long, capillaceous. spreading-erect. Involucre (7—)9—11 x 1-1.5 mm at anthesis. Phyllaries 3-4-seriate; outer ones 6-8, ovate, subimbricate. with broad, scarious margins, inner ones 7 -1 1 \ 1—1.5 mm, with narrow scarious mar­ gins Ligules bright yellow. 6.5-9 x 1.5-1.9 mm; anthers without basal appendages, 2.1-3.2 mm long. Cypselas brown to dark-brown, heteromorphic, 2-3.6 x 0.4-0.7 nun, outer ones curved and compressed, sub-fusifonn but more tapered toward the apex, 5-ribbed. transversely sharply wrinkled, apex cuspidate to shortly rostrate; inner sub-cuneate to columnar. 4-angled, 4-ribbed. smooth to slightly wnnkled Pappus dimorphic. 4-6 mm long, per­ sistent, of numerous dow ny and smaller number of erect bnstles. Woodland or grassland on limestone or black clay, of­ ten between rocks of sandstone, granite or basalt; 1700-2400 in. EW EE SU AR SD HA; Somalia. Djibouti and N Kenya. Mooney 5551; Burger 2029; Thulin 1563 Launaea pseudoabyssinica was kept as a synonym of L. massauensis Jcf. Jeffrey in Kew Bull. 18. 474 (1966); Cufodontis (1967:1183)|. However. Kilian (1997) wrote that “ .. from L. massauensis it is clearly distinct through its perennial habit, general appearance, the somewhat larger capitula. and. as a particularly reliable diagnostic feature, through its distinctly larger florets with longer an­ ther tubes (2.1-3 2 mm versus 0.8-l(-1.2).” As regards variation within this species, Kilian w rote “by far the most gatherings from this region have typically linearlanceolate to ribbon-like and only sinuate-dentate cauline leaves like the type collections of L. pseudoabyssinica from Eritrea In other regions of the distribution area (i.e . Mega and Harar area. Ethiopia, and N Somalia) the plants liave mostly pinnaiifid leaves like the type specimen of L boranensis However, no conelation with other cliaracters could be found, and both leaf shape are moreover linked by intermediate forms.” 9 L. taraxacifolia (Willd.) C. Jeffrey ( 1966); Sonchus taraxacifolius Willd. (1803); Lactuca taraxifolia (Willd.) Schumacher ex Homem. (1819) tvpe: Ghana/Benia l(ex-Danish) Guinea). Isert s.n. (B-W1LLD no 14538 holo ). Microrhynchus pentaphvllus Hochst., nom. n u d . quoad Kotschy 337 (Sudan, Seimaar, BM K), Lactuca pentaphylla Sch. Bip.. nom. nud.. quoad Kotschy 337 (BM); Sonchus pentaphyllus Schweinf. & Asch. (1867). nom. nud., quoad Kotschy 337, 321 and 153 (BM); Lactuca kotschyi Sch. Bip., nom. nud.

145. ASTERACEAE: 23. Launaea, 24. Rcichatdia

63

Lrect, perennial herb, up to 1 m high. Stem glabrous, erete, glaucescent, branched at apex. Leaves mostly iasal, glabrous, runcinately pinnatifid often with broadly deltoid triangular lobes, narrowed into a broadly-winged ■etiole, up to 20 x 6 cm, margins denticulate with sharp allous-thickened teet, cauline leaves sessile, amplexiaul. Capitula c 20-flowered, each on a short peduncie and everal grouped together in paniculate cymes, peduncle • ith few bracts. Involucre cylindric-campanulatc, 10-11 2.5-3 mm at anthesis. Phyllaries 3-4-seriate, glabrous. ;ght-green, outer ones continuous with bracts on i -eduncle. deitoid to ovate-lanceolate, 1.5—4.5 x 1—1.5 im, with broad scarious margins, tliickened at base, inner ones c 5, linear-oblong, 9-9.5 x 2-2.5 mm, with narrow Scarious margins. Ligules golden yellow, 6-7.2 x 1.6-2.2 mm; anthers with basal appendages, 2.4-3 I mm long. Cypselas black, heteromorphic, narrowly elliptic, taperi ig to a short, slender, pale beak. 3.3-3 5 x 0.6 mm. t ansversly rugulose. glabrous, c 5-ribbed on each face. Pappus white, dimorphic, of 7-7.5 mm long, erect, t aibellate bristles and slender hairs.

apex. Pappus dimorphic, of erect, barbellate bristles and soft liairs. white, 8-8.5 mm long. Fig. 145.40.12-19.

Moist grassland, weed of irrigated fields and cultiv ations; 550-1800 m. WU GJ GG SU IL; Senegal. Sierra 1 s>ne, Ghana, Benin. Nigeria. Sudan. Tanzania. Pichitermolli 2145; Parker 523; Coady K23.

Erect, perennial herb, up to 30 cm high. Stem branched only in the inflorescences. Leaves both basal-rosulate and cauline. oblanceolate, simple and coarsely toothed to pinnatifid with closely lobulate-serrate segments, gla­ brous, narrowed at base into a winged petiole, margins with callose-thickened teeth, cauline leaves with tabulate-serrate, auriculate-amplexicaul base. Capitula c 15-flowcred, in open paniculate cymes; peduncle gla­ brous. with several lobed bracts. Involucre campanulate. 9-10 x 4.5-5 mm at anthesis, up to 6 mm wide at base in fruit. Phyllaries 3-4-seriate, glabrous or with whitish powder, dark-purplish at base and at apex; outer ones ob­ long with acuminate, callose-thickened apex, 2-3 x 1-1.5 mm; inner ones oblong with acuminate apex, 7-9 x 1.3—1.5 nun, callous-thickened at base in fruit. Ligules bright yellow, dorsally greyish with blackish veins, 7-10 x 1.8-3.2 mm, anthers with basal appendages, 2.2-2.8 (-3) mm long. Cypselas heteromorphic, columnar, greybrown with brown annular truncate apex, 4-4.5 x 0.5-0.6 nun. glabrous or puberulous. Pappus dimorphic, of white smooth bristles and soft, flexible liairs. 7-8 mm long. Fig. 145.41.

Engels and Goettsch [in Engels, J.M., Hawkes, J.G. & I lelaku Worede (eds.). Plant Genetic resources o f Ethio/ ia: 175 (1991)] reported the use of this plant as salad in t -e Konso region of Ethiopia. See note above fordistinct on from L petitiana.

1 ). L. hafunensis Chiov. (1929) - type: Somalia, Hafun, Puccioni & Stefanini 12 (FT holo., K photo). L. cufodontii Lanza (1939) - type: SD. Mega. Cufodontis 650 (FT holo., K photo). V oody, perennial heib, up to 30 cm high (up to 50 cm in Somalia). Stem densely dichotomously branched, branches spreading, light green (of current season’s s owth) or light purplish brown (in older parts), glaucous. L eaves semi-succulent, deeply pinnatifid, glaucous, up to 7 \ 5 cm, glabrous, lobes coarsely dentate, 2-4 mm wide, h near, sometimes bilobed at base. Capitula c 20-flowered, a ranged in laxly branched cymes. Involucre cylind lc-campanulate, 8-9 x 2.5-3 mm at anthesis, c 4 mm v de and cartilaginous thickened at base in fruit. Pits llaries 3-4-seriate, light green with white puberulous h iirs. outer ones deltoid-ovate, 1.6-2.7 x 0.6-1.2 mm, some with sessile glands, concolonous, apex with tufted v :ite hairs inner ones oblong-linear, 10-11 x 1.5-2 mm. s metiines puberulous only at apex or glabrous, purp : -tinged and narrowly scarious at margins, calU ns-thickened at base in fruit, inflexed. Ligules bright y :lJow, flushed reddish on outer surface, 7-10 x 1.9-2.5 mm; anthers without basal appendages. 2.9-4 8 mm long; s igmas black. Cypselas grey-brown, slightly heteromcrphic, oblong with convex outer surface, 3.9-4 x 0 S-0.9 mm, c 6-ribbed on each face, papillose, truncate at

Lowland and montane grassland; SD HA; 500-2000 m; Somalia Socotra Kenya Tanzania Yemen, Oman. Friis et al. 8471; Popov 1091; Gilbert 2131. Launaea hafunensis is extremely polymorphic as re­ gards foliar morphology. The basal leaves are entire and linear-spathulate to deeply pinnatifid or bipinnatifid with irregularly and shallowly sinuate-dentate margins. Cauline leaves are found at the lower nodes only; the low­ ermost are similar to the basal leaves but the upper ones are linear to almost ribbon-like and entire.

IL L . m ucronata (Forssk.) Muschl. (1912); Leontodon mucronatum Forssk. (1775) - type: Egypt, Forsskal in Herb. Forssk. 1401 (C syn.) & 1402 (C syn.). subsp. m ucronata

Dry sandy wadis, coastal plains on sandy soil in cultivations; 10-70 m. EE; Chad, Sudan, M auritania Mo­ rocco, Libya Egypt, Iraq. Iran, Jordan. Bahraia Saudi Arabia. Parkin 20; Bally 6872, 6874. Kilian (1997) recorded this as an “annual (to occasion­ ally pauciennial) herb” and also wrote that L. cornuta subsp. cassiana (Jaub. & Spach.) N. Kilian has similar distributional range as the type subspecies but it is ‘vmore restricted” ecologically.

24. REICHARDIA** Roth (1787) Picridium Desf. (1799) Erect, glabrous, annual or perennial herbs. Leaves alter­ nate or radical, entire to pinnatisect with dentate or denticulate margins. Capitula homogamous. ligulate. on

64

145. ASTERACEAE: 23. Launaea. 24. Reichardia

Figure 41.

LA UNAEA MUCRONA T A X - upper stem part. 2 portion o f lower stem with leaves; 3 capitulum , 4 - cypsela x \2X/2. 1-3 specimens not given jn orig.; 4 from Plowes 2068. 1-3 drawn by Ruth Koppel (reproduced from FI. Palaestina, vol. 3, PI. 7 2 8 ,4 drawn by Eleanor Catherine (reproduced with permission from FI. Zamb ., vol. 6(1). Tab. 40E (as L. cornuta).

long, apically swollen peduncles. Involucre cylindrical to campanulate. Phyllaries imbricate, several-seriate, with scarious margins; the outer ones continuous with bracts on peduncle. Receptacle epaleace. naked. Florets pale to bright yellow. Ligules sometimes grey or purplish on outer surface, truncate. 5-dentate at the apex. Anther base sagittate. Style bifurctate. branches slender Cypselas brown, glabrous or tuberculate. 4-5-ribbed. transversely rugose, slightly constricted but not beaked at apex Pappus white, of numerous, smooth, bristles, connate at base and deciduous together. A genus w ith 8 species native of the Mediterranean area and the Middle East; 1 species in the Flora area.

R. tingitana (L.) Roth (1787); Scorzonera tingetana L. (1753); Sonchus tingetanus (L.) Lanv (1792); Picridium tingetanum (L.) Dcsf. (1799) - type: “Habitat in Tingide [Tangier|.” Linnaean Herbarium No. 947.10 [LINN, lecto., se­ lected by Jeffrey in Kew Bull. 18 477 (I966)[. Picridium abyssinicum Hochst.. nom. nud . R. tingitana (L.) Roth var. abyssinica (Hochst.) Chiov. in Ann. R. 1st. Bot. Roma 8. 199 (1904) - type. Ethiopia jexact region not known], Schimper 2107 (FT holo., K iso.). Scorzonera orientalis L. (1763); Reichardia orientalis (L.) Hochr. (1904); Reichardia tingitana (L.) Roth var. orientalis (L.) Fiori in Fiori & Paoletti. Flora Analitica d ’ltalia 3: 425 (1904) - ty pe: “Habitat

145. ASTERACEAE: 24. Reichardia, 25. Sonchus

65

[inOnente. Hasselquist.]” Herb. Hasselquist, No. 672 [UPS lecto., selected by Jeffrey in Kew Bull. 18: 477 (1966)]. :rect, glabrous, annual herb, up to 75 cm high. Leaves iblong, oblong-ovate or lanceolate or the lower ones oovate, upper leaves cordate-auriculate, semi-amplexiaul, entire to pinnatifid, with dentate-denticulate to ieeply and irregularly serrate margins, up to 1 5 x 4 cm, liabrous, glaucous, sometimes violet or purple-tinged. Inolucre 14-15 x 15-16 mm at anthesis. Phyllaries broadly v ate to lanceolate, outer ones 6-8 x 4-5 mm, with broad carious margins; inner ones 12-15 x 4-5 mm. Florets ellow. Cypselas grey-brown, oblanceolate, 1.6—1.8 x 5-0.6 mm, transversely rugose or tuberculate, truncate i apex. Pappus white, of 3-7 mm long, smooth or mi­ nutely barbellate bristles. Fig. 145.42. Roadside margins, waste ground, weed in farmland, nver banks; 200-2400 m. EW TU GD WU SU SD AR TA; a Mediterranean species distributed from the Canary slands eastwards to Polynesia, and southwards to Australa; introduced in Sudan, Somalia, Socotra, Kenya and Tanzania. Ash 2141; Parker 505; Thulin 1296.

25. SONCHUS L. (1753) B oulos in Bot. Not. 125: 2 8 7 -3 1 9 (1972); ibid. i 56-196 (1973); ibid. 127: 7 -3 7 & 402-451 (1974).

126:

Annual or perennial heibs. Stems sometimes woody at the :>ase. Leaves radical or alternate, simple to bipinnatisect; .auline ones often auriculate-amplex ligulate, pedun­ culate. Receptacle epaleate. Involucre cylindric or campanulate. Phyllaries multiseriate, glabrous or glandu­ lar hairy Florets yellow, often tinged red or pink on jutside; corolla tube hairy near the junction with ligule. Anthers sagittate at the base. Style bifurcate, branches densely papillose. Cypselas more or less flattened, elliptic to oblanceolate, ribbed, sometimes transversly rugose, glabrous or spiculate. Pappus multiseriate, dimorphic, ■ith fine flexible liairs and erect bristles. A genus of 60 species with world-wide distribution but mainly in Africa and the Canary Islands; 9 species in ihe Flora area. 1. -

Plants annuals. 2 Plants perennials. 5 Cypselas rugose or verrucose, oblanceolate. often under 1 mm wide, each surface with 3 main ribs; tube of ligule equal to or shorter than limb. 3 Cypselas smooth, oblong-elliptic, appearing winged, often 1.4-1.8 mm or more wide, each surface with many and variable number of ribs; tube of ligule much longer than the limb. 4 Leaves pinnately lobed. terminal segment ovate or deltoid and widest; limb of ligule equal to the tube. 7. S. oleraceus Leaves bi-pinnately lobed, terminal segment linear or linear-lanceolate, same width as the others or less wide: limb of ligule much longer than tube. 8. S. tenerrim us

Figure 42. REICHARDIA TINGITANA. 1 & 2 - habit, 2 is a depauperate form usually encountered in drier and/or degraded areas; 3 - phyllary x 114; 4 - cypsela with detached pappus x 4'/2. I from G. Aweke & Gilbert 828; 2 -4 , specimen not given in orig. 1 drawn by Damtew Teferra; 2 -4 drawn by Ruth Koppel. (2-4 reproduced from FI. Palaestina , vol. 3, PI. 745.)

4. Involucre of flowering heads 10-12 x c 8mm; leaves simple to pinnatifid. 5-12 cm wide in wid­ est part, terminal segment deltoid. 4. S. asper - Involucre of flowering heads c 9 x 5 mm; leaves serrate to lobulate-dentate, l-4(-4.5) cm wide. 9. S. gigas 5. Involucre of flowering heads 4-5 mm wide; outer phyllaries densely tomentose with interspersed, erect, purplish, gland-tipped hairs, apices often black. 6 - Involucre of flowering heads 6-12 nun wide; outer phyllaries pilose, if tomentose, only at base, liairs cglandular but sessile glands may be present, concolorous or purplish tipped. 7

iff------

66

—

145. ASTERACEAE: 25 Sonchus

-

Leaves pinnatisect or bipinnatisect, ovate-elliptic in outline; outer phyllaries cartilaginousthickened at base, all or most inflexed and cover­ ing receptacle in fruit. 3. S. luxurians

10 Leaves up to 23 x 3.5 cm; involucre 11-17 x 6-12 mm 1. S. schweinfurthii Leaves up to 15 x 1.5 cm; involucre 10-11 x 5-6 mm 2. S. bipontini

1 S. schweinfurthii Oliv. & Hiern (1877) type: Rep Dem. du Congo, Schweinfurth 3460 (K holo ). Erect, perennial herb, up to 1.5 m high Stem often un­ branched, glabrous, hollow, light green, lower parts up to 10 mm in diameter Leaves simple, linear-lanceolate to oblanceolate, sessile, semi-amplexicaul, auriculate with acute or apiculate auricles, glabrous, glaucesent, up to 23 x 3.5 cm; margins regularly serrate with recurved callose-tluckened teeth. Capitula arranged in subumbelliform cymes; peduncle up to 3 cm long. Involucre campanulate, 11 17 x 6-12 nun, densely white tomentose below Phyllanes 3-4^seriate. thinly pilose with or with­ out gland-tipped hairs, grey-green, purple tinged; inner phy llaries up to 13 x 2.5 mm. Florets yellow. Cypselas grey-brown, smooth or rugulose, 2 .8-3 x 0 .5-0.7 mm, ob­ long-elliptic, 4-stnate on each face. Pappus white, 6-7 mm long Fig. 145.43.

Figure 43 SONCHUS SCH W EIN FU RTH II I - habit x «/«o. 2 -3 - m id'caulinc leaves x */3; 4 - cypsela x c 10; 5 - cypsela with pappus x c 1 1 from Brummitt 9164; 2 from Richards 452; 3 from Fries et al 2432; 4 & 5 from Richards 1245. Drawn by Eleanor Catherine. (Rearranged and reproduced with permission from FI. Zamb., vol. 6(1), Tab. 41 p.p.)

6.

-

Plants 10-15 cm high; stem 6-12 mm thick with few thick, short branches near ground level; leaves 3-6 x 1-8 cm, terminal segment obtuse at apex. 5. S. ohtusilohus Plants 15—45 cm high; stem up to 5 mm thick, pro­ fusely branched with slender branches; leaves 4-8 x 1-2.2 cm, terminal segment acute to elon­ gate-attenuate at apex. 6. S. melanolepis

7. -

Cypselas smooth, grey or brown. Cypselas rugose, rugulose or verrucose, brown.

8 9

8.

Involucre of fruiting head c 17 x 12 mm; leaves lin­ ear lanceolate, 8-15 times as long as broad.

-

Involucre of fruiting heads up to 12 x 8 mm; leaves 3-4 times as long as broad. 4. S. asper

9.

Leaves entire to lobulate-dentate, linear-lanceo­ late; outer phyllaries membraneous, totally re­ flexed in fruit exposing receptacle. 10

1. S. schweinfurthii

Tall grassland, nver banks, marshland, mountain slopes, roadside banks. 1310-2000 m. SU WG KF; Su­ dan. E Africa. Cameroun. Nigeria, Guinea, Central Afri­ can Republic, Rep Dem. du Congo, Burundi, Malawi, Angola. Zambia. Zimbabwe. Gilbert & Thulin 618; Ash 1987; Mesfin T & Kagnew 2476. 2. S. bipontini Asch. (1867); S. lactucoides Sch. Bip. ex A. Rich (1848), non Bunge (1835), nom. illegit. - type. GD, Simen Mts., Dschenausa. PI abyss. II, Schimper 1299 (P holo., BM K iso ). S. bipontini forma elata polycephala ... Vatke in Lmnaea 39. 517 (1875), nom. non rite publ. - based on: GD, Gaffat (near Debra Tabor], 1863, Schimper 1239 (B holo destroyed). S. bipontini forma minor pauciflora ... Vatke in Lmnaea 39: 517 (1875), nom. non rite publ. - based on: TU. Mt. Hedscha. 1862, Schimper 355 (B holo. de­ stroyed) S. bipontini Asch. var. glanduligerus auct.. non (R E Fries) Robvns, FI. Sperm. P.NA. 2, Symp.: 596 (1947). Boulos(J 974. 412). Perennial herb, 0.5-2 m high. Stem often tinged red, wood> at base and hollow in upper parts, branches strag­ gling through bushes and shrubs. Leaves simple, linear to narrowly elliptic, up to 15 x 1.5 cm, tapering to a narrow, winged petiole, or sessile with auriculate or sagittate base; margins minutely denticulate with recurved callosethickened teeth. Capitula homogamous, ligulate, most in densely aggregated cymes; peduncle absent or up to 6 mm long, rarely up to 22 mm, glabrous or glandular-setose. In-

145. ASTERACEAE: 25. Sonchus

v ilucre cylmdric, 10-11 x 5-6 nun. densely white toment< se or woolly. Phyllaries 3-seriate, glabrous, woolly at b tse, glandular or glandular-setose; outer ones 2-5 x 1-1.5 mm, linear or ovate-lanceolate with broad scarious n argins; inner ones 10-12 x 1-1.5 mm, w ith narrow s arious margins. Florets yellow; corolla tube hairy. Cyps las yellowish brown, narrowly elliptic, 2.5-3 x 0.6-0.8 n m, transv ersely rugulose, glabrous. Pappus white, c 5.5 mm long, caducous. Derived evergreen bushland and degraded forest, s ream banks, plantations of Eucalyptus, upland grassl;jid, dry evergreen woodland, rarely weed in gardens: 1700-3330 m. TU GD GJ WU SU AR WG IL KF SD BA I- A; Sudan. E Africa. Rwanda. Rep. Dem. du Congo. Ma­ lawi. Greathead 115; Friis et al. 1472; Mesfin T. & Kagnew 1664. : S. luxuriant (R. E. Fries) C. Jeffrey (1966); S. bipontini Asch. var. pinnatifidus Oliv. & Hiem forma luxuarians R.E. Fries in Acta Hort. Berg. 8(6): 102 (1925) - type: Tanzania. Mt. Kilimanjaro. Marangu, 1500 m, Volkens 753 (B holo. destroyed. BM lecto., G isolecto ). S. bipontini Asch. var. pinnatifidus auct., non Oliv. & Hiem (1877): R.E. Fries (1925). Densely branched perennial herb, rhizomatous. 0.5-2 m high Stem branched, herbaceous, cylindric. finely stri. le-sulcate. Cauline leaves glabrous, 5-20 x 2-9 cm. ellipit in outline (those toward the apex triangular), 'innatisect, lobes triangular, rarely elliptic, margins .entate, denticulate or rarely entire, bases auriculate. auri. les mostly elliptic or circular. Capitula in dense groups. .*•hite tomentose at base. 12-15 x 5-7 mm, subscssilc or iicdunculate, peduncle 1-20 mm long, densely white to­ mentose and also glandular hairy. Phyllaries c 25. outer V-13, 3-8 x 1.5-2 nun, thick and cartilaginous at base in ' ruit; inner 8-12. 12-15 x c 1.5 mm. Florets yellow. Limb bfiligules 4-6 nun long, tube 8-12 mm long. Anthers c 2.5 mm long. Cypselas 3.2-3.8 x 1-1.4 nun. creamy yellow, parrowly elliptic, transversely ridged. 4-5-ribbed on each lace. Pappus c 10 mm long, caducous. Weed at margins of cultivated fields: 1650 m. SU; E Afnca. Rep. Dem. du Congo, Rwanda. Burundi. Mogk 208. This weedy species seems to be under-represented in collections of Ethiopian plants. 4. S. asper (L.) Hill (1769): S. oleraceus L. var. asper L., Sp. PI.: 743 (1753) type: From Europe. Herb. Burser VI: 14 [UPS lecto.. selected by Boulos in Taxon 47(2): 368 (1998)]. E rect, annual or short-lived perennial herb, up to 1 m high. Stem hollow, branched toward the apex, glabrous or glan­ dular setose in upper parts, densely lcafv throughout. L eaves simple and spathulate with narrowly winged petiiTe, to runcinately pinnatifid. sessile with rounded iunculate-ainplexicaul base, up to 20 x 6 cm, margins denticulate with usually long and short callose-thickened

67

teeth intermixed. Capitula in open corymbose cymes; peduncle up to 6 cm long, glabrous to densely glandularsetose. Involucre campanulate, 10-12 x 7-8 mm, callosethickend at base. Phyllaries 3-seriate, glabrous to densely glandular-setose, callose-thickened at base and medially in lower half; outer ones 2-6 x 1-2 mm. inner ones 9-12 x 0.7-1.3 nun. Florets lemon-yellow, corolla tube hairy. Cypselas yellowish brown to orange-yellow, smooth, gla­ brous. 4-4.2 x 1.3-1.4 mm. 3-ribbed on each face, outer surface convex, elliptic. Pappus white. 6-7 mm long. Fig. 145.44. Waste ground, stream margins, grassy lawn, forest margins; 1050-2850 m. EW TU GJ SU KF BA: wide­ spread in tropical and South Africa. Mesfin T. 3119; Ev­ ans & Leakey 141; Schimper 495. As the available material from the Flora area is insuf­ ficient. subdivision of the species has not been possible. 5. S. obtusilobus R. E. Fries (1925) - type: TU. Erarta Mt.. 3200 m. Schimper 614 (BM lecto.. K isolecto ). Perennial herb, only up to 15 cm high, with a thick, w oody base. Stem 6-12 mm wide at base. Leaves crowded at apex of the woody base of the stem, pinnatifid with rounded lobes. 3-6 x c 0.8 cm, glabrous, margins dentate, apex obtuse, rounded. Capitula several at the apex of the stem: peduncle 1-4 cm long. Involucre oblong. 10-11 x 6-6.5 mm, brown-woolly at base. Phyllaries 3-seriate, 4-11 x 0.8-1 mm. grcy-grcen or blackish with black setae medially. Cypselas brown, c 3.5 mm long, striatc-sulcate. shiny, glabrous. Pappus 3-5 mm long. Habitat not known; c 3200 m. TU: not known else­ where. Schimper 614. Known only from the type collection. 6. S. melanolepis Fresen. (1839) - type: GD. Simen Mts.. Riippell s.n (FR holo ). Perennial herb, usually found hanging from rock crevices, densely branched with up to 40 cm long tangled branches. Leaves linear to runcinate-pinnatifid. glabrous, glauccsccnt. sessile, auriculate at base, up to 8 x 2.2 cm; seg­ ments linear or sometimes the ultimate segments rounded and much wider than the remaining ones. Capitula solitary or 2-3 at the apex of branches; epedunculate or peduncle short, up to 5 mm long. Involucre oblong-campanulate. 9-10 x 4-5 mm. densely brown-tomentosc at base. Phyllaries 3-seriate. grey-green or sometimes blackish grey: outer ones 2.5—3 x 0.5-1 mm. ovate, glandular-setosc with black hairs; inner ones with whitish mar­ gins. 8-10 x 1-1.5 mm. linear, glabrous. Florets pale yel­ low. Cypselas brown or black. 2.5-3.5 x 0.8-1 0 mm. transeversly rugulose, glabrous. Pappus w hite, c 6-7 mm long. Usually hanging from weathered volcanic rocks on steep mountain slopes, and in protected sites in rockcreviccs; 2400-4100 in. EW TU GD GJ WU SU AR BA;

68

145. ASTERACEAE: 25 Sonchus

Figure 44 SONCHUS A SPER 1 habit x 12 . 2 cypscia with pappus x 5. 3 outer phyllaries x 4; 4 inner phyllary x 4 (No specimens cited in orig.) Drawn by Bhola Ram. (Repro­ duced with permission from Flora of India, vol. 12. fig. 82.)

?Ycinen. Mesfin T. 8426; O. Hedberg 5638; W. de Wilde 9029.

E:rect. annual herb, up to 1 m liigh. Stem striate-sulcate, glaucescent. branched above, hollow. Leaves simple to deepK pinnatifid. glaucous-green above, much paler be­ low. oblanceolate or oblong in outline, up to 23 x 10 cm. 7. S. oleraceusL. (1753) margins denticulate with sharp callose-thickened teeth; type: “Habitat in Europae cultis." Linnaean Her­ terminal segment often triangular or deltoid, wider than barium No. 949.6 [LINN lecto.. selected by Boulos in the remaining lobes: cauline leaves usually simple, Bot. Notiser 126: 155 (1973)). auriculatc-amplexicaul. sagittate at base. Capitula homoS. macrotus Fenzl. (1844). gamous. arranged in corymbose cymes; peduncle gla­ S. schimperi A. Br. & Bouche (1857); S. brous to setose-glandular, up to 6 cm long. Involucre oleraceousxai. schimperi (A. Br. & Bouche) A Br. & campanulate. 8-9 x 5-6 mm. callose-thickened at base. Bouche in App. Ind. Sem. Hort. Berol. (1857) - ty pe: a Phyllaries 3-senate, grey or glaucous-green, purpleplant raised from seeds, sent by Schimper in 1856 from the Flora area, cultivated in Berlin Bot. Garden tinged, glabrous, with sessile glandular excretions and/or (B holo., not seen). gland-tipped setae outer ones ovate-lanceolate, tapering

69

145. ASTERACEAE: 25. Sonchus

Figure 45.

SONCHUS OLERACEUS 1

habit x '/*; 2 - flowering capitulum x c 3; 3 series o f phyllaries from the outer (left) to the innermost x c 5; 4 - floret with­ out ovary x 7; 5 stamens with tailed anther lobes x 13; 6 - cypsela with pappus partially removed x 9; 7 - cyp­ sela x 9. All from Vaughan in MAU 13452. Drawn by Eleanor Catherine. (1 -6 reproduced from FI. Mascareignes 109. C om posees, PI. 12, and 7 with permission from FI. Zamb.. vol. 6(1), Tab. 41C.)

ward the apex, 6-7 x 1-1.5 mm, thickened medially in nver half; inner ones oblong, attenuate at apex, 7-9 x 1.5-2 mm, margins scarious. Florets yellow; corolla tube lairy. Cypselas brown, oblanceolate, 3.3-3.7 x 0.8-1 nun. errucose with transverse ridges, multi-ribbed. Pappus tv itite, c 5 nun long. Fig. 145.45. Roadside margins and grassy Fields; 1650-2440 m. V TXJ GD GJ WU SU SD HA; a cosmopolitan weed na­ ive of Eurasia and N Africa; also found in Sudan. Soma­ lia. Sococtra. De Lotto 17/831; Schimper \292\ Mesfin T. -032.

lanceolate in outline, margins lobed to trilobed, up to 15 x 6 cm. upper cauline leaves auriculate-amplexicaul with lobed bases. Capitula in corymbose cymes; peduncle gla­ brous to densely glandular-setose, up to 5 cm long. Invo­ lucre cylindric-campanulate, 8-9 x 4-5 mm at anthesis. glabrous and callose-thickened or densely white-tomentose at base. Phyllaries 3-seriate, glabrous or glandularsetose; outer ones 1-4 x 1-1.5 mm. inner ones 10-12 x 1.5-2 mm, medially thickened at base and in lower half. Florets yellow, corolla tube hairy. Cypselas orangeyellow, oblanceolate, slightly compressed. 2.5-3 x 0.6-0.8 nun, glabrous, transversely rugulose. Pappus white, 5-8 nun long, persistent. Fig. 145.46.

H S. tenerrim us L. (1753) Montane grassland. EW; native of the Mediterranean type: “Habitat Monspelii. Florentiae.” Linnaean area and W Africa, widely distributed in central Europe Herbarium No. 949.9 [LINN lecto., selected by and eastwards to Pakistan. Schweinfurth & Riva 1647; Boulos in Bot. Notiser 126: 158 (1973)]. Terraciano & Pappi 1982. : -ect annual herb, up to 75 cm high. Stem branched in the Schweinfurth & Riva 1647 has sometimes been identi­ n florescence. Leaves deeply pinnatifid to bipinnatifid fied as S. oleraceus. I ith linear or narrowly ovate-lanceolate segments, ob­

70

145. ASTERACEAE: 25. Sonchus, 26. Tolpis

Weed in cultivated Fields and roadsides. 1200-2500 m. EW GJ SU tL BA; Senegal, Sudan, Rep. Dem. du Congo, Angola, Zambia, Mozambique, Namibia, South Africa, Madagascar. Boulos 9300; Mesfin T.& SebsebeD 4028; Mesfin T. 5592.

26. TOLPIS A dans. (1763) Erect annual or perennial herbs. Leaves mostly basal, sim­ ple to pinnately dentate or lobed. Capitula homogamous. ligulate, arranged in branched, cymose panicles; terminal capitulum superseded by lateral ones; peduncle slender, glabrous except near receptacle, bracteate. Receptacle flat epaleate. Involucre campanulate. Phyllaries 2-3seriate, outer ones unequal, inner ones subequal, linearlanceolate, acute with pale margins. Florets yellow, inner ones usually much shorter and purplish-brown. Cypselas mono- or dimorphic, quadrangular, ribbed, glabrous, brownish, truncate at apex. Pappus setaceous; setae often one-seriate, unequal, distinct to base, 4-17, glabrous or barbellate. A genus with 20 species found mostly in the Macaronesian Islands; 1 species in the Flora area.

Figure 46. SONCHUS TENERRIM U S I - upper part o f stem; 2 - mid-stem leaf; 3 - cypsela with pappus x 6. (No specimens cited in orig.) Drawn by Ruth Koppel. (Reproduced from FI. Palaestina, vol. 3, PI. 735.)

T. virgata (Desf.) Bertol. (1803); Crepis virgata Desf. (1792) - type: ? Algeria, Desfontaine s.n. (P holo ). Crepis altissima Balb. (1804); Tolpis altissima (Balb.)Pers. (1807). Crepis ambigua Balb. (1805); Schmidtia ambigua (Balb.) Cass. (1827). Tolpis abyssinica Sch. Bip. ex A. Rich. (1848); Schmidtia abyssinica (Sch. Bip. ex A. Rich.) Schweinf. & Asch. (1867) - type: TU, Memsach. PI. abyss. I, Schimper 1 (P holo., K iso.).

Erect perennial herb, up to 75 cm high, with a stout tap­ root. Stems solitary or several, each with its cluster of leaves al base, hairy or even tomentose at base, glabrous or glabrescent above, branched at base or in the inflores­ 9. S. gigas Boulos ex Humbert (1963) cence, branches erect or spreading. Leaves mostly basal type: Zambia, Muckle Neuk, Robinson 904 (K or rosulate, simple, often 5-18 x 1-3 cm, oblanceolate, holo). sparsely ciliate along veins to densely hairy all over, apex acute, margins denticulate to dentate, base attenuate end­ Erect, annual or short-lived perennial herb. 0.4-1.5 m ing in a narrowh winged, up to 2 cm long petiole; stem high. Stem glabrous or glandular-setose, usually un­ leaves few. linear or narrowly elliptic, uppermost bract­ branched, leafy at base. Leaves simple to pinnatifid, ob­ like. Capitula cylindric-campanulate, 2-2.5 cm wide at lanceolate or ovate-lanceolate, up to 23 x 4.5 cm, margins anthesis. in lax paniculate cymes; peduncle 1-18 cm long, irregularly serrate with spine-like, pointed, and calloseglabrous except sometimes near the capitula where thickened teeth. Capitula in open corymbose cymes; puberulous. Involucre campanulate, 5-6 x 4-5 mm at peduncle up to 5 cm long, glabrous or glandular-setose. anthesis, not much altered in size in fruit, cartilagiInvolucre 9-10 x 5-6 mm at anthesis, up to 1 2 x 9 mm in nous-thickened at base in fruit. Phyllaries 2-3-seriate, fruit, sparsely to densely white tomentose at base, espe­ green, densely white-pubescent especially in the lower cially in young capitula. Phyllaries light green, often with half; outerones 2.5—3 x 0.3-0.5 mm, apex acute, cartilagiseveral brown sessile glands, outer ones linear, 3-4 x nous-thickened, linear-lanceolate, 18 or more and contin­ 0.8-1 mm, inner ones oblong-linear with whitish margins. uous with bracts on peduncle, inner ones 7-8 x c 1 mm, 10-13 x 1.2-3 mm. Florets yellow. Cypselas brown, ob­ lanceolate with acuminate apex, 20 or more, margins long-elliptic, flat, 2.8-3.8 x 1—1.3 mm, with 3 main ribs on scarious. Florets yellow. Cypselas dark-brown, 2-2.2 x each face, appearing winged. Pappus white, 7-8 mm long, 0.8-1 mm, oblanceolate, quadrangular, 8-ribbed, finely caducous. rogulose, glabrous. Pappus of 2-rowed bristles, outer row

145. ASTERACEAE: 26. Tolpis, 27. Helminthotheca, 28. Picris

71

mall, more or less equal, inner 4-17. longer, unequal. :>rownish yellow, 0.5-5.5 mm long, antrorscly barbellate. Fig. 145.47. Erica-Myrica-Olea scrub, montane grassland: 200010 m. TU GD GJ SU WG KF SD HA; NW Africa (Tuusia, Libya. Algeria), S Europe, Palestine, W Syria, Ye­ men, Saudi Arabia. Mooney 5017A; Friis et al. 1252; Mesfin T. & Kagnew 2187. Similar to T. capensis in many morphological fea„res, and differing from it in pollen ultrastructural details Jarvis, 1986).

27. HELM INTHOTHECA** Vaill. exZinn (1757) ■I W. Lack in Taxon 24: 111-112 (1975). \nnual or perennial herbs with erect to decumbent stems, rriumentum mostly of glochidiate 2-4(-6) hooked hairs, eaves both basal and cauline; basal leaves oblanceolate. tcute at apex, margins sinuately dentate, rarely entire, at„*nuate and sub-petiolate at base; cauline leaves ovate or blong. sessile and auriculate or cordate at base. Capitula to several, cymosely arranged, homogamous. ligulate. Receptacle epaleacous. Involucre ovoid-urceolate. livllaries 2-seriate. outer ones foliaceous. spreading, vate to cordate, inner ones linear-lanceolate, green. FIojts yellow. Cypselas obovoid or oblong, transversely igose, long-beaked. Pappus white, plumose. A genus with 4 species native of the Mediterranean sa and S W Asia. The following weedy species was re( eptly introduced in the Flora area.

Figure 47. TOLPIS VIRGA T A X - habit, x '/*; 2 - cypsela x 4'/2. I from Mesfin & Kagnew 2187. I draw n by Damtew Teffera; 2 redrawn from FI. Palaeslina , vol. 3, PI. 693.

1 1echioides (L.) Holub (1973);

Picris echioides L. (1753) - type: “Habitat in Angliae. Galliae sylvis caeduis. aggeribus.” Linnaean Herbarium No. 948.1 [LINN lecto.. selected bv Lack in Taxon 24: 113 (1975)1. .inual or short-lived perennial herbs, up to 75 cm high. Stem moderately to profusely branched, branches spread­ ing Whole plant scabrid-hispid. covered with 2-4t irbed. glochidiate hairs. Leaves basal or cauline. simple, oblanceolate, 5-13 x 1-2.5 cm. margins entire to sinuately dentate or denticulate, upper leaves smaller and sessile. ( ipitula c 2.5 cm wide at anthesis. solitary or in open cymes: peduncle up to 2 cm long. Involucre c 10 x 6 inin at a ilhesis. Outer phyllaries 5. green, ovate-lanceolate. 9-12 \ 3-4.5 mm. margins and mid-vein densely glochidiate. I mer phyllaries 8. greyish green, oblong-lanceolate. 1 )—14 x 2.5—3.5 mm. mid-vein densely glochidiate. li idurate in fniit. surfaces with fine liairs. Florets yellow, li i*ules c 1 cm long, usually light purplish on lower surf ce. Cypselas orange, oblong, c 2.1 x 0.6 mm. trans\ rsely rugulose. glabrous, with a long (c 3 mm), narrow, yellow beak expanded into a disc at summit bearing p ippus-hairs. Pappus white, 4.5-5.5 mm long, plumose. F ;g 145.48.

Weed of wasteland and cultivated fields; 2000-2450 in. SU; native of Europe, SW Asia and N Africa; intro­ duced elsewhere (N and S America, S Africa and Austra­ lia). Parker 125; Mesfin T. 3685; W. de Wilde 6386.

28. PICRIS L. (1753) W. Lack in Pl. Syst. EvoL 131 35-52 (1979). Annual or perennial herbs with one to several, erect, or decumbent stems. Indumentum mostly of glocliidiate liairs with 2-4 barbs. Leaves mostly basal, simple to pinnatifid. oblanceolate. obtuse or acute at apex, attenuate at base; upper leaves oblong or elliptic, sessile. Capitula homogamous. ligulate. solitary at apices of branches. Re­ ceptacle epaleate or setose. Involucre campanulate. Phyllaries 2-3-seriaie. linear-lanceolate, imbricate, the inner ones subequal, longer than the outer ones. Florets yellow. Cypselas homo- or heteromorphic, cylindric and narrowed at each end or slightly attenuate toward the

72

145. ASTERACEAE: 27. Helminthotheca, 28. Picris

1

Upper part of corolla tubes with short liairs; cypseias 4-5 mm long, not beaked. I P. abyssinica Upper part of corolla tubes with long liairs; cypselas 6-8 mm long, beaked 2. P. xyiopoda

1. P. abyssinica Sch. Bip. (1839) - type: TU/GD, between Halei and Temben, Ma> 1832, Riippell s.n. (FR holo., P iso.).

Erect, perennial herb, up to 30 cm high, with a woody deep-seated tap root. Stems solitary or several, branched at the base or in the inflorescence. Leaves mostly radical, up to 15 x 2.5 cm, obovate to oblanceolate, simple, scabrid-hispid with 2-barbed glochids. obtuse at apex, at­ tenuate into a broadly or narrowly winged petiole; mar­ gins denticulate to pinnately shallowly lobed; cauline leaves few. similar to radical leaves or linear and sessile, the uppermost ones bract-like. Capitula solitary or in lax paniculate cymes. 2.5—3 cm wide at anthesis; peduncle up to 11 cm long, stnate-sulcate, hispidulous throughout or white-pubescent near the apex. Involucre campanulate, 8-10 x 6-7 nun at anthesis, up to 12 x 10 mm in fruit. Phyllaries 2-3-seriate, often white-tomentose with 2-barbed glochids all along the midrib; outer ones 7-8. linear-lanceolate. 2-5 x 0.5-1 mm. glabrous at margins; inner ones 12-14, lanceolate. 10-12 x 2-2.5 mm. gla­ brous and yellowish at margins. Ligules 13-14x2-3 mm. yellow, 5-dentate at apex or 2-dentate each with 2 or 3 teeth; tube densely pubescent in upper parts. Cypselas light brown, cylindrical, attenuate towards both ends, 4-5 (-6) x 0.8-1 mm. 5-ribbed, ribs smooth to transversely rugose. Pappus yellowish wliite, persistent. 5-8.5 mm long, plumose. Fig. 145.49.6. Montane slopes with degraded bushland and grass­ land. 2100-2200 m EW TU, Yemen. Schweinfurth & Riva 1296; Schimper 163; Pappi 2857. In his revision of the genus Picris in tropical Africa, Lack (1979:40) maintained that "P. scahra is definitely not identical with P abyssinica” and also stated that “due to Forsskal s poor description and the absence of adequate material from Yemen, it is not possible at the moment to interpret this name (i.e., P. scabra) properly and. conse­ quently, P. scabra is treated here as a doubtful name.”

Figure 48 HELM INTH O TH ECA ECH IOID ES 1 - upper part o f stem; 2 - cypsela with pappus. (No specimens cited in orig.) Drawn by Ruth Koppel. (Reproduced from FI. Palaeslina , vo) 6, PI. 711.)

apex, transversely rugulose. Pappus bristles numerous, plumose, or outermost shortly scabrid. unequal. A genus of about 40 species native of the Mediterra­ nean area. Lack (1979) recorded two species for the Flora area and this is maintained here until more material is available (see comments below).

A depauperate specimen. Mesfin T. & Sebsebe D. 3769 obtained from the Mereb Valley in TU at 1350 m. lias beaked cypselas that are 6-7 inm long, thus weaken­ ing the distinction between P. abyssinica and P. xyiopoda Further collections might show the true relationships be­ tween these taxa. Engler (1891 452) cited a Schimper specimen, no. 334 from “Hedscha um 3000 m”. The present author lias not seen this specimen and therefore the higher altitudinal record for the species in the Flora area cannot be authenti­ cated. 2 P. xyiopoda Lack (1979) -ty p e: Nigeria, Naraguta. Coombe 44 (K holo ). Perennial herb, up to 50 cm high, with a long, thick, deep-seated taproot. Stems solitary or several, branched

73

145. ASTERACEAE: 28. Picris, 29. Tragopogon

Figure49. PICRISXYLOPODA 1 - habit x Vi; 2 - outer phyllary x 3: 3 - inner phyllary x 3; 4 - ray floret without ovary x 3; 5 - cypsela with pappus x 3. P. ABYSSINICA 6 - habit x V4. 1-5 from Tewolde B.G.E. 91113-05; 6 from Ryding 1447. Drawn by Birhanu Mihretu.

: the inflorescence. hispid with long glochidiatc hairs, eaves in basal rosettes, simple, held erect. 5-35 x i 5-4.2 cm. narrowly oblanceolate. acute at apex, i ntate-dencticulate or sinuately-dentate at margins, at. -mate into a narrow or broad winged petiole at base, both iurfaces pilose with glochidiatc long hairs; cauline leaves iimilar to basal ones, except being narrower and reduced n si/.e. Capitula solitary at the apex of stems or branches. » a few in very open corymbose cymes, c 2.5 cm wide at mthesis; peduncle up to 22 cm long, sparsely pilose ex­ cept near apex where glochidiatc hairs arc denser. Involu:re 9-10 x 6-7 mm at anthesis. up to 1 5 x 7 nun in fruit. Phyllaries 2-3-seriate, sparsely white-tomentose with 2-barbed glochids. all along the midrib, outer ones 6-7. inear-lanceolate. 4-5.5 x 1—1.2 mm. glabrous at margins, nner ones 12-14. lanceolate. 10-13 x 1.5-2 nun. gla­ brous at margins or sometimes throughout. Ligules 16-18 < 2.5—3 mm. 5-dentate at apex, tube 7-8 nun long, pilose.

Cvpselas orange-brow n. 6-8.5 x 0.8-1 mm. narrowed at apex, transversely rugose, c 20-ribbcd. ribs spiculate; apex truncate, expanded into a disc. Pappus yellowish white. 3-8.5 nun long, unequal, plumose. Fig. 145.49.1—5. Dry hillside slopes and on plains along hike shores. 1500-1800 m. SU (mainly in the Rift Valley around the lakes); N Nigeria. B oulos 9800; I'otova 216 6 ; Tewolde B. (}.! be real, but could also be due to undcrcollccting.

29 TRAGOPOGON* / (175^)

Erect, annual or perennial herbs Stems simple or branched, leafy Leaves simple, entire, usualh grass-like.

74

145. ASTERACEAE: 29 Tragopogon

pappus. Pappus homomorphic or heteromorphic, of nu ­ merous plumose bristles. A genus with about 110 species found mainly in southwest to central Asia. Tragopogon dubius Scop, with lemon-yellow florets may also occur in the Flora area but no specimen has been seen

T. porrifolius L. (1753) type: Herb. Burser XV(2): 69, centre specimen [UPS lecto., selected by Diaz de la Guardia & Blanca in Taxon 41(3): 549 (1992)]. Erect, annual herb, up to 1 m high. Stem branched, gla­ brous. Leaves linear-lanceolate, up to 15 cm long and 1 cm at the base, glabrous, simple, sessile; margins entire. Capitula solitary at the apex of branches; peduncles swol­ len below the capitula. Involucre cylindric-urceolate. Phyllaries 5-9, equal to or longer than the florets. Florets lilac or purple. Cypselas yellowish brown, 2.5-5 cm long (including beak), with 10 longitudinal rows of short scales Pappus of 8-13 mm long bristles, those o f outer cy pselas barbellate, those of inner ones plumose. Fig. 145.50 Occasional weed in and around cultivated fields; 1900 m SU: native of the Mediterranean area; commonly introdued in Europe and N Africa and widely naturalized in N America. Parker 4235. A recent introduction in the Flora area.

Tribe 4 V E R N O N IE A E Cass. (1819) C Jeffrey in Kew Bull. 43: 195-277 (1988).

Figure 50 TRAGOPOGONPORRIFOLIUS: 1 - habit x '4; 2 fused phyllaries x Vi; 3 - cypsela with pappus x 2'/j. (No speci­ mens cited in orig.) Drawn bv John H. Rumely. (Reproduced with permission from Vascular Plants o f the Pacific Northwest, part 5. 1955. p. 331 p.p.)

rarely broadly oblong, broader at base and semiamplexicaul. Capitula homogamous, ligulate, solitary on long peduncles. Phyllaries 2-3-seriate, herbaceous, equal in length, lanceolate, enlarged in fruit, free to the base. Re­ ceptacle epaleate. Florets purple or yellow. Cypselas light brown, fusiform, beaked, 5- or 10-ribbed, glabrous, ribs muricate; beak long, distinct, constricted at apex into a brownish annulus before expanding into a disc bearing the

Herbs, shrubs or small trees. Indumentum o f simple or symmetrically or asymmetrically T-shaped, short- or long-stalked hairs, rarely globose trichomes on phyllaries and cy psclas. Leaves simple, alternate, rarely whorled or tcmate. entire, serrate or dentate Inflorescence simple or paniculate (rarely scorpioid) cymes or corymbs. Capitula separate or syncephalous in bracteate glome rules, one- to many-flowered, cy lindric, campanulate, turbinate or hemisplicncal, homogamous. Receptacle naked, rarely paleatc. foveolate or fimbrilliferous. Phyllaries numerous, few - to many-seriate, usually free, linear to ovate, acute, obtuse, or with a terminal appendage. Florets bisexual, pmk. purple, yellow, orange, violet or white. Corolla tu­ bular or campanulate. or gradually dilated above, 5-lobed, lobes usually equally cleft, rarely asymmetrical. Anther base sagittate or obtuse, apex with a linear or lanceolate, free connective Sty le bifurcate, branches elongately Un­ ear, papillose throughout. Cypselas columnar, cylindric or obovoid, terete, angled or ribbed, or costate, glabrous or hairy tliroughout or in longitudinal bands, in many species globose trichomes common between the ribs or costae. Pappus absent or present; when present one- to sev­ eral-seriate, white; outer series of bristles or scales usually

145. ASTERACEAE: 30 Vernonia

shorter than the inner, rarely coroniform; inner series of b< rbellate bristles persistent or caducous.

7.

A. tribe with about 70 genera in both the Old and the N : w World 7 genera in the Flora area with Vernonia having the largest number of species.

-

A large well-defined tribe, common in tropical parts of th world. "Its subdivision into subtribes and genera is stilJ very uncertain” Jeffrey, (1988). The most problemat c member is the genus Vernonia. In a review of the Virnonieae in East Tropical Africa, Jeffrey (1988:196) st; ted that, because of the currently available data differeri.:es in chemistry , cytology and palynology between the 0 d and New World taxa of Vernonia, none of the Old W rid species should properly be referred to the genus Vernonia.” On the basis of leaf vein numbers, floret col01 \ anther base features and pollen wall characteristics. R htnson [in Proc. Biol. Soc. Wash. 99: 493-501(1986)) se .'.regated Distephanus from Vernonia. Pollen morphol­ ogy seems to support this [cf„ Mesfin Tadesse in Comp. N< ws. 22: 17 (1992)]. Robinson [in Proc. Biol. Soc. Wash. 103 248-253 (1990)] also resurrected Baccharoides M tench and Cyanthillium Blume by transferring a few of t h : Old World species from Vernonia. However, until a th :rough revision of the genus is made, all taxa occurring in the Flora area, except those recognized in Distephanus, a r : here treated within Vernonia.

8.

1. Capitula grouped together in globose, terminal, bracteate, solitary or cymose heads, 4-5- flow­ ered; phyllaries 8, decussate; pappus one-seriate, widened at the base. 36. Elephantopus Capitula not grouped together in bracteate, cymosely-arranged heads, separate or axillary, 1-many-flowered; phyllaries more than 8, spi­ rally imbricate; pappus one- to several-seriate, uniform throughout. 2 2 -

Leaves whorled, usually temate. 34. Bothriocline Leaves alternate, rarely sub-opposite. 3

3. Pappus present, of bristle and scales. 4 - Pappus absent or represented by an apical rim of corona. 7 4.

-

Pappus of numerous, persistent, white bristles and scales, cypselas usually daik-brown or black, cylindric, terete or ribbed, usually shortly uniformly 5 villous, with or without sessile glands. Pappus falling early; cypselas light brown, angled or ribbed, if terete, obovoid and glabrous except for the sessile trichomes near the apex and base. 6

5. Florets yellow or orange; leaves 3-veined from base of blade; anther base tailed. 31. Distephanus - Florets purple, reddish or white; leaves usually with a single mid-vein; anther base sagittate, not tailed. 30. Vernonia 6.

-

Hairs on undersides of leaves symmetrically Tshaped; corolla glabrous or sparsely hairy; cyp­ selas (3-)5-10-ribbed. 32. Gutenbergia Hairs on undersides of leaves simple; corolla usu­ ally densely pubescent; cypselas 3-4-angled or ribbed. 33. Eriangea

-

75

Hairs on stems and undersides of leaves symmctrically T-shapcd; corolla glabrous or sparsely hairy; cypselas (3-)5-10-ribbed. 32. Gutenbergia Hairs on stems and undersides of leaves simple; co­ rolla densely pubescent or sparsely papillose in lower parts, cypselas 2-6-ribbed. 8 Cypselas 0.8-2 mm long, surfaces between ribs usually longitudirally pigmented, often also with globose trichomes and short white hairs: corolla sparsely papillose in lower parts. 35. Ethulia Cypselas 2.5-3.2 nun long, glabrous except for the few globose tnchomcs at apex; corolla densely pubescent. 33. Eriangea

30. VERNONIA Sch re h. (1791). nom. cons. Linzia Sch. Bip. ex Walp. (1843). Wehbia DC. (1836). Triplotaxis Hutch. (1914). Herbs, shrubs with erect, scandent or scrambling branches, or trees Indumentum of simple, 1-seriate and multicellular. or T-shaped hairs. Leaves alternate or rarely sub-opposite, simple, entire, serrate or dentate. Inflore­ scence cvmose. scorpioid. corymbose or paniculate, rarely capitula solitary. Capitula homogamous. 1- to many -flowered; involucre cylindric, campanulate, urceolate. hemispherical or turbinate. Receptacle epaleate. fovcolatc or finibrillifcrous. Phyllaries green, herbaceous, numerous, fcw-to many-seriate, usually free to base, lin­ ear, lanceolate, ovate, apex acute, obtuse, or with an ap­ pendage Florets bisexual, purple, reddish or white. Co­ rolla tubular or campanulate. or dilated in upper parts, usually equally 5-lobed. Anther-base sagittate, apex with a linear or lanceolate free connective. Cypselas black or dark-brown, cylindric or narrowed below or turbinate. 4-5-angled, 4-20-ribbed or costate, glabrous, uniformly hairy, or hairs in longitudinal series, or sessile-glandular between the ribs, with or without a basal callus. Pappus (l-)2 - to many-seriate, white; outer series usually much shorter than the inner, bristly, squamifonn or coroniform; inner series usually setaceous, barbellate. persistent or caducous. A genus with about 500 species (Bremer. 1994) found mainly in tropical and wanner parts of North and South America, tropical Africa. Madagascar and in SE Asia. Many of the neotropical species previously placed in Vernonia are now reclassified under other closely related g en era, e.g.. B accharoides Mocnch. Cynnthillium Blunic, and Distephanus Cass. It is most probable tliat not all of the species recorded from the Flora area will be retained in Vernonia in the future but in the absence of a worldwide revision of the genus, all of them are treated in this genus. Vernonia is noted for its sesquiterpene lactones, a class of bioactive substances tliat also occur in other mem­ bers of the Asteraceae. Vernonia amygdalina Del., which is quite commonly used in Ethiopia in the preparation of local beer (TELLA in Amharic, the official language of Ethiopia) and also as fumigant, lias been reported as being chewed by sick cliimpanzees in Tanzania. Currently, sci-

76

145 ASTERACEAE 30 Vernonia

entists think that the chimpanzees may be using the plant to cure themselves of schistosomiasis - a disease also af­ fecting many people in tropical Africa and Asia (cf. NAPRECA Newsletter, vol. 13(2): 3 (1996)).

-

Leaves petiolate, petiole (0.5-)l-6 cm long; peren­ nial heib or shrub 1.2-4 m high. 12

11. Most leaves 8-17.5 x 2.5-6 cm, margins denti­ culate; corolla 14-15 mm long; stem unbranched except in the inflorescences, up to 0.5 m high 38. V. didessana Most leaves up to 5.5 x 2.5 cm, margins serrate; co­ rolla 16.5—17 mm long; stem profusely branched, 0.5-1 m high. 16. V. printzioides

So far about 49 species are recorded from Ethiopia. A few of the names previously listed under Vernonia by Cufodontis (1967) are not accounted for here due to lack of specimens and/or inadequate information. 1.

2. 3.

4. 5.

-

Leaves ± succulent entire and sessile; corolla pur­ ple or turquoise. 22-23 mm long; phyllaries coriaceous, glabrous. 30. V. wakefiefdii Leaves not succulent other characters not as com­ bined above. 2 Plants annual with the roots not surviving beyond the life span of the aerial stems. 3 Plants perennial with thick rootstocks, shrubs or small trees. 7 Outer and middle phyllaries appendaged; append­ ages usually dark green, narrowly lanceolate and often reflexed; corolla 13-18 mm long. 4 Phyllaries not appendaged, usually the upper part darker, corolla 5-9 mm long. 5 Corolla 13-14 mm long, sessile-glandular. 32. V. galamensis Corolla 17-18 mm long, glabrous or thinly pu­ bescent. 33. V. ituriensis Leaves (20-)30-46 mm wide, margins senate or dentate; capitula 6-6.5 x 5-5.5 mm; longest phyllaries 4—4.3 mm long; corolla pilose. 26. V. cinerea Leaves 2-17 mm wide, margins entire or inrolled; capitula 12-16 x 8-12 mm; longest phyllaries 8-12 nun long; corolla sessile-glandular 6

6. Leaves oblanceolate to elliptic, 14-17 mm wide; petiole up to 10 mm long; longest phyllaries 8-9 nun long; corolla 6.5-7 mm long. 49. V. ambigua - Leaves narrowly linear to narrowly oblanceolate. 2-4 mm wide, sessile; longest phyllaries 10-12 mm long; corolla 8-9 nun long. 43. V. perrotetti 7

-

Phyllaries appendaged, i.e., with an abruptly dis­ tinct darker green, whitish, purplish or pinkish, ascending, spreading or recurving, ovate to lan­ ceolate upper part; corolla 11-25 mm long. 8 Phyllaries not appendaged. the upper part merely darker coloured and/or acuminate-apiculate or at­ tenuate, only gradually and/or indistinctly dis­ tinct from the lower part, corolla often 5-11 mm long. 17

8. Capitula 25-40 x 20-40 mm. - Capitula 8-25 x 6-20 mm.

9 10

9. Leaves sessile or subsessile, blade 6-7.5 x 1.5-2 cm; perennial herb up to 0.4 m tall. 37. V. buchingeri - Leaves with up to 2 cm long petioles, blade 10-20 x 3-6 cm; shrub or perennial herb up to 2.5 m tall. 42. V. adoensis 10. Leaves sessile or subsessile; perennial herb, 0.5-1 m high. 11

12. Perennial heib, 1.2-2 m high, apex of phyllaries and appendages dark green, turning conspicu­ ously dark brown upon dry ing; cypselas 3-9 mm long. 13 Shrub, 1-4.5 m high; phyllaries uniformly col­ oured or appendage only indistinctly distinct from the lower part; cypselas 2.5^t mm long. 15 13. Cypselas 7-9 x 1.2-1.3 mm; longest pappi 11-13 mm long; phyllaries 7-8-seriate, coriaceous. 33. V. ituriensis - Cypselas 3-4.5 x 0.8-1 mm; longest pappi 8-9 mm long; phyllaries 3-5(-6)-seriate. 14 14. Cypselas c 20-ribbed; middle leaves 30-50 x 8-10 cm, petiole up to 0.5 cm long. 36. V. schimperi - Cypselas c 10-ribbed; middle leaves 12—15(—25) x 3—5(—7) mm. petiole up to 2 cm long. 39 V. filigera 15. Capitula 8-10 x 6-8 mm; cypselas 2.3-2.5 x 0.8-1 mm. 41. V. lasiopus Capitula (10-) 12-25 x 10-20 mm: cypselas 3-4 x 0.&-1.3 nun 16 16. Cypselas 10-ribbed; usually densely sessileglandular, corolla 11-13.5 mm long, lobes with sessile glands; phyllaries linear, 1.5-2 mm wide 39 V. filigera - Cypselas 20-ribbed, usually glabrous, sometimes with few sessile glands; corolla (13-) 15-20 mm long, lobes eglandular. phyllaries 2-4.5 mm wide, linear to narrowly ovate-lanceolate. 40.V. hymenolepis 17. -

Cypselas 10-20-ribbed. Cypselas 4-6-ribbed. 4-5-angled orsmooth.

18 34

18 -

Cypselas 9-10 x 2-2.3 mm. Cypselas 2-5 x 0 .5 -1(-2) mm.

19 20

19 Capitula solitary terminating branches and/or main stem, surrounded by stem leaves, c 25-35 x 15-25 mm; corolla 25-30 mm long. 34. V. gilbertii - Capitula in axillary and/or terminal, usually py­ ramidal or elongated oblong panicles, clear of the foliage, c 20 x 10 mm: corolla 12-15 mm long. 35. V. congolensis

145. ASTERACEAE: 30. Vernonia

2 2 -

Involucre 2-7 mm wide, oblong, cylindric or campanulate. 21 Involucre 8-12 mm wide, campanulate or hemi­ spheric. 30 I Shrub with scandent. scrambling or lianoid branches. 22 Shrub with erect branches, or trees. 23

21 Capitula 22-24 mm tall at anthesis; cypselas 4-4.5 mm long; florets pale purple, violet or pinkishviolet 29. V. tewoldei - Capitula 8-10 mm tall at anthesis; cypselas 3-3.3 mm long; florets white or lilac. 3. V. urticifolia 2 -

Phy llanes glabrous except for the ciliate margins and sessile-glandular spot near the apex. 24 Phyllanes sparsely to densely pubescent with or without sessile glands, floccose or tomentose. 27

2>i Trees or large shrubs, 2-10 m high; stem usually 10-40 cm in diameter, leaves petiolate. petiole 5-30 mm long, cuneate at base. 25 - Shrubs or perennial herbs. 1-3 m high, stem 0.5-3 cm in diameter; leaves sessile, expanded and auriculate. or semi- amplexicaul at base. 26 2:

-

2(

2' 2i

-

29 -

Capitula 10-20-flowercd; leaves glabrescent to puberulous and sessile-glandular, ashy-green; phyllaries light green with dark green and ses­ sile-glandular blotches near the apices, coriaceous. 1. V. amygdalina Capitula 4-6-flowcrcd; leaves glabrous, dark green; phyllaries usually dark purple, glabrous except for the ciliate margins of the outer ones. 12. V. rueppellii Leaves pallid beneath, with crispatc tomcntum, of­ ten rusty-brown upon drying. 8. V. ischnophylla Leaves greea glabrous to subglabrous beneath or on both surfaces. 7. V. thcophrastifolia Leaves 3-40 x 2-16 mm; involucre 6-7 mm high; stem 0.5-2 cm in diameter. 20. V. cincrascens Leaves (50-)80-250 x 30-110 mm; involucre 9-12(-24) mm high; stem 2-10 cm in diameter. 28 Capitula 22-25-flowered, 22-24 mrn long at anthesis; involucre 5-6 nun wide; phyllarics membranous. 29. V. tewoldei Capitula 1-5-flowcrcd. 10-12 mm long at anthesis; involucre 1.3-3 mm wide; phyllaries coriaceous. 29 Longest phyllaries 1-3(—4)-flowered. Longest phyllaries 2-5-flowered.

3—4 5-6

mm

long; capitula 10. V. auriculifera mm long; capitula 9. V. m yriantha

30 Stem with T-shaped liairs. Stem with simple, erect or flagellifonn hairs. 31

-

31 32

Capitula 15-17 x 8-10 nun; outer phyllaries acuminate, tips reflexed; inner phyllarics oblong or oblong-elliptic, whitish, hispid with sparse, sessile glands. 28. V. friisii Capitula 6-10 x 5-8 mm; outer phyllaries acutcapiculate, erect; inner phyllaries linear-lanceo­ late. apex purplish or greyish, densely glandular. 27. V. hochstetteri

77

32. Capitula surrounded by leaf-like bracts; leaves sca­ brous; stem glabrescent to thinly setulosescabrid. 31. V. pu rpu rea Capitula without leaf-like bracts; leaves glabres­ cent to tomentose especially beneath; stem greybrown, densely tomentose or softly puberulous. 33 33. Cypselas c 20-ribbed; phyllaries herbaceoustipped, middle series oblong to ovate-oblong, sessile-glandular toward the apex; middle stem leaves 8-10 mm wide. 36. V. schimperi Cypselas c10-ribbed, phyllaries long-apiculate. appearing spinescent, middle series glabrous or sparsely pubescent; middle stem leaves 15-35 mrn wide. 2. V. yabelloana 34. Small to large shrubs (0.5-4 m liigh) with erect. scrambling or scandent branches, or small trees. 35 Perennial herb, up to 2 m liigh often with thick rootstock. 54 35. Plant scandent or with scrambling or scandent brandies growing through and over shrub clumps, sometimes creeping. 36 - Plant with erect branches, or trees. 42 36. Cypselas 4.9-5.2 mm long; involucre 13-15 x 10-12 mm. leaf margins entire. 17. V. dalettiensis Cypselas 1.8-3 mm long; involucre 5-10 x 3-8 mm; leaf margins crenate. serrate, denticulate, sinuate-dentate to subcntirc. 37 37. Stem up to 0.5 m high; leaves densely tomentose. oblong to oblanceolate. 46. V. popeana Stem (0.7-)2—4 in high; leaves thinly pubescent with liairs mainly on veins or pilose, ovate, ovatc-lanccolatc or elliptic. 38 38. Corolla 6-7 mm long, lobes setose on outside or sessile glandular. 39 - Corolla 9-10 mm long, lobes glabrous or sessileglandular. 40 39. Outer phyllaries 1-1.5 mm wide, sparsely sessileglandular near the apex; corolla sessile-glandular; hairs on lower leaf surface simple. 25. V. hildebrandtii - Outer phyllaries 0.2-0.4 mm wide, setose, corolla lobes setose; hairs on lower leaf surface T-shaped 24 V. wollastonii 40. Leaves green and glabrescent above, silvery-green and sericcus beneath to densely wliite tomentose; cypselas not ribbed. 22. V. sm ithiana - Characters of leaves and cypselas not as combined above. 41 41. Middle leaves (5—)7—11 x 3—5(—7) cm; hairs on brandies and leaves simple; corolla densely ses­ sile glandular (at apex in bud) in lower parts. 6. V. brachycalyx - Middle leaves 2.5-5.5 x 1-2.5 cm; hairs on stem and branches asymmetrically T-shaped; corolla glabrous or sparsely sessile glandular. 4. V. biafrae 42. Involucre 2-6(-7) x 1-5 mm; capitula 3-12flowercd. 43 - Involucre (7—)8—15 x (4—)5—10 mm; capitula (10-) 12-25-flowered or more. 45

78

145 ASTERACEAE: 30. Vemoma

43 Capiiula 3-5-flowercd: cypselas 1.8-2 mm long. glabrous or with a few glands. 5. V. thomsoniana Capiiula (7-)]()-]2-flowered. or more; cypselas 2-3 ram long, hispid. 44 44 Stem with T-shaped hairs. 2 1 V . phillipsiae - Stem with simple, erect or flagelliform liairs. 6. V. brachycalyx 45. Phyllaries glabrous except for the cilialc margins, dark purple; capitula oblong. 2-2.5 times as long as wide ll.V . cylindrical Phyllaries sparsely to densely pubescent with or without sessile glands, floccose to tomentose. light to dark green, or purplish brown; capitula campanulate. 1.5-2 times as long as wide. 46 46 Stem with simple, erect or flagelliform liairs Stem with T-shaped liairs.

47 49

47 Involucre 12-14 x 8-10 mm; capitula in axillary and terminal scorpioid cymes; phyllaries densely arachnoid-tomcntose. purple in upper half with dark-purple mid-vein. 23. V. thulinii Involucre 6-10 x 3-5 mm; capitula in divaricate, richly branched, terminal cymes; phyllaries pu­ bescent on margins and toward the apex 48 48 Leaves ovate to ovate-lanceolate, 4—15(—20) x 3-4 (-7) mm. margins entire to irregularly 1-2-lobed; capitula 10-12-flowercd; corolla 8.5-9 mm long. 2 1. Leaves obovate-oblong or the upper ones linearoblong. 35-45 x 4-7 mm. margins entire; capitula 20-25-flowcred. corolla 6-7 mm long 48. V. piumhaginifolia 49. Leaves 40-155 x 8-70 mm; shrubs. 0.6-2 in high, rarely woody perennial herbs. 50 Leaves 2-25 x 2-7 mm; low shrubs or shrublets. 0.1-0.5(-0.75) m high. 52 50. Style 8-10 nun long, capitula subscssilc. in termi­ nal or axillary scorpioid cymes onwidely diver­ g ed branches. 15 V.karaguensis - Style 4-7.2(-8 7) mm long; capitula shortly pedunculate, in con mbose or paniculate cymes on closely held branches. 51 51 Leaves thinly pale brownish pubescent with the epidermal surface visible; liairs predominantly T-sliapcd. with unequal short arms. 14 V. leopoldi - Leaves densely whitislv-tomcntose beneath with the epidermal surfacc not visible, hairs predomi­ nantly T-shaped with subcqual to equal arms. 13 V. bipontini 52. Erccl shnib. ofien 0.5-0.75 m high with spreading branches. 2 1 V . phillipsiac Low shrubs or perennial herbs, up lo 0.4 m high; stem often dichotomously branched from near the base. 53 53. Cypselas 1-1.5 mm long, densely appressed hispid; leaves trullate or rhomboid, margins 1-3lobcd at middle or near the apex, lobes very shod and the margins appearing rather toothed. 45. V. arahica

-

Cypselas 2-3 mm long, villous, setae long, leaves ovate-lanceolate or linear to linear-elliptic, mar­ gins entire or deeply 3-lobed. 21. V. phillipsiac

54 Cy pselas not ribbed; leaves silvery-green and senceus to densely white tomentose beneath; capitula 4-5 x 4-6 mm. 22. V. smithiana - Cypselas 4-6-ribbcd. 4-6-angled; leaves not sil­ very and senceus but variously pubescent to to­ mentose; capitula 8-13 x 5-10 mm. 55 55 Stem with T-shaped hairs. 56 Stem with simple erect, or flagellifonn hairs. 58 56 Plant scandent with slender branches growing through and over shrub clumps 24. V. wollastonii - Plant with erect, independent branches. 57 57 Leaves sessile, blade 7 - 15 mm wide; capitula 8-10 x 5-7 nun: plant 0.2-0.5 m high. 44. Leaves pcholalc. petiole up to 1.5 cm long, blade 20-65 m i wide, capitula 11-13 x 8-10 mm; plant 1.5-2 m high. 18 V. turbinatu 58 Corolla 7-10.5 nun long, leaves 10-20 x 3-8.7 cm; plant 0.5-2 m high. 59 - Corolla 5-5.5 mm long; leaves 5-10 x 1.5-4.5 cm: plant 0.3-0.5 m high. 60 59 Plant often 0.5-1 m lugh; phyllaries 4-5-senate. apex rccuned: inner pappus 4-4.5 mm long; co­ rolla glabrous, 8.5-10.5 mm long. 19.V. unionis V. in phillipsiac Plant 1.5-2 high; phyllaries 6-8-seriate. apex pointed, ngid. red-violet; inner pappus 6.5-7.5 mm long; corolla glabrous or sessile-glandular. 7-8 mm long 18. V. turhinata 60 Leaves oblong lo oblanceolate; longest phyllaries 7 5-8 5 nun long, hairs on stem simple, llagcllifonn 46 V. popcana Leaves broadly ovate to ovate-lanceolate, longest phyllaries 4- 4.5 mm long. 47. V. uncinata I

V. amygdalina Del. (1826); Decnneurum nmygdolinum (Del.) DC. (1836); Gymnanthemum amygdalinum (Del.) Walp (1843) type. Sudan/Ethiopia |GJ/WG). Fazughli {Fazoglj. Cailliaud s.n (MPU holo ). Cheilusia nbyssinica Sch Bip . nom nud.. quoad Schimper 31 386. and 1986 (UPS); Gymnamhemum abyssinicum Sch. Bip. ex Walp (1843) - type: Til. Genma, Mensach Distr (near Adua|, It. abyss. 1. Schimper 31 jcitcd as 34. in error!, on p. 948) (B lecto.. BM K. UPS iso ). Vernonia adenosticta Fenzl. ex Walp. (1846) tvpc: Sudan/Ethiopia (GJ/WG1, Fazughli [Fazohel), 1837-38. Koischy 479 (B holo.. EA K iso.). 1' eritreana Klatt (1896) - type: EW/EE. Mailam-Dembelas. 1894. Kaiser s.n. (Z holo., K photo). Bracheileina paniculatum R.Br. ex Sail (1814). nom nud.

Slinib or small tree usually branched from near the base. 2-10 m high, mink c 10-40 cm in diameter; bark rough, longitudinally fissured; slash cream with dense black streaks; branch stems densely lenUcellate. terete, striate.

79

145. ASTERACEAE: 30. Vernonia

Figure 51.

VERNONIA AMYGDALINA: 1 up­ per part ot' stem; 2 capitulum; 3 phyllary; 4 - floret; 5 anthers; 6 style arms; 7 cypsela with pappus. No magnifications and no specimens given in orig. Drawn by W. E. Trevithick. (Reproduced with permission from FI. IV. Trop. Afr . . vol. 2, ed 2. fig. 260.)

g abrous or puberulous or rarely densely pilose with dark b own hairs in terminal parts. Hairs simple. Leaves altern ite, elliptic, lanceolate or ovate-lanceolate, me mb ra­ il us, glabrescenl to puberulous and sessile-glandular v th denser hair along veins, (3—)5—22.5 x (1.5—>2—9.5 c : . base cuneate, sometimes asymmetric, margins entire ti serrate; apex acute, subsessile to petiolate; petiole up to 2 :m long, puberulous to pilose. Involucre 4-6 x 3-4 mm. c impanulate. Capitula in dense, richly branched globose o pyramidal terminal panicles, 10-20-flowered. 9-11 x 3 -4 mm at anthesis, pedunculate; peduncle up to 5 mm It ng, sparsely pubescent to glabrous, flattened or angular, b acteate. Receptacle epaleate, flat, areolate. Phyllaries ( -)4(-5)-seriate, continuous with bracts on peduncle, light green with daik-green and sessile-glandular spot n ear apex, coriaceous, glabrous except for the ciliate mar­ gins, not appendaged; outermost ovate, 1.5-2 x 0.5-1 mm, persistent; innermost oblong-elliptic, c 4.5 x 1.2 mm, acute, often tinged purple in upper half, deciduous. Florets v ute to light purple. Corolla densely sessile-glandular in li V'er parts. 7-7.5 mm long, 5-lobcd, lobes c 2.5 mm long, glaorous. Anthers c 2.5 mm long, base sagittate, obtuse. 0 ::-0.3 mm long. Style c 10 min long, papillose near fork, b inches c 2.5 mm long, densely papillose throughout. Cypselas light-brown, c 10-ribbed. terete, oblanceolate.

3-3,2 x 0.8-1 mm. sparsely setulose and sessileglandular. Pappus l(-2)-seriate. setae more or less equal, some shorter than others, barbellate. 5-6 nun long, white to light brownish-yellow. Fig. 145.51. Podocarpus or Aningeria forest, usually in open spots near streams, or in fringe of glades, secondary forests, evergreen woodland and bushland, roadsides, wasteland, also grown in backyard gardens; (650-) 1200-3000 in; EW TU ?WU GD GJ SU WG IL KF GG SD BA HA; Su­ dan. E Africa and Yemen. Mooney 5633; Mesfin T. 879. Friis et al. 544. The leaves are used together w ith those of R ham nus p rin o id es L'Herit. in the p re p a ra tio n of local beer (TELLA) but more commonly they are used as fumigants. The bark is used as a febrifuge. 2. V. vabelloana Mesfin (1997) type: SD. Yavello. 5700 ft. Bally 9234 (EA holo.. K iso ). Tall, shrubby herb. Old stem greyish-brown young branches densely tomentose. Leaves simple, alternate, green and pilose above, densely white-toinentosc beneath with 4-5 conspicuous secondary veins, broadly elliptic.

80

145. ASTERACEAE: 30. Vemonia

Figure 52. I E R N O N IA YA B E L L O A N A 1 up­ per stem portion with a few leaves stippled to indicate their discolorous nature x Vy, 2 5: phyllaries from out­ side to inside (2 ,4 & 5 x 4 ; 3 x 5'/^), 6 floret x 5; 7 anther lobes with upper part o f filament x 22; 8 style-arm branches with dense covering o f stiff hairs x 22, 9 pappus bristles x 5Vi. All from Bally 9234. Drawn by Mesfin Tadcsse. (Reproduced with perm is­ sion from Kew Bull. 52. 1997. p. 586. fig. 2.)

3-7.5 x 1.5—3.5 cm. margins serrate, teeth apiculatc. base rounded or cuncatc. apex acute-acununate, petiolate. peti­ ole densely tomentose. up to 1 cm long. Capitula campanulate. 15-17x8-12 mm at anthesis. terminal, cymosc. c 25-30-flowered. sessile, or peduncle only up to 5 nun long, densely tomentose. Phyllanes 6-7-seriate. not appendaged; outermost continuous with bracts on pcdunclc. densely tomentose. lanceolate. 3-1 x 1-1.5 nun. acute-acununate at apex, margins scarious. middle scncs obovate with a scarious rotuided or tmncatc. fimbriate shoulder and long apiculatc (up to 2 mm) spinescent apex, sparsely pubescent and purplish in middle. 6-8 x 2-3 nun; innermost oblong-lanceolate, margins scarious. fimbriate or lacerate, apex acute-acuminate, glabrous. 1()—13 x 1.7-3 nun. Florets mauve. Corolla tubular, sessile-glandular in lower parts. 4-6 mm long. 5-lobcd. lobes c I nun long, papillose. Anthers 2.5—3 mm long. Style 4 5-6 nun long, bifurcate, branches 1.5-2 nun long, papillose throughout.

Cypselas (only immature seen) obovatc-cy lindnc. c 2 x 0.5 nun, brown, c 10-ribbcd. villous. Pappus biscriate. brownish-white. barbellate; outer scries 0.5-0.8 mm long; inner 4.5-5 5 nun long. Fig 145.52 Presumably Acacia Commiphora bushland on red sand; c 1740 in SD; not known elsewhere. Bally 9234. The most distinctive feature of I '. vahelloana is the long apiculatc scarious middle phy llanes with the spme-likc extensions projecting from a rounded or trun­ cate apex. 3. V. urticifolia A. Rich. (1848) type SU. ‘C hoa\ Petit s.n. (P holo.. K photo) |as I ’ urticaefolia]. I '. Jlageflaris Vatke (1875) - type: GD, Debre Ta­ bor. 1863. Schimper 1499 (B destr., BM K syn ); GD. Gaffat. 1863. Schimper 1501 (B destr.. BM K sy n ).

145. ASTERACEAE: 30. Vcrnonia

V. antinoriana Avetta (1889) - type: SU, WoinaDega. Let Marefia,Antinori s.n. (FT holo., K photo). Scrambling or scandcnt slirub with lianoid or flexuous 'ranches. up to 4.5 m liigh. Stem striate, terete, glabrous o pubescent, liairs simple. Leaves alternate, ovate, narowly elliptic or lanccolatc; base cuneate; margins vcrrulatc. crenatc-dcnticulatc or subcntirc; apex acute or cumulate; surfaces scabrid above and thinly pubescent >eneath, mainly on veins, or rarely glabrous and shiny, up o 14.5 x 7.5 cm; petiole up to 7(—15) mm long. Capitula ampanulate, 8-10 x 5-6 mm at anthesis. c 15-flowered. n congested or loose axillary- and terminal paniculate vines; peduncle up to 7 mm long, sparsely pubescent. Receptacle epaleate. Phyllaries 5-scriatc, thinly floccose uth hairs mainly along margins on the outer ones; apex cute, mucronulate. purple-bordered or tinged, not tppendaged: outermost ovate. 2-2.5 x 1.0-1.2 mm, innernost 5-6 x 1.5-2 mm long, oblong. Florets white, rarely ale pink or lilac. Corolla tubular, glabrous. 10-11 mm ong. 5-lobed. lobes 2.5-4 mm long. Anthers c 3 mm long, brownish-yellow, base sagittate, obtuse, c 0.5 mm long. Style bifurcate, c 13 nun long, glabrous, branches 3.5-4 nm long, papillose; stigma blackish. Cypselas light>rown. 10-ribbed. c3.2 x 1 mm. thinly hirsute to glabrous. : .ippus 2-senate, white, barbellate; outer series c 2 mm long, inner c 6-7 mm long. In montane forest or moist scmb border, in steep ra. ines usually hanging from slopv ground, in underbush; f '00-2600 m. TU/GD GJ SU AR KF SD BA HA; Kenya, i.ep. Dem. du Congo. Mesfin T. 756. 5703; Mooney 8854. 5134. 4 V. biafrae Oliv. & Hiem (1877) type: Cameroun. Mt. Cameroun, Mann 1296 (K holo.). V. rhodocalymna Chiov. (1940) - type. SD. Wondo. Marchetti 40 (FT holo.. K photo). A climbing slinib. up to 4 m high. Stem terete, striate. Miberulous with long-stalked asymmetrically T-shaped h;urs. Leaves alternate, ovate to ovate-lanceolate, mcm'•'ranous, 2.5—5.5 x 1-2.5 cm. petiolate, base subtruncate broadly cuneate; surfaces thinly pubescent especially on veins and minutely sessile-glandular; hairs simple: margins sinuate, dentate, denticulate to subentire; petiole lender, up to 1.2 cm long. Capitula campanulate. 12-15llowered. c 10 x 5 nun, in open, terminal, divaricating, paniculate cymes; peduncle up to 2 cm long, slender, stria.ic. pilose, with 3—4 scattered bracts. Receptacle epaleate. Phyllaries 4-5-seriate, pubescent and glandular outside. . mceolate to narrowly elliptic (innermost), margins -arious, not appendaged; outer 1-2 x 0.7-1 mm, with a iieen or dark-purple stripe at middle, persistent, acute or s iibapicuJate; inner6—7 x 1—1.5 mm. purple in upper parts, iducous. obtuse at apex. Florets purple. Corolla funid-shaped. glabrous or sparsely sessile-glandular. 9-9.5 nm long. 5-lobed. lobes 2-2.5 nun long. Anthers c 2.5 mm long, base sagittate, acute, c 0.5 nun long. Style c 10 Him long, setose near fork, branches c 2 mm long, tapillose. Cypselas light brown, c 3 x 0.8 nun.

81

oblancelate, 5-6-ribbed with white hairs and sessile glands in between the ribs. Pappus 2-seriate, white, barbellate; outer series c 0.5 mm long; inner c 7.5 mm long. Riverine forest; 1500-2410 m. IL SD. west and cen­ tral tropical Africa. Sudan. E Africa. Malawi, Zambia. Ensermu K. et al. 917; Mesfin T. 6912.

5. V. thomsoniana Oliv. & Hiern ex Oliv. (1873) - type: Uganda, Madi, 1862, Grant s.n. (K holo ). Shrub or small tree' with well defined trunk, up to 3 m high; brandies striate, terete, puberulous. Leaves alter­ nate. thin, stiff, ovate, lower ones up to 18 x 11.5 cm, often deciduous, cordate at base, branch leaves usually 5-8 x 2—3.5 cm. attenuate at base into the narrowly winged, up to 2.5 cm long petiole, puberulous to scabrid above, thinly tomentose beneath. Capitula oblong-campaiiulatc. c 5 x 2 mm, (3-)4(-5)-flo\vered. crowded in corymbose terminal panicles; peduncle c 2 nun long, pilose/hirsutc. Recepta­ cle epaleate. Phyllaries 4-5-seriate. glabrous except for the ciliate margins, not appendaged; outermost ovate, c 0.5 x 0.8 nun. greenish; middle series c 1.2 x 1 mm. light-green in middle; innermost elliptic, c 3-3.5 x 1 mm. pale yellowish-green, margins scarious. tinged purple, ob­ tuse. erase or denticulate at apex with minute sessile glands. Florets white, blue-purple or lilac, fragrant. Co­ rolla funnel-shaped, glabrous to sparsely sessileglandular. Cypselas light-brown. 4-5-angled, glabrous except for the minute scattered sessile glands on some parts, c 1.8-2 x 1 nun. Pappus 1-seriate, barbellate. c 3 nun long, white. Stereospermum-Combre turn wooded grassland, open woodland with Acanthus, Dalbergia. etc; margins of rem­ nant Podocarpus forest; 1700-2200 m; GJ SU IL KF SD BA; Uganda. Tanzania, Sudan, south-central and south­ ern Africa to Zimbabwe and Angola. Friis et. al. 2279; Gillett 14849; Mesfin T. 7636.

6. V. brachycalyx O. Hoffm. (1895) types: Tanzania. Kwa Mshusa. Holst 9096 (BM K syn ): Kenya, Asi. Fischer 356 (B syn.. destr.); Volkens 524 (B syn., destr.); Kenya. Teita. Hildebrandt 2466 (BM K syn.. selected as lectotype by Beentje, 2000.) & 2499 (B syn., destroyed); Rep. Dem. du Congo, Stuhlmann 3051, Pogge 1290. 1304. and 1306. (B syn. destroyed). V. jodopappa Chiov. (1935), nom. illeg.. non Sch. Bip. (1845); V. jodopapposa Chiov. cx Lanza (1939) type: Mt. Kenya, Nycri, Balbo 428 (TOM holo, not seen. FT iso). The collector's name does not appear on the isotype at FT. V. jodopapposa Chiov. ex Lanza forma ere eta Chiov. ex Lanza in Miss. Biol. Borana 4, Racc. Bot. : 224 (1939)-type: SD. Mega. 1800 in. ( 'ufodontis621 (FT holo.). V brachycalyx O. Hoffm. var. megana Cufod. in N. Giorn. Bot. ital., n.s. 50: 102 (1943) - type: SD. Mega. Corradi 1742-1744. 1760 (FT syn.).

82

145. ASTERACEAE: 30. Vemoma

Shrub or perennial herb, 0.75-4 m liigh; young branches slender, flexuous or scandent, densely appressedtomentose; hairs brownish, simple. Old stem glabrous, lenticellate, dark-brownish. Leaves alternate, ovate to ovate-lanceolate, (5—)7—11 x 3-5(-7) cm, petiolate, densely sessile-glandular, sparsely pubescent above, pu­ bescent to thinly tomentose beneath with denser brown­ ish, simple liairs along veins; base broad, abruptly decurrent on petiole; margins denticulate-serrate; apex acute-acuminate; petiole appressed tomentose. up to 2 cm long. Capitula campanulate, 5-6 x 3-3.5 mm. in dense terminal and axillary coiymbose cymes; peduncle sessile glandular, pilose, up to 3 mm long. Receptale epaleate. Phyllaries 3-seriate, green with darker green mid-vein, thinly pubescent especially on margins and near apex, sparsely sessile-glandular, not appendaged; outermost c 1 x 0.3 mm, linear; innermost 3-4 x 0.8-1 mm. linear-lanceolate, often purple at apex or on mar­ gins. Florets mauve, blue to light purple. Corolla densely sessile glandular (at apex in bud) in lower parts, fun­ nel-shaped with a slender (c 4 mm long) tube, c 9.5 mm long, 5-lobed, lobes c 2.5 mm long. Anthers c 2.5 mm long, base sagittate, acute, c 0.4 mm long. Style c 10.5 mm long, papillose near fork, branches c 2.5 mm long, papillose. Cypselas light-brown, obovate, curved, incon­ spicuously 5-ribbed with few scattered hairs on ribs and near apex, otherwise glabrous, 2-2.2 x 0.8-1 mm. Pappus 2-seriate, white, barbellate, outermost series c 0.5 mm long; inner c 4-5 mm long. Open Combretum-Terminalia woodland; open bushland with scattered Acacia, Lippia and Grewia; TecleaAcokanthera scrub; Tarchonanthus camphoratus scrub replacing Juniperus forest; grassland; 1300-1850 m. SD. E Africa Rep Dem. du Congo, Rwanda Burundi. Ma­ lawi, Zambia, Angola. Mooney 7376; Gilbert & Sebsebe D. 8797; Mesfm T. et al. 3358'.

7. V. theophrastifolia Schweinf ex Oliv. & Hiern (1877) type: Sudan, Nahambosso. Schweinfurth 2990 (K holo.). V. myriocephala A. Rich. (1848), nom. illegit.. non DC. (1836); Cacafia richardiana O. Kuntze (1891); Vernonia richardiana (O. Kuntze) Pic.Semi. (1950) type: GD, Gondar, Quartin-Dillon (P holo.). V. macrophylla Chiov. (1911), nom. illegit., non Less. (1831) - type: GD, Tzellemti. Chiovenda 768 (FT syn, K photo), Chiovenda 3184 (FT syn, K photo). V. seretii De Wild. (1907) - type: Rep. Dem. du Congo, Niangara to Gumbari. Seret 438 BR syn); Amadi to Suronga Seret 344 (BR syn.). Shrub or perennial herb, up to 3 m high. Stem profusely branched above, especially in the inflorescence, hollow, branches spreading. Leaves alternate, thin, up to 30 x 9 cm. glabrous to thinly puberulous on lower surface, acuminate towards the sessile, auriculate and amplexicaule base; margins denticulate with narrow acute teeth terminating the slender veins Capitula campanulate. 7-9 x 3^1 mm, in open divaricately branched terminal or axillary cymes; peduncle up to 5 mm long, thinly pubes­

cent. Receptacle epaleate. Phyllaries 4-5-seriate. coria­ ceous, glabrous except for the ciliate margins in outer and middle ones, not appendaged, outermost ovate or suborbicular. apiculatc. c 1 x 1 mm, persistent, innermost ob­ long-lanceolate, 5-6 x 2-2.5 mm, caducous, purplebordered or appearing dark-green upon dry ing, obtuse. Florets pale violet. Corolla funnel-shaped, glabrous. Cyp­ selas light brown (only immature seen), c 3.5-4 x 1 mm. oblanceolate, hispid, c 10-ribbed. Pappus wliite, sev­ eral-seriate, unequal, barbellate. outer series c 1 mm long; innermost 6-7 mm long. Forest margins, riverine forests, mountain grassland with scattered scrub; 900-2000 m. TU GD SU IL KF GG: Sudan, Uganda. Kenya, Rep. Dem. du Congo. Nigeria Friis et al. 6011; Tekle H. Hagos 11; Mesfin T. 8039. 8. V. ischnophylla Muschl. (1911) - type: .Rep. Dem. du Congo. Kivu-Rugege. Mildbraed 1073 (B holo.. destoyed, BR iso ). Shrub up to 2.5 m high with numerous stems from the rootstock. Stem unbranched except in the inflorescences, striate-sulcate. leafless in lower parts, puberulous to glabrescent. Leaves alternate, oblong-elliptic, erect (held at an acute angle to the main avis), up to 20 x 3 cm. pale green and glabrous above, paler and whitish crispatetoinentose beneath (often rusty-brown upon drying), coriaceous, narrowed into a sessile auriculate base, mar­ gins denticulate-serrate. Capitula oblong-campanulate, c 10 x 5 mm. 7-10-flowercd. in dense terminal paniculate cymes, peduncle up to 7 mm long. Receptacle epaleate Phyllaries 4-5-senate. glabrous to thinly floccose. green­ ish in upper parts (appearing spotted upon dry ing), not appendaged; outermost ovate or orbicular, c 0.5-1 x 1.5 mm; innermost oblong-elliptic, c 5-6 x 2 mm, apiculatc. margins scarious. Florets deep purple; corolla glabrous, funnel-shaped. Cypselas brownish, c 10-ribbed. densely setulose, c 3.5-4 \ 1 mm. oblanceolate. Pappus 2-seriate. wlute. barbellate; outermost series c 1 nun long; inner 8-9 mm long. Woodland, gallery forests, thicket border, grassland, roadsides; 1450-2(KK) m. GD GJ WG KF SD BA; Uganda. Rwanda, Burundi. Rep Dem. du Congo. Moo­ ney 5477; Ash 1449; Mesfm T. 5228. Material of this species lias often been misidentified as V. seretii De Wild 9. V. m yriantha Hook. f. (1864) ty pes: Cameroun. Mt. Cameroun. Mann 913 (K sy n ); Bioko |Femando Po|. Mann 622 (K syn.). I', podocoma Sch. Bip. ex Vatke (1875); I. oliveriana Pic Semi. (1950). nom. illegit. - type: Ethi­ opia |exact place unknown). “Clioa and Oudgeratc', Petit s.n. (K lecto ). V ampla O Hoffm. (1901) - type: Tanzania Goetze 1286 (B holo.. destroy ed. BR. P. iso ). I ’, chiarugu Pic.Semi. (1951) - type: GJ. Amba Covina, near Bahir Dar. Pichi-Sermolli 2426 (FT holo. K photo).

145. ASTERACEAE: 30. Vernonia

83

V. cvlindrica auct., non Sch. Bip. ex Walp. (1843): A. Rich. (1848), p.p.

0.5 nun. Pappus several-seriate, white, barbellate: outer­ most series c 1 nun long; inner c 5-6 nun long.

S nub or small tree, up to 6 m high, with widely spreadir ii branches; branch stems puberulous to tomentose. densely lenticellate. Leaves alternate, ovate-elliptic to e lptic, narrowed to an obtuse or subcordate-base, g ;ibrescent to puberulous above, thinly whitish tomentose beneath, margins denticulate-serrate; middle ones 2 i- 25 x 8-11 cm, lower up to 42 x 16 cm; petiole of midd p leaves 4-6 cm long, puberulous. Capitula oblong or c Imdric, up to 1 2 x 3 mm, (3-)4-5-flowered, in dense, spreading, terminal and axillary panicles; peduncle up to 0 5 mm long, tomentose. Receptacle epaleate. Phyllaries 5~t-seriate, coriaceous, not appendaged; outermost p iberulous to tomentose, ovate-rotundate, c 1-1.5 x 1.5 mm. apex acute; innermost oblong-elliptic. 5-6 x 1-1.5 ran, glabrous or sometimes ciliate on the scarious and p jrplish margins, apex obtuse, mucronate, often darkpurple. Florets pale lilac, lavender, white, blue or re J-purple. Corolla funnel-shaped, glabrous. Cypselas light-brown, 10-ribbed, thinly pilose with small sessile giands in the intercostal grooves, c 3 x 0.8 mm. Pappus 2-seriate, brownish-white, barbellate; outer series 1-2 n m long: inner 8-9 mm long.

Forest margins, moist woodland or grassland; 1200-2200 m. WG IL KF SD ?BA; E Africa. Nigeria. Cameroun, Rep. Dem. du Congo, Burundi, Angola. Ash 1407; Friis et al. 2083; Mesfin T. 7902.

Riverine forest, stream banks, moist woodland, sec­ ondary scrub on moist ground; 1750-2900 m. GJ SU AR KF GG SD BA HA, E Africa, Sudan, Guinea, Sierra Le­ one. Cameroun, Rep. Dem. du Congo, South tropical Afn.ia. Burger 3641; Ash 1851; Aiesfin T. 8304. Material of this species lias often been named V. subuligera O. Hoffm. 1' i V. au ricu lifera Hiem. (1898) - types: Angola. Golungo Alto & Cazengo. Welwiisch 3258 (BM syn.., K isosyn.); Humpata. Wetwiisch 3262 (BM syn., K isosy n.). SLrub or small tree, up to 6 m high. Stem densely bfanched, branches striate, lenticellate, glabrous to hairytc mentose in terminal parts. Leaves alternate, elliptic to oblanceolate. up to 22 x 7 cm (upper), lower ones deciduous when the shrub/tree is in flower, cuneate or narrowed ai base and shortly decurrent on petiole: margins serrate w ith apiculate teeth: apex obtuse or acute; surfaces gla­ brous. light yellowish-green above and paler below, or thinly pilose on lower surface: young developing leaves o r :n wliite-tomentose beneath; petiole c 2-3 cm long, oftc p. auriculate at base. Capitula oblong-cylindric, 1-2 ( : )-flowered, sessile, in dense spreading terminal and a\illary panicles, up to 10 x 2 mm. Receptacle epaleate. Phyllaries 5-seriate. coriaceous, not appendaged; outer­ most ciliolate on margins and thinly on surfaces, c 0.5 x 0 ; mm. ovate: middle c 3 x 1.3 nun. ovate-elliptic; inneru ost about twice as long as middle ones, greenish and p irple-bordcrcd at apex, c 5 x 1.2 nun. elliptic with acuminate apex. Florets pale pink, mauve, whitish or Hue-violet Corolla funnel-shaped, glabrous. Cypselas greyish-brown, c 10-ribbed. thinly hirsute with small biownish sessile glands in the intercostal grooves, c 3 x

11. V. cylindrica Sch. Bip. ex Walp. (1843) type: TU, Zadda Kolia [NE of Adual, 1800 m, PI. abyss. II, Schimper 640 (BM K isosyn.). Wehbia serratuloides SckBip. nom. nud.. quoad Plow den s.n Shrub up to 3 m high. Stem puberulous to glabrescent. subterete, reddish-purple. Leaves alternate, ovate, coria­ ceous, scabrid. up to 1 7 x 7 cm, base rounded, decurrent on the petiole, margins minutely serrate, apex acute; peti­ ole up to 7 mm long. Capitula oblong- campanulate to cy­ lindric, up to 1 5 x 6 mm, in open corymbifomi terminal panicles; peduncle pilose with pinkish liairs, up to 2.5 cm long. Phyllaries (5-)6-7-seriate, closely appressed, coriaceous, shiny, purplish, glabrous except at margins where ciliate, obtuse at apex, not appendaged; outermost c 2 x 2 mm, ovate; innermost 8-9 x 2.2-3.5 nun, oblong or oblanceolate. Florets purple. Corolla glabrous, cylindric. Anther base obtuse. Cypselas light-brown. 5(-6)-ribbed or angled, minutely glandular, glabrous, oblanceolate. c 3 x 1 mm. Pappus 2-seriate, white, barbellate; outer series c 1 nun long; inner c 5 mm long. Scrub on dry hillsides, often a characteristic species in undergrowth of bamboo thicket of Oxvtenanthera: Combretum-Terminalia woodland: 1300-1700 m. TU GD GJ WG, not known elsewhere. Friis et al. 7885: Pichi-Sermolli 2438. Mesfin T. & Kagnew 2278. 12. V. rueppellii Sch. Bip. ex Walp. (1843) -type: TU, between Halei & Tembein. June 1832. Ruppell s.n. (P holo., K photo). V. francaviliana Oliv. & Hiem (1877) - type: GD. Gondar. D 'Hericourt s.n. (P holo., K plioto). V podocoma Sch. Bip. ex Vatke var. glabrata Fiori in N. Giorn. Bot. Ital. 47:43 (1940) - type SD. between Wondo (Uondoj and Agare Salaam. Dec. 1937, Saccardo, s.n. (FT holo.). Shrub or small tree with a rounded crown, up to 6 m high. Stem with a diameter of up to c 40 cm at base; branches subterete, brown or purplish-brown. Leaves alternate, narrowly elliptic, up to 1 5 x 5 cm, glabrous, attenuate at the base into the petiole, dark green above, paler beneath: margins serrate to doubly-serrate with sharp teeth, firmly membranous; petiole up to 3 cm long. Capitula oblong campanulate, 8-9 x 3-4 mm. (4-)5(-6)-flowered. sessile or subsessile, in dense terminal corymbose panicles: peduncle, when present, up to 1.5 nun long, pilose. Re­ ceptacle epaleate. Phyllaries 4-5-seriate, glabrous except for the ciliate margins of the outer ones, usually dark pur­ ple, not appendaged: outer ovate c 1 x 1 nun, purplish at the acute apex; inner oblong-lanceolate, purple-bordered, c 7 x 2 mm, margins scarious, apex acute. Florets pale pur­ ple to white. Corolla funnel-shaped, glabrous. Cypselas

84

145. ASTERACEAE: 30. Vemonia

grey or dark-brown, c 5x1 mm, c 10-ribbed, setulose. ob­ lanceolate. Pappus 2-seriate, unequal barbellate. white to yellowish white, outer c 1 mm long; inner 8-11 mm long

-

Outer phyllaries 2.3—5.3 mm long, bracteoles on peduncles equal to or exceeding them; leaf mar­ gins indistinctly to very finely serrate. 2

Forest margins, grassland with evergreen scrub. Croton-Calpurnea woodland on montane slopes; 2150-3000 m. EW GD TU WU SU AR KF SD BA HA; not known elsewhere Gilbert et al. 7941; W. de Wilde 10037; Mesfin T. 9192.

2

Phyilaiy-like bracteoles and outermost phyllaries densely tomentose, apex obtuse or acute; inner­ most phyllaries shorter than 6 mm. var. gonderensis Phyllary-likc bracteoles loosely tomentose except at the acute-apiculate to acuminate apex: inner­ most phyllaries longer than 6.3 nun var caccaensis

13 V. bipontini Vatke (1875); Gyanopis hvpoleuca Sch. Bip. ex Walp. (1846). nom. illcgit.. non DC. (1836); I ernonia leopoldi (Sch Bip. ex Walp.) Vatke var. hypoleuca Chiov. in Malpighia 35. 59 (1939) - type; TU. near Axum. PI. abyss. Ill,Schimper 1542 (B holo.. BM K P UPS iso ). Vemonia leopoldi (Sch. Bip. ex Walp.) Vatke var incana Avetta in N. Giorn. Bot. Ital. 21: 349 (1897) - type: SU, |exact locality not known|. Dec.-Jan 1893. Antinori s.n. (FT holo ). Cyanopis leopoldi auct.. non Sch Bip ex Walp (1846-47), quoad Schimper 246. Shrub, or rarely erect to straggly, woody herb. 0.6-1 5 m high Indumentum of densely apprcssed. greyish-white to brownish-grey. T-shaped liairs with equal to subcqual filiform amis Leaves pilose-scabrous or tomentose above, densely whitish tomentose to velvety beneath. 4.5—14 x 0.8-4.5(—6.2) cm, base dccurrent. petiole 2-10 (-25) mm long. Capitula 8-14 mm long at anthesis. 5-11 mm long when fruiting: peduncles 2-15 mm long, densely brownish-white velvety tomentose. Involucre 4-9 x 4-6 nun. Phyllaries not appendaged: outermost linear. linearlanceolate to narrowly triangular. (1.1—)1.5—3.4 (-5.3) x 0.3-0.5(-0.9) mm, apex attenuate-apiculatc. reflcxcd; middle phyllaries ovate-oblong, or elliptic-lanccolatc to oblanceolate. 2.5-5.3(-6.5) x 0.5-1.1 (-1.7) mm. margins sometimes minutely toothed, outer surface densely appressed whitish-grey tomentose on all exposed parts, inner surface papillate, apex rcflexed. apiculatc: inner phyllaries 4.3—6(—8.5) x 1.0—1.2(—1.6) mm, acute to apiculate at the apex. Corolla glabrous. (3.5-)5.6-8.4 (-10.8) x (0.6-)0.8-l. I mm: anthers ( 1.5—>2—3.8(—4.2) mm long, style (3-)4.7-7.2(-8 7) mm long. Cy pselas (1.3-) 1.6-2.2 (—2.6) x (0.3-)0.4-0.7(-0.9) mm. 4-5(-6)-ribbcd. Pappus: outer series coromfonn rim, 0.1-0.4 mm long, inner bristles (3.6-)3.9-6.0(-7.0) nun long. Figs 145.53.1-7; 145.54. 5-17. This species is highly variable in its morphology. It is easily confused with I', leopoldi (Sch Bip ex Walp.) Vatke. to which it is closely affiliated. It is, however, dis­ tinct from the latter species, yet the intra-specific variation in I bipontini is so large tliat one is tempted to ascribe specific rank to many of the most deviating populations. This high degree of variability within the species seems to be attributable to its wide ecological amplitude. Three va­ rieties have been rccognizcd |S. Wattimah & Mesfin T.. in Nordic.). Bot. 14(2): 209-219 (1994)|. 1

Outermost phyllaries 1.1—2.1 nun long, bracteoles on peduncle not exceeding them, leaf margins distinctly serrate orcrcnate. var. bipontini

var. bipontini Stems unevenly wlutc-tomcntose. several from the root­ stock. woody , 0.3-1 m high. Leaves ovate to ovatelanceolate, margins serrate or crenate. upper surface sub-scabrous with multiseriate. multicellular. glandular and T-shapcd hairs. Outermost phyllaries 1.1-2.1 ram long, lanceolate Innermost phy llaries 5.2-7.6 mm long. Cypselas 1.6-2 2(-2 6) 111111 long. Inner bristles of pappus 4-6 mm long Fig 145.53.8-14. Dry shrubby broad-leaved woodland, bushland and disturbed areas; 1000-2900 m. EW TU WU GD GJ SU; Yemen and Saudi Arabia. Friis et al. 1530; Mesfin T &. Kagnew 1955; Mooney 6282 var gonderensis .S' Wattimah A Mesfin in Nordic J. Bot. 14(2): 218(1994) - type: GD. Gondcr. Mesfin T. & Kagnew 1835 (ETH holo.). Stems unevenly whitc-tomcntosc, several from a woody rootstock, c J 111 high. Leaves ovate to ovate-lanceolate, margins sub-entire with minute even serrations, upper surface subscabrous with both multiseriate. multicellular and 1-senate, glandular liairs and T-shaped hairs Capitula subtended by long phyllary-like, filifonn bractcoles tliat arc densely white-feltish tomentose. 2.3-5 3 nun long, almost the same length as the involucre. Innennosi phy llaries lanceolate. 5.3-5.8 111m long. Cy pselas 1.3-2 1 111111 long Pappus bristles 4.0-5.1 mm long. Fig 145.53.17 & 18. Along river banks or in cultivation, rarely in open bushland; 2090-3300 111. GD SU; not known elsewhere Mesfin T & Kagnew 1835 and 1919; Tewolde B.GlF. 1189. Friis et al 3825. from a veiy wet seepage area in open grassland at 2600 m. is probably also this variety . var caccaensis .S' Wattimah & Mesfin in Nordic J. Bot. 14(2): 219(1994) - type. AR, Mt. Cacca. Mooney 5250 (ETH holo . EA FT K iso ). Stems unifonti/v pale-whitish tomentose. several from a woody rootstock. 1-1.5 111 high Leaves ovate-lanceolatc, margins minutely serrate with the bases of the teeth indis­ tinct. upper surface pilose with 1-seriate, glandular and T-shapcd liairs. lower surface silvery to greyish-white to-

145. ASTERACEAE: 30. Vernonia

85

mentose. Phyllaries: outermost and middle elongated to a tout half the length of the involucre, apices purple, f iform, gradually extended into reflexed tips. Cypselas 1 9-2.4 mm long. Pappus bristles 5.4-5.8 nun long. Fig.

1 45.53.15 & 16. Open grassland on shallow soil or in rocky crevices on r icky slopes; 2700-3350 m. SU AR KF; not known else\ here. Mooney 5250; Ash 2333; Friis et al. 1429. 1 - V. leopoldi (Sch. Bip. ex Walp.) Vatke (1875); Cyanopis leopoldi Sch. Bip. ex Walp. (1843) type: TU, Adwa, Mt. Scholoda, PI. abyss. I, Schimper 9 (P lecto.. BM K UPS isolecto.). Vernonia natalensis auct.. non Sch. Bip. ex Walp. (1843): Cufod. (1967: 1073), quoad Mooney 5250. S lirub or rarely woody herb, 0.5-2.5 m high. Indumentum brownish-grey, with short appressed T-shaped hairs with sic ut unequal arms. Leaves glabrescent. pilose or shortly strigose above with denser hair especially on the veins ber :ath, 9-15.5 x 5-7 cm. base obtuse to decurrent, petiole (2 )3-6(-8) mm long. Capitula 8-11 mm long at anthesis. 6-8 mm long in fruit. Involucre 3-7 x 4-6 mm. Phyllaries r )t appendaged; outermost usually appressed. linear lancjolate to ovate lanceolate. (1.2—)1.5—2.7(—3.2) x (0.3-) ( 4 - 0.6(-0 8) mm, inner surface glabrescent, papillate, apex acuminate; middle series appressed, oblong lanceo­ late, oblanceolate, or elliptic lanceolate. (2.5—)3.5—5.5 (|5.8) x (0.6—)0.8—1.4(—1.6) mm. margins entire to mi­ nutely toothed at the apex, outer surface tomentose at the ap-cx. inner surface glabrous or glabrescent. papillate; innermost (3.7—)4.2—6.6 x 0.6-1.6 mm. Corolla glabrous, 4 -6.6 x 0.4-0.9 inin; anthers 2.3-3.2 nun long; style 4 ; -6.5 mm long, branches 1.5-2.9 nun long. Cypselas (1 )-)l.3-2.3 x 0.5-0.7 mm, (4-)5(-6)-ribbed. Pappus: outer coroniform rim 0.1-0.4 nun long, inner bristles 3 3-5.6 mm long. Fig. 145.54.1—4. Forest margins. Acacia wooded grassland with scrub of Maytenus, Rosa abyssinica and Carissa on shallow soil, often in ravines, roadside thickets, wasteland; 1830-2850 in. TU GD GJ WU SU WG KF HA GG; not k own elsewhere. Mesfin T. & Kagnew 2043; Mooney 5i 15; W. de Wilde 8956. i : V. karaguensis Oliv. c£ Hiern (1873) type: Tanzania, Korogwe, Grant s.n. (K holo ). V. campanea S. Moore (1916)- type: Uganda, 160 km NW of Kampala. Brown in Dummer 2652 (BM holo., K iso ). V. nielanacrophvlla Cufod. (1943) - types: GG/SD El Dire, Corradi 1805, 1806. 2035. 2036. 2117 (FT syn.. K photo) and between El Baiuio and El Dire. Corradi 2057. 205 (FT syn., K photo). V. nielanacrophvlla Cufod. var. hispida Cufod. in V. Giorn. Bot. Ital. n.s. 50: 103 (1943) - type: SD. Negelli. Corradi 1984 (FT holo.). V. nielanacrophvlla Cufod. var. pseudoblumeoides Cufod. in N. Giorn. Bot. Ital. n.s. 50: 103 (1943) - types. SD. Mega, Monte Pelato. Corradi 2065, 2066, 2072 (FT syn.).

Figure 53. VERNONIA BIPONTINI. 1- part o f upper stem , 2 6: leaf types x '/3; 7 capitulum x 1. var. BIPO NTINI 8 10 outermost phyllaries x 5; 11 - middle phyllary x 5; 12 14 - in­ nermost phyllaries x 5. var. CACCAENSIS. 15 - outermost phyllary x 5. 16 - innermost phyllarie x 5. var. GONDERENSIS. 17 - outermost phyllary x 5; 18 - innermost phyllary x 5. 1 from Mercier 2778; 2, 10 & 14 from Afeworki & Mehret 12: 3 from Friis et al. 3820; 4, 8, 9, 11-13 from Mesfm & Zerihun 2835; 5 from Mesfin 7268; 6, 7, 17 & 18 from Sehsebe 416; 15 & 16 from Ash 2713. Drawn by Damtcw Teferra. (Combined and reproduced with permission from Nordic J. Bot. 14, 1994, pp. 2 1 6 ,2 1 7 , figs 7 & 8.)

V. melanacrophylla Cufod. var. pseudoprintzioides Cufod. inM Giorn. Bot. Ital. n.s. 50: 103 (1943) -types: SD. Mega. Corradi 1728, 1799. 1874, 1882 (FT syn.). V. leopoldi Vatke var. angustifolia Chiov., nom. nud., quoad Vatova 130, 228. 2177.

86

145. A S T E R A C E A E 30 Vernonia

Figure 54 I E R S O M A LEOPOLD / I capitula \ 2 ‘ 2 . 2 4 phyllaries \ c 5. out­ ermost (2). middle (3), innermost (4). I . B IP O X T IS I 5 9 phyllaries x c 5. out­ ermost (5). middle (6 8). innermost (9); 10 cypsela with pappus x c 5. 11 capitula x 2' 2 . 12 floret x c 5. 13 sta­ men x c 5. 14 17 hairs x 150: 14 T-shaped hair with more or less equal stout arms: 15 longhorn T-shaped hair with round, subcqual and filiform arms. 16 & 17 T-shaped hair with round, stout unequal arms. I from Mesfin A Kagttew 2049; 2 4 from Mesfin T. 7259; 5 from Evans 436; 6 11 from Friis et al 3820; 12 13 from A/eworki Meheret 12. Drawn by Damtcw Teferra. (Rearranged and reproduced with permission from Nordic.!. Bot 14. 1994. fig. 2. p. 210 & fig.6 . p. 215.)

Erect, stiff perennial herb or shmb up to 2 m high. Stein terete, striate, covered with short crisp pubesccne or to­ mentose. Hairs simple (mostly on upper surface of leaves) and long stalked, symmetrically T-shaped Leaves alter­ nate, light green, ovate, narrowly elliptic, lanceolate. 4-7.5 x 2-4 cm. subsessile to petiolate; base broadly cuneate. slightly decurrent on petiole: margins serrate to serrulate with acute-apiculate teeth, upper surface scabrid. lower sparsely to densely pubescent, sometimes sub-tomentose. densely resinous-dotted, apex acute; peti­ ole up to 1.5 cm long, slightly wider at base; pubescent. Capitula subturtnnate. many-flowered. 7-12 x 4-7 mm al anthesis. subsessile on the divergent branches of terminal, scorpioid or corymbose cymes. Receptacle epaleate. Phyllaries (4-)5-6-seriate, oblong-lanceolate, pubesccnl and sparsely sessile-glandular except in areas of overlap, purplish above with daiker mid-vein and tip. not appendaged: outermost 0.5-1 x c 0.5 mm; innermost 3—6 x c 1 mm. with scarious margins; apex acute or subacutc. sometimes recurved Florets red-violet or purple Corolla narrowly funnel-shaped, sessile-glandular. 5.5-9 inm long. 5-lobed. lobes c. 1.5-3 mm long. Anthers 2-3.5 nun long, base sagittate, acute, c 0.2-0.5 mm long. Style 8-10

mm long, glabrous, branches 2.5—3 nun long, sparsely to densely papillose. Cypselas light-brown. 5-ribbed. setulosc and sessile-glandular in the grooves between the ribs, oblanceolate. c 1.5 x 0.5 mm. Pappus white. 2-seriate: outer series c 0.5 mm long, squamulous, fimbriate: inner 3—1 mm long, barbellate. deciduous. Forest edges and clearings of Podocarpus and Podocarpus .Juniperus forest. Combretum woodland, grassland with scattered evergreen bushland. limestone slopes in Acacia ( 'ommiphora bushland: 1400-2750 in. GJ WU SU AR WG IL KF GG SD BA HA; Sudan. E Af­ rica. Nigeria Rep Dem. du Congo. Rwanda. Burundi. Mozambique. Malawi. Zambia. Zimbabwe. Gillett 14492: Gilbert & Tewolde B.G.F. 2518; Mesfin 7. et al. 8026.

16. V. printzioidcs Muschl. (1911) type: SD. Giaribule. Riva 1371 (1341) (B holo destroyed. WU iso.. K photo).

var. printzioidcs

145. ASTERACEAE: 30. Vernonia

P :)fusely branched bushy perennial, 0.5-1 m high. Stem striate-sulcate, pubescent to thinly appressed tomentose on branches. Leaves alternate, ovate to ovate-lanceolate. 2.5-5.5 x 1.0—2.5 cm, branch leaves always much smaller than main stem leaves, densely sessileglandular, pubescent to thinly tomentose especially on lower surface, subsessile to shortly petiolate, petiole c 2 : mm long; margins irregularly serrate; apex acute. Hairs on leaves simple. Capitula hemispheric, both a n a i y and terminal, in paniculate corymbose cymes, 1. 15 x 10-12 cm at anthesis. peduncle densely tomen­ tose, up to 1.5 cm long. Phyllaries 5-6-seriate, continuous with bracts on peduncle, ciliate on margins and sparsely on surfaces, or glabrous and sparsely sessile-glandular, apex appendaged; outer 5-6 x 0.5 mm, appendage green. c : 5-4.5 mm long; middle c 8 x 2 mm, appendage c 4 mm long, densely sessile-glandular at base; innermost 10-11 x 2 : mm, appendage 1—1.5 nun long. Florets white, pur­ ple-tinged toward apex of corolla. Corolla c 16.5-17 mm long, 5-lobed, lobes c 2-2.2 mm long with yellow sessile glands in bud. glabrous or sparsely sessile-glandular at anuiesis. Anthers c 5 mm long, base sagittate, c 1 mm long, obtuse. Style c 20 mm long, glabrous, branch c 3 mm long, sparsely papillose. Cypselas black, obconic, indis­ tinctly c 10-ribbed. thinly hispid, liairs light brown. 4 2 ^ .5 x 1.8-2 mm. Pappus multiseriate. flat, yellowisl -white, ciliate on margins; outermost c 1 mm long; in­ ner 9-10 mm long.

87

9-10 nun long, lobes c 2.5 mm long. Anthers c 3.5 mm long, appendage c 0.8-1 mm long, base sagittate, c 1 mm long, acute. Style c 11.5 mm long; branches c 3.2 mm long, densely papillose throughout. Cypselas brownishyellow, sessile-glandular, glabrous, arcuately oblanceo­ late. 4.9-5.2 x 0.8-1.1 nun. 4-5-angled. 5-6-ribbed. Pappus 2-seriate. wliite; outer 0.5-1 mm long, fimbriate: inner 6-7 nun. barbellate. Mixed woodland and scrub on hillslopcs; c 1300 m. HA; not known elsewhere. Burger 3076. 18. V. turbinata Oliv. & Hiern (1873) - type: Uganda, Madi, Dec. 1862. Grant s.n. (K holo.).

Erect, perennial herb, 1.5-2 m liigh. Stem densely branched, branches striate-sulcate. puberulous to densely wliite tomentose. hairs both simple and long-stalked and symmetrically T-shaped. Leaves alternate, up to 17 x 6.5 cm. scabrid and yellow-green above, paler and sparsely to densely woolly tomentose beneath, oblong-elliptic, broadly cuneatc at base and narowly decurrent on petiole; margins serrate-denticulate (entire in upper ones), apex acute; petiole up to 1.5 cm long. Capitula turbinate. 11-13 x 8-10 mm at anthesis. in elongated scorpioid. paniculate cymes; peduncle absent or up to 3 cm long. Receptacle epaleate. Phyllaries (6-)7(-8)-seriate. moderately to densely tomentose except in the dark-green or purple mar­ gins. glabrous, pointed, rigid, red-violet or purple in upper Hilly grasslands. 1800-2200 m. SD |Mega. Wadcra|. exposed parts, apex with subpatent apiculus. not m t known elsewhere. Ballv 9203; Gillett 14413; Mesfm appendaged; outermost linear-lanceolate. 3—3.5 x 0.3-0.6 T et al. 3343. nun. usually greenish: innermost oblong-elliptic. 6-7.5 x 1.2-1.8 mm. scarious at middle. Florets mauve or var tomentosa Cufod. in N. Giorn. Bot. Ital. n.s. 50: 102 red-violet. Corolla 7-8 mm long, funnel-shaped, glabrous < 1943) - ty pe: SD. Mega, Corradi 1863 (FT holo.). or sparsely sessile-glandular. 5-lobed, lobes c 2-2.3 mm long, shortly setose. Anthers 2.5-3 mm long, base Di ffers from the type variety by its larger, densely tomensagittate, acute. 0.5-1 mm long. Style 8-9.5 111111 long, tose leaves with irregularly crisped margins, and short shortly setose near fork, branches 2.5-3 mm long, setose. bi anches with numerous capitula. Cypselas (only slightly immature seen) orange-brown, c 3 No details on habitat. SD; known only from the type. x 1 mm. (4-)5-anglcd and ribbed, glabrous to sparsely pu­ bescent between the ribs. Pappus 2-seriate. white, 1 V. dalettiensis Mesfm (1997) barbellate; outer narrowly squamifonn. c0.5 111111 long; in­ - type: HA. W of Daletti. Burger 3076 (K holo.). ner c 7 mm long, slightly expanded at apex. Sfirub with ascending branches, up to 2 m liigh. Old stems C'ombretum- Terntinalia woodland with remnant stnate-sulcate, dark grey, young stems light brown. Oxythenanthcra: open woodland with Svzygiunt. d< nsely appressed-pubescent. especially near inflorescenNaudea, Grewia, etc.; dense Oxvthenanthera thicket; ces; hairs asymmetricaly T-shaped. Leaves alternate, grassland; 1300-1860 111. GD GJ WG IL; Sudan. Uganda. b: aadly elliptic, up to 8.5 x 4 cm, densely resinousKenya. Rep. Dem. du Congo. Fergusson A sh 2202'. pi nctate, scabrid on upper surface, liairs simple, sparsely Mesfm T. & Kagnew 2236. tc densely pubescent on lower surface, hairs long-stalked. T shaped with equal filiform arms', petiole densely 19. V. unionis.S'c/7. Bip. ex Walp. (1846) a|: pressed-hispid. 0.5-1.5 cm long. Capitula hemispheric. - type: TU, near Gafta. PI. abyss. 11. Schimper 1 -15 x 10-12 mm, in paniculate, subscorpioid cymes 1223 and 1212 (P K isosyn.). te minating branches; peduncle appressed-pubescent. up I ". bipontini auct., non Vatke (1875): Schweinf. & tc 2.5 cm long. Receptacle epaleate. Phyllaries 5-6Aschers. (1867). quoad Schweinfurth & Riva 1553. s< nate, dark-purple near the apex where densely white puErect, perennial herb, up to 1 111 high, often from a tomenb« scent apex spinous, not appendaged; margins ciliolate; tose-tufted rhizomatous rootstock, with slender roots. 0 ter oblong. 2-2.5 x 1 nun; inner oblong-elliptic, c 8-9 x Stem herbaceous, terete, striate, thinly tomentose espe­ 1 5 mm, sometimes sessile-glandular. Florets bright purcially above, or glabrescent. leafy throughout: liairs sinip j-Iavender. Corolla funnel-shaped, sessile-glandular.

88

145. ASTERACEAE: 30. Vemonia

pie. Leaves alternate, ovate or ovate-elliptic, upper lan­ ceolate, scaberulous above, thinly appressed tomentose or glabrate beneath, 10-20 x 3-8.7 cnr. base cuneate, mar­ gins dentate-serrate; apex acute, petiole narrowly winged, up to 1 cm long. Capitula hemispheric, 11-13 x 7-9 mm, unilateral or scorpioid on the spreading branches of a broadly corymbiform terminal panicle, peduncle densely pilose, up to 2 cm long. Receptacle epaleate. Phyllaries 4-5-seriate, continuous with bracts on peduncle, thinly to­ mentose, recurved or subpatent above, apex acute or acuminate, not appendaged; outer subulate. 3-5 mm long; inner 5-6 mm long, oblong-lanceolate. Florets red-violet. Corolla tubular, glabrous, 8.5-10.5 mm long, 5-lobed, lobes c 2.5 mm long. Anthers c 3.5 nun long, base shortly sagittate, c 0.5 mm long. Style bifurcate, c 10.5 mm long, papillose near fork, branches c 2.5 mm long. Cypselas brownish-yellow. 4(-6)-ribbed, with minute intercostal setae, 2.5-2.8 x 0.8—1 mm. Pappus white. 2-seriate; outer squamulose. 0.5-1 mm long, inner 4.5 -5 mm long, baibellate. Wooded grassland, on steep hill or mountain slopes; 700-2010 m. EW GD TU SU; Sudan. Ash 2691; Schweinfurth & Riva 1553; Gilbert & Thulin 432. Vernonia unionis can be distinguished from V. karaguensis on its larger capitula, its phyllaries, which are continuous with the bracts on the peduncle and the larger and wider membranous leaves.

20. V. cinerascens.S’c/7. Bip. in Schweinf. & Asch.(\%61) - type; TU/GD, Gageros [Agau. in the Tacazze Valley], 1854. Schimper 2133 (BM K iso ). V. spaihulata Hochst., non Sch. Bip. in Schweinf. & Asch. (1867), quoad Schimper 2133. Bushy, perennial herb or shrub. 0.5-2 m high. Stem terete, covered when young with short-stalked. T-shaped, white appressed liairs mixed with small sessile glands, some­ times brownish- or purplish-grey. Leaves alternate, some­ times fascicled, obovate-spathulate, greyish-green, (3-) 8-27(-40) x (2-)4-10(-16) ram, densely resinous dotted, sessile, attenuate at base, entire to serrate, rounded or ob­ tuse at apex, scabrid to densely appressed pubescent. Capitula campanulate. c 10-flowerei 6-7 x 4-5 mm, in Miffuse axillary and terminal cymes; peduncle densely appressed pubescent, up to c 10 mm long. Receptacle epaleate. Phyllaries (3-)4(-5)-seriate. lanceolate to ob­ long, sparsely to densely pubescent, mid-rib conspicu­ ously dark purple or greenish, margins near apex purple, apex apiculate, not appendaged; outermost c 1-1.2 x 0.8 mm; innermost c 3.5 x 0.5 mm. Florets deep purple or ma­ roon. Corolla cylindric. with sessile glands. Anther base sagittate, obtuse. Cypselas light-brown, c 10-ribbed. with ascending setae and numerous light brown sessile glands in the grooves between the ribs, c 2.5 x 0.8 mm. Pappus 2-seriate, white, barbellate: outer c 0.5 mm long, inner 3-4 mm long. Fig. 145.55. Acacia-Commiphora bushland on dry rocky ground, dry semi-desert grassland; 500-1450 m. EW EE TU AF WTI SU GG SD BA HA; Sudan, Somalia, E Africa, Sene­

gal. Arabia and eastwards to NW India. Mesfin T. & Voilesen 4292. 4157; Mesfin 7'. & O. Hedberg 1064. A recent record, Schloeder & Jacobs 441. from the Oino National Park, 500 m. KF. is probably also this spe­ cies. 21. V. phillip.viae S. Moore (1900) - types: Somalia, Upper Sheik, 1897. L. Phillips s.n. (BM syn., K isosyn ); Wagga Mountains 1897. Lort Phillips s.n. (BM syn ). Low shrub or shrublet, 0.1—0.5(— 0.75) m high. Stems and branches dark purple below, grey-green in upper parts, sparsely pubescent, hairs both short-stalked, T-shaped and simple. Leaves alternate, ovate to ovate-lanceolate, (4-)6-15(-20) x 3-4(-7) mm, densely resinous punctate, narrowed to base and apex, sparsely pubescent, hairs mostly simple, margins entire to irregularly 1-2-lobed; apex acute-apiculate or obtuse; axillary shoots suppressed and hence leaves look fascicled. Capitula turbinatecampanulate, c 10-12-flowered, 7-10 x 3-5 mm at anthesis, in open corymbose cymes terminating branches and main stem; peduncle up to 3 cm long. Receptacle epaleate. Phyllaries 4-5-seriate, continuous with bracts on peduncle, light purple in upper parts, yellowish green in lower parts, thinly pubescent in upper half or on mar­ gins, apex of outer and middle phyllaries sharp and pointed (point up to 1 mm long), not appendaged; outer ovate, c 2-2.5 x 1 mm. inner oblong-elliptic, c 6.5-7.5 x 1.5, obtuse or apiculate. Florets purple. Corolla 8.5-9 ram long, narrowly funnel-shaped, glabrous or sparsely sessile-glandular, 5-lobed, lobes c 3 mm long. Anthers 3 .5-4 mm long, base sagittate, c 0.5 mm long. Sty le bifurcate. 10.5-11 mm long, papillose-hairy near fork, branches c 2 mm long. Cypselas light brown, thinly villous, ses­ sile-glandular, c 3 x 0.5 mm. 4-angled, columnar to slightly oblanceolate. Pappus 2-seriate, light brown; outer fimbnate, 0.3-0 5 nun long: inner c 5-6 nun long, baibellate Open Combretum Terminalia-Acacia woodland or bushland on rocky outcrops, 1450 m. SD, Somalia. Friis et al. 3279; Gilbert et al. 7775; Mesfin T. 4470. 22. V. smithiana Less. (1831); Webbia smithiana (Less.) DC. (1836) - type: Rep. Dem. du Congo. Congo R., Smith s.n. (BM K P iso ). I'ernonia chiesii Chiov.. nom. nud., quoad Negri 1069. Erect herb up to 0.7 m high, woody at base. Stem solitary or several from woody base, each often 3-5 mm wide at base, simple or slightly branched above. Leaves alternate, ascending, narrowly elliptic, oblong-lanceolate or ovateelliptic. subsessile. narrowed at base, obtuse or subacute at apex, glabrcscent and medium green above, silverygreen and sericeous to densely white tomentose beneath, up to 6 x 2 cm. Capitula hemispheric, up to 5 x 6 mm, in small but dense terminal panicles; peduncle up to 10 mm long, densely white tomentose. Phyllaries 3-4-seriate, narrowly elliptic-oblong, mucronulate, with dark purple

145. ASTERACEAE: 30. Vernonia

89

tip dcnsel> white tomentose. not appendaged. 3-5 x O .i-1 mm. Florets pale violet. Corolla campanulatec> indnc. 6.5-8 mm long, thinly white pubescent. 5-kibed. lobes c 2.8 mm long. Anthers c 2.5 mm long, ba c sagittate, obtuse. 0.3-0.4 min long. Style c 9.5 mm loi g densely bristly near fork, branches c 2 mm long. C\ pselas stngose. 2-3 mm long, not ribbed, light brown. Pa:>pus 2-scriate: outer 1—1.5 mm long; inner c 4 mm loi t white to slightly yellowish, barbellate. Wooded grassland on dry rocky hillside, growing after gn ss fire; 1350-1830 m. SU GG SD ?BA; Sudan. Rep. D< m. du Congo. Rwanda. Burundi. Cameroun, Gabon. l/< nev 5640; Snieds 680; Gilbert & Jefford 4346. 23 V. thulinii Mesfin (1997) - tvpe: WG, E slopes of Didessa R. valley, Gilbert & Thulin 733 (ETH holo., K UPS iso.). Shmb up to 3 m high. Stem slender, terete, glabrous exce H in the inflorescence branches; liairs simple. Leaves all mate, lanceolate, up to 12 x 2.6 cm. light green and sc. brid above, densely white tomentose except on midve n beneath, veins prominent below, base cuneate; margi is entire to denticulate-recurved; apex acute-acuminate. C; oitula campanulate, 12-13 x 8-10 mm at anthesis. in ax llarv and terminal scorpioid cymes; peduncle up to 5 m n long, densely white tomentose (including infloresc nee branches). Receptacle ?epaleate. Phyllaries 5-6se late, densely arachnoid-tomentose, linear to oblongla ceolate, purple in upper half with dark purple mid-rib, ap 2.x acute, not appendaged; outer c 2-2.5 x 1 mm, inner 7. -8 x 1.5-2 mm. Florets purple. Corolla glabrous or sp irsely sessile-glandular in the middle, 8.5-9.5 mm long, 5-lobed, lobes c 2.2-2.5 mm long. Anthers c 3 mm lo ig. base sagittate, acute c 0.5 mm long. Style c 11 mm lo tg. setaceous near fork, branches c 2.5 mm long, se .aceous throughout. Cypselas (only immature seen) lij rit brown, 6-ribbed. sessile-glandular and setose espec l l l v in grooves, c 2 x 0.8 mm. Pappus 2-seriate, brownis -white; outer series c 0.5 mm long, ?squamulous; inner 4 - 5 mm long, barbellate. Wooded grassland (Didessa River valley) on slopes: A i acia-Lannaea bushland; 1000-1200 m, GJ WG; not ki own elsewhere. Gilbert & Thulin 733; Friis et al. T :*4. 2‘ V. wollastonii S. Moore (1908) - type: Uganda, Mt Ruwenzori, 26 Jan. 1906. Wollaston s.n (BM holo.). V. heterocarpa Chiov. (1929) - type: SU/AR, in the Lake Ziway [Zuai] basin. Basile in Duca di Abruzzi 36 (TO holo ). Pi ^nnial herb or shrub with slender scandent branches gi :*wing through and over shrub clumps, up to 2.5 in high. S i:m striate, green, thinly pubescent. Leaves alternate, oftc with small, leafy, axillary brandies, ellipticla ceolate, ovate-lanceolate. 5-9 x 1.5-2.5 cm, membi inous, light-green, thinly pubescent on veins, hairs lo i^-stalked, symmetrically T-shaped; base cuneate and

Figure 55 VERNONIA CINERsiSCENS 1 - part o f upper stem x W, 2 - outer phyllary x 512, 3 cypsela with part o f pappus x V/2 . (No specimens cited in orig.) Drawn by Bhola Ram. (Reproduced with permission from FI. India . vol. 13. fig. 115.)

narrowly decurrcnt on petiole, rarely cordate (in forms from drier sites); petiole up to 1.5 cm long, slender; mar­ gins widely denticulate-serrate. Capitula campanulate. 6-8 x 4-5 mm. c 20-flowered, in loose terminal or axillary paniculate cymes; peduncle slender, up to 10 mm long, appressed-pubescent. Receptacle epaleate. Phyllaries 4-5-seriate. graded monomorphic, lengthening inwards, densely pubescent with interspersed sessile glands, linear to linear-lanceolate, light green with darker green or dark purple mid-rib in upper parts; apex acute, purple, not appendaged; outer c 1.5 x 0.3 mm; inner c 4 x 1 mm. slightly navicular (boat-sliaped) in lower half. Florets pur­ ple. Corolla funnel-shaped, 6.5-7 mm long. 5-lobed. lobes c 2.5 mm long, white-setose. Anthers c 2.5 mm long, base sagittate, obtuse, c 0.5 nun long. Style c 7 nun long, papillose near fork, branches c 2.5 nun long, papillose. Cypselas greyish-brown, glandular-setose, 4-5-anglcd. ribs inconspicuous, c 2 x 0.5 mm. Pappus 2-seriate. white, barbellate: outer series 0.5-1 nun, inner 4—4.5 mm. Montane rain forests and Podocarpus-Juniperus for­ est. often along forcst-tracks. sometimes in deep shade, forest margins, derived evergreen bushland: 1500-2435 m. SU WG KF GG SD BA; Sudan. E Africa. Mozam-

90

145 ASTERACEAE: 30. Vernonia

biquc. Malaw i. Zambia Zimbabwe. Mesfm T. et al. 8232; Mesfin T. 6974; Friis et aI. 3481. 25 V. hildebrandtii I aike (1875); Gongrothamnus hildebrandtii (Vatke) Oliv. & Hiem (1877); Cacalia hildebrandtii (Vatke) O. Kuntze (1898) - type Tanzania. Zanzibar, Hildebrandt 1020 (‘>WAVU holo.. K iso.). Slrnib. 1-1.5 m high. Stem terete, striate-sulcate, sparsely hairy, hairs simple. Leaves ovate or ovate-elliptic, petiolate or pseudo-petiolatc. base cuneate, rarely sagittate. 3-5.5 x 1-2.7 cm. upper surface sparsely pilose, lower surface with denser hair. margins crenate-serrate. petiole up to 1 cm long. Capitula cylindric at anthesis. 2-3 x 1.5-2.5 mm. up to 7 x 4 mm in fruit, in dense corymbose cymes, peduncle 2-5 mm long, densely pubescent. Outer­ most phyllaries ovate or ovate-oblong. 2-2.5 x 1-1.5 mm. glabrous except for a few liairs near the apices, densely sessile-glandular along the thickened mid-nb near the api­ ces. apex spinulosc. middle and inner series similar to outer except in si/e and pubescence: middle up to 7 x 3.5 nun; innermost up to 10 x 1.5 mm. all sessile-glandular near the apices. Florets white or pale purple. Corolla 6-7 nun long, tubular, sessile-glandular. 5-lobed. lobes c 1.5 mm long. Anthers 3.5^4 mm long, connective lanceolate. c 0.6 mm long Style bifurcate. 6.5-8 mm long, brandies c 1.5-2 111111 long Cypselas light brown. 4-angled and 4(-6)-ribbed. densely covered with dark brown sessile glands, sparsely upwardly pilose, 2-2.5 x 0.6-1 mm. oblong-oblanceolate. Pappus 2-seriate, outermost scries 0.5—1 mm long, innermost 5-7 mm long, white, barbellate. No habitat and altitudinal data from the FEE area [in Somalia: Acacia-Commiphora bushland on alluvial soil along rivers. 25-230 m|; SD (Neghelle); Somalia (where commmon). Corradi 1958a (FT). 26. V. cinerea (L.) Less. (1829); Convza cinerea L. (1753) - type: “Habitat in In­ dia.” Herb Hermann 3: 16. No. 419 jBM lecto.. se­ lected by Grierson in Dassanayake & Fosberg (ed ), Rev. Handb. FI. C'evlon 1: 133 (1980) and Jeffrey in Kew Bull. 43: 224 (1988)). var. cinerea Erect annual herb, up to 0.75 m liigh. Stem stnale. branched above, shortly pubescent Leaves alternate, ovate to ovate-elliptic, up to 7 x 4.6 cm. dark-green above, paler beneath, sparsely puberulous on both surfaces or subscabnd above, liairs both simple and T-shaped with unequal arms; base cuneate. attenuate at base into a short petiole, or blade extending into petiole, margins grossly serrate-dentate or subentire; apex acute or rounded. Capitula campanulate. 6-6.5 x 5-5.5 nun, c 20-flowered. in axillary and terminal many-headed cymes; peduncle slender, densely pilose, up to 2.3 cm long. Receptacle epaleate. Phy llanes 4-seriate, linear-lanceolate, mucronulate, mid-vein prominent densely pilose with minute

glands, not appendaged; outermost 1.2-2 mm. innermost

4-4 3 x 1 mm Florets pale purple. Corolla tubular, lobes pilose Anther base shortly sagittate. Cypselas light to dark-brown. 1.5-2 x 0.5 mm long, subterete to 4-5ribbed, unifomih setulose with short white hairs, glandu­ lar or glabrous Pappus 2-seriate, white, barbellate, caducous: outer series c 1.5 mm long, inner c 4-4 5 mm long. Disturbed sites in riverine and lowland forests; 150—J 700 m. EW EE IL ?BA; Somalia, Sudan, Kenya, Tanzania, Burundi. Rep. Dem. du Congo, Central African Republic, Gabon. Cameroun Friis et al. 2458; Gilbert & Friis 8373; Pappi 73. Var. lentil (O. HofTm.) C. Jeffrey occurs in Uganda and Kenya (cf. Jeffrey, in Kew Bull. 43: 224 (1988)]. J 27 V. hochstetteri Sch. Bip. ex Walp. (1843); Amhassa hochstetteri (Sch. Bip.) StecLz (1864) type: TU. Ml Kubbi. near Adwa PI abyss. 1. Schimper 255 (BM BR K P UPS iso ). var hochstetteri Vernonia hochstetteri Sch. Bip. ex Walp. var rubicunda Sch Bip. ex Walp., Repert. Bot. Syst. 2. 945 (1843) - type: TU. near Adwa PI. abyss. I. Schimper 413 (BM K iso ). I'. kostlinu Hochst. (1841), nom. nud., quoad Schimper 413. V. jugalis Oliv. & Hiem. (1877) - type: Sudan. Monbuttu, Schweinfurth 3434 (K holo.). • Shrub up to 2 5 m high. Stem terete, slightly stnate. shortly and closely purplish-brown tomentose: hairs longor short-stalked. T-shaped Leaves alternate, thin, ovatelanceolate or lanceolate, upper surface thinly scabnd. lower pubescent-tomentose with denser hair along veins, up to 20.5 x 9 cm; base cuneate with blade extending into the petiole; margins serrulate, apex acute-acuminate; peti­ ole up to 1.2 cm long, brownish pubescent-tomentose. sometime dilated and clasping at base. Capitula campanulate. 6-10 x 4-6 mm. in dense terminal corym­ bose cymes; peduncle up to 2 cm long, pubescenttomentose, most with several small bracts. Receptacle epaleate. Phyllaries 5-6-seriate. continuous with bracts on peduncle, purple with darker spots or lines near apex, sparel} to densely sessile-glandular and sparsely liairy. margins fimbriate, linear-lanceolate to lanceolate, apex acute-apiculate. not appendaged. outer c 2 x 0.5 mm; in­ ner c 7.5 x 2 mm. Florets purple. Corolla cylindric. densely sessile-glandular in lower parts, c 9 mm long, 5-lobed. lobes 2—2.5 mm long; anthers c 2.5 nun long, base sagittate, c 0 5 mm long. Style c 12.5 mm long, gla­ brous at fork, branches c 4.5 nun long, glabrous or with few papillae. Cypselas light-brown, c 10-ribbed. hispid, c 3 x 1 mm Pappus 2-seriate, brownish-white, outer series c 1 nun long, fimbriate or laciniate, inner c 7 mm long, barbellate. Fig. 145.56. Margins of Jutuperus-Podocarpus forests, often in cleared parts or along footpath in forests; roadside thick­ ets. 1200-2500 m. TU GD GJ SU WG IL KF GG SD BA

91

145. ASTERACEAE: 30. Vemonia

Figure 56.

VERNONIA II OCHS TETTE R ! var H OCHS TETTEIU : 1 part o f plant x 2/3. with detail o f leaf underside: 2 capitulum x 3: 3 outer phyllary \ 6: 4 inner phyllary x 6: 5 floret, cypsela and pappus removed x 4; 6 anther tube opened out x 10: 7 style x 10:8 imma­ ture cypsela with pappus x 6: 9 cypsela (inner pappus rem oved) x 6. All trom Davies 179. Drawn by Fat Halliday. (Re­ produced with permission from /*'/. Trop E. Afr. C om positae. fig 43.)

H/ Sudan. E Africa, Rep. Dem. du Congo. Burger 3175; Friis et al. 1719; Mesfin T. & Kagnew 2510. Var. kivuensis (Humb. & Starter) C. Jeffrey occurs in inda, Rwanda. Burundi and Rep. Dem. du Congo, wl ile var. dekindtii (O. Hoffm.) C. Jeffrey occurs in Cen­ tra1 Tropical Africa (Angola to Central African Republic an : Cameroum) [cf. C. Jeffrey , in Kew Bull. 43: 226 ( 1* * 8) 1.

28 V. friisii M esfin (2004) type; GG, 3 km N of Ghidole. 27 Dec. 1997, Friis, Bidgood, Melaku W. & Gashaw G-H 8833 (K holo., C ETH iso.). Pe ennial herb or shrub, about 0.7 m high. Stem round, so: Jy tomentose. hairs long-stalked, mostly T-shaped, ig it brown. Leaves simple, up to 15 x 6 cm, pcliolate, ba e cuneate, blade broadly elliptic, upper surface thinly ha i\'. hairs mostly T-shaped, some hairs simple, others

appearing T-shapcd but one arm missing, liairs denser on the lower surface with also sessile y ellow ish glands, mar­ gins inconspicuously serrate, petiole up to 4 cm long. Capitula narrowly hemispheric, in loose paniculate cymes. 15-17 x 8-10 nun at anthesis; peduncle up to 3 cm long, densely pilose. Phyllaries 5-6-seriate. outermost and middle series linear-lanceolatc. pilose and denselv sessile-glandular. 5-8 x 1—1.5 mm. greenish or dark grey ish-purplc on upper half and pale brown or whitish in lower parts, apex long-acuminate, refle.xcd; innermost phyllaries oblong or oblong-elliptic. 10-11 x 2.5-3 nun. whitish, thinly pilose with sessile glands on upper half near the apex. Florets purple. Corolla 8-9 mm long, pu­ bescent and sessile-glandular in middle part, lobes c 3.5 nun long, glabrous. Anthers c 2 mm long, conncctivc c 0.2 mm long. Style 9.5-10 mm long, branches peniccllate. 1.5-2 mm long. Cypselas (immature) light brown. 10-ribbed. c 3 x 0.5 mm. upwardly hispid in grooves. Pappus 2-scriatc. brownish-white: outer scales oblong.

92

145. ASTERACEAE: 30. Vernonia

0.5-0.6 mm long, hispid on margins; inner bristles 6-8 mm long, barbellate. Lowland woodland; fallow fields; 900-2200 m. GG; not known elsewhere. Friis et al. 8833; Fukui 17, 327, 241. Vernonia friisii is allied to V. hochstetteri and V muellerii subsp. integra C. Jeffrey of East tropical Africa. 29. V. tewoldei Mesfin (1997) - tvpe: KF, Bonga, Friis, G. Aweke, Rasmussen & Vollesen 2150 (ETH holo., K iso.). V holstii auct., non O. Hoflm. (1894): Cufod. (1967), quoad Meyer 7752 (K). Shrub or shrubby liana, 1.5-3(-4) m high; branches slen­ der, striate, light to dark brown, sparsely to densely appressed tomentose; hairs brown, mostly simple with few scattered T-shaped ones on peduncles. Leaves alter­ nate, elliptic. (8—>J5—21 x (3.2-)4.7-6.5 cm; upper sur­ face sparsely pubescent lower surface pubescent to sub-tomentose with denser hairs along veins; margins ser­ rate with sharp apiculate teeth; base cuneate: apex acute-apiculate; petiole up to 1 cm long, tomentose. Capitula turbinate-campanulate, 22-24 x 5-6 mm at anthesis. in axillaiy or terminal few-headed cymes; c 22-25-flowered; peduncle densely brown tomentose, up to 2 cm long. Receptacle epaleate. Phyllaries (4—)5(—6)seriate, purple in upper half, membranous, continuous with bracts on peduncle, apex not appendaged; outer 2-3 x 1-1.5 mm, ovate-lanceolate, pubescent (including mar­ gins) with loose, long hairs and sessile-glands; middle thinly pubescent near apex, with scattered sessile glands; inner oblong-lanceolate, c 11-12 x 1 mm, glabrous or glabrescent and with few scattered sessile glands. Florets pale purple-violet, or pinkish-violet. Corolla narrowly funnel-shaped, c 10-11 mm long. 5-lobed, lobes 3.5-4 mm long, sparsely sessile glandular. Anthers c 4 mm long, base sagittate, acute, c 2.5-3 mm long. Style bifurcate, 13-15 mm long, setose near fork, branches 2.5-3 mm long, setose throughout. Cypselas (only immature seen) brown, c 10-ribbed. c 4-4.5 x 1 mm, densely sessileglandular (white), sparsely hispid, oblong-cylindric. Pappus 2-seriate, white: outer 1—1.5 mm long, ciliatefimbnate at margins; inner 7-8 mm long, barbellate. Glades in evergreen montane forest along stream in forest; 1600-2150 m. KF BA; not known elsewhere. Friis et al. 2150; Mooney 8457; Gilbert & Rankin 4198. 30. V. wakefieldii Oliv. (1887) - type: Kenya, Nyika, Wakefield 1 (K syn ); Tan­ zania, Moshi, Johnston s.n. (K syn.). Shrub up to 2 m high. Stem scrambling, striate, glabrous, succulent, arising from inflated tuberous base, branched from ground level. Leaves alternate, oblong-elliptic, lan­ ceolate to oblanceolate, (4-)6-8 x 1.9-2.8 cm, glabrous, slightly succulent, narrowed at base and sessile; margins entire; apex obtuse or acute. Capitula heinisphericcampanulate, 2.4-2.7 x 1—1.5 cm at anthesis, solitary or in few-headed cymes terminating branches or main stem;

peduncle glabrous to thinly floccose, bracteate. up to 2.5 cm long. Receptacle epaleate. Phyllaries 5-6-seriate. con­ tinuous with bracts on peduncle, coriaceous, glabrous or with scattered white hairs in outer ones, margins densely white ciliate; apex puiple or purple-bordered, not appendaged. outer oblong-ovate, 4-5 x 2.5-3 mm; inner 12-13 x 2-2.5 mm. Florets pale purple with dark blue style and stigma. Corolla glabrous, tubular, 5-lobed. 22-23 mm long, lobes c 9 mm long. Anthers c 5.5 mm long, base sagittate, obtuse, c I nun long. Style c 23 mm long, setaceous near base of fork, branches 7.5-8 mm long, externally hispid-papillose throughout. Cypselas brown (only immature seen), c 5.5 x 1.2 mm. hispid, c 10-ribbed, densely sessile-glandular, oblong-cylindric. Pappus multi-seriate, light brown, barbellate. outermost c 1.5 mm long; inner c 12-13 mm long. Acacia-Commiphora bushland on grey sandy soil. 1100-1200 m SD; Kenya (where common), Tanzania. Mesfin T. & Vollesen 4159; Friis el al. 8705. 31. V. purpurea .Sc/7. Bip. ex Walp. (1843) type. GD. near Gafat. PI. abyss. II. Schimper 1197 (P holo., K photo). V. schimperi Sch. Bip. (1842), nom. illegit. non DC. (1838); V inulifolia Steud. ex Walp. (1843) type: TU, Mt. Sholoda, PI. abyss. 1, Schimper 221 (BM K P UPS iso ). V jacaeoides A. Rich. (1848) - type. TU. Chire, Quartin-Dillon et Petit s.n. (P holo.. K photo). V. negrii Chiov., nom. nud.. quaod Negri 1057 (FT). Erect perennial herb, up to 1.5 m high. Stem rigid, simple or branched above, terete, striate, glabrescent to thinly setulose-scabnd. leafy to the apex. Leaves alternate, ellip­ tic to oblong-elliptic, erect-patent or ascending, scabrous, obscurely pellucid-punctate, narrowed at base; margins serrulate; apex acute-apiculate; middle ones 6-11 x 1. 5—4 cm; subsessile to shortly petiolate. Capitula hemispheric, 10-16 x 8-16 mm. solitary' or in few-headed terminal cymes, surrounded at or near the base with leaf-like bracts, equalling or usually much over-topping the involu­ cre, epedunculate or peduncle only up to 2 mm long. Re­ ceptacle epaleate Phyllaries 5-6-seriate, ovate to oblong, narrowed to a rigid, acute apiculus. arachnoid-pubescent or tomentose and matted, especially when young, apex purplish, not appendaged; outermost 2-3 x 0.5-1 mm; in­ nermost 8-13 x 1.5-3 mm. Florets red-violet, bluishpurple to blue. Corolla narrowly campanulate, sessileglandular, lobes setaceous at apex. Cypselas oblong, grey-brown, strigose, 10-13-ribbed. c 3 x 1 mm. Pappus multiseriate. baibellate. light-brownish, persistent stiff, outer irregular. 1.5-2 mm long, inner c 4.5 mm long. Combretum Terminalia woodland, grassland, open bushland. rarely waste ground, 950-2600 in. EW TU GD GJ SU WG IL KF GG SD BA; Kenya, Uganda, Tanzania, Sudan. Malawi. Mooney 8698; Friis et al. 2308; Mesfin T. & Kagnew 2188 A species tliat can easily be distinguished by the stiff and scabrid leaves with a single capitulum subtended by

145. ASTERACEAE: 30. Vemonia

93

leaves, or numerous and compact terminal capitula that are clear of the foliage. When growing in moist situations, as n a riverine forest, the capitula are held loosely and the inflorescence approaches a scorpiod cyme, e.g.. Friis et al. 2308. 7904.

Degraded Acacia-Commiphora woodland or bush­ land on rocky, granitic slopes with thin soil; 1650 m; SD GG. Sudan. Chad. Cameroun, Nigeria, Benin. Ghana. Ivory Coast. Sierra Leone, Niger, Bourkina Fasso, Guinea Bissau. Mali. Gambia. Senegal. Friis et al. 8345, 8840.

32. V. galamensis (Cass.) Less. (1829); Centrapalusgcilamensis CasS. (1817)- type: Sen­ egal. Herb. Galam 80 in Jussieu (P holo.. K photo). I Conyza pauciflora Willd. (1803); Vernoma oauciflora (Willd.) Less. (1829), nom. illegit.. non Poir. (1817).

var. petitiana (A. Rich.)M. G. Gilbert in Kew. Bull. 41(1). 25 (1986); Vernonia petitiana A. Rich. (1848) - type: Ethio­ pia 1?WU/SU). Yedjou. Petit s.n (P holo ).

sutsp. galamensis

Dwarf Commiphora Lannaea woodland. Acacia Commiphora scrub. Acacia drepanolobium woodland on black cotton soil: 700-1700 m. EW TU SU SD; Sudan. Somalia, E Africa. Gilbert & Sebsebe D. 8666; Schimper 1724; Mesfin & I ollesen 4309.

Annual herb, c 0.75-1.5 m high (up to 3 m high in cultiva­ tion) Stem striate or narrowly ribbed, adpresscdpuberulentto coarsely spreading-pilose. Leaves alternate, sessile, up to 25 x 5 cm. elliptic to lanceolate; base nanrowly cuneate. apex acuminate; margins irregularly demate, senate to serrulate, upper surface aspcrous to spatsely pilose, lower surface sparsely to densely pilose, liairs both simple and short-stalked T-shapcd. Capitula hemispheric. 8-25 x 8-12 mm. in open terminal leafy cymes or corymbose cymes; peduncle pubcsccnt. up to 10 mm long. Reccptaclc epaleate. Phyllaries 4-6-scnate. verv sparsely pubescent to woolly, apex appendaged. outer c 8-12 nun long, linear, often with well defined mi Jrib; inner 10-22 mm long, oblong to narrow ly lanceo­ late with scarious margins and acuminate tip. Florets bn*ht blue to almost white, sometimes flushed pale yel­ low- or green, narrowly funnel-shaped. Corolla 13-14 mm long, sessile glandular. 5-lobed, lobes c 5 mm long. Aniheis c 4 mm long, base sagittate, c 0.6 mm long, obtuse. St} te c 16 mm long, setose near fork, branches c 4.5 mm loig Cypselas dark brown to black. 5-8 x 0.8-1.2 mm. 10-nbbcd, strigosc. Pappus barbellate. white to light brown; outer up to 1.5 mm long, inner up to 8 mm long. M.G.Gilbert [inKew Bull. 41: 22 (1986)1 rccogni/.cd 5 subspecies within this spccics; only tlic type subspecics occurs in the Flora area I galamensis subsp. galamensis. w hich occurs in West Afnca. in Sudan and the Flora area, ana through East Africa south to Zimbabwe, lias again been subdiv ided into 4 varieties. Specimens referable to 2 of iliese varieties, var. petitiana and var. ethiopica. have been seen with ccrtainty from the Flora area; material which approaches tlie third variety, var. gal amen sis. lias also been seen. 1. -

Involucre 17-22 mm long. Involucre 8-16 mm long.

2.

Leaves often over 11.5 111111 wide; involucre (1 1-) 13-17 mm long. var. galamensis Leaves usually under 11.5 111111 wide; involucre 8—12(—16) mm long. var. pctitiana

-

var. ethiopica 2

va galamensis Leaves often over 11.5 111111 wide: involucre (11—) 13—16 (-17) mm long; florets often bright blue to purple.

Leaves often under 11.5 mm wide; involucre 8—12(—16) mm long; phyllary tips (0.3-)0.4-0.5(-0.6) mm wide; flo­ rets pale blue to mauve.

var. ethiopicaM. G. Gilbert in Kew Bull. 41(1): 26 (1986) -type: HA. rock-vallcy in Harar-Jijiga road. 1700 m.M.G. endaged: outerc 4 x 0.8 mm; innerc 17 x 1.5 mm. Florei • red to deep blue-violet. Corolla campanulate. sparsely ha r \ . Anthers shortly sagittate. Style pubesent in upper pa ts. Cypselas grey-brown. 10-ribbed. 9-10 x 2-2.3 mm. de isely and sub-appressedly hirsute. Pappus inanysei rate, light brownish or greyish, barbellate; outer c 1.5 mm long; inner c 12-13 mm long. Grassland with scattered bushes; 1600-2300 m. EW T l GD GJ SU WG KF; not known elsewhere. Friis et al. 43n W. de Wilde 8786; Mogk 413. Two other subspecies (subsp. congolensis and subsp. lot tvflora C. Jeffrey) occur in central and tropical East Africa. 36 V. schimperi DC. (1836) - type: Yemen, Cara, Schimper 868 (K iso.). V. abvssinica Sch. Bip. ex Walp. (1843); Stengelia controversa Sch. Bip. ex Walp., nom. non rite publ. ; 1843) - tvpe: TU, near Adoa, PI. abyss. I. Schimper 389 (BM K UPS iso.). V. polymorpha Vatke var. microcephala Vatke in ' innaea 39: 477 (1875) - type: TU, Anadehr [near AduaJ, 20 Oct. 1862. Schimper 581 (BM iso., K photo).

95

V. abyssinica Sch. Bip. ex Walp. var. vestita Cufod. in N. Giorn. Bot. Ital., n.s. 50: 102 (1943) types: SD. Neghelle, Corradi 1996, 1976 and 1982 (FT syn., K photo). Teichostemma fruticosum R. Br. in Salt (1810). nom.nud.. quoad Salt 1 (BM); Vernoniafruticosa Sch. Bip. in shedae. nom. nud. quoad Salt 1 (BM). Perennial herb or shmb. 1.5-2 m high. Stem branched, terete, often light brown, conspicuously striate, softly puberulous. Leaves alternate, thin, narrowly elliptic to lanceolate, glabrescent to tomentose. middle ones 30-50 x 8-10 mm; base cuneate: margins serrate-dentate: apex acute; petiole up to 5 mm long with decurrent blade. Capitula campanulatc. 15-17 x 10-12 nun. in terminal cory mbose panicles, peduncle up to 3 cm long, pilose, with 1 or 2 bracts at middle or near apex. Phyllaries 4-5seriate. apex appendaged; outermost oblong, herba­ ceous-tipped. tip green and spreading, c 3 x 0.5 mm. sparsely hairy at base: middle ones oblong to ovateoblong. narrowed rather abmptly into a lanceolate ap­ pendage. hairy with sessile glands at constriction and lower parts of appendage: innermost ones lanceolate, greenish-yellow, glandular near apex, c 9-10 x 1.5 mm. Florets purple or violet. Corolla campanulate. sessileglandular in middle part. Anther bases acute. Cypselas ob­ long-cylindric. c4 x 1 mm. hirsute, with basally glandular liairs. c 20-ribbcd. Pappus white. 2-seriate. barbellate: outer c 1 nun long; inner c 8-9 mm long. Combretum Terminalia Croton woodland, bushland with Acacia. Rhus and Fuclea. open grassland: 1050-2400 m. EW TU/GD TU WU SU SD BA HA: Kenya (Kl). Djibouti. Egypt. Yemen, Saudi Arabia. Smith & Andemeskal 4532; Gilbert et al. 7408; Mesfin 7 et al. 8205. In notes on herbarium sheets (Smith & Andemeskal 4531. 4532 and 4536) the oil from this species is stated to be 19.4%. 21.5% and 15.8% while the epoxy acid is 67.8%. 67.1% and 67.6%. respectively. 37. V. buchingeri (Sch. Bip. ex Steetz in Peters) Oliv. & Hiem (1877); Ascaricida buchingeri Sch. Bip. ex Steet/ in Peters (1864): - type: Ethiopia (exact region not known|. Schimper 386 (BM K iso.). Vernonia argut idens Cliiov. (1911) - type: GD. Uoghera (=Wogera). C’hiovenda 1006/1000 (FT holo.. K photo). Erect, perennial herb, up to 0.4 in high, with a tluck rootstock. Stem leafy throughout, striate-sulcate. thinly pubescent. Leaves alternate, oblong-lanceolate, at closely spaced intervals, thickish. or slightly coriaceous, sessile or subsessile. 6-7.5-1.5-2 cm; margins denticulateserrate; surfaces with thinly scattered minute crispid hairs, densely resinous-dotted: apex acute. Capitula hemispheri­ cal. few. in terminal cymes, 30-38 x 35-40 mm; peduncle short, up to 6 nun long, sulcate. pubescent. Phyllaries 5-6-seriate, coriaceous, thinly pubescent on outer surface, apex appendaged; outermost lanceolate, purplish. 3-5 x 3.5-4 mm, appendage green. 8-10 x 4-5 mm; middle

96

145 ASTERACEAE 30. Vernonia

8-13 x 5-6 min. ovate-lanceolate: innermost lanceolate, c 19 x 5 mm. Florets ?purple. Corolla c 25 mm long, 5-lobed. lobes 3.5-4 min long, funnel-shaped with long (c 17-18 nun) tube, slender, sparsely sessile-glandular al middle. Anthers c 7 mm long, base sagittate, acute, c 1 nun long. Style c 27 nun long, setose near fork, branches c 3 mm long. Cypselas (only immature seen) c 6.5 mm long, densely silky pubescent. Pappus c 4-seriate. flat, barbellate. inner ones slightly expanded at apex, reddishbrown (whendry); outerc 2 nun long; innerc 15 mm long. Known only from the types cited above: GD: Schimper 386, Chiovenda 1006. 38. V. didessana Mesfin ( 1997) - type: GJ, 74 km on Injibara-Guba road, 16 km W of Chagni, Thulin & Hunde 3994 (K holo.. ETH UPS ISO).

Erect, perennial herb up to 0.45 m high, with a woody rootstock. Stem striate-sulcate. densely brown-pubescent to sub-tomentose. unbranched except in the inflorescen­ ces. liairs simple. Leaves alternate, deciduous below, scabnd-pilose above, liispidulous with denser hair along veins, densely sessile-glandular, oblanceolate, obovateblong to elliptic. 8-17.5 x 2.5-6 cm. narrowed to base, sessile or subsessile: margins denticulate: apex subobtuse. apiculate. Capitula campanulate-hemispherical. 20 x 10 mm at anthesis, solitary and terminal or in few-headed cymes: peduncle up to 5.5 cm long, pilose. Receptacle epaleate. Phyllaries 4(-5)-seriate. apex appendaged: outer herbaceous. 6-12 x 1-2 mm. densely pilose, linearoblong; middle 12-13 mm long, appendage 8-9 mm long, densely pilose and sessile-glandular, purplish; inner 12-13 x 3—4 mm. appendage c 1.5-2 nmi long, glabrous, purplish. Florets pink, violet or pale lilac. Corolla sparsely sessile-glandular at middle, c 14-15 mm long. 5-lobed. lobes c 2.5 111111 long, sessile-glandular at apex, tubular and abruptly narrowed to a slender lower part. Anthers 4-4.3 111111 long, base sagittate, c 0.8 nun long. Style c 18 nun long, liispidulous near fork, branches c 3 mm long, liispidulous throughout. Cypselas (only immature seen) black, liispidulous. Pappus several-seriate, squamulous/ flat with ciliolate-barbellate margins, spirally twisted, shinv. 8-9 mm long, light yellowish-brown. Tall grassland with scattered trees, including Piliostigma, C.omhretum, Schefflera. Entada. on sandy soil: 1220-1700 111. GJ WG, not known elsewhere. Thulin & Hunde 3994, Ash 3080. W. de Wilde 10742. 39. V. filigera Oliv & Hiern (1877) - type. GD. Gaffat. 1863, Schimper 1530 (K BM). V hvmenolepis auct.. non A. Rich. (1848); Vatke (1875). Shrub or rarely perennial herb. l-2.5(-4) m high, with a tluck rootstock. Stem branches terete, striate, leafy' tliroughout. minutely and closely ashy-tomentose. Leaves alternate, ovate-oblong, elliptic or lanceolate, 12—15(—25) x 3-5(-7) cm, grey-green above with hoary inid-vein. pale grey-whitish with dense sessile resinous glands or

silvery tomentose beneath, petiolate; base cuneate, nar­ rowed into the petiole; margins unequally acutely dentate to serrate denticulate; apex acute; petiole up to 2 cm long, densely and closely tomentose, slightly dilated at base. Capitula hemispherical, (10-) 12-20 x 10-15 mm at anthesis, in corymbose terminal panicles; peduncle (1-) 2—5(—8) cm long, densely and closely tomentose. Recep­ tacle epaleate. Phyllaries (3-)4-6-seriate, apex append­ aged; outer greea linear-subulate, 5-15 x 0.5-1.5 mm, of­ ten densely tomentose. appendage (2—)5—12 x 0.5-1.5 mm, often pilose at base and also half-way up; middle and inner ones (3-)5-18.5 x 1.5-2 mm, sparsely pilose or cili­ ate at m a tin s and base of appendage, appendage (2-) 3.5-8.5 x 1.5—2 mm, white, sometimes pale-pink or pur­ ple-tinged. sometimes recurved; inner oblong, c 4-8 x 2 mm. appendage 1-3 mm long, sometimes sessileglandular, erect or recurved. Florets white, pale pink, li­ lac. Corolla cyhndnc above, narrowly tubular below, gla­ brous or with sessile glands at middle, c 11—13.5 mm long. 5-lobed, lobes 1.5-2 mm long, densely glandular at apex. Anthers c 3.8-4 mm long, base sagittate, acute. c0.5 mm long Stylej c 18 mm long, papillose near fork, branches c 4 mm long, papillose. Cypselas black or brownish-black, c 10-ribbed, densely sessile-glandular in the intercostal grooves, not hairy, 3^4 x 0.8—1 mm Pappus several-seriate, brownish-white, slightly expanded near apex; outer c 2 mm long, inner c 8-9 mm long. Wooded grassland, scrub with Protea, Maesa and Syzvgium, seepage area and rocky slopes near lakes and streams: 1935-2850 m. TU/GD GD SU KF SD; not known elsewhere FriiseI al. 1435 & 3826; Mooney 5321. According to a herbarium sheet (Smith, Desta & Sesai 4564 (K) the fraction of oil obtained in this species is 5.4% and the epoxy acid 56.4%. Tlus species is similar in habit and general capituiar features to V. hvmenolepis A. Rich, but differs from it in shape of phyllary appendages, only c 10-ribbed cypselas, setaceous pappus and in the inflorecesences (strictly corymbose in V filigera), generally smaller capitula. and in its ecology (seems to be associated with slopes near lakes and streams) It is particularly common at lakes Arakit and Awassa. and at the Gefersa reservoir. 40. V. hvmenolepis A Rich. (1848) type. Ethiopia (exact region not knownj, Yedjou and [probably SU] lC hoa\ Dillon & Petit (P syn ); ibid.. Petit (K isosyn.). V. rothii Oliv. & Hiem (1877) - type; SU, Ankober, Roth 346 (K holo.). V. leucophylla auct., non O. Hoffm. (1901); Cufod.( 1967), quoad Meyer 8902 (FT). Shrub up to 4.5 m high, several plants growing together and often forming a dense thicket; branch stems clothed with pale compact tomentum and often with blackish lenticels, liairs simple. Leaves alternate, ovate-lanceolate, (10—) 13—30 x (3-)4-8 cm, petiolate; base broadly cune­ ate, narrowly decurrent on the petiole; margins serratedenticulate; apex acute; upper surface sparsely pubescent or midrib hoary, often sparsely or densely resinous-

145. ASTERACEAE: 30. Vemonia

d:>tted; lower surface thinly to densely tomentose, some­ times densely resinous-dotted; petiole densely tomentose, (!: — )3—6 cm long. Capitula hemispheric, 1.5-2.5 x 1.2-2 c n. in dense terminal c o r y m b o s e cymes or elongated cynose clusters; peduncle densely tomentose, up to 2.5 cm 1 *ng. Receptacle epaleate, convex. Phyllaries 8-9-seriate. c maceous. spreading or reflexed, apex appendaged; outem ost longest, often on expanded portion of receptacle. I 4-24 x c 2 mm. sparsely to densely hispid outside, ap­ pendage 9-20 x 1.5-2. linear to lanceolate, sess le-glandular or pubescent at base; middle series ( -)14—20 x (1.5—)2—4.5 mm, glabrous, ciliolate at mar­ gins to sparsely pubescent, appendage (5—)8—16 x 2-4.5 n im, usually white or pink-tinged, ovate-lanceolate, multinerved, submembranous, often reflexed, apex acute. Flo­ a ts pale purple, pink, or lilac. Corolla cylindric above, a inaptly narrowed into a slender tube, sessile glandular in D iddle, 19-20 mm long, 5-lobed, lobes c 3 mm long. Ani ers c 5 mm long, base sagittate, obtuse, c 1 mm long. 5 ty le c 23 inm long, papillose near fork, branches 3-4 mm 1ccose; outer ones 1.5-2.7 x 0.7 -1.8 mm: inner ones 6 5-7 x 2-2.5 mm. Florets bright yellow , 4-6. Corolla tub ilar-campanulate, 5-6 mm long, glabrous. 5-lobed. lobes 1.5-1.8 mm long, sessile-glandular on outside. An­ ti ers light brown. 1.5-1.8 mm long, base sagittate, c 0.5 n m long, connective 0.5 mm long. Style bifurcate. 7-8 n m long, branches 1.6-1.8 mm long, papillose on outs Je. Cypselas light brown, strigose, c 2 x 0.7 mm, o ilanceolate, subquadrangular in upper half, narrowed at b ise. Pappus of white or brownish narrow scales, c 4 mm long, uni- or sub-biseriate, 30-40, some slightly wider than others, smooth to inconspicuously barbellate.

Capitula homogamous, many-flowered; involucre obconic campanulate to hemispheric: few to many-seriate, imbricate. Receptacle epaleate. Florets bisexual, purple, pink, rarely white. Corolla tubular or campanulate. 5-lobed. Anthers sagittate at base, obtuse, connective lan­ ceolate. Cypselas oblong or obovoid, (3—)5—10-ribbed or angled, ribs equal or alternately narrower, glabrous, ses­ sile-glandular or pubescent. Pappus absent or of caducous bristles.

Deciduous bushland on hilly ground, river banks with d -use mixed evergreen and deciduous vegetation; c 1700 n GG SD. Kenya. Tanzania. Rep. Dem. du Congo. Mo­ zambique, Malawi, Zambia. Zimbabwe. Botswana. Ang ila, Namibia. South Africa (Transvaal). Mesfin T. et al. 34X9; M.G &.S.B. Gilbert 1532.

1.

32. GUTENBERGIA Sch. Bip. (1840) Triplotaxis sensu Cufod. (1967), non Hutch. (1914). A unual or perennial, branched herbs. Indumentum of s it rt-stalked, long-armed, symmetrically T-shaped hairs o stems and leaves. Leaves alternate or the lower ones s b-opposite, simple, linear to ovate, entire. Inflorescence o open, dense or sparse paniculate or coiymbose cymes.

A genus with about 20 species confined to tropical Af­ rica. 3 species in the Flora area.

-

2.

-

Cypselas (7-) 10-ribbed, with more prominent ribs alternating with less prominent ones, or not obvi­ ously angled and almost or quite terete, c 2 mm long, yellowish or light brownish; pappus usually absent 1. G. rueppellii Cypselas (3-)5(-6)-ribbed or angled, flat between the ribs or angles, c 1-1.5 nun long, dark brown; pappus present or absent. 2 Leaves ovate-lanceolate or ovate, largest 15-30 (-40) mm wide; involucre 5-7 mm long; phyllaries with deeply laciniate-pectinate, mem­ branous margins. 2. G. boranensis Leaves linear-lanceolate or narrowly elliptic to lan­ ceolate. largest up to 15 nun wide; involucre 3-4 mm long; phyllaries membranous but not laciniate-pectinate. 3. G. somalensis

104

145. ASTERACEAE 32 Gutenbergia

1. G. rueppellii Sch. Bip. (1840); Ethulia rueppellii (Sch. Bip.) A. Rich. (1848) type: GD/TU between Temben and Simien. 6 July 1832. Riippell s.n. (P holo., K photo). Gutenbergia abyssinica Sch. Bip. ex Walp. (1846), nom. nud. var. rueppellii Erect, or rarely decumbent or spreading, annual, or short­ lived perennial, herb Stem branched in upper parts, thinly appressed tomentose. Leaves alternate or the lower oppo­ site or subopposite. 2-10 x 0.5-1 5(-2.5) cm. linearlanceolate to linear-oblanceolate. narrowed at base: mar­ gins entire or obscurely denticulate: apex acute or broadly pointed: upper surfacc green, sparsely pilose with asym­ metrically T-shaped liairs. lower appressed tomentose Capitula 4-5 x 4-5 cm. in open corymbosc cymes: peduncle up to 2 cm long. Phyllaries 3-4-seriatc. imbri­ cate. ovate-lanceolate. 2-4 x 1-1.5 mm: outer acute, densely appressed-pubesccnt. inner usually purplish above, scarious at margins. Florets purple. Corolla 3-4 111111 long. 5-lobed. glandular and pilose all over Cypselas yellowish to light brown. (7-) 10-ribbed with more promi­ nent ribs alternating with narrower ones, or not obviously angled and almost terete, c 2 x 1 nun. Pappus usually ab­ sent. Fig. 145.61.1—3. Boswellia Acacia Sterculia woodland, rock) slopes with scrub of Euclea, Carissa and Calpurnea, rolling grassland, fannland on black soil, weed in fields, waste­ land: 1200-2300 in. EW TU WU GD GJ SU AR WG KF SD BA HA. Somalia. Sudan. E Africa. Bunindi Mesfin T. 7342. Friis et al. 3062; Mooney 7604. Var.fischeri (RE. Fr.) C. Jeffrey fin Kew Bull 43 254 (1988)| is found in E Africa. 2. G. boranensis (S. Moore) A/.G. Gilbert (1981); Erlangea boranensis S. Moore (1902) - typc: Ethi­ opia [?SD]/Kenya, Boran. Donaldson Smith 320 (BM holo.). E. cordifolia (Benth. ex Oliv.) S. Moore var. fimbriata O. Hoffm. & Muschl in Bot. Jahrb. Syst. 46: 65 (1911). - type: SD. Ciaffa. 1893. Ruspoli & Riva 606/1153 (FT holo.). E. boranensis S. Moore var. major Lan/a in Miss. Biol. Paese Borana 4. Racc Bot.: 244 (1939) - tvpc: SD. Javello. “ Po/y.i Acacie". ('ufodontis 421 (FT holo). E. cordifolia (Benth. ex Oliv.) S. Moore var hutmhs Cufod. in V Giorn. Bot Ital n.s. 50: 101 (1943)- tvpc: SD. Mega. Corradi 1713. 1755. 1756. 1872. 20.31 (FT syn.). Erect, branched, annual or short-lived, bushy perennial herb. 20-50 cm high. Stem leafy throughout, striate, pu­ berulous with appressed. short-stalked T-shapcd hairs with unequal arms. Leaves alternate, simple, narrowly ob­ long to ovate. (1.5—)2.5—8 x 0.5-3 cm. narrowed below into a short (up to 5 nun long) petiole, green above, thinly pilose, liairs both 1-seriate, multicellular and T-shapcd. whitish below' with dense floccosc tomcntum and

appressed T-shapcd liairs on veins; margins entire; apex acute or obtuse. Capitula hemispheric, 5-7 x 6-9 mm. few in open terminal cymes; peduncle up to 4 cm long. Recep­ tacle epaleate. Phy llaries multiscnale. imbricate, ovate to ovate-lanceolate. 3-7 x 2-3 nun. light green w ith lacerate scarious margins ;md purple or violet apex, sparsely hairy near apex, acute or pointed. Florets pink to purple rarely white. Corolla glandular pubescent especially in lower parts, tubular, c 7 nun long. 5-lobed. lobes c 2 mm long, setose outside. C>pselas dark-brow n, obovate. c 0.8-1 x 0.7 nun, 5-ribbcd. sessile-glandular and sparsely liaiiy in between nbs. Pappus absent in mature cypselas. (3—)5(—7) on ovaries and immature fruits. 1.5-2 mm long, barbellate. Fig. 145.61.4— 6. Woodland on sleep slopes with Acacia. Combretum, Termmalia and hnada; Acacia drepanolobiun: woodland on black cotton soil. Acacia bushland: 1130-1850 m SD GG BA: Kenya Mesfm T. & Vollesen 4112; Moonev 7311; Friis et al. 3278. Gute'nbergia arenosa S. Moore, misspelt as ‘G. araneosa' and now considered a synonym of G. cordifolia Oliv.. occumng in East Afnca. has. according to Cufodontis (1966: 1062). been recorded from SD. Mate­ rial lias not been seen, and the record probably represents a misidcntification of material of G. boranensis

3. G. somalensis (O. Hoffm.) M G . Gilbert ( 1981); Herderia spmalensis O Hoffm. (1906); Triplotaxis somalensis (O Hoffm.) Hutch. (1914) - types: Ethiopia |?HA|. between Dagage & Gobeli. Ellenbeck 1013 (B svn destroyed); 7HA/BA. Ogaden. Dec. 1892. Ruspoli (V Riva 19 (FT svn.). Bush>. perennial lierb or shmblct. 10-40 cm high Young siem slnatc. quadrangular, appressed tomentose with sliort-stalked asymmetrically T-shapcd hairs. Leases alter­ nate. linear-lanceolate to elliptic. 3.5— 6(—9) x 0.4-1.5 cm. narrowed below into a short petiole, green above, thinly floccosc or appressed tomentose. whitish below with dense floccose tomcntum and long stalked, symmetrically T-shaped liairs on veins; margins crcnulatc. nanowly revolutc: apex acute-acuminate Capitula campanulas. 3-5 \ 3 4 nun, numerous in terminal paniculate cymes: peduncle up to 1 5 cm long. Reccptacle epaleate. Phy llanes 4-5-scnate. imbricate, linear to lanceolate. 1-2.5 x 0.5—1 nun. densely appressed pubescent with wliite silky liairs. light green with purple pointed lips. Florets bright purple, or magenta. Corolla sessile-glandular and densely white pubesccnl on one side, c 7 nun long. 5-lobcd. lobes c 2 mm long Cypselas brown, obovate to oblong, c 1.2-1.5 \ 1 mm. (3-) 5(-6)-nbbcd or angled, more or less flat between the nbs or angles, sessile-glandular and pubesccnl Pappus absent. Fig. 145.61.7—9. Acacia Balanites bushland on low-rolling hills on limestone. Acacia drepanolobium bushland on black cot­ ton soil. Acacia ( ommiphora bushland: 870-1860 m. SD HA; Somalia. N Kenya. Gilbert 2026; Mesfin T. & I ollesen 4186. Friis et al. 3 192.

145. ASTERACEAE: 32. Gutenbergia

1 C, x E

105

aure 61. G U TE N BE R G IA R U E P P E L L II var. R U E P P E L L II 1 - habit x c 1; 2 phyllary x 24; 3 im m ature cypsela x 12. B O R 4 N E N S IS 4 - habit, x c 1 . 5 - phyllarv x 24; 6 - cypsela x 12. G. S O M A L E N S IS : 7 - habit, x c 1; 8 - phyllary x 24 9 - c> psela 2 .1 -3 from Friis , Mesfin T. & Vollesen 3062; 4 -6 from Mesfin T. & Vollesen 4305; 7 -9 from I'm s, Mesfin T. & Vollesen 3192. awn by Victoria G. Friis.

106

145. ASTERACEAE: 33. Eriangea, 34. Bothriocline

33. ERJLANGEA Sch. Bip. (1853) C. Jeffrey in Kew Bull. 43: 255-257 (1988) Annual or perennial herbs. Indumentum of 1-senate. multiceLlular liairs with apically elongated cells on vege­ tative parts. Leaves alternate, simple, sessile, linearoblong to linear-lanceolate, penninerved. Capitula homo­ gamous. many-flowered, pedunculate in tenninal cymes, or solitary , tenninating branches; involucre hemispheric. Phy llaries herbaceous or conaceous. 2-4-seriate Recep­ tacle epaleate. Florets purple Corolla tubular. 5-lobed. lobes wilh stiff hairs. Anthers shortly sagittate. Cypselas 3-6-angled, glabrous, truncate at apex. Pappus 1-senate, of short barbellate bristles, or absent. A genus with about 60 species in tropical Africa: 2 species in the Flora area. I -

Cypselas 4-angled; epappose; phyllaries appendaged. 1. E. centauroides Cypselas 3-angled; pappus bnstles barbellate; phyllaries without appendage. 2. E. .smithii

1 E. centauroides (.S'. Moore) S. Moore (1902); Slephanolepis centauroides S. Moore (1900); Bothriocline centauroides (S. Moore) M G. Gilbert (1981)-type: Somalia. Upper Sheik, 1897. Lort Phil­ lips s.n (BM holo ). Perennial herb with several spreading branches, stem 15-100 cm long. Leaves simple, alternate, oblanceolate. 10-60 x 3-30 mm, glaucous, densely resinous-punctate, sparsely liairv. subsessile. margins undulate-dentate Capitula discoid, solitary. 12-15 x 8-10 mm, peduncle 2-3 cm long, appressed pubescent. Phy llanes 4-6-senatc. dark green or purple toward the apex, outer ones lanceo­ late, 1-2 mm long, middle ones round or deltoid with wrinkled or lacerate margins, inner ones oblong, c 5 mm long. Florets purple or blue-violet. Corolla tubularcampanulatc. 8-10 mm long. 5-lobed. lobes c 3 mm long, papillose on outside Anthers sagittate at base, c 2 mm long, connectivc triangular, c 0.3 mm long Cypselas light brown, c 2.5-2.8 x 1.3-1.5 mm. 4-angled, glabrous except forthe sessile glands at apex. Pappus absent. Fig. 145.62. Acacia wooded grassland or scmb. in damp depres­ sions; 1360 m. HA (Ogadcn). Somalia. Gilbert 4006: Ash 1185: Burger 3008a. 2. E. smithii .S'. Moore (1902); Bothriocline smithii (S. Moore) M G. Gilbert (1981) - type: GG/SD. (Lake Stephanie or Chew Balur|. Donaldson-Smith 359 (BM holo ). Rhizomatous perennial, up to 30 cm high. Stem branched toward the apex. Leaves alternate, sessile, oblonglanceolate. 4-6 x 0.9-1 8 cm. light green, sparsely pubes­ cent on both surfaces. Capitula homogamous. few in open pedunculate cymes or solitary tenninating branches; peduncle up to 2 cm long, apprcsscd-pubcsccnt. Phy 11arics 5-6-seriate. imbricate, ovate to lanccolete. all charactenstically purple-tipped, outer 1.5—2 x 1-1.5 nun: sparsely appressed pubescent; inner 5-6 x 2-2.5 mm.

sparsely appressed-pubescent and densely sessile glandu­ lar. Florets dark purple. Corolla tubular, c 5 mm long. 5-lobed. lobes pappillose. Anthers sagittate at base, c 2 mm long, connective c. 0.4 nun long. Cypselas light brown. 3(-4)-angled or ribbed, sparsely sessile glandular. 3—3.2 x 1-2 mm. Pappus white, c 1.2 mm long, barbellate. Acaciaseval woodland on black cotton soil; 440-1700 m. EW GG SD Somlia, Kenya and Tanzania. Gilbert & Sebsebe I). 8752; Baldrati 1541; Bonnefille 159. Imperfectly known o r excluded taxa Cufodontis (1957) recorded the follow ing names for matenal from the Flora area. One of these is a inrssapplied name while the other has not been verified because the type specimen lias not been found. E. schebelensis .S' Moore (1914) - type: HA or BA. Ucbi-Sccbclli. DonaldsonSmith s.n. (?BM holo.. not traced). The taxon cannot be identified due to lack of adequate charactcrs in the protologue and lack of type specimen. E. tomcntosa {Oliv. & Hiem) S. Moore (1908) - type: Tanzania. Kilimanjaro. New s.n. (K holo ). The type of tlus name was examined and was found to be­ long to Bothriocline. Jeffrey (1988) has placed it as a synon> m of B. long fees (Oliv. & Hiem.) N.E. Br. (1894).

34.BOTHRIOCLINE Oliv. ex Benth. (1877) Annual or perennial herbs or slmibs. Leaves alternate, op­ posite or rarely whorled. penninerved. Inflorescence of corymbose cymes. Capitula homogamous. manyflowered: involucre hemispheric or broadly campanulate Phyllaries few- to many-seriate, imbncate. unequal, conaceous. Florets bisexual. Corolla tubular. 5-lobed. Anthers sagittate and obtuse at base, connective lanceo­ late. Style bifurcate, branches thinly pappillose. Cypselas oblong, obovoid-oblong. obovoid-ellipsoid. rounded to truncate with a low apical rim. or a distinct apical cup. or with crcnate cartilaginous rim. nanowly 4-5 (-9)-ribbed. glabrous and smooth between the ribs or densely covered with 1-celled mcliomes. Pappus of caducous white barbellate bristles A genus witll about 14 species in tropical Afnca: I species in the Flora area. B. schimperi Oliv. A- Hiem ex Benth. (1877); Eriangea schimperi (Oliv. ct- Hiem ex Benth.) S. Moore (1902)-type: GD. Gaffat |near Debra Tabor|. 8100 ft {2700 m|. Schimper 1497 (BM holo.). Bothriocline angelinii Fiori (1940); Eriangea angelinii (Fiori) Cufod. (1966) - type: KF. near Bonga. Saccardo 18 (FT holo., K photo). Shrub. 1-3 m high, or nchly branched perennial herb, up to 2 in high. Stem more or less woody and glabrous below;

145. ASTERACEAE: 33. Erlangea, 34. Bothriociinc. 35. Ethulia

107

ha rs simple. 1-seriate. Leaves opposite orwhorled. rarely alt mate, lanceolate or ovate-lanceolate, base rounded or dil led; petiole broad, ciliate. up to 2 cm long, joined by a na row band around the stem; surfaces dark green and sp rsely pubescent above, pale green and thinly pilose-tomentose beneath; margins regularly serrate with po rued teeth: apex acute, 2.5-18 x 1—5.5 cm. Capitula bn adly campanulate. 7-10 x 5-7 mm. in paniculate co vinbose cymes; peduncle up to 2 cm long, tomentose. R ejp tacle foveolate. flat. Phyllaries 4-6-seriate. ovate to ob jng-lanceolate. 3-7 x 1-1.2 mm. firm, green or pur­ plish upwards with a broad scarious. whitish serrulate m; rgin and apex; apex acute and pointed or obtuse; sur­ faces bristly-pubescent near apex, hairs with glandular base, innermost linear-oblong, wholly scarious. Florets pa : or dark lilac. Corolla tubular, glabrous or sparsely se iile-glandular. c 8 mm long. 5-lobed. lobes c 2 mm lo: t Anthers 3-4 nun long. Cypselas obovid or ellipsoid, c Ip - 1 .8 x ! mm, glabrous, with 4(-6) prominent, white, ro nded ridges alternating with black narrow bands; apex tn iicate. Pappus (when mature) caducous; in young fruits an : ovaries c 1.5 mm long, barbellate. Fig. 145.63. Thick wet evergreen montane forest margin, in damp de :ressions in Combretum-Terminalia wooded grassland wi 1 tall grass etc; grassy roadside thickets along stream ba iks. in dry woodlands, 1300-2820 in. GD GJ SU AR W ] IL KF GG SD BA; not known elsewhere. Mesfin T. & Kt znew 2424; Thulin 1641; IV. de Wilde 9379.

35. ETHULIA L ./(1 7 6 2 ) H oehnelia Schweinf. (1892)

M d Gilbert & C. Jeffrey mKew Bull. 43 165-193 0988). Ai :iual or short-lived perennial, rarely rhizomatous he bs. Indumentum of simple liairs and globose trich mes. Stem striate, cylindric. Leaves alternate, simple, se:sile to petiolate, ovate to linear-lanceolate, base cu eate to decurrent, margin subentire to serrate or de tate, apex acute, rarely obtuse; surfaces glandularpu ictate, glabrous to densely pubescent. Inflorescence a tei ninal cy me. Capitula small, homogamous; peduncle sk rider with small filiform bracts. Receptacle naked, flat or >hghtly convex. Phyllaries several-seriate, imbricate, m; rgins scarious. Florets pink, lilac or purple, bisexual, ex erted. Corolla narrowiv campanulate. 5-lobed. sp rsely papillose in lower parts. Anther base obtuse. St le branches subulate. Cy pselas cylindrical to obconic. with a prominent pale apical rim contiguous with 2-6 Iongn idinal ribs, surface between ribs usually pigmented and wi li globose trichonies. rarely also a few' short white ha re. Pappus absent.

Figure 62. ERLANGEA CENTAL!ROIDES. 1 - part o f stem; 2 outer phyllary, 3 floret: 4 corolla; 5 corolla split open to show style and anthers; 6 anther; 7 - cypsela with a cross-section, 8 - pappus bristle. All from Lor! Phillips s.n. No magnifications given in orig. Drawn by S. Moore. (Reproduced with permission from Journ. Bot. 38, 1900, Tab. 409:A. as

Steplianolepis centauroides.)

A genus with 18 species of which 15 are restricted to Al ica and Madagascar; 2 species in the Flora area. 1

Capitula mostly pedunculate, if subsessile then al­ ways solitary; corolla 2.2-2.5 mm long; involucre hemispherical in fruit, fruits hidden by the incurved phyllaries. 1. E. gracilis

-

Capitula usually in sessile or subsessile groups; co­ rolla 1.2-1.5 mm long; involucre round in fruit, fruits easily visible en mass from the side. 2. E. conyzoides

108

145. ASTERACEAE: 34 Bothnoeline. 35 Ethulia

Figure 63.

B O T IIR IO C U N E SCH IM P E R I 1 upper pari o f plant x 'v. 2 capitulum \ 11a. 3 outer phyllary x 3. 4 inner phyllary x 3; 5 floret without ovar> and pappus x 3: 6 cypsela with pappus x 3. All from Mesfin / ’ & Sehsehe I). 1271 Drawn by Birhanu Mihrctu

1 E. gracilis Del. (1826); E. conyzoides L. f. var. gracilis (Del.) Asch & Schweinf.. III. Flore d ’Egypte: 84 (1887) - type: Su­ dan. el-Qerebyn. Cailliaud(MPU holo ). Erect, annual herb, 0.5-1.5 m high. Leaves narrow ly lan­ ceolate to elliptic, up to 1 2 x 2 cm. sessile or petiolate. sparsely appressed-pubescent; base narrowly cuneate: margins irregularly serrate-dentate; apex acute; petiole up to 1.5 cm long, with narrow ly decurrent leaf base, pilose Capitula campanulate to urceolate, up to 2 x 1.5 mm at anthesis. usually pedunculate, with peduncle up to 8 mm long, appressed-pubescent. if sessile, capitula solitary Fruiting capitula 2.5-3 mm wide with fruits often hidden by the inclined phvllaries. Phyllaries 3-4-seriate. 1.5-3 x 0.5 nun. lanceolate to elliptical, basal lialf to two-thirds light greenish-yellow, smooth, glabrous or with few scattered wlute hairs, apical portion foliaceous. often purplish-violet, and with globose trichomes. margins scarious. ciliate; apex acute or acuminate. Florets pale lilac to mauve. Corolla 2.2-2.5 mm long Cypselas yel­ lowish brown. 0.8-1.7 x 0.8-1 mm, obovate (2-)4-7ribbed. with globose trichomes between ribs, otherwise glabrous, apex with narrow rim forming a shallow cup Pappus absent. Disturbed river banks, margins of pools, weed in crop fields; wet roadside ditches: 300-2150 m. EW TU GD GJ SU IL SD BA; Sudan. Uganda. Chad. Central African Re­ public, Cameroon. Mesfin T. & Kagnew 1696; Thulin & Hunde 4056; Bigazzi & Tardelli 81.

2. E. conyzoides L f (1712) - type not designated Original material a plant grow n from seeds sent by van Royen from Sn Lanka (Ceylon), not localized. subsp conyzoidcs Erect, annual herb, up to I in high. Stem glabrous below, appressed pubcsccnt in upper parts. Leaves elliptic to lan­ ceolate. up to 1 2 x 2 cm. subsessile to shortly petiolate; thinly appressed pubcsccnt: petiole with narrow decurrent leaf base, up to 6 mm long, appressed pubescent: base cuncate: margin serrate especially from middle to apex, apex acute to acuminate. Capitula campanulate at anthesis. 2-2.5 \ 2-2.5 mm at anthesis. mostly sessile (2 to 3 together) or. in some, peduncle up to 5 111111 long Fruiting capitula 3.5-4 5 111111 w ide, rounded, fruits easily visible en mass from the side. Phy llaries 2-3-senate. lan­ ceolate. 1-2 \ 0.5 nun. glabrous from base to middle, green in upper half and with globose tnchomes: margins scanous. ciliatc. apex acutc. Florets inauvc or purple, rarely white. Corolla 1.2-1 5 mm long, tubular with nar­ rower tube. Cypselas forming smooth hemispherical mass when fully mature, yellowish-brown, obconical. 1.2-2 x 0.8-1 mm. (3-)4-5(-6)-ribbed with laterally expanded apical rib and globose trichomes between ribs, otherwise glabrous; apex truncate. Pappus absent. Fig. 145.64. Seasonally marshy areas, weed in field, bank of rivers; 1620-2100 in. GD GJ SU: widespread in Egypt, most of tropical Africa and N Mozambique; tropical and warm temperate Asia (e.g.. Vietnam, Thailand. India. Sri Lanka.

145. ASTERACEAE: 35. Ethulia, 36. Elephantopus

109

Syn a) (cf. Gilbert & Jeffrey 1988). Gilbert & Getachew A . >48; Lundstrom 50; Mercier 1593. Subsp. kraussi (Walp.) M.G. Gilbert & Jeffrey is found in Mozambique, South Africa (Natal) and Brazil.

36. ELEPHANTOPUS* L. (1753) Perennial herbs, sometimes scapose. Leaves cauline. Caoitula small, few-flowered, aggregated in secondary hei is surrounded by foliaceous bracts; secondary heads sol lary or variously arranged in corymbs, panicles, or spi ;es. Phyllaries usually 8, in four decussate pairs. Flo­ ret white, pink, or blue, slightly bilaterally symmetrical an< :;>seudoligulate. more deeply cleft between two inner lot:s. Cypselas oblong-obovoid, somewhat flattened. 10 nbbed, setose and often glandular between the ribs. Pa pus of 5 to many, scabrid-barbellate, basally widened bn ties, rarely reduced to a scaly corona. A genus with about 32 species in tropical parts of the world but most common in South America; the following wi lespread species known from the Flora area. E. mollis Kunth (1818) - type: Venezuela, Caracas, Humboldt & Bonpla n d 627 (P holo.). Er cl, perennial herb, up to 50 cm high, stem hirsute with wl i.te ascending liairs. Leaves alternate (middle ones), pa ifcgreen up to 16x 6 cm, obovate-oblanceolate to ellip­ tic sub-sessile, or petioles up to 15 mm long with narrow de urrent leaf base and clasping stem; base cuneate; margii s cnenate-serrate; apex acute or acuminate, scabrid ab >ve, soft-puberulent and densely resinous-punctate be­ ne ilh. Capitula in axillary' and tenninal corymbose gli merules, up to 2 cm wide, surrounded by 3-4 ovatelai ceolate. densely hirsute bracts, shorter than the git merules; each 4-5-flowered. Phyllaries 4-5-seriate, gr en, narrowly lanceolate and spinous, glabrous or sp irsely hirsute towards apex; outer c 4x1 mm; inner c 10 x 1.5 mm. Florets white. Corolla c 7 mm long, 5-lobed, loi >es c 2 mm long, narrowly tubular below, funnelsh iped above, glabrous. Cypselas c 4 mm long, greybr wn. 9 - 10-ribbed, setulose between the ribs. Pappus br 5tles c 5-6, c 4.5-5 mm long, barbellate except for the di ;ited base Fig. 145.65 In shade in gallery forest; 1600 m. WG; native of the m ist neotropics and widely naturalized in the Old World, in l u d i n g West, Central. East and South-Central Africa. H de Wilde 8891. This species is rather similar to the Old World E. sc i er L„ which is distinguished by its stiff, coriaceous, ot ong-oblanceolate leaves tliat are not soft-puberulent be leatli.

Tribe 5. A R C T O T E A E Cass. (1819) Annual or perennial herbs. Stem erect or decumbent, sometimes absent and plant scapigerous. Leaves alternate

Figure 64. ETHULIA CONYZOIDES subsp. CONYZOIDES 1 - habit x 4/5; 2 - cypsela x 12. (No specim ens cited in orig.) (Reproduced with permission from Flora o f India, vol. 13. fig. 103.)

or radical and more or less rosulate, simple to pinnatifid. Capitula heterogainous and radiate, campanulate. hemi­ spheric. ovoid or subglobose, pedunculate, usually large and showy. Receptacle naked or fimbriate-alveolate,

110

145 ASTERACEAE: 36 Elephantopus

Figure 65.

ELEPHANTOPUS MOLLIS

1 -

habit x '/2; 2 - glom erule o f capitula x c 2 ;3 capitul um x 53/4; 4 outer and inner phyllaries x 53/4; 5 - floret x c 7; 6 stam ens x 22; 7 - style arms x c 20(7); 8 - cypsela with pappus x c 7 1, 3 -4 from Boivin 1128; 2. 5 -7 from Ca­ det 2217; 8 from Gueho (in MAU 20351). Drawn by Eleanor Catherine (Reproduced from FI. Mascareigttes. C o m p o se s . PI. 5.)

rarely paleate Phyllaries multiseriate. free or fused at base, margins and tips scarious or spinescent. Ray florets feinale or neuter, with or without stanunodcs. styles filiform, sty le brandies linear-lanceolate or elliptic. Disk florets yellow, bisexual, sometimes the innermost func­ tionally male. Corolla actinomorphic. 5-lobcd. tubular or campanulate and usually wider in upper parts Anthersentirc or sagittate at base, the auricles obtuse or acute, rarely failed. Style teretc or flattened. apicaJly cleft, portion be­ low the cleft to the area with the fringe of papillae tluckencd. apex of branches truncate or acute, inner surface papillose. Cypselas glabrous or hairy, often ribbed. Pappus of scales or barbellate setae. 2- to several-seriate A tribe of 16 genera with some 200 species naUve to the Old World and centered in South Africa. 3 indigenous and 2 introduced genera in the Flora area. 1

Ray florets female; phyllaries more or less free, apices of especially the inner ones obtuse, scanous. 2

Ray florets sterile; phy llaries connate at the base or higher up. apices acute or acuminate, often spinetippcd 3 Cypsela surface cliambered or grooved on one side, convex and smooth on the other, inner pappus scales oblong to oblong-ovate, irregularly dentate at margins, apex obtuse. 37. Arctotis Cypsela surface obscurely longitudinally ribbed, not grooved or cliambered; inner pappus scales linear with sharp acuminate apex. 38 H aplocarpha Phyllarics spiny, connate at the base only. 39. Berkheya Phy llaries not spiny, mostly bristly, connate up to near middle forming a cup-like involucre. 4 4

Plants more or less bristly-setose throughout or glabrescent. pappus dimorphic, of elliptic to ovate-lanceolate scales, the outer whorl much longer than the inner, overiapping, sessile; the in­ ner scales stipitate. 41. Hirpicium

145 ASTKRACKAK: 37 A rctohs. 38. I laplocarpha

Plants nol hispid-setose. arancosc or glabrous; pappus inonoinorphic. of subequal slender linear scales 40. Ga/.ania

37. ARCTOTIS* A. (1753) Pe cnnial herbs. Stems erect or decumbent. branched. Leaves alternate. entire to pinnateh lobed. often whitish toi icntosc Capitula radiate, solitary , long-pedunculatc. Rc.cptacle flat, pitted. bristly Involucre heinisphenc. Ph llanes 4-5-scnate. imbricate or overlapping. Ray flo­ ret- yellow. 15-25. female. Disc florets yellow, numerou| bisexual or innennost fcmalc-slcrilc. Anther bases minutely sagittate, connective ovate-triangular. Style sic ider below. thickened above the minutely liairy node, an i' very short. Cypselas obovoid-obconical. outer sur­ face 3-5-ribbcd. sometimes fonning two dorsal furrows at m< unity. inner surface smooth or nigose and without nbs. gl< fctous lo pilose, often with a basal tuft of long hairs. Pappus of two rows of overlapping scales. A genus confined to southern Africa occurring clucfly in South Africa. Namibia and Angola and introduced or cu uvated elsewhere. So far one species is found as a gar­ de i ornamental in the Flora area. A. stoechadifolia P. Berg. (1767) - type: South Africa. Berguis s.n. (SBT holo ). Pe cnnial lierb. c 1 m high. Stem terete, densely whitewoolly Leaves obovate to oblong in outline, pinnateh loi cd. lobes entire or irregularly toothed, densely whitcwc oily especially on lower surface, sessile. l-10cm long, loi cs 4-7 Him w ide. Capitula 60-80 x 10-15 mm at an iicsis. solitary Peduncle striatc-sulcaic. 10-25 cm loni white-woolly. Ray florets unifonnly yellow or white ib )Ve and violet below, c 20 Disc florets violet or purple. Cc idla tubular below, campanulate in upper parts. 5-1 obed. Cypselas obovoid. 3-5-ribbed w ith 2-3 longitu­ dinal grooves on one side, base with many straight hairs. Pa ipus 2-seriate. outer scales small, inner scales larger th; it cypselas and its hairs. Grown mostly as an ornamental garden plant, but also known as a rare escape from cultivation: 1800 m. SD (Y rga Muda). a native to South Africa S um & Faris 568.

3X H A P L O C A R P H A I.css. (1831) Landtia Less. (1832). Oliv. sp africanus (H ook.f.) Ortiz & Paiva in Bot. J. Linn. africanus. The epithet subsp. africanus (1995) lias prior­ Soc. 117:44 (1995); ity at tliat rank over subsp. buchwaldii (1996). A. Telekia africana Hook. f. (1864); Anisopappus chinensis subsp. chinensis occurs in Madagascar and in africanus (Hook, f.) Oliv. & Hiem (1877); - type: Asia to China, while a third subspecies, A. chinensis Cameroun, Mt. Cameroun, Mann s.n. (K holo.). subsp. lobatus (Wild) Ortiz & Paiva occurs in Zambia and A. chinensis (L.) Hook. & Am. subsp. buchwaldii Zimbabwe. (O. Hoffm.) Ortiz., Paiva & Rodr.Oubina in Anal. Jard. Bot. Madrid 54: 383 (1996); Sphacophvllum buchwaldii O. Hoffm. (1898); Anisopappus buch­ 2. A. holstii (O. Hoffm.) Wild (1964); waldii (O. Hoffm.) Wild (1964) - type: Tanzania. Sphacophyllum holstii O. Hoffm. (1895) - type: Usambara Mts., Buchwald 221 (B holo., destroyed; Tanzania. Usambara, Holst 3915 (B holo., destroyed); BM COI K iso.). Usambara, Kwa Mshuza, Holst 8898 (K nco., P Anisopappus inuloides Hutch. & B.L. Burtt (1932) isoneo., selected by Beentje 2002). - type: Congo, Quarre 2847 (BR holo., K iso.). A. suborbicularis Hutch. & B.L. Bum (1932) - Erect annual herb. 15-25 cm high. Stem slender, c 1 nun type: Congo, de Witte 482 (BR holo.. K iso ). wide, sparsely branched in upper parts, striate-sulcate, Telekia niamniamensis Schweinf., nom. nud. pilose. Leaves simple, alternate, petiolate, blade ovate, 1.5—4 x 0.6-2 cm, membranous, hairy on veins of particu­ E ecL stiff perennial herb, c 1 m high; stem glabrcscent to larly the lower surface, glabrous or glabrcscent above, d nsely crisped-pubescent. Leaves alternate, up to 6.5 x 3 margins serrate, petiole up to 10 mm long, densely ciliate. cm, deltate to ovoid-oblong, light-green, 3-5-veined from Capitula campanulate, 3-5 x 3-4 mm at anthesis, not tl . base, w ith only the midvein being prominent, sparsely much altered in fmit, in open terminal or axillary cymes, s*. atorid above, densely pubescent below with denser hair peduncle up to 2 cm long, pubescent. Phyllaries whit­ in the prominent reticulate veins, base cordate-hastate, ish-green with dark green mid-rib, folded inwards, 1-1.5 m argins crenate except near tip; apex obtuse or acute; x 0.3-0.5 mm, pilose with liairs mostly on mid-rib and to­ p< iiolate, petiole 1-1.5 cm long, crispid-pubescent. ward the apex, apex apiculate. Paleae scarious, folded flat C ipitula solitary at the end of branches, or several corymor folded. 1.5-2 mm long, apex apiculate, similar to b 'sely arranged on up to 10 cm long, densely scabridphyllaries. Ray florets yellow, female, rays 1-3 mm long. pi lose, peduncles; involucre depressed-hemispheric. Disc florets yellow. Corolla tubular-campanulate, c 1.5 P:ryllaries 3-4-seriate. up to 10 x 1 mm, light green, mm long, glandular in lower parts. 5-lobed, lobes c 0.3 subequal, linear-oblanceolate, tapering to an acute apex; mm long. Style bifurcate, c 1.4 mm long. Anthers yellow, densely pubescent on outside. Ray florets yellow, 12-14 1

Capitula 10-25 mm in diameter, ray florets 12-14 mm long; perennial herbs, c 1 m high. 1. A. chinensis subsp. africanus Capitula 3-5 mm in diameter; ray florets 1-3 mm long; annual herbs, c 15-25 cm high 2. A. holstii

120

145 ASTERACEAE 42. Anisopappus. 43 Geigeria

Figure 69.

ANISO PAPPU S C H IN E N SIS subsp AFRICANUS: 1 flowering branch. 2 outer phyllary; 3 ray floret. 4 -d is c floret and palea. 5 inner phyllary . 6 two anthers. 7 - style arms. No magnifi­ cations given and no specimens cited in original publication. Drawn by W.E Trevithick. (Reproduced with perm is­ sion from FI. W. Trop. Afr., vol. 2 ,1963, fig. 255, as/1, africanus.)

c 0.6 nun long. Cypselas black, columnar, c 10-ribbed. c 1 x 0.3 nun, glabrous. Pappus absent. Woodland with Acacio, Combretum, Entada, Sclerocarya and Zanthoxvlum on steep slopes with abandoned cultivations. 1550 m. GG: E Africa. Gilbert et al. 8971 Until recently this species lias only been known front East Africa; the specimen cited was collected from an area between Jinka and Konso (GG). It is not known whether it represents a natural population or a recent introduction But the relatively recent collections of other species, such as Vanilla roscheri Rchb.f. jOrchidaceae. Bidgood & Cnbb in Kew Bull. 54: 378 (1999)) from the same general area with similar distributional gaps, might indicate that tins is a natural component of the vegetation. Further col­ lections in Gamo Gofa might also bring forth other re­ cords showing similar vicariance.

43. GEIGERIA Griess. (1830) Diplostemma Hochst. et Steud. ex DC. (1838). Araschcoolia Sch. Bip. ex Benth. & Hook. f. (1873). 11 M erxm uller in Mitt. Bot. Staatssamml. Mtinchen 1, 7 239-316(1953). Bushy herbs, up to 50 cm liigh with prostrate or erect branches, or rosulate acaulescent herbs; stem winged or wingless. Leaves simple, opposite or alternate, linearlanceolate to linear-elliptic, glabrous to woolly. Capitula sessile to shortly pedunculate, often concealed amongst the leaves, terminal and solitary or aggregated, or in the leaf axils or forks of the branches, heterogamous, radiate Involucre ovoid or campanulate. Phyllaries multiseriate. imbricate, outermost leafy, inner scarious or leathery. Re­ ceptacle flat or convex, paleate. Ray florets uni-seriate, fe­ male. tubular below; rays oblong to linear-oblong. 3-fid or 3-lobed at apex. Paleae conduplicate, fimbriate. Disc florets yellow, tubular, fertile, 5-iobed. Anthers sagittateauriculate at the base, with long caudate-acuminate tails;

style branches linear-lanceolate, apices obtuse to sub­ acute Cypselas indistinctly 3-4-angled, hairy . Pappus of 2-senate, similar scales, or the inner series longer and bristly. A genus w ith about 25 species mainly found in south­ ern Africa; 2 species in the Flora area. 1.

-

Leaves glabrous or glabrescent, glandular, shiny; stem up to 50 cm high or. if acaulescent, leaves glabrous and shiny. 2. G. aiata Leaves densely whitish-araneose or cottony ; acau­ lescent herbs. 1. G. acaulis

1. G. acaulis Benth & Hook.f. ex Oliv. & Hiern (1877); type: Sudan. Cordofan. Arasch-Cool. Kotschy 104 (K holo.. M P Z iso ). Araschcoolia acaulis Sch. Bip., nom. nud.; Diplo­ stemma acaulis Sch. Bip., nom. nud. Acaulescent perennial herb, with slender, up to 15 cm long taproot, rarely annual, stem rarely up to 2 cm long, araneose. Leaves crowded at base, sessile or subsessile, up to 8.5 cm long, 5-9 mm wide, narrowly oblanceolate, tapenng to the base, densely grey-white araneose or cot­ tony. margins entire, apex subobtuse, mucronate. Capitula solitary, sessile in the crowded leaf axils, villose, c 10 mm in diameter, ovoid. Phyllaries 2-3-seriate, subequal, 5-7 mm long, lanceolate, tapering to an acuminate apex, ciliate at margins near apex, densely cot­ tony at base. Paleae c 2.5 mm long. Ray florets female, pale yellow, rays up to 5 inm long, oblanceolate, apex Vfid. Disc florets pale yellow, c 2-3 mm long, cylindric campanulate. with short tube; corolla 5-lobed with long narrow lobes, papillose. Cypselas c I mm long. narrowly cylindric, hairy. Pappus bi-aristate; outer series palea­ ceous. c 1 mm long, oblong, obtuse at apex; inner series aristate, longer than the outer series. Fig. 145.70. Alluvial flats in open Acacia bushland w ith numerous water courses. 325-650(-1900) m. EW SD GG; Sudan,

145. ASTERACEAE: 43. Geigeria

121

Figure 70. G EIGERIA ACAU LIS 1 habit x 2/ 3 ; 2 - capitulum , cross section (simplified) x 6 ; 3 - inner phyllary x 6 ; 4 - palea x 12; 5 - outer floret x 10; 6 style o f outer floret x 18; 7 - inner floret x 10; 8 - stamens x 18; 9 - style o f inner floret x 24; 10 - cypsela x 12. 1 from Gilbert 4731; 2 -1 0 from Polhill & Greenway 452. Drawn by Pat Halliday. (Reproduced with permission from FI. Trop. E. Afr .. C om p o sitae, fig. 6 8 .)

U anda, Kenya. Tanzania. Angola. Zimbabwe. Namibia. S< ,lh Africa (Transvaal). Mooney 9535; Gilbert et al. 8 41; Sebsebe D. & Ensermu K. 2 758. 2 G. alata (DC\) Benth. & Hook. f. ex Oliv. & Hiern 1877); Cichorium a Ialum Hochst. & Steud.. nom. nud.; Diplostemma alatum DC. (1838) - type: Yemen. Mt. Sedder. Schimper 853 (K M). Annual herb, up to 60 cm high; stem dichotomously benched, glabrous or sparsely pubescent, often 3w tnged. Leaves oblong-lanceolate, bright yellow-green, sessile, decurrent, up to 5 x 1.2 cm. glabrous to sparsely pu rescent, densely glandular-punctate, lateral veins fused

and running parallel to margins, margins entire; apex acute-mucronatc. Capitula sessile, solitary or several at the forks o f the branches, u rc c o la te. 1 0 -1 2 -flo w e rc d . Phyllaries imbricate. 5-6-seriatc. c 4.5-5.5 mm long, pu­ bescent. glandular-punctate, ovate and terminated by a spinous, reflexcd awn. Ray florets bright yellow, female. c 4 mm long, tube c 2.5 mm long, glandular-punctate. Disc florets yellow, campanulate. densely glandularpunctate. c 3.5 mm long; corolla 5-lobed. Cypselas c 1.5 mm long, densely sericeous. Pappus c 1.8 mm long. 2-seriate. outer series paleaceous, c 1 mm long, inner sc­ ries similar but with elongate aristae. River-bed in dry bushland with Boswellia. Commi­ phora. and Acacia, open dry busliland. subdesert with lava;

122

145 ASTERACEAE: 43 G eigena, 44 Inula

60-1500 m. EE SD HA; Egypt. Sudan, Yemen. Saudi Ara­ bia. Mali. Niger. Chad. Somalia, Djibouti, Kenya, Namibia. Ellis 221; Gilbert et al. 8189; Gillett 15092.

44. INULA L. (1753) Petrollina Chiov. (1911) Perennial herbs or shrubs. Leaves simple, alternate, entire or serrate, sessile to shortly petiolate, seabrid-pubeseent or tomentose. Capitula usually in terminal corymbose cymes, occasionally on short lateral branches, hetero­ gamous. radiate; involucre hemi-spherical to turbinate. Receptacle plane or slightly convex, areolate orfoveolate. Phyllaries multiseriate, imbricate, herbaceous or scarious. Florets yellow; ray florets female or absent, one- to multi-seriate. Female florets few or many. 1-2-seriate. tu­ bular 3-fid at apex. Disc florets bisexual, tubular, corolla 5-lobed; anthers sagittate and tailed at base. Cypselas cy­ lindric, slightly constricted at apex, angular or 4—10ribbed. Pappus one- to several-seriate, persistent, un­ equal. scabrid.

late, peduncle 0 5-1.5 cm long, villous. Receptacle naked, flat c 8-10 mm across. Phyllaries multiseriate, imbricate, more or less equal, 9-10 x 1.5—2.5 mm. apex acute, cili­ ate; outennost senes foliaceous, densely villous; middle lanceolate, villous, successively wider, inner linear shortly hairy. Florets yellow, outermost with strap-shaped rays. Cypselas oblong. 1.5-2.0 mm long, costate, with erect patent short setae near the apex. Pappus brownish. 1-seriate, barbellate. Erica arboreo scrub; 3200-3600 m. GD; not known elsewhere. Schimper 1061; Lemma G.S. 690; PichiSermolli 2621. Inula arbuscula is very closely related to the more

widespread /. confertiflora, the differences being mainly in the size and shape of the phyllaries and the type and de­ gree of indumentum on the leaves and phyllaries. More collections from the type locality of I. arbuscula and from Wello and Goj[jam might further reveal the affinities be­ tween them.

A genus with about 100 species found in Europe Asia and Africa. 4 species in the Flora area.

2 . 1, confertiflora A. Rich. (1848) - type: SU. ’Choa'. Petit s.n. (P holo. not seen)

1

Shnib. 1.5—2.5 m high, stem much branched above, branches terete, stnate, white tomentellous and leafy at extremities. Leaves densely fonned towards apex of branches, sub membranous, elongately lanceolate. 3—13.5 x 1— 4.5 cm. white tomentellous beneath, nearly glabrous above, reticulately veined, narrowed at base, shortly petiolate. margins denticulate, teeth gland-tipped, apex acute. Involucre sub-hemisphenc. 10-18 mm across. Capitula many-flowered, in dense corymbose or rounded terminal cymes, pedunculate, peduncle with foliaceous bracts, up to 2.5 cm long. Receptacle flat, naked. 3-5 mm across. Phyllaries multiseriate. unequal, imbricate, ciliatc. outer successivcK shorter, appendaged. more obtuse lan­ ceolate and ham at the back; inner linear, acute. Florets yellow, outermost with strap-shaped rays. Cypselas ob­ long. 1.5-2.2 mm long, terete, c 10-ribbed. with short, erect-patent setae near the apex. Pappus 1-senate, c 3 mm long, pale brow nish, minutely scabrid. Fig. 145.71

-

2

Leaves sessile to shortly petiolate. cuneate al the base; inflorescence closely surrounded by leaves; capitula hemispheric, usually 10-20 mm wide at anthesis. Cauline leaves cordate-aunculate. amplexicaul; in­ florescence carried above leaves, capitula ob­ conic to turbinate, usually 4-7 mm wide at anthesis. Phyllaries nearly equal in length, densely shaggy; leaves dark-green, slightly paler beneath

1 . 1, arbuscula

3.

Phyllaries unequal, outennost c half as long as the innennost. becoming glabrous towards center, leaves light yellowish green, silvery beneath. 2 . I. confcrtiflora lnflorecence orange-yellow; phyllaries uniformly light-green, lanceolate to linear-lanceolate, taper­ ing to an acute apex; plants of usually dry wood­ land. 3 . 1. paniculata Infiorescencc purple or yellow; phyllaries darkbrown at apex, obtuse; plants of usually forest margins. 4. I. mannii

1 I. arhuscula Del (1843) type: Ethiopia |region not known|. Ferret et Galmier s.n. (P holo ). / fruttcosa Sch. Bip., nom nud. quoad Schimper 664. Slirub. up to 3 m high; branches terete, striate and pubemlous below, villous, leafy and brownish above. Leaves ovate or lanceolate. 5-15 x 2.5-6 cm wide, nar­ rowed at the base into a short petiole, pale tomentose beneath, less so above, margins denticulate, apex acute or subacute, mucronulate; upper leaves much smaller. In­ volucre hemispheric, c 2.5-3 cm wide. Capitula manyflowered. in dense corymbose, tenninal cymes, peduncu­

Margins of and cleanngs in Juniperus Podocarpus forest. Erica arborea scrub, along stream banks in Euca­ lyptus plantations, montane grassland on slopes with scat­ tered Juniperus. Hagenia. Hypericum, etc.; 2500-3730 m. WU SU AR BA HA; not known elsewhere Mesfin T. 5828: Friis et al 5705; Mooney 6449.

3

I. paniculata (Klatt) Burtt-Da\>y (1 9 3 5 ); Monactinocephalus paniculatus Klatt (1 8 9 6 ) ty pe: South Africa. Transvaal, Rehman 6 0 6 8 (K iso ). Inulaster macrophvllus Sell Bip. ex A. Rich (1 8 4 8 ); Inula macrophvlla (Sch. Bip. cx A Rich ) Schweinf & Asch. (1 8 6 7 ). non Kar. & Kir (1841). nom. illegit.; I. decipiens E. A. Bruce (1 9 4 7 ) - type: TU. Mt. Scholoda. PI. abyss. II, Schimper 912 (K syn.. P isosy n ): ibid.. PI. abyss. II, Schimper 9 3 8 (K synl. P. isosyn.).

145. ASTERACEAE: 44. Inula. 45. Pentanema

123

E nect, perennial herb, up to 75 cm high; rootstock thick, p Je-brown, villous; stem sub-tetragonal or cylindric, stria ; pubescent. Leaves both radical and cauline; radical h aves petiolate, up to 60 cm long, petiole up to 10 cm It ng. clasping at the base, semi-circular and more or less c inaliculate or flattened above, sparsely liirsutc or g abrescent, blade up to 45 x 15 cm. ovate-elliptic to 0 ilanceolate, base cuneate and shortly decurrent on the p.mole, forming narrow wings, margins coarsely and irr gularly dentate, scabrous with tuberculate hairs above, p ibescent to browny-tomentose beneath, reticulate venat on prominent, especially on lower surface; cauline 1 aves smaller, gradually diminishing upwards, at first s . ssile with cuneate base but higher up becoming cordate, a iricled and amplexicaul. and passing into the bracts of t * inflorescence. Inflorescence a dense, tenninal. corymt : se cyme. Capitula obconic to turbinate, 4-6 mm wide; peduncle up to 1 mm long, scabrous-pubescent. Phyllaries 4-seriate. outer lanceolate, 2-3 mm long, inner progres5ively longer, c 5 mm long, linear-lanceolate, tapering to acute apex, ciliate on the margins, pubescent outside, metimes purple-tipped. Florets all bisexual, or: nge-yellow. Corolla nanowly tubular below, widening A funnel-shaped in the upper parts, 4.5-5.5 mm long, -lobed; lobes glabrous. Cypselas cylindric, pubenilous, -1.2 x 0 2-0.4 mm. Pappus 1-seriate. 3.5-4 mm long, dab rid. Penmsetum schimperi grassland, broad-leaved decidous woodland with Combretum, Terminalia, Faurea, i tc., grassy glade in moist, semi-evergreen forest; 1400; 300(-2600) m. TU GD SU AR WG KF GG SD BA HA; jdan, E Africa Zambia, Zimbabwe, Malawi, South Afza (Transvaal). Mesfin T. & Kagnew 2439; Ash 673; Mooney 5021. 4 I. mannii (Hook, f ) Oliv. & Hiem (1877); Vernonia ?mannii Hook. f. (1864) - type: Cameroun, Mann 1933 (upper part of sheet! (K syn.); ibid., Mann 1314 (radical leaf onlyl (K syn.). Loggera heteromalla Vatke (1875); Petrollinia heteromalla (Vatke) Chiov. (1911) - type: GD. near Debra Tabor. 26. Nov. & 15. Dec., 1863, Schimper 1528 (B holo. destroyed, K lecto., WU iso.). reef robust, perennial herb, up to c 2.5 m high; stem and tranches dark purplish red, angular ribbed, greyomentose. Leaves both radical and cauline; radical leaves io to 60 x 20 cm, elliptic, attenuate at the base into a short petiole, sinuate or entire and denticulate, glandular-pilose on both surfaces, daric-green above, paler beneath. auline leaves up to 50 x 15 cm decreasing in size acropetally, lanceolate-elliptic, sessile, with cordate-auricuite base, margins denticulate; apex acute, mucronate; virface light green often with reddish midrib above, nlose, grey-green tomentose beneath. Inflorescence of ]ense corymbose cymes on both stem and branch apices. ' apitula cylindric to turbinate, c 5-7 mm wide at anthesis: .eduncle up to 1 cm long, densely hirsute. Receptacle ^reolate or smooth. Phyllaries 4-5-seriate, imbricate, light -zreen with dark brown apex on upper half, linear-

Figurc 71 I N V L 1 CO NFERTIFLOR j\ 1 - upper part o f stem. 2 -cap itu lu rn in longitudinal section x 2V2\ 3 - ray floretx 5. (No specimens cited in orig.) Drawn by W. Burger. (Reproduced with permission from Burger: Families o f Flowering Plants in Ethiopia, 1967, fig. 68:4.)

lanceolate, grey-hairy ; the inner linear and tapering to an acute or acuminate apex, densely hirsute. Florets yellow or reddish purple. Corolla 5-6 mm long, cylindric but slightly wider in upper parts. 5-lobed. Cypselas 1.2-1.5 mm long, oblong-cylindric, 8-10-ribbed, glabrous. Pappus 3.5—4 mm long, scab rid. Evergreen forest margins, montane slopes with de­ rived evergreen bushland; 2100-2600 m. TU/GD GD SU KF WG; Cameroun. Congo (Kinshsa), Rwanda. E Africa. Malawi, Zimbabwe. W. de Wilde 9533: Friis et al. 498; Benedetto 238. Mooney 6132 from ‘marshy ground in grassy valley' and Anderberg 1638 from 'large bog' may belong to this species.

45. PENTANEMA* Cass. (1818) Vicoa Cass. (1829). Inula Sect. Vicoa O. Hoffm. (1890). Mesfin Tadesse in Kew Bull. 50: 401-408 (1995). Annual or perennial herbs. Leaves simple, alternate, rarely opposite. Inflorescence a lax cymose panicle. Capitula small, hemispheric, heterogamous or homo-

124

145. ASTERACEAE: 45. Pentanema, 46. Blumea

Erect, annual herb. 0.4-1 m high. Stem simple or branched above, branches sparsely pubescent. Leaves thin, alternate, sessile, 15-80 x 5-15 mm, linearlanceolate to narrowly lanceolate or narrowly elliptic, ta­ pering to an acute apex, auriculate and amplexicaul at base, puberulous, margins entire to serrate-crenate, midvein prominent below. Capitula homogamous, discoid, 10-15 x 4 -5 mm at anthesis. not much altered in height in fruit, arranged in lax cymose panicles; peduncle puberulous, up to 4 cm long, with small (3-10 mm long) linear auriculate bracts which increase in size downwards and pass into the upper leaves. Receptacle epaleate. pitted. Phyllaries multiseriate, unequal, 2-6 mm long, increasing inwards, with or without orange glands; outer green, linear-acuminate, sparsely white-hairy, spreading or re­ curved; inner linear, tapering to an acute apex, 1-veined, margins hyaline Florets 40-85, bisexual, yellow, 4-6 mm long. Corolla 3.8-4.9 mm long, narrowly cylindrical, gla­ brous. 5-lobed, lobes densely orange-glandular, c 0.5 mm long. Stamens 4.5-5.2 mm long; anthers c 3 mm long (incl. tailed connective); tail c 0.5 mm long, slightly ex­ ceeding filament-collar joint, connective c 0.3 mm long, ovate Style c 4.8-5.2 mm long, bulbous at base, bifurcate at apex, branches c 1 mm long, stigmatic surfaces mar­ ginal, confluent at apex. Cypselas light green, c 1-1.3 mm long, narrowly oblong-elliptic, pubescent, base hyaline, annular shiny. Pappus of 3—3.2 mm long scabridulous bristles. Fig. 145.72. Figure 72. PENTANEMA INDICUM: Habit x V2. (No specimen cited in orig.) Drawn by Bhola Ram. (Reproduced with permis­ sion from FI. India, vol. 13, fig 9.)

Grassland w ith patches of bare, pale brown silty soil; 500-800 m. EW IL (Gambella); West Africa, Sudan, Uganda, Tanzania Mozambique, Malawi, Zambia Zim­ babwe, Angola; Pakistan, India. Sri Lanka. Gilbert & Friis 8365; Pappi 6280; Terracciano & Pappi 2785.

gamous. Phyllaries multiseriate, imbricate, unequal, nar­ rowly linear, tapering to an acute or acuminate apex, mar­ gins scarious. Receptacle epaleate. Florets numerous, ra­ diate or discoid (in the Flora area). Corolla narrowly tubu­ lar 5-lobed; anthers sagittate and tailed at base; style bifur­ cate, stigmatic branches wide, truncate or obtuse at apex. Cypselas narrowly oblong-elliptic, smooth Pappus of 1-seriate, barbellate bristles.

Cufodontis (1967: 1117) cites Beccari s.n., from EW, Mai mentai, as tliis species, but the specimen has not been seen by the present author.

A genus with about 12 species centered in the Middle East and Asia. The following cosmopolitan species is found in the Flora area.

P. indicum (L.) Y. Ling (1965); Inula indica L. (1763) - type: “Habitat in India orientali.” Linnaean Herbarium No. 999.16 [LINN, lecto., selected by Anderbeqj in Taxon 32:652 (1983)]. Vicoa auriculata Cass. (1829). - type: Sri Lanka, Merat s.n. (P not traced). Inula leptoclada Webb. (1849); Vicoa leptoclada (Webb.) Dandy (1956) - type: Cape Verde Us, Webb s.n. (P holo ). Inulaster kotschyi Hochst. (1841), nom. nud. quoad Kotschy 108 (Sudan); Varthermia kotschyi Schweinf. & Asch (1867), nom. nud., quoad Cienkowsky s.n (Sudan); Conyza kotschyi Schweinf. & Asch. (1867), nom. nud.

46. BLUMEA DC. (1833) A. J. Randena in Blumea 10: 176-317 (1960); A.A. Anderberg in PI. Syst. & Evol.: 176: 105 (1991) & Willdenowia 25: 19-24 (1995). Annual or perennial herbs. Indumentum of 1-seriate, multiseriate and capitate glandular hairs. Stems simple or branched, usually erect, sometimes ascending, terete, gla­ brous. pubescent or tomentose. Leaves simple, alternate, linear-elliptic to obovate-lanceolate, margins serrate, dentate, denticulate or pinnatifid, sessile with rounded semi-clasping base or petiolate with tapering base, pubes­ cent to villous, apex acute-apiculate. Capitula hetero­ gamous, campanulate, disciform, pedunculate or sessile, in lax or compact terminal and axillary panicles or cory mbs. Receptacle epaleate, flat, convex, areolate or alveolate. Phyllaries 4-5-seriate, imbricate, increasing in length inwards, herbaceous, linear to linear-oblong, densely pubescent to villous. Florets purple, pink or vio­ let, numerous, outermost female, innermost bisexual or female. Corolla of female florets filiform. 2-4-lobed or

145. ASTERACEAE: 46. Blumea

bijabiate, glabrous; of bisexual florets tubular-funnelsh ped with the tube expanding from the base upwards, so] iclimes with sessile glands in upper parts, 5-lobed, lot es papillose. Stamens 5; anthers sagittate with tailed ba c connective triangular with obtuse apex. Styles ex erted, bifurcate. Cypselas brown, angular ribbed, pubt .cent. Pappus white, I-seriate, barbellate. A tropical and subtropical genus with about 100 spe­ cies, many occurring in SW Asia. A few species are native tc Africa, 3 species in the Flora area. In a note on the relationships between Blumea and 0 .her genera, Randena [in Blumea 10: 191 (1960)1 wrote t ial "Loggera and Blumeopsis perhaps have the closest a Fmities ...” with Blumea and also provided the different aJ characters. Wild (1969), in the Compositae of the 1 k>ra of Zambesiaca area, argued that Laggera can not be laintained as a distinct genus and thus reduced it to syn­ onymy under Blumea. Although Laggera and Blumea ;hare some characters the author here follows the tradilonal definition of these genera and keeps them apart. Blumea aurita (L. f.) DC., transferred to Laggera by v ilanderia (1960: 298) is now Pseudoconyza viscosa (Mill.) D'Arcy [in tribe Plucheeae (Benth.) A. Arvderb.]. The anther tails in Pseudoconyza viscosa (syn. B. aurita) io not reach up to the joint of the filament-collar. but are distinctly longer than in members of Laggera. However, anther tails with unequal lengths were observed in some specimens of P. viscosa. The female florets are bilabiate n P. viscosa, whereas in Laggera they are distinctly 4 t-5)-lobed. Recently, Andeiberg (in Willdenowia 25: 19-24, 1995) ’’resuscitated” the genus Doellia Sch. Bip. ex Waip. (1843) and included in it two of the species that were kept in Blumea by Randeria and others, i.e., B. bovei (DC.) Vatke and B. caffra (DC.) O. Hoffm. The major reason for this is the presence of “conspicuous resiniferous ducts ap­ pearing as longitudinal, red strips in the mature cypselas.” According to this and and also according to an earlier study, Doellia was placed in the tribe Plucheeae, close to the “Laggera clade”. Blumea appears to be a poly phy letic genus but monophyly. based on a single or a few characters and particu­ larly taking only a tiny segment of the genus, must not be employed for recognition of genera. In relating Blumea to other genera in Inuleae s.l., Randeria wrote “further divi­ sion of the Plucheinae into genera has been achieved on the basis of differences in the pappus and other individual characters of the capitula, florets and phyllaries. The same general trend for delimiting genera by single or very few characters seems to chracterize the entire Compositae and other families like Gramineae, Cruciferae and Umbelliferae." Randeria studied 49 of the species in the genus rather welL but left about 49 others as “taxa and names of uncer­ tain status”, an unusually large number of names to be un­ treated in a work that purports to be a revision of a genus. Anderberg (1991:105) reiterated this and in his latter work (Anderberg 1995:19), where he transferred two of

125

the species to Doellia, also wrote ’’more work remains to be done before the taxonomic situation in Blumea is un­ derstood in detail,...” The name Doellia is not typified, as is clearly stated by Anderberg (1995:20), but is being proposed for use with an amended description because of its association with Blumea abyssinica Sch. Bip. ex A. Rich. (-B. bovei). This may be sufficient to warrant the recognition of Doellia as a genus but due to the reservations expressed above, and also due to the large number of names that still need to be addressed in Blumea, including the widespread B. axillaris that occurs in the Flora area, the author prefers to maintain the three African species in Blumea until a more comprehensive treatment is obtained. 1.

2. -

Plant up to 75 cm liigh, leaves linear or ob­ lanceolate, rounded or clasping at base: capitula pedunculate, peduncle up to 4.5 cm long. 2 Plant up to 1.5 cm high, leaves obovate to elliptic, often narrowed or tapered to the sessile or petiolate base; capitula sessile or peduncle only up to 1 cm long. 3. B. axillaris Capitula 3-8 mm wide; corolla lobes of female flo­ rets glabrous. 1. B. bovei Capitula 8-15 mm wide; corolla lobes of female florets with 1-4 erect bristles. 2. B. caffra

1. B. bovei (DC.) Vatke (1875); Convza bovei DC. (1834) - type: Egypt. Sinai. Bove 118 (G-DC holo., K iso.). Blumea abyssinica Sch. Bip. ex A. Rich. (1848) type: EW. near Aguar [AgerJ. Medat [Modal) Valley, PI. abyss. II, Schimper 1006 (P syn.. K isosyn.) & 1032 (P syn.). Perennial herb, up to 50 cm liigh. with a woody rootstock, rarely annual with a long taproot. Stems solitary' or sev­ eral. each ascending from the apex of the rootstock, branched or unbranched, terete, light green to dark purple, sparsely pubescent. Leaves thick, linear-oblong to ob­ long-lanceolate, 15-80 x 2-8 nun, sparsely pubescent on both sides to glabrous, sessile-auriculate with rounded base, margins sharply denticulate, apex acute-apiculate. Capitula heterogamous, campanulate, 5-6 x 3-4 nun at anthesis, solitary or in open, branched corymbs, pedunculate, peduncle up to 4.5 cm long, bracteate, sparsely to densely pubescent. Receptacle flat, alveolate. Phyllaries c 4-seriate, purple-tinged, linear, densely pu­ bescent with both glandular and eglandular liairs; outer c 4 x 0.5 mm; inner c 5-6 x 0.5 mm. margins scarious. Florets purple. Corolla of female florets filiform, c 2.7-3 .1 mm long, 2-3-lobed; of bisexual florets 4.0-4.8 mm long, 5-lobed, lobes papillose and with stiff bristles, funnelshaped. Cypselas c 0.7 x 0.3 mm, sparsely pubescent, subtetragonal. Pappus white. 2.5-3 nun long, barbellate. Fig. 145.73. Wet montane grassland with seasonal streams; river banks; 1200-2200 m. EW TU GJ SU HA; Egypt, Sudan, Somalia; Yemen, Saudi Arabia, Oman, Palestine. Afganistan. Schweinfurth & Riva 1097; Schimper 111; Mesfin T. & Sebsebe D. 3764.

126

145. ASTERACEAE 46 Blumea

monomorphic, linear to linear-lanceolate with narrow scarious margins, densely pubescent on outside mainly along the gre> -green (dark purple when dry) median striae, 3-6 x 0.5-1 mm. apex long acuminate. Florets pur­ ple. Corolla of female florets filifonn, 3-4-lobed. c 3 mm long, glabrous: style c 4 mm long, corolla of bisexual flo­ rets tubular, funnel shaped above, c 3.5 mm long, 5-lobed. style c 4.5 mm long. Cypselas black, densely white pubes­ cent. c 1 x 0.5 nun. Pappus white, pubescent, 3-4 mm long Riverine scrub with Acacia, Taniarix, etc.; saline hab­ itats and hot springs; 1400-1600 m. SU; Kenya, Tanzania, Mozambique, Zimbabwe. Botswana. Namibia. South Af­ rica. and Madagascar. Friis et aJ. 713; Gilbert & Gilbert 3965.

F igure 73. BLUMEA BO VE I 1 upper stem portion. 2 female flo re tx 4 l' j ; 3 -b isex u al floret x4>/3:4 cvpselavvith pappus \ c 7 (No specimens cited in orig.) Drawn bv Ruth Koppel (R epro­ duced from FI Palaestma. vol 3. PI. 500.)

2 B. caffra (DC.) O. Hoffm. (1889): Conyza caffra DC. ( 1836) - type. S Africa. Cafra. Key River. Drege s.n. (G-DC hoio.. K fragment) Erect perennial herb, c 50-75 cm high; stems several from the woody rootstock, terete, glabrous to sparsely ciliate. dark -purplish or reddish-brown at base. Leaves sessile, yellowish-green, oblong to oblong-lanceolate alternate, diminished in size upwards. 3-6 x 0.5-1 cm. w ider at base, sparsely hairy on veins beneath. Capitula solitary at leaf axils or several in open corymbose cymes, pedunculate. 8-15 x 6-7 mm. peduncle up to 4 cm long, pubescent, bracteate Phyllaries multi-senate, graded

3 B. axillaris (Lam.) DC. (1836); Conyza axillaris Lam. (1786) - ty pe Maurice (Mauntjus). ('ommerson s.n. (P-LA holo.. P -JU 8615; P iso ). B. Iacera (Bunn, f.) DC. (1836); Conyza lacera Bunn. f. (1768) - type: Java, Burmann s.n. (G-DC. not seen). B. lacera forma elata robust a Vatke in Linnaea 39 485 (1875). nom. non rite publ.. quoad Schimper 810. Blumea drageanoides Sch. Bip. ex A Rich (1848) - types: TU/DG. near Ferrokoba. Schimper 633 (P syn., K isosyn.), GD, Wogera. Schimper 1297 (P syn., S isosyn ); TU. Chire. Ouartin-Dillon s.n. (P syn ); Choa. Petit s.n. (P syn., K isosyn ). B. lacera forma humilis fa ccid a Vatke in Linnaea 39: 485 (1875), nom. non nte publ.. quoad Schimper (1862) 804 & (1862) 182. B. lacera form elata robustior Vatke in Linnaea 39 485 (1875). nom. non rite publ.. quoad Schimper 810. B. lacera au c t. non (Bunn f.) DC. (1834). quoad Schim per 810. B. solidaginoides (Poir.) DC. (1836). B. hieracifolia auct non DC (1836). quoad Negri 519 (FT). B. lacera (Burnt f.) DC. var. glabrescens Cluov.. nom. nud.. quoad Watova 397 (FT). Annual or short-lived perennial herb, up to 1.5 m high, with a long taproot. Stem usually solitary. sometimes sev­ eral from the base, smaller and narrower titan the central one and ascending, striate, terete to angular often densely tomentose to velutinous with silky pale brownish hairs, sometimes also with glandular hairs. Leaves obovate to clliptic-obovate. up to 12 x 3.5 cm. progressively dimin­ ishing in size upwards, narrowed to the sessile base or shortly petiolate. petiole up to 3 cm long, tomentose to velutinous on the upper surface, tomentose to woolly on the lower surface, margins serrate, apex acute or obtuse. Capitula heterogamous. campanulate. 4-6 x 3-4 mm. ses­ sile to pedunculate, in axillary and tenninal dense pani­ cles: peduncle up to I cm long, densely tomentose or velutinous; liairs dark purple Receptacle flat or convex, alveolate. Pliyllariesc 4-seriate, linear densely velutinous, also with glandular hairs; outer 2-2.5 x 0.3-0.5 nun, pur-

127

145. ASTERACEAE: 46. Blumea. 47. Pegolcttia

Figure 74 BLUM EA AXILLARIS: 1 - part o f (lowering plant x 2 i. 2 capitulum. longitudinal section x 9: 3 4 phyllaries x 9: 5 outer tlorct (pappus removed) x 18, 6 inner flo­ ret (pappus removed) x 18; 7 sta­ mens x 18; 8 style o f inner floret x 18; 9 —cypsela x 36; 10 pappus bristles x 18. All from Gueho in MAIJ 15199. Drawn by Pat IIalliday (Reproduced from FI. Mascareignes . 109. C om posees. PI 19.)

pic unged; inner 5-6 x 0.3-0.5 inm. margins scarious. Floi els pale purple, whitish at base of corolla. Corolla of fen ale florets filiform, c 3.4 mm long, glabrous, those of bisL' ual florets funnel-shaped, c 4.6 nun long. 5-lobcd. lob s papillose. Cypselas light yellowish-brown, soinewh i angular, sparsely pubescent on the ribs, oblong, shij \ . c 8.0 x 0.3 mm. Pappus white. 3.0-3.5 mm long. Fig 145.74. Disturbed places in rocky hills with remnants of tall . libnia woodland, dense bushland of Carissa and Pteroloh uni, Acacia bushland. grassland with Acacia on allu­ via soil, stream or river banks, near waterfall, waste plalep; 850-2575 m. EW TU GD SU KF GG SD HA: widespread in Africa and SE Asia. Burger 3710; Gilbert 46961 Mesfin T. & Sebsebe I). 3773.

47. PEGOLETTIA Cass. (1825) Slmibs. subshrubs or herbs with erect or spreading brandies; old stems glabrous, smooth, brown, reddish, yellowish brown, or greyish; young branches green, stri­ ate. glanduJar-pubcsccnt. Leaves alternate, filiform, lan­ ceolate. oblong or spathulate in outline, simple to pinnatifid. glandular-pubescent. Capitula discoid, homogamous. solitary to loosely corymbosc. terminal. Phyllaries leathers. lanceolate to narrowly oblong or nar­ row In obovate. glandular margins ciliate. apex acute to acuminate. Receptacle epaleate. flat. Florets bisexual. yelloNv. turning purplish with age; corolla glandularliairy. 5-lobed. lobes narroNvly triangular, glandular base Nvidened into a ring-shaped foot: anther base sagittate, tails branched: style bifid at apex and swollen at base.

128

145. ASTERACEAE: 47. Pegolettia. 48 Pulicaria

Erect, annual herb. 8-60 cm high, stem sparsel) to densely branched, green, glandular-pubescent. Leaves simple, 5-40 x 1 10 mm, narrowly linear to lanceolate or spathulate, glandular-pubescent, sessile; margins serrate; apex obtuse with a small mucro. Capitula 5-10 x c 5 mm, in loose corymbs, rarely solitary. Phyllaries c 3-senate, spreading at anthesis. apex acute, inner longest Florets yellow or purplish (with age), 14-30. Corolla 6-8 x 1-1.5 mm. zygomorphic with one large lobe with bristly hairs at apex and the rest minute. Anthers c 2 mm long, base sagittate with long tails. Style 5-6 mm long, branches c 1.5 mm long. Cypselas 3 ^ .5 x 1-1.5 mm. thinly pubes­ cent, 12-16-ribbed, ribs with protrusions toward the apex. Pappus 1-seriate, with more or less equal plumose bristles and deeply incised scales externally. Fig. 145.75. Grassland with scattered Acacia. grassland on loamy soil, rocky hills with Acacia and Balanites. subdesert: 260-2000 in. EE SU GG SD HA; Senegal, N igena Chad. Algeria. Egypt. Sudan. Somalia Kenya. Tanzania, Zim­ babwe. Angola. Zambia. Mozambique. Botswana Swazi­ land. Namibia, S Africa, Cape Verde Island: Yemen. Saudi Arabia. Oman. Pakistan. India. Mesfin T. & O. Heclberg 1116; Gilbert et al. 230; Sue Edwards et al. 2763

48. PULICARIA Gaertn. (1791) E. (iamal-Eldin in Phanerogam. Monogr 14: 1-311 (1981).

Figure 75 PEGOLETTIA SE N EG ALEN SIS 1 habit x ? x 2 4 phyllaries (outer, middle and inner) x 3; 5 - cypsela with p a p p u sx 6 :6 outer pappus x 10:7 corolla with style branches x 6. 8 - stvle branches x 10; 9 stamen with tailed anthers x 10. (No specimens cited in orig.) Drawn by M.J Vesque & D Godot de Maurov. (Rearranged and reproduced from Flore de Mada­ gascar et des Comores. C om posees. Tome II. tig 110.)

Cypselas narrowly cylindrical, tapered to base. 12—20ribbed. ribs with knob-like protrusions toward apex, glan­ dular-pubescent. Pappus of one to several-seriate barbellate or plumose bristles and small or large, entire or in­ cised bristle-like scales. A genus with 9 species mainly occurring in S Africa: one species in the Flora area. P. scncgalensis Cass. (1825) - type: Senegal. Gay 86 (P holo.. K iso ). KuhniaarabicaDC. (1838)- type: Yemen. Gesscr Mts. Schimper 863 (G lecto.. FT K M NY iso ).

Perennial or rarely annual herbs, or shrubs; whole plant usually sparsely to densely hairy . Leaves simple, alter­ nate. auriculate or cordate, sessile, rarely petiolate. mar­ gins denticulate, entire or serrate. Capitula hetcroganious. radiate, hemispheric or rarely campanulate (in P. soma­ lensis). solitary or in paniculate cymes. Receptacle epaleate. flat or convex, arcolate. Phyllaries few- to many-seriate, sparsely to densely pubcsccnt and glandular to tomentose. outer usually herbaceous and shorter Flo­ rets yellow. Ray florets female, with short or long rays, glabrous, pubescent or glandular. Disc florets bisexual Corolla tubular 5-lobed. lobes acute. Anther base sagittate, tailed Style bifurcate, branches obtuse. Cypselas c 10-nbbed. cylindric to slightly angular, sparsely setose, light to dark brown Pappus diinorpliic and 2-seriate; outer cup-shaped or scale-like. 0 1-0 7 nun long: inner setaceous or nat and wider toward apex (in P. somalensis). 2.5-5 mm long, barbellate. or usually smooth toward tlic base. A genus with about 54 species in Europe. Asia Mid­ dle East. North, tropical and South Africa; 11 species in the Flora area. 1.

Leaves petiolate: petiole narrowly w inged, up to 1.5 cm long; blade ovate to lanceolate. 8 P. petiolaris - Leaves sessile w ith cordate or aunculate-amplexicaul base or narrowed from about middle to base, blade obovate-spathulate. oblanceolate. oblong or linear. 2 2. Outer pappus scale-like, all or most separate. 3

145. ASTERACEAE: 48. Pulicaria

-

Outer pappus cup-shaped with irregularly cut mar­ gins. 4

3.

Cypselas 0.7-1 mm long; leaf margin serrate and undulate, often folded or crumpled; inner pappus setaceous, uniformly wide. 10. P. undulata Cypselas 1.8-2 mm long; leaf margin entire or rarely with few lobular dentations, noi folded or crumpled; inner pappus scaie-Iike and wider to­ ward the apex. 9. P. somalensis

-

4

5

-

6

-

Involucre 15-22 mm wide; lower surface of ray floret pubescent at least half way, or sparsely glandular. 5 Involucre 8-12 mm wide; ray floret glabrous. sometimes tube sparsely glandular. 8 Phyllaries equal, densely woolly, linear to ob­ lanceolate. green and herbaceous; outer pappus c 0.7 mm long. 7. P. schimperi Phyllaries lengthening inwards, sparsely to densely pubescent with dense sessile glands, at least mar­ gins and apex scarious or chartaceous; outer pappus c 0.1—0.3 mm long. 6 Ray florets glabrous or sparsely covered w ith ses­ sile glands; inner phyllaries with acuteacuminate. hairy apex, c 3 mm long, cypselas c 1-1.2 nun long. 4. P. dysenterica Ray florets pubescent at least half way up on out­ side: inner phyllaries with scarious. white, brownish or silvery glabrous apex. 4.5-9 nun long: cypselas c 1.5 mm long. 7

7

Outer and middle phyllaries oblong or oblonglanceolate with obtuse to truncate, spinescent apex; inner phyllaries 7-9 mm long; ray florets 4-5 mm long. 6. P. incisa Outer and middle phyilaries linear or linearlanceolate with acutc-acuminatc apex, inner phyllaries 5-6 mm long; ray florets 6-7 nun long. 5. P. jaubcrtii

£

Ray florets densely pubescent at least half-way up on outside; phyllaries densely woolly, or sparsely appressed pilose and densely glandular toward apex or in middle, herbaceous. 9 Ray florets glabrous or sparsely sessile-glandular; phyllaries sparsely to densely pubescent with erect or spreading long liairs, at least margins scanous. 10

9

Phyllaries densely woolly, herbaceous; ray florets 8-8.5 nun long. 7. P. schimperi Phyllaries sparsely appressed-pilose but densely glandular toward the apex or in middle; ray flo­ rets 4-5 mm long 6. P. incisa Ray florets 1.5-2.5 nun long; inner phyllaries 3—3.5 nun long; leaves up to 5 mm wide. 2. Ray florets 4-7 nun long; inner phyllaries 4-6 nun long; leaves (2—)5—15 mm wide. 11

l ! Erect annual herbs; disc floret corolla 3.8-4.5 nun long. 12 Rliizomatous perennial herbs; disc floret corolla 2.6-3 8 mm long. 13

129

12. Phyllaries and peduncle villous (with long liairs); ray floret lamina glandular; corolla lobes setose. 1. P. arabica - Phyllaries densely capitate glandular with sparse silky hairs; ray floret lamina glabrous; corolla lobes glabrous. 11. P. attenuata 13. Capitula 4-5 x 10-12 nun; cypselas 1.3-1.5 nun long; leaf often oblanceolate. narrowed from middle to base. 3. P. scabra - Capitula 7-8 x 12-17 nun; cypselas 1-1.2 nun long, leaf oblong-lanceolate. 4. P. dysenterica

1. P. arabica (L.) ('ass. (1826); Inula arahica L. (1767) - type: "Habitat in Arabia. Forsskal; India.” Linnaean Herbarium No. 999.20 [LINN, lecto.. selected by Lack in Rechinger (ed). FI. Iranica 145: 115 (1980)j. Erect, annual herb, up to 50 cm liigh; stem branched, sparsely to densely pubescent, with erect or spreading long liairs. and glandular. Leaves oblanceolate or linearoblong (in upper parts). 10-50 x 2-10 nun. sessile, cordate-amplexicaul to sub-auriculate, sparsely to densely pilose and glandular on both surfaces, margins entire or undulate, apex acute. Capitula hemispheric, hcterogamous. radiate. 6-7 x 10-12 nun. solitary , termi­ nating branches, or few in open cymes: peduncle leafy, up to 4 cm long. Receptacle flat, epaleate. areolate. Phyllarics c 4-seriate. linear-lanceolate, sparsely to densely liairy and glandular; outer 3-4 x c 0.4 mm; inner 5-6 x 0.4-0.5 mm: apex acuminate and sharply pointed, curled. Ray florets yellow, 5-6 x c 1 nun. glabrous or rarely sessile-glandular on tube, apex 3-fid. Disc florets yellow, c 5.5 mm long. Corolla c 4.5 nun long, 5-lobed. lobes setaceous and glandular. Cypselas oblong-elliptic, brownish. 1-1.2 x c 0.4 111111. sparsely liairy and glandular at apex. Pappus 2-seriate. wliite; outer cup-sliaped. 0.2-0.3 nun long, margins incised; inner setaceous. 3-3.5 nun long, minutely barbellate on the upper half. Figs 145.76: 145.77.4 & 10. Gamal-Eldin (loc.cit.) cited “ 1839. Schimper (S)" and “Dec. 1874. Schimper (GOET)" under P. arabica for Ethiopia. These specimens have not been seen by the au­ thor. As the species is quite weedy in Sudan and Egypt, it is included liere. The above description is based 011 mate­ rial from N Africa and Turkey. 2. P. inuloides (Poir.) DC. (1836); Erigeron inuloides Poir. (1817) - type: Morocco. Mogador. Essaouna. Broussonet s.n. (FI holo.. P G-DC). Pulicaria longifolia Boiss. (1856); cf. Chiov. (1904:180). P. inuloides Perennial herb. 15-20 cm high; roots several, slender; stem terete, divaricately branched, sometimes pinkish or purplish, faintly striate, sparsely hairy and glandular. Leaves oblanceolate. linear-oblong to elliptic (in upper parts), up to 4 x 0.5 cm. light yellow-green, densely glan­ dular-punctate. sparsely pubescent 011 young branches

130

145. ASTERACEAE: 48. Pulicana

Figure 76 PULICARIA ARABICA 1 upper part o f stem; 2 - capitulum x 2V2; 3 base o f cypsela and cypsela with pappus x c 3; 4 enlarged top o f cypsela showing 2 types o f pappus x c 13. (No specimens cited in orig.) Drawn by Ruth Koppel. (Reproduced from FI. Palaestina , vol. 3, PI. 532.)

and developing leaves with denser hairs along veins, hairs white; leaf margins entire or denticulate near apex, apex acute or obtuse, inucronulate; sessile, cordate, semiamplexicaul. Capitula hemispheric, heterogamous. radi­ ate. 5-6 x 8-9 mm. in tenninal pedicellate cy mes: pedi­ cels up to 2 cm long, hairy with denser hair at base of the involucre. Receptacle epaleate. areolate. Phy Hanes c 4-seriate. linear with apiculatc apex, densely glandular and pubescent, liairs white, rather long; outer 2-3 x c 0.5 mm: inner often with curv ed or curled, pinkish or purplish apex. 3—3.5 x c 0.5 mm. Ray florets pale yellow, female. 1.5-2.5 mm long, glabrous. Disc florets yellow. 2.5-3 mm long. Corolla glabrous, c 2.5 mm long. 5-lobed. lobes densely papillose. Cypselas brown, c 10-ribbed. hairy, c 1.2 x 0.3 mm. shiny, cylindrical. Pappus 2-seriate. white: outer cup-shaped, c 0.2 nun long, inegularly incised; in­ ner setaceous. c 2.7 mm long, minutely barbellate toward the apex. Fig. 145.77.3. Reeky, stagnant pools; 2000-2745 m. ?EW TU SU. Morocco, Algeria. Tunisia. Libya. Niger, Cliad, Egypt. Somalia. Sinai. Saudi Arabia, Yemen. Oman. W. de Wilde 10892; Ash 1633.

3. P. sea h r a (Thunb.) Druce (1917); Erigeron scoher Thunb (1800) - type: South Afnca. Cape of Good Hope | ‘Cap. b Spei'|. 1774. Thunherg (’'UPS holo.. S). Erect, perennial herb, up to 50 cm high; roots several to numerous, slender from the rootstock; stem unbranched except in the inflorescence, terete, sparsely pubescent, striate. Leaves (in upper pans) oblanceolate to oblong. up to 8 x 1 cm. sparsely pubescent with long erect lairs denser on the veins, densely glandular sessile, nanowed from about middle to the base, margins denticulate in the upper half, apex acute. Capitula hemisphenc. hcterogamous. radiate 4-5 x 10-12 mm. pedunculate in termi­ nal cymes: peduncle up to 3.5 cm long, pubescent with long spreading hairs. Receptacle epaleate. flat, areolate Phyllaries 4-5-seriate. pale-green: outennost linear 2.5-3 5 x 0.3-0.5 pun. densely hairy on outside with ba­ sally multicellular and expanded hairs: middle and inner oblong-linear 4-5 x c0.5 mm. pubescent in the upper half Ray florets yellow, female, c 5.5 mm long, glabrous; apex 3-fid Disc florets yellow', c 4 nun long. Corolla c 2.6 nun long, glabrous. 5-lobed. lobes papillose. Cy pselas brown.

145. ASTERACEAE: 48. Pulicaria

131

Figure 77. PULICARIA spp. 1-9: leaves showing variations in size, shape or indu­ mentum; 10-14: phyllaries from outer to inner; 1 - P. SCABRA, 2 - P D YSEN­ TERICA, 3 - P . IN U LO ID E S ; 4 & 10 P. ARABIC A, 5 & 13 - P . INCISA, 6 & 12 - P. P E TIO LA R IS ; 7 & 14 - P. UN DU LA 7/1; 8 - P. SO M ALENSIS: 9 & 11 - P. JAU BERTII. Magnifications: 1 -6 ,9 x 2/ 3 ; 7 x c 3 ;8 x 1; 1 0 -1 4 x c 6. (No specimens cited in orig.) [Combined and reproduced from Revision der G altung Pulicaria (C om positae. Inuleae) fiir Afrika, M akaronesien und Arabien. P barter ogamarum M onographie 14, (1981) figs 1, 4 -6 , 9, 12, 13, 21 (as P. crispa), 34, 61, 65, 68, 69 and 76 (as P. crispa)].

shi iv, sparsely to densely pubescent. 1.3-1.5 x c 0.3 mm ob ;mg-elliptic. Pappus 2-seriate, whitish or yellowish; ou 0.1—0.3 mm long, cup-shaped, with lacerate marg i i i n n e r setaceous. 2.5-3.5 mm long, barbellate in the up ter half, barbs on lower half not clearly distinguished Fi* 145.77.1 In grass along stream margins; 2200-2400 m. EW SU; Su :an. Zambia. Zimbabwe, Namibia, Botswana, South Ai ica. Rvding & Kassa 2086; Pavlov & Petelin 105; IV. de Wilde 6056. 4. Vdysenterica (L.) Bernh. (1800); Inula dysenterica L. (1753) - type: "Habitat in Europae fossis subhumidis.” Linnaean Herbarium No. 999.14 [LINN, lecto., selected by Lack in Rechinger (ed ), FI. Iranica 145: 113 (1980)1. RJ i/.omatous, perennial herb, up to 75 cm liigh; stem te i. te, striate, sparsely to densely pilose, usually branched in upper parts. Leaves oblong-lanceolate, pubescent on bo h surfaces, glandular especially on lower surface. 2-6 x ( 5-1.5 cm, those on branches usually much smaller and

narrower, sessile to cordate-auriculate at base, margins entire or denticulate, apex acute. Capitula hemispheric, heterogamous, radiate, 7-8 x 12-17 mm, pedunculate, in teminal corymbs. Receptacle epaleate, areolate. Phyllaries 4-5-seriate, linear pubescent and glandular 3-5.5 x 0.2-0.4 mm. Ray florets yellow, female, 4-7 mm long, glabrous, rarely sessile glandular, apex 3-fid. Disc florets yellow, c 4 mm long. Corolla 5-lobed, 3-3.8 mm long. Cypselas oblong, c 10-ribbed, 1-1.2 x c 0.3 mm. sparsely pubescent especially in upper parts. Pappus 2-seriate; outer cup-shaped, c 0.2 mm long; inner setaceous, 2.5-3.5 mm long, minutely barbellate. Figs 145.77.2; 145.78. Gamal-Eldin (1981:125) recorded one specimen “Ab­ yssinia, 1862, Franqueville (K)” from Ethiopia. This specimen has not been located. It is perhaps a specimen collected by A. Petit. The species is widespread in Europe and N Africa, and has perhaps been introduced in the Flora area. The above description is drawn from speci­ mens of North Africa, Turkey and Iran where the species is widespread.

132

145. ASTERACEAE: 48. Pulicana

Cypselas brownish yellow, c 1.5 x 0.5 mm, c 10-ribbed, sparsely pubescent to glabrescent, oblong to obovate. Pappus 2-seriate, white; outermost cup-shaped, c 0.3 mm long, irregularly incised; inner setaceous, c 3.5 mm long, barbellate except near apex where sub-plumose. Fig. 145.77.9 & 11. Woodland; bank of Wabe Shebelle River, 305-1070 m. ?EW HA; Somalia, Socotra, Saudi Arabia, Qatar Ye­ men. Oman. Michel Corra s.n.; Ash 1153 (or 1158); Anderberg 1728. Gamal-Eldin (loc. cit.) cited a cultivated plant (Fiori 1805. FT) which is probably from Eritrea. This species differs from P. schimperi, with which it is usually confused, by its indumentum and the shape of es­ pecially the middle and inner phyllaries. The hairs in P. jaubertii are smaller, sparser and not densely matted (flexuous). The middle and inner phyllaries are whitish at apex and much wider than those of P. schimperi.

5 P. jaubertii Gamal-Eldin (1981) type: Arabia, Muscat, Aucher-Eloy 4739 (P holo.).

6. P. incisa (Lam ) DC. (1836); Inula incisa Lam. (1789) - type: Senegal, Roussilon (P-LAM.). P. undulata DC. var. abyssinica Chiov. in Ann. R. 1st. Bot. Roma 8 : 180 (1904) - type: Ethiopia, without locality, Schimper s.n. (FT syn.) [12 other syntypes are cited in the protologue]. P. suffrutescens Schweinf. (1892); P. incisa (Lam.) DC. subsp. suffrutescens (Schweinf.) GamalEldin in Phanerogam. Monogr. 14: 170 (1981) - type: EW, Sageneiti, Schweinfurth & Riva 1282 (FT iso ). P. incisa (Lam.) DC. subsp. denticulata GamalEldin in Phanerogam. Monogr. 14: 168 (1981) - type: Ethiopia [region not known], Schimper 673 (S holo., WU). P. aromatica R. Br. in Salt (1814), nom. nud. P. undulata auct., non (L.) C.A. Meyer (1831): Ohv. & Hiem (1877); Cufod. (1967). P. suffrutescens Schweinf. ex Penz. (1893), nom. nud.. quoad Schweinfurth & Riva 714.

Erect perennial herb, up to 40 cm high, with a long tap root and spreading branches, aromatic, whole plant smell­ ing of aniseed; stem sparsely to densely pubescent and or­ ange-glandular. striate, terete, profusely branched. Leaves oblanceolate, (only upper seen) up to 4.5 x 0.8 mm, softly pubescent with long flexuous hairs on both sides, densely covered with orange glands, sessile, auriculate: margins widely dentate or denticulate from middle to apex; apex acute. Capitula hemispheric, heterogamous. radiate, 6-8 x 15-22 mm. pedunculate; peduncle up to 8 cm long, densely pubescent. PhyLlaries c 4-seriate; outer oblanceolate-spathulate. 4.5-5.5 x 1-1.5 mm, densely hairy with dense orange glands: middle and inner linear-oblong, pu­ bescent mostly in the middle and toward apex, 5-6 x 0.6-0.8 mm. upper one-fourth shiny-white, glabrous; apex acute-acumiate. Receptacle epaleate, areolate. Ray florets orange yellow, female, 6-7 mm long; tube and outer surface of lamina pubescent. Disc florets yellow, c 5.5 mm long. Corolla c 4.5 mm long, glabrous, 5-lobed.

Erect or semi-erect, richly branched, aromatic, viscid an­ nual or short-lived perennial herb, 10-50 cm high, with a long tap root; stem densely pubescent with long some­ times intertwined white hairs, sessile-glandular stnate. Leaves oblanceolate or upper oblong to linear up to 5 x 2 cm, sessile-auriculate. densely sessile-glandular and pu­ bescent to tomentose with cottony hairs on young leaves and branches, margins irregularly serrate, apex acute or obtuse. Capitula hemisperic, heterogamous, radiate, 6-10 x 10-30 mm, pedunculate; peduncle up to 4.5 cm long, bracteate Receptacle epaleate, areolate. Phyllaries 4-5seriate; outer oblong to linear, hairy and densely glandu­ lar, 3.5-5 x 0.5-1.4 mm, margins ciliate, apex apiculate to spmose; middle series similar in pubescence, oblong with truncate, lacerate, scarious but spiny apex, or lanceolate with acuminate and scarious apex, 4.7-6 x 1-1.6 mm; in­ ner oblong-elliptic. 7-9 x 0.6-0.8 mm, sparsely glandular and pubescent along the green or herbaceous median outer surface, apex scarious, lacerate, or margins incised. Ray florets yellow. 4-5 x c 1 mm; female; lamina 3-fid at

Figure 78. PULICARIA D YSE N TE R IC A : 1 - part of upper stem; 2 - cypsela with pappus x 12. (No specimens citcd inorig.) Drawn by Ruth Koppel. (Reproduced from FI Palaestina . vol.

3. PI. 530).

145. ASTERACEAE: 48. Pulicaria

133

Gamal-Eldin (loc.cit.) recognized 3 subspecies within P mcisa but as some of the specimens were cited either ui ier two subspecies or under other species of Pulicaria, th : subdivision of P. incisa is not accepted here.

8. P. petiolaris Jaub. & Spach (1852) - type: Yemen, Botta s.n. (P). Vicoa divaricata Oliv. & Hiem (1877) - type: EE. Eilet, 1820-1826, Ehrenberg s.n. (K holo.). Kiliana chrysopsioides Sch. Bip. (1841), nom. nud. Pulicaria chrysopsioides Schweinf. (1867), nom. nud. ?Allagopappus chrysopsioides Schweinf. (1867), nom. nud. Pulicaria petiolaris Jaub. & Spach var. pentachaeta Hochst., nom. nud., quoad Schimper 2301 (K BM). P. petiolaris Jaub. & Spach var. pentachaeta Hochst. forma major, nom. nud., quoad Schimper 2301a (K).

7 P. schimperi DC. (1838) type: Arabia, near Dscheddam (=Jeddah), Schimper 908 (K iso.). P. rueppellii Sch. Bip. ex A. Rich. (1848) - type: EW, Modat, Schimper 1041 (K S iso ). P. orientalis Jaub. & Spach (1852) p.p.; P. undulata DC. forma orientalis (Jaub. & Sp.) Terracc. in Ann. R. 1st. Bot. Roma 5: 121 (1894); P. undulata DC. var orientalis (Jaub & Sp.) Chiov. in Ann. R. 1st. Bot. Roma 8: 179 (1904)-ty p e: Plate 343 in Jaub. & Spach.. III. PI. Orient. 4: 65 (1852). P. involucrata R. Br. (1814), nom. nud.; P. undulataDC. var. ehrenbergii Penz. inAtti Congresso Bot., Genoa 1892: 315, 346 (1893), nom. nud.; P. orientalis DC. var. ehrebergiana Engl. (1910), nom. nud.

Subshrub or bushy perennial herb. 0.3-1 in high; stem, leaves and branches glandular puberulous, sticky, shortly appressed pubescent; branches irregularly divaricate, terete and leafless below, striate, angular and leafy above. Leaves alternate, ovate to lanceolate, acute or subacute at apex, cuneate or truncate at base, narrowed into slightly winged petiole, membranous, serrate or dentate; blade up to 2.5 x 1.5 cm; petiole up to 1.5 cm long. Capitula subhemispheric, up to 7 x 5 mm at anthesis, solitary on long, slender, axillary and terminal pedicels or loosely corym­ bose, radiate. Receptacle flat, punctate. Phyllaries 3-4seriate, unequal, imbricate, thinly glandular-pubescent, linear or subulate with acute or acuminate apices. Florets yellow. Cypselas elliptic, contracted at the apex below the pappus, thinly pubescent, c 1 x 0.2 mm. Outer pappus short, connate at base; inner setaceous, barbellate. Fig. 145.77. 6 & 12.

ap :x. Disc florets yellow, c 6.7 mm long. Corolla 5-lobed, c 5 5 mm long. Cypselas brownish-white, oblong-elliptic to blanceolate, c 10-ribbed, pubescent, 1.2-1.5 x 0.4-0.5 mri. Pappus 2-seriate, white, outer fused, scale-like, cu '-shaped. dentate, c 0.5 mm long; inner setaceous, 15 -17, 2-4.5 mm long, baibellate. Fig. 145.77.5 & 13. Remnant Acacia bushland on rocky slopes; 135-2100 m EE EW TU; Sudan, Djibouti, Chad, Cameroun, Nige­ ria. Ghana, Niger, Egypt. Mesfin T. 8774; Bally 6645; Zorn 9(808).

Erect, bushy annual herb, pleasantly aromatic (with smell o:aniseed), 10-60 cm high, from a long tap root; stem p lie green, densely white woolly to densely glandular hispid pubescent. Leaves obovate-spathulate, (10-)20-50 x 2 0 ( —30) mm, pale blue-green, densely woolly on both surfaces to densely glandular-pubescent, sessile, auricled ard clasping the stem, or narrowed at base into a winged petiole, margins denticulate, apex acute. Capitula radiate, b ::nisphenc, heterogamous, 7-12 x 7-20 mm at anthesis, solitary, terminating branches; peduncle densely woolly, f tiyllaries 2-3-seriate, green and herbaceous, more or less equal, c 3.5-5(-13) x l-1.5(-2.7) mm, densely woolly and glandular linear to oblanceolate. Receptacle flat, e idleate. Ray floret orange-yellow, female, c 8.5 mm 1 mg, densely pubescent on outside; tube c 4 mm long. 1 iniina c 4-5 x 0.4 mm long, 3-fid at apex. Disc floret yel1 vw, c 6 mm long. Corolla c 4 mm long, 5-lobed, lobes f lpillose. Cypselas bright yellowish brown, shiny, c 2—1.4 x c 0.4 mm, sparsely hairy to glabrous, terete. I 3.ppus 2-seriate; outer cup-shaped, c 0.7 nun long, irreg­ ularly dentate; inner setaceous, white, 2.5-4 mm long, I arbellate Acacia-Commiphora bushland on limestone, low Acacia scrub, subdesert with volcanic ash, subdesert with l andy soil (Red Sea Hills); 50-1900 m. EE EW TU AF : U HA BA; Sudan, Djibouti, Somalia; Saudi Arabia, Yei ien, Oman. Gilbert et al. 7413; Mooney 8577; Sue Ed­ wards et al. 3792.

Open secondary bushland on steep lull slopes, usually between rock crevices; 80-2600 m. EE EW TU/GD; Egypt, Sudan, Djibouti, Somalia; Saudi Arabia Yemen. Rvding & Sileshi N. 1892; Tekle H. Hagos 152; Bally 6797. 9. P. somalensis O. Hoffm. (1906) - type: Somalia Dabad. 1900, Ellenbeck 162 (B syn. destroyed). Zeila, Bricchett 137 (RO syn ). subsp somalensis Shrub. 0.5-1.5 m high; stem woody, densely branched, grey and glabrous below, young branches sparsely to densely pubescent and pale green. Leaves grey-green, oblanceolate to linear (in upper parts), up to 4.5 x 0.5 cm. densely pubescent to tomentose on both surfaces and glandular, narrowed to the sessile base, margins entire or dentate, apex acute. Capitula campanulate, heterogamous, radiate (some discoid), 5-6 x 7-10 mm, pedunculate, in terminal open cymes; peduncle 1—2(—3) cm long, pubes­ cent and glandular. Receptacle epaleate, convex, areolate. Phyllaries pale green, c 4-seriate, linear to lanceolate, leathery, sparsely hairy but densely glandular; outer 2-2.5 x 0.6-0.7 mm; inner 4-5 x 0.6-0.7 mm. Ray florets yel­ low (sometimes absent), 5-6 x c 1 mm, female, glabrous. Disc florets yellow, 4.5-5 nun long. Corolla 4-4.5 mm long, 5-lobed, lobes glabrous to sparsely glandular. Cyp­ selas light brown. 1.8-2 x 0.7-0.8 mm, c 10-ribbed,

134

145. ASTERACEAE: 48. Pulicaria, 49. Iphiona

sparsely pubescent oblong. Pappus 2-seriate, white; outer of irregularly cut scales, 0.3-0.4 nun long; inner of flat setae, wider near apex, margins barbellate, 4-5 mm long. Fig. 145.77.8. Open bushland on stony soil; c 980 m. 7AF/HA; So­ malia, Djibouti, Saudi Arabia, Yemen. Seegeler 2695; Negri 1356. Subsp. schweinfurthii Gamal-Eldin (loc.cit.) occurs in Aden (Yemen). 10. P. undulata (L.) C.A. Mey. (1831); Inula undulata L. (i767) - type: “Habitat in Aegypto.” Linnaean Herbarium No. 999.15 [LINN, lecto., selected by Lack in Rechinger (ed.), FI. Iranica 145:120 (1980); review of usage by Hind & Boulos in Kew Bull. 57(2): 495 (2002)]. Aster crispus Forssk. (1775); Inula crispa (Forssk.) Pers. (1807); P. crispa (Forssk.) Benth. ex Oliv. (1873) - type: Egypt, Kahira [Cairo], at the pyra­ mids, 1762. Forsskdl in Hert>. Forssk. 990 (C lecto ). P. crispa (Forssk.) Benth. ex Oliv. var. macrocephala Mattei in Lanza & Mattei in Boll. R. Ort. Bot. Palermo 8:49 (1910) - type: EW/EE, Filfil, Senni 356 (PAL holo.). P. scabra auct., non (Thunb.) Druce (1917): Cufodontis (1967: 1121), quoad Burger 1823. Bushy annual or short-lived perennial heit> or subshmb, aromatic, viscid, 0.5—1 m high; stem and branches white tomentose. sometimes glabrescent, terete, striate, leafy. Leaves obovate-oblong and narrowed below the middle to linear or lanceolate (in upper branches), up to 5 x 2 cm sessile, auriculate and clasping stem, upper surface sparsely to densely glandular thinly floccose, linear lower surface sparsely to densely tomentose, with dense sessile glands; margins serrate and undulate, often folded or crumpled, apex acute-apiculate. Capitula radiate, hemi­ spheric, heterogamous, 5-7 x 8-12 mm, in dense paniculate corymbs. Receptacle flat, epaleate. Phyllaries 4-5-seriate, continuous with bracts on peduncle, linear with sharp erect or slightly curved apex, sparsely pubes­ cent but densely glandular margins ciliate, 2.5-4 x c 2 mm, increasing in length inwards, greenish yellow. Ray florets bright yellow, female, c 5.5 mm long; tube c 2 mm long, glabrous or sparsely hairy; lamina c 3.5 x 0.3 mm, 3-fid at apex. Disc florets yellow, c 4.2 mm long. Corolla c 3 mm long, 3-lobed. Cypselas grey-brown, glabrous or sparsely hairy, c 0.7 x 0.3 mm, shiny, oblanceolateoblong. Pappus 2-seriate. white; outer squamulose. c 0.4 mm long, margins fimbriate: inner setaceous, c 3.8^t nun long, barbellate to sub-plumose toward the apex. Figs 145.77.7 & 14; 145.79. 1. Branches sparsely leafy; leaves markedly curly. subsp. undulata Branches densely leafy; leaves not curly. subsp. tomentosa subsp. undulata Leaves linear-oblong or narrowly oblanceolate, only up to 6 mm wide.

Open short grassland on red clay overlying limestone, Acacia-Commiphora bushland, Acacia-Terminalia wooded grassland on limestone; 1200-2350 m. EW SD HA; North Africa from Morocco to Egypt, Sudan, Soma­ lia; Sinai, Saudi Arabia, Yemen, Oman. Thulin et al. 3471; Friis et al. 3021; Mesfin T. & Sebsebe D. 4037. subsp. tomentosa (Gamal-Eldin) Hind & Boulos in Kew Bull. 57: 497 (2002); P. crispa (Forssk.) Benth. ex Oliv. subsp. tomen­ tosa Gamal-Edlin in Phanerogam. Monogr. 14: 274 (1981) - type. EW, Keren, Bogos, along the Alseba River, Fiori 1792 (FT holo.). Leaves spathulate, lower ones often 10-15 mm w ide, densely white tomentose on both surfaces or on the lower surface, especially in young and upper ones. Open short grassland, at margins of fields cleared for cultivation, Tecceze valley, sometimes weed in town; 700-2350 m. EW TU SD HA; Sudan, Somalia G. Aw eke & Gilbert 936; Mesfin T. 8758; Ryding 1137. 11. P. attenuata Hutch. & Burtt (1933) type: Sudan, Darfur, Jebel Meidob, Michelmore s.n. (K holo.). Erect, annual herb, 15-30 cm high Stem branched at base and in upper parts, covered with flexuous hairs, striate, terete. Leaves spathulate-oblanceolate, (2—>4—5.5 x 0.5—1 cm. attenuate at base into a narrowly winged petiole, grey-green, covered with white flexuous hairs, denser on lower surface, median vein prominent, margins entire to shallowly serrate toward the acute apex. Capitula broadly campanulate, 5-6 x 10-12 mm, solitary, terminating branches, peduncle densely glandular with sparse flexu­ ous hairs, 3-12 cm long. Receptacle epaleate, convex. Phyllaries 3-seriate, linear, outer greenish, 4-4.5 x 0.3-0.5 mm, densely sessile-glandular with sparse whit­ ish silky hairs, inner 3.5-4 x 0.4 mm, thickened and con­ vex in middle, yellowish white with a median green band, sparsely sessile-glandular. Ray florets yellow, female, 4-4.5 x c0.4 mm, glabrous, apex 2-3-fid. Disc florets yel­ low, c 5.5 mm long. Corolla c 3.5 mm long, glabrous, 5-lobed. Style c 4.5 mm long. Cypselas oblong-elliptic, light yellowish brown, 1-1.2 x c 0.4 mm, sparsely setose. Pappus 2-seriate, white; outer cup-shaped, c 0.1 mm long, margins incised, inner setaceous, 4-4.5 mm long, mi­ nutely barbellate on the upper half. Habitat not known; 1000-1300 m. EW; Sudan, Soma­ lia. Jones 103 (K). The collection from the Flora area is from Eritrea, 'N. Sahil Awraja, close to the border with Sudan’. The species may occur at lower elevations in the rest of Ethiopia as it is quite common in Somalia.

49. IPHIONA Cass. (1817) A. Anderberg in Nordic J. Bot. 5: 169-194 (1985); Mesfm Tadesse in Kew Bull. 50. 401-408 (1995).

145. ASTERACEAE: 48. Pulicaria, 49. Iphiona

135

Figure 79. PULICARIA UNDULATA. 1 - habit; 2 - upper stem portion; 3 cypsela with pappus x 9; 4 - cypsela with outer pappus x 18. (No specimens cited in orig.) Drawn by Ruth Koppel. (Reproduced from F/. Palaestina, vol. 3, PI. 534, as P. crispa .)

Br nched, perennial herbs or subshrubs, up to 1 m high; sten terete or angled, striate, old stems light-brown to gn v ish, smooth or cracked, young ones green, smooth. gl< :>rous or glandular-pubescent. Leaves alternate, simple to pinnatifid, linear to lanceolate, sometimes fleshy or sp ny. Capitula radiate or discoid, solitary, or several arrat 2ed in loose corymbs. Phyllaries several-seriate, lan­ ce late, elliptic to broadly ovate, leathery, spreading or en ct at anthesis, yellowish, usually with a dark median sti ation, sessile-glandular near the acute to cuspidate ap Receptacle epaleate. foveolate, flat. Ray florets pr sent or absent. 1-seriate. neuter, yellow, linear to obio ig, 3-5-veined, glabrous to glandular-pubescent; apex ec.ire or 3-fid. Disc-florets bisexual. Corolla yellow, 5- nbed, lobes acute, glandular or glabrous; anther base

sagittate; style bifid, branches obtuse at apex and with few papillae. Cypselas narrowly cylindrical to elliptic, 10-costate, glandular-pubescent. Pappus of one- to sev­ eral-seriate barbellate unequal bristles with an outer row of free or basally connate scales. A genus with 12 species mostly confined to the middle and Near East and Arabia; 1 species in the Flora area. L scabra DC. (1834) -type: Egypt, Tor, Bove 104 (G-DC holo., K iso.). Subshrub with erect to spreading branches, up to 60 cm high; old stems yellowish white to grey, smooth to rough, glabrous to glandular-pubescent; young stems green, striate, glandular to glandular-pubescent. Leaves

136

145. ASTERACEAE: 49. Iphiona, 50. Iphionopsis

simple to pinnatifid, basally with filiform, soft or stiff lobes, 10-50 x 1-7 mm; surface flat to longitudinally folded along the mid-rib, glandular to glandularpubescent; apex spiny and pungent. Capitula solitary or 3-10 in loose corymbs, discoid, 5-12 x c 6 mm. Phyllaries oblong to triangular, glabrous or glandular, spreading at anthesis; margins hyaline; apex apiculate to cuspidate. Ray florets absent. Disc florets 8-30. Corolla 5-9 x 0.5-2.5 mm. Anthers 2.5-3.5 mm long. Style 2.5-7 mm long, branch apices obtuse. Cypselas elliptic, terete to slightly flattened, 1-3 x 0.5—1.5 mm. sparsely to densely pubescent, sometimes with few glands at apex. Pappus of several-seriate, unequal, barbellate. and spreading bristles. Fig. 145.80. Stony and rocky deserts, sandy areas, sea level-50 m. EE; Israel, Jordan, mainly in areas near the Red Sea in Egypt, Sudan, Saudi Arabia Yemen, extending to Oman Defiers 1163; Russel s.n.

Receptacle epaleate. Phyllaries (3-)4(-5)-seriate, contin­ uous with bracts on peduncle, outer loosely imbricate, mid-vein purplish brown throughout or from near the base to the apex, linear to narrowly ovate-lanceolate, 15-2 5 x c 1 mm, glabrous to pubescent at the base or in middle, in­ ner conduplicate. 5-6 x c 1 mm, linear-lanceolate, gla­ brous to sparsely pubescent in upper half. Florets 10-15. white, pink, purple or reddish, rarely yellowish. Corolla glabrous, c 8 x 1.2 mm, 5-lobed, lobes 0.5-0.8 mm long Anthers c 3.5 mm long, appendage c 0.5 mm long, base sagittate, narrow, c 0.3 mm long, acute. Style 10-11 mm long, branches c 2.5 mm long, papillose at base. Cypselas light brown, 2 8-3.2 x c 0.5 nun, oblong, 4-angled, with ascending white bristles and sessile glands. Pappus 2-seriate. barbellate, brownish white, outer c 1.5—2 mm long, inner c 7-8 mm long. Fig. 145.81. Rocky bushland with Acacia and Caesalpinia. Commiphora bushland, Terminalia-Cordia woodland; 300-700 m. SD HA; Somalia, Djibouti, Kenya (Kl) Gilbert 2078; Simmons 541; Gilbert et al. 8105.

50. IPHIONOPSIS A. Anderb. (1985) A Anderberg in Nordic J. Bot. 5: 51-56 (1985) Shrubs with spreading to erect-patent branches. Leaves alternate, simple and leathery', round to spathulate or lan­ ceolate, glabrous to sparsely haiiy, shortly petiolate; mar­ gins denticulate to dentate; apex acute or obtuse. Capitula discoid, 10-15 mm long, corymbose. Phyllaries sev­ eral-seriate, leathery, yellowish with green to brown striations, subcordate. ovate to narrowly oblong or lan­ ceolate, glabrous or glandular. Receptacle epaleate. flat. Florets 8-15, bisexual, white to reddish or yellowish. 6-9 mm long. Corolla 5-lobed. lobes obtuse. Anthers c 3 mm long, base sagittate with long tails. Style 7-10 mm long, branches c 2.5 mm long, papillose in upper half or throughout. Cypselas terete, smooth, obconical to ellip­ soid, 1-2 x 0.5-1 mm. densely hairy and often also glan­ dular. Pappus of several-seriate, equal, baibellate bristles. A genus with two species, one of which occurs in Ethiopia, Somalia and Kenya and the other one in Mada­ gascar. I. rotundifolia (Oliv. & Hiem) A. Anderb. (1985); Iphiona rotundifolia Oliv. & Hiem (1877) - type: Somalia without further locality, Playfair s.n. (K holo.). Shrub or shrubby herb with brittle stems and branches, often growing in shelter of other bushes, 0.5-2.5 m high, aromatic, glaucous; old stems light brown to greyish, smooth, glabrous to bristly; young stems light brown, gla­ brous to sparsely hairy; hairs simple. Leaves alternate, pale-greea elliptic, oblanceolate, spathulate to rounded, 10-50 x 5-25 mm, surfaces with short scattered bristly hairs, or glabrous, base narowed into a distinct, up to 5 mm long petiole; margins sharply denticulate with sharp glandular teeth; apex obtuse. Involucre narrowly cyathiform. Capitula campanulate, c 20-flowered, 8-10 x 4-6 mm, up to 30 arranged in corymbose cymes terminating branches; peduncle sparsely pubescent, up to 10 mm long.

Tribe 7. P L U C H E E A E (Benth )A . Anderb. (1989) Inuleae Plucheinae Benth. (1873). A Anderberg in Plant. Syst. Evol. 176: 145-177 (1989). Perennial herbs or shrubs, rarely annual herbs. Leaves simple to pinnatifid. alternate, often decurrent, sparsely to densely pubescent, sometimes scabrid. Indumentum of 1-seriate or, basally multiseriate, erect, patent or rarely flexuous hairs and sessile glands. Capitula heterogamous. disciform, campanulate to hemispheric, solitary to vari­ ously corymbose or glomemlate. Receptacle flat convex, globose, ovate or linear-elliptic, epaleate or paleate and hairy (in Pseudoblepharispermum). Phyllaries cartilagi­ nous. rarely herbaceous, 3-10-seriate. Female florets on outside, in many series, filiform and often narrowed to­ ward the apex; corolla bilabiate or 2-4-lobed; style exserted, bifurcate or rarely undivided, and apically notched. Bisexual florets often functionally male, corolla tubular or tubular in lower parts and funnel-shaped above, 5-lobed; anthers and style branches shortly exserted. Antherbase sagittate and tailed or lobed. rarely truncate, con­ nective triangular Style bifurcate; stigmatic area apically confluent; outer surfaces of branches, and also upper part of style below the point of bifurcation, with obtuse sweep­ ing hairs Cypselas ellipsoid, turbinate or triquetrous, gla­ brous or hairy Pappus of barbellate bristles, rarely absent. Anderberg (1989) included about 22 genera in this tribe and also provided a few more that are difficult to classify; 6 genera are recorded from the Flora area. Doellias which was recognized by Anderberg (in IVilldenowia 25: 19-24 (1995)] on the basis of the elongate resiniferous ducts on the mature cypselae. is not included here (see notes under Blumea). The monoty ptic Litogyne Han' has not been found in the Flora area but as it is

145. ASTERACEAE: 49. Iphiona

137

Figure 80. IPHIONA SCABRA. 1 part o f plant from A den x 14; 2 - stamen x 10; 3 - style x 10; 4 - part o f pi ant from Oman x Vi; 5 - floret x 7>/3; 6 - part of plant from Sinai x 14; 7 & 8 - phyllaries, outer left, inner right, x 5. 1 from Grier­ son 100; 2, 3, 5 -7 from Neergaard s.n.; 4 & 8 from Bornmiiller. Drawn by A. Anderberg. (M odified and reproduced from Nordic J. Bot. 5(2), 1985, p. 188, fig. 14, with perm ission from the author/ artist.)

known from both Sudan and Somalia, it might as well turn p here. Hence, it is included in the key and a full descripi«n is provided of the one species that is quite commonly ound in NE Africa. Receptacle paleate, paleae hairy. 55. Pseudoblepharisperm um Receptacle without paleae. 2 2.

Capitula crowded into globose, ovoid, or cylindri­ cal heads, cypselas without pappus, corolla swol­ len and suberized (sclercnchymatous) below. 56. Sphacranthus Capitula solitary or, if crowded, not aggregated into heads; cypselas usually with 1-seriate pappus, corolla uniform or, if slightly wider be­ low, not suberized, parenchymatous. 3

3.

Pappus absent in female florets, absent or up to 3 smooth in bisexual florets; style undivided or mi­ nutely notched at apex. 57. Litogyne Pappus of 1-seriate. barbellate. usually 2-15 bris­ tles; style bifurcate at apex. 4 4. Pappus 2-5 from an annular corona, rarely absent, subshrubs or shrubs with prostrate or trailing and mat-fonning branches and stems. 51. Delamerea - Pappus 10-17; erect perennial or rarely annual herbs, or shrubs with erect branches. 5 5. Annual or short-lived perennial herbs; outer phyllaries herbaceous. 6 - Shrubs, subshrubs or perennial heitos, rarely annu­ als; outer phyllaries entirely, or at least in lower parts, leathery; corolla of female florets 2-4 (-5)-lobed. 7

138

145. ASTERACEAE: 50. Iphionopsis, 51. Delamcrea

Figure 81.

IPH IO NO PSIS ROTUND I FOLIA 1 portion o f plant x Vi; 2 - floret x c 8: 3 stamen x 9'/2; 4 - style x 9'/2; 5 & 6 phyllaries, short ones outer, long ones in­ ner, x 4. 1-3, 6 from Ironside Wood S /73/165; 4 from Glover & Gilliland 387; 5 from Gillen 13202. Drawn by A. Anderberg. (Reproduced from Nordic J. Bot. 5, 1985, p. 53. fig. 2, with perm is­ sion from the author/artist.)

6.

-

7.

Stem often winged, wings continuous or inter­ rupted; leaves decurrent; corolla of female florets bilabiate or 2-4(-5)-lobed. 53. Laggera Stem not winged, leaves slightly amplexicaul: co­ rolla of female florets with or without minute lig­ ules. 52 Pseudoconyza Anther base truncate or shortly sagittate with equal or unequal, up to 0.1 mm long, lobes. 53. Laggera Anther base sagittate with the Jobes/tails reaching filament-collar joint. 54. Pluchea

51. DELAMEREA S. Moore (1900) Shrub or sub-shrub with trailing or prostrate branches, villous. Leaves small, alternate, sharply serrate. Capitula small, campanulate, solitary , terminal on branch apices, hcterogamous. disciform. Receptacle flat, epaleate. Phyllaries few-seriate, imbricate, increasing in length in­ wards. outer firmly membranous, inner narrower and lon­ ger. Florets numerous, outer female, inner bisexual, nearly all fertile. Corolla of female florets filiform. 4-5-lobed at apex with equal or sub-equal lobes; of bisex­ ual tubular-campanulate. 5-lobed; style glabrous, more or less equal to corolla in length. Anther base shortly caudate, tail fused, apex lanceolate. Style of bisexual flo­ rets filiform, papillose, bifurcate with linear apicallv rounded branches. Cypselas short, compressed; those of female florets glabrous, those of bisexual florets setose. Pappus of 2-5 setae, soon falling.

A monotypic genus restricted to drylands in Southern Ethiopia and Northern Kenya. S. Moore [in Journ. Bot. 38: 457 (1900)] placed this genus in the ‘group of genera including Epaltes Litogvne) and Denekia’ and pointed to its affinity with Denekia.

D. procumbens S. Moore (1900) 1898, type. Kenya, near Lake Marsabit. Delamere s.n. (BM holo.). Prostrate shrub with long trailing stems, and clumps mea­ suring up to 6 m in diameter, some branches erect; stem striate, sparsely pubescent with soft hairs. Leaves oblanceolate to spathulate. 5—10(—25) x 3—6(—15) mm, covered with dense white soft liairs. sessile or narrowed into the short (up to 8 nun) petiole, margins toothed or blade appearing 3-(5-8)-lobed, apex obtuse. Capitula campanulate. hcterogamous. 6-8.5 x 4-5 mm. sessile or sliortly pedunculate, solitary and terminating branches or axillary, usually subtended by one or a few narrowly oblanceolate leaves. Phyllaries 4-5-seriate. leathery, liairy at margins or near the apex: outer and middle 2-4 x 1-2 m m ovate; inner 5-7 x 1—1.5 mm. ovate-lanceolate, sometimes tinged pmk. Florets lavender, c 58-64; 39^13 female, c 6 nun long; 18-21 bisexual, c 9.5 mm long. Co­ rolla of female florets 3 .5-4 mm long, c 0.7 mm wide at base; style c 5 mm long, ovary epappose or rarely with 2 bristles. Corolla of bisexual florets 5-5.5 mm long, c 1.5 mm wide near lobes; stamen c 7.5 mm long, anthers c 3 mm long; style c 7.5 mm long, papillose in upper parts, branches c 0.7 mm long, rounded at apex. Cy pselas light

145. ASTERACEAE: 51. Delamerea, 52. Pseudoconyza

139

Figure 82.

DELAMEREA PROCUMBENS 1 part o f flowering plant; 2 - capitulum; 3 - outer female floret; 4 - upper parts of style; 5 - bisexual floret; 6 anthers. No m agnifications given and no specimens cited in original publication. Drawn by H. Morgan. (Reproduced with perm is­ sion from Jounx. Bot., vol. 38, 1900. Tab. 416.)

trow n (only immature seen), 1.2-1.5 mm long, angular v :th ascending white bristles. Pappus white. (0-)2-5 from an annular corona, c 4 mm long, barbellate towards apex. Fig 145. 82. Seasonally flooded clay flats besides standing pools, in bare areas in open bushland; c 40() m; GG (near Lake 1 uikana and Chew Bahir); Kenya (K1 and 2). Carr 780. 844; Corradi 2034, 1853 (FT); Vatova 1309 (FT).

52. PSEUDOCONYZA** C.uatrec. (1961) uinual or short-lived perennial herbs; stem with resin caials. Leaves simple, alternate, margins dentate to basally tiallowly lobed. slightly amplexicaul. Capitula discirm, heterogamous, terminal, solitary or few together, jceptacle epaleate, convex in fruit, marked with remains of cypsela scars. Phyllaries several-seriate, graded monomorphic with the innennost the longest separate. Outer orets female, filifonn. ligules minute or absent; sty lei ranches conspicuously exserted; central florets bisexual, orolla tubular 4-5-lobed; anthers caudate-auriculate or lagittate at base, connective oblong, obtuse; stylci ranches linear, glandular-papillose outside. Cypselas ci­ rsoid or fusifonn, silky-strigose. Pappus of free, yellow­ ish-white. barbellate bristles. A monotypic genus found in Central America, Africa, md Asia The single species in tliis genus was long re­

ferred to Conyza in the Astereae. The nanower and more puberulent styles, the caudate-auriculate or sagittate an­ ther base, the fewer pappus-bristles. the elliptic or fusi­ fonn cypselas and the graded-monomorphic phyllaries suggested that the species is out of place in Conyza and probably also out of place in the Astereae. Some authors, e.g., D'Arcy. although he originally made the combina­ tion in Pseudoconyza [in Phytologia 25: 281 (1973)j later included it [in D ’Arcy et al.. Ann. Miss. Bot. Card. 62; 1034 (1975)] in Blumea, following Badillo in Revta Fac. Agron. (Maracay) 7: 9 (1974)], while others, e.g.. Merxmiilleret al. (in Hey wood et al. p. 578, (1977)] considered this doubtful and recognized Pseudoconyza Cuatrec. un­ der the Inuleae. Since then this view lias been accepted by many sy nantlicrologists. although it has been placed in the Plucheeae rather than the Inuleae, e.g., Anderberg {in Bremer. Asteraceae - ( 'Iad istics die Classification'. 302. (1994)]; M c V a u g h [in Flora Novo Galiciana 12: 781 (1984)]. The same taxon was addressed by G. Nessom Jin Sida 10: 30-32 (1983)] wherein he discussed its evolu­ tionary origin, but within Blumea. See also discussion un­ der Blumea.

P. viscosa (Mill.) I) A rev (1973); Conyza viscosa Mill. (1768); Blumea viscosa (Mill.) Badillo (1974) - type: Mexico, Houston s.n. (BM holo.). Conyza aurita L. f. (1781); Laggera aurita (L. f.) C. B. Cl. (1876); Blumea aurita (L. f.) DC. (1834) -

140

145. ASTERACEAE: 52. Pseudoconyza. 53. Laggera

type: a cultivated plant grown from seeds obtained from India (not traced). Conyza lyrata H.B.K. (1820); Pseudoconyza lyrata (H.B.K.) Cuatrec. (1961) - type: Ecuador, Guayaquil, Humboldt & Bonpland s.n. (P holo.). Blumea pappi Gand. (1918) - type: EW, Debre Nehsii, 9 Sept. 1905. Pappi 6401 (FT holo.). Erect, annual or short-lived perennial herb, 0.5-1 m high, with a taproot. Stem richly branched from the base or above, sometimes densely stipitate-glandular and usually with silky, eglandular hyaline or whitish hairs. Leaves al­ ternate, obovate to obovate-oblong, pilose and glandular on both surfaces, margins coarsely and sharply dentate or serrate, 2-5(-9) x 1—2(—3.5) cm, lower leaves subpetiolate, usually with 1-4 pairs of narrow spreading lobes just above the base, or decurrent on the stem below the base of the blade, apex acute, upper leaves sometimes rounded at the sessile base, amplexicaul. Capitula solitary or 3-10 together, often arranged in subcorymbose pani­ cles which are one-third to one-half the height of the plant, peduncle 5-15 nun long. Involucre cup-shaped, greenish white, truncate at base, 7-8 x 6-7 mm, graded-monomorphic. about 4-seriate. Outer phyllaries lanceolate, acute, green or with green or purple tips, mostly hairy and glandular on the outer surface, 2-5 x 0.3-0.5 mm; inner phyllaries sub-equal, subscarious, pale or sometimes pur­ plish. linear-subulate, 5-7.5 x 0.4-0.7 mm, tapering to acute pilose tips. Florets pale lilac, violet or purple. Fe­ male florets numerous; corollas 4-4.8 mm long, filiform, the rays minute or none; style-branches filiform, conspic­ uously exserted. bisexual florets 10-15. Corollas 5.2-5.6 mm long, with slender cylindrical throat shorter than the filiform tube. 4-5-lobed; anthers pale brown, 1.5-1.8 mm long, caudate-auriculate or sagittate at base; style 5.5-6.5 mm long, branches linear glandular-papillose outside. Cypselas fusiform or elliptic, dark brown, c 0.8 x 0.4 m m silky-strigose. Pappus bristles 8-10, yellowish white. 2.5-5 mm long, baibellate. Fig. 145. 83. Rare in grassland and open fields; 580-900(-1600) m. EW EE TU SU IL GG SD; widespread at lower altitudes in tropical and North America, Arabia. India, tropical Asia. Ash 750; W. de Wilde 10616; Mesfin T. 5855.

53. LAGGERA Sch. Bip. ex Benth. & Hook, f (1873) Perennial herbs or subshrubs, rarely annual herbs; stem simple or profusely branched, often winged, wings con­ tinuous or interrupted. Leaves simple, alternate, oblong, lanceolate or elliptic, sessile or decurrent, surface scabrid to densely tomentose, hairs both simple, 1-seriate and ses­ sile-glandular, margins senate to denticulate. Capitula campanulate to hemispheric, heterogamous, disciform, outer female, usually numerous, inner bisexual, fewer, ar­ ranged in axillary and/or terminal panicles. Receptacle epaleate. flat or convex. Corolla of female florets filiform, bilabiate or 2-4(-5)-lobed. lobes more or less equal. Co­ rolla of bisexual florets tubular expanded from about 2/3 of the way up, 5-lobed. Anther base truncate or sagittate, lobes often unequal and c 0.1 mm long, filament collar c

0.3-0.5 mm long. Style of bisexual florets bifurcate at apex with branches c 0.5-0.6 mm long, papillose with papillae extending to 1/3 of the way down. Cypselas light brown, c 5-ribbed, thinly pilose along the ribs, narrowly oblong-elliptic, base annularly thickened. Pappus white, 1-seriate, faintly barbellate. A genus of about 17 species confined to the Old World tropics. About 6 species occur in the Flora area. The dis­ tinction between Blumea and Laggera has not been satis­ factorily worked out. Some species have constantly been moved between the two genera e.g., Laggera decurrens (Vahl) Hepper& Wood [in A'eiv Bull. 38: 83-86 (1983)], Blumea decurrens (Vahl) Merxm. [in Merxmiiller & Rossler. Mitt. Bot. Staatssaml. Mtinchen 20: 1-9 (1984)], Blumea gariepina DC., Laggera gariepina (DC.) Randeria [in Blumea 10: 298 (I960)], etc. L. decurrens is quite common in Somalia and it might occur in the south­ eastern comer of Ethiopia. Hence it is included in the key For distinction from Blumea, see note under that genus in the Inuleae. 1

Branch or stem intemodes with interrupted, tooth­ like wings. 4 L. crispata - Branch or stem intemodes with continuous narrow or broad wings, sometimes wings absent and leaves only shortly and narrowly decurrent. 2 2. Capitula large, c 14-20 mm wide. 3 Capitula small, c 5-10 mm wide. 4 3. Capitula sessile or shortly pedunculate, in terminal, simple or slightly branched leafy spikes or racemes; leaves thinly scabrid-pubescent. 5 Capitula on up to 2 cm long peduncles, 3-4 termi­ nating axillary' branches, these arranged in termi­ nal, elongate panicles; leaves thinly floccose above, softly tomentose below with denser hair on the reticulate venation. 6. L. elatior 4 Bushy perennial herbs or subshrubs; leaves grey or ashy-green, silky or woolly to floccosetornentose especially beneath; outer and middle phyllaries tomentose with or without sessile glands. 5 - Erect perennial with few branches near the apex, not bushy, leaves light to bright-green, usually scabnd pubescent, rarely tomentose-subscabrous; outer and middle phyllaries puberulous to scabnd-pilose. usually with numerous sessile glands 6 5. Florets pink or purple; leaves floccose-tornentose. often 5-10 x 1—1.5 cm; capitula c 12-15 mm long 1. L. tomentosa Florets yellow; leaves silky villous, up to 3 x 0.5 cm; capitula c 5-7 mm long. L. decurrens 6.

Capitula c 8-10 nun wide; phyllaries continuous with bracts on peduncle, outermost usually with green, herbaceous, ovate-lanceolate appendages, densely scabnd on both surfaces. 3. L. alata Capitula c 5-6 mm wide; phyllaries not continuous on peduncle, glandular-puberulous or glabrescent, often acute, rarely herbaceous and slightly wider. 2. L. crassifolia

141

145. ASTERACEAE: 52. Pseudoconyza, 53. Laggera

Figure 83.

PSEUDOCONYZA VISCOSA . 1 - part o f flowering plant x 14; 2 - stamens x 9; 3 - cypsela with pappus bristles x 9. All from D 'Arcy 4097. Drawn by J. L. W il­ son. (Reproduced with permission from Ann. Miss. Bot. Gard. 62, 1975, p. 1036, fig- 40.)

1. L. tomentosa (Sch. Bip. ex A. Rich.) Oliv. & Hiem . 1877); Blumea tomentosa Sch. Bip. ex A. Rich. (1848) :ype: TU, Gennia. Mensach district, Schimper 236 (P holo., K S iso.). B. petitiana A. Rich. (1848); Laggera petitiana (A. Rich.) Schweinf. & Asch. (1867) - type: SU, ‘Choa’, Petit s.n. (P holo.). Bushy, perennial herb or subshrub, c 0.5-1.2 m high, aron ;ttic; stem sub-terete, narrowly winged, wings continuo is. c 1—1.5 mm wide, densely tomentose. Leaves grey or a hy green, linear-oblong, lanceolate or oblong-elliptic, 3 10 x 0.5-1.5 cm, floccose-tomentose especially ben :ath, decurrent, margins denticulate to sub-entire, apex a ute or subacute. Capitula broadly campanulate, hetero2 iimous, 12-15 x 8-10 mm, solitary or in few-headed cymes terminating the axillaiy branches, in some forming loose panicles; peduncle up to 3 cm long, densely toment >se. Receptacle epaleate, flat. Phyllaries 6-7-seriate; c uter and median 3-5 x 0.7-1 mm, sparsely to densely tor rentose with sessile glands, lanceolate, spreading or rec urved at tip, often bristle-pointed, inner c 10 x 1 mm, pu­

bescent at middle with sessile glands, purple at apex, mar­ gins scarious, linear-lanceolate; innermost sparsely liairy near apex, c 10 x 0.5 mm, linear. Florets pink or purple, outer female, c 9 mm long, inner bisexual, c 12.5 mm long. Corolla of female florets c 7.5 mm long, c 4-5-lobed, of bisexual florets 10 mm long, 5-lobed, lobes papillose. Anthers c 2.2 mm long. Style of bisexual florets c 10.5 mm long, branches c 0.8 mm long. Cypselas light brown, c 1.4 x 0.4 mm, c 5-ribbed, pubescent on ribs, ob­ long-elliptic. Pappus white, 1-seriate, barbellate, 5—5.5 mm long. Dry hill and mountain slopes; 2345-2950 m. TU GD GJ WU SU HA; not known elsewhere. Mooney 7847; Mesfin T. 3696; Pichi-Sermolli 2628. This is a well-known and frequently cultivated medic­ inal plant; it is also used as a fumigant and in cleansing milk containers. 2. L. crassifolia (Sch. Bip. ex A. Rich.) Oliv. & Hiern (1877); Blumea crassifolia Sch. Bip. ex A. Rich. (1848) types: TU, ‘inter Adoa et Gennia’, Schimper 939 (K

142

145. ASTERACEAE: 53. Laggera

lobed, of bisexual florets c 7.6 mm long, 5-lobed. Anthers c 2.5 mm long. Style c 8.7 mm long, bifurcate. Cy pselas brown, c 5-ribbed. ribs pubescent, 1.2-1.5 x c 0.3 mm. Pappus white, c 5 -6 mm long. Stony dry valley in Acacia woodland; 1500-2000 in. EW TU GJ SU SD HA; Sudaa Rep. Dem. du Congo. Schimper 939. 1996; Wellby s.n. (Awash & Meki rivers); Pichi-Sermolli 924 3 L. alata (£>. Don) Oliv. (1873); Erigeron alatum D. Don (1825); Conyza alata (D. Don.) Roxb. (1814, nom. nud.; 1832); Blumea alata (D. Don.) DC (1836) - type: Nepal, Narainhetty, Hamilton s.n. (BM holo.). Blumea alata (D Don) DC. var. natalensis DC. in Prodr. 5: 448 (1836); Laggera alata (D. Don) Oliv. var. natalensis (DC.) Chiov. in Ann. R. 1st. Bot. Roma 8: 173 (1904) - type: S Africa, Natal, Gariepina, Drege 4996 (G-DC). Laggera involucrata Schweinf., nom. nud., quoad Schweinfurth & Riva, 761 and 767. L. appendiculata auct., non Robyns (1943): Cufodontis (1967: 1095), quoad Burger 3410.

Figure 84. LAGGER4 A L A T A : Part o f flowering plant x c I (No specimen cited in orig.) Drawn by Bhola Ram. (Reproduced with permission from FI India . vol. 13, fig. 46.)

WU syn.); TU. Gennia. Memsach, Schimper 154 (K S syn.); TU. Quartin-Dillon s .a (?P syn.). Laggera (Blumea?) crassifolia Sch. Bip.. nom. nud.. quoad Schimper 154 and 1996. Pluchea (Laggera) crassifolia Sch. Bip., nom nud.. quoad Schimper 939. Perennial herb or subshrub, 0.5-1 m high; stems solitary or several from the base, often branched, terete, striate, upper narrowly winged and glandular-pubescent, wings entire, c 1 mm wide. Leaves narrowly elliptic, oblong or oblanceolate. narrowed into the decurrent base, glandular and scabrid-pubescent. 5-12 x 1-3.2 cm margins denti­ culate, apex acute or obtuse. Capitula campanulate, heterogamous, 8-10 x 5-6 nun; pedunculate, in oblong or pyramidal terminal panicles; peduncle up to 1 cm long. Receptacle epaleate, c 3.5-3.7 mm wide, convex, smooth/ areolate. Phy llanes 6-7-senate; outer c 2-3 x 1 nun. puberulous and glandular lanccolate. coriaceous below, appendaged. appendage green, herbaceous and slightly wider, middle 6-7 x c 1 mm. puberulous. coriaceous, in­ ner 7-9 x c 0.5 mm. linear, glabrous, scarious. Florets pur­ ple, outer female, c 7.3 nun long, inner bisexual, c 9.5 mm long. Corolla of female florets c 5.7 mm long, 4(—5)-

Perennial herb, 0 7-1.5 m high, rarely annual herbs; stem branched, scabrid to densely glandular-pubescent, terete, winged, wings up to 4 mm wide, continuous and entire. Leaves oblong-lanceolate to narrowly elliptic, 3-8 x 1-2 cm (upper), sessile ;ind clasping stem (in branches) or nar­ rowed into the decurrent base, scabrous to tomentosesubscabrous on one or both surfaces, margins denticulate to broadly dentate-denticulate, apex acute. Capitula broadly campanulate, heterogamous, 10-12 x 8-10 mm. nodding, in wide pyramidal terminal panicles, peduncle up to 3 cm long Receptacle epaleate. flat. Phyllaries 8-9-senate; outer 3-4 x 1.5-2 mm, continuous with bracts on peduncle, spathulate or ovate-lanceolate, with green, herbaceous upper part and leathery lower half, scabrid-pubescent, densely glandular, middle lanceolate, 4-5 x c 1 mm, leathery, sparsely pubescent in middle; in­ ner linear-lanceolate. 7-9 x c 1 mm. leathery, often pur­ plish at apex. Florets pink to red-violet, purple-mauve. Fe­ male florets c 7 5 mm long, corolla c 6.1 mm long, 4(-5)-lobed, bisexual florets c 9.7 mm long, corolla c 8 mm long, 5-lobed. lobes papillose. Anthers c 2.6 mm long. Style c 8.9 mm long, bifurcate, branches c 0.9 mm long. Cypselas light-brown, c 1.4 x 3 mm. c 5-ribbed, pu­ bescent on ribs, oblong. Pappus white, 5-6 imn long, barbellate. Fig. 145.84. Open woodland with Combreium, Syzygium and Piliostigma on red sandy clay, Albizia-Acacia wooded grass­ land, Carissa-Maytenus evergreen scrub; 1400-2700 in. EW SU KJF GG SD BA HA. Mooney 8429; Friis et al. 1564; Mesfin T. 5658. This species is currently excluded from Flora ofTropi­ cal East Africa (cf. FTEA, Compositae. 2) noting that the type has "recurved phyllaries with a hairy adaxial surface, similar to L. brevipes, and this taxon does not occur in East Africa ...” Much of the material from the region, which was previously identified as L. alata. is now kept in

145. ASTERACEAE: 53. Laggera

143

L nspata. In the field L. alata and L. crispata are distinct ar d can easily be distinguished. The material from the F ira area may represent a distinct entity but until the genus is revised in whole, the present author prefers to mainta n it as L. alata.

4 L. crispata [Vahl) Hepper & Wood (1983); Conyza crispata Vahl (1790); Blumea crispata (Vahl) Merxm. (1984) - type: Yemen. Hadie. 1763. Forsskal in Herb. Forssk. 1140 (C holo., microfiche 32: I. 5,6). Blumea pterodonta DC. in Wight (1834); Laggera pterodonta (DC.) Oliv. (1873) - type: India, Wallich 3039b (K holo.). Blumeapurpurascens Sch. Bip. ex A. Rich. (1848) -type: TU, near Adwa,Schimper 153 (Pholo., K iso ). Serratula polygyria A. Rich. (1848) - type: Ethio­ pia [region not known], Aikhe, plate 62 in A. Rich.. Tent. FI. Abyss., Atlas. (1850). Laggera (Blumea) purpurascens Sch. Bip.. nom. nud.. quoad Schimper 153. E ect annual or short-lived perennial herb, often 1—2(—3) m iigh. aromatic; stem branched from middle upwards ai d hence pyramidal in outline, sticky' owing to glandular h; s, terete, striate, narrowly winged with teeth or unin­ terrupted wings. Hairs both simple and gland-tipped. L ; ves elliptic to lanceolate (upper), membranous, thinly h; ry especially on veins on underside, sessile or nar­ rowed into the decurrent base, margins dentate or dt ticulate. apex acute; upper stem leaves 5-7 x 1.5-2 c i i middle branch leaves up to 17 x 7.5 cm; lower cauline lea es up to 30 x 12 cm, sometimes shortly petiolate. C ipitula broadly campanulate, heterogamous, 10-12 x 8 10 mm, in densely branched, tenninal. pyramidal pani­ cle . peduncles up to 4 cm long, densely glandularpnrescent, bracteate. Receptacle epaleate. areolate. Phy llaries purplish-green, c 10-12-seriate, unequal; outer aad middle series linear-lanceolate, 3-6 x 0.5-0.8 mm, d. sely glandular-pubescent, spreading or recurved; inru 9-10 x 0.7-0.9 mm, scarious, glabrous, linear with at i :e-acuminate apex. Florets pale lilac or purple, outer feiuale, c 7.7 mm long, numerous, inner bisexual, c 8.8 mm long, few. Corolla of female florets filiform, c 6.1 mm loiiiZ, 2-lobed. of bisexual florets c 8.8 mm long, tubular. e: iranded from middle to apex, 5-lobed, lobes sparsely p; rillose; ovary papillose. Anthers obtuse at base, c 2-5 mm long, connective oblong-lanceolate. Style of bisexual florets c 9.J mm long, bifurcate, papillose in upper tliird. b; i iches c0.6 nun long. Cypselas brown, oblong-elliptic, nbbed, sparsely pubescent. 11-12 x 0.2-0.3 nun. Pappus w m e. 1-seriate. 5-8.5 mm long. Fig. 145. 85. Margins of montane scrub or grassland, river or st ream banks, also weed of cultivation and waste places, et (600-) 1600-2650 in. EW TU GD SU AR WG IL KF G SD BA HA; widespread in tropical and South Africa and in tropical Asia. Mooney 5240; W. de Wilde 9238. 8593; Friis et al. 1448. The low altitudinal record is from Gambella (IL).

Figure 85. L/XGGER/X CRISPATA. Part o f flowering plant x c Vi. (No specimen cited in orig.) Drawn by Bhola Ram. (Repro­ duced with permission from FI. India , vol. 13. fig. 47.)

5. L. braunii Vatke (1875) -type: GD, Dembea. Woin Arek, 1900 m, 9. Nov. 1863, Schimper 1348 (B holo., destroyed, K iso.). Erect, annual herb, c 50 cm high; stem terete, sparsely pu­ bescent, winged, wings up to 5 mm wide and with entire margins. Leaves elliptic to oblong-lanceolate (upper), thinly scabrid-pubescent. narrowed into the decurrent base. 10-12 x 3-5 cm; margins serrate or serrulate, apex acute. Capitula hemispheric, heterogamous. 13-15 x 15-18 mm, subsessile or shortly pedunculate in a terminal simple or slightly branched leafy spike or raceme. Reccptacle epaleate, flat, areolate. Phyllaries multiseriate. linear acuminate and purple at apex, puberulous. 4-14 x 0.5-1 mm; middle series with recurved apex. Florets ?purplc. outer female, c 7.8 mm long, inner bisexual c 9.7 mm long. Corolla of female florets c 6 mm long. 2-lobed. of bisexual florets c 7.3 nun long. 5-lobed. lobes papillose. Cypselas brown (only immature seen), c 1.7 x 0.3 mm. sparsely pubescent, ribbed. Papppus white. 1-seriate. 5-6 nun long. Open Annona senegalensis, Syzygium, Nauclea and Grewia woodland, Oxvtenanthera abyssinica thickets; 1300-1800 m. TU GD GJ SU WG; Sudan (Jcbel Marra). Friis et al. 7865 & 923 1; Schimper 1348.

144

145. ASTERACEAE: 53. Laggera, 54. Pluchea

6. L. elatior R.E. Fries (1928); Blumea elatior (R.E. Fries) Lisowski (1989) type: Kenya, Mt. Kenya, Forest station, 2300 m, R.. E. & Th. C. E. Fries 425 (UPS holo., BR K iso.).

On the basis of a specimen, Senni 773/bis from ‘D olo\ southern Bale. Cufodontis (1967:1096) cited P. arguta Boiss. for the Flora area but this species is restricted to Pakistan and NW India.

Erect perennial herb, up to c 1.5 m high; stem winged, terete, wings c 4 mm wide, continuous, tomentose. Leaves (upper) lanceolate, upper surface floccose, lower softly tomentose with denser hairs on the reticulate veins and on the mid-rib, hairs brownish, broadly decurrent margins senate or denticulate with sharp teeth, apex acute. Capitula hemispheric, heterogamous. nodding, 12-15 x 14-20 mm at anthesis. 3-4 terminating axillary branches, arranged in terminal, elongate panicles; peduncle up to 2 cm long. Receptacle epaleate. flat. Phyllaries multiseriate, linear to linear-lanceolate with narrower terminal, re­ flexed, or variously curled appendages, margins ciliate; outer and middle series 4-7 x c 1 mm, sparsely to densely pubescent with sessile glands throughout or on upper half; inner linear c 10 x 1 mm, sparsely pubescent and glandu­ lar, apex acuminate, curled or reflexed, purple; innermost glabrous, with acuminate apex. Florets ‘red’ or purple, outer female, c 7.4 mm long, inner bisexual, c 11.6 mm long. Corolla of female florets c 6.7 mm long, 4(-5)lobed, of bisexual florets c 10 mm long, 5-lobed, lobes glabrous or sparsely papillose. Anthers c 2.5 mm long. Style c 10 mm long, bifurcate, branches c 0.6 mm long. Cypselas brown, c 1.4 x 0.3 mm, c 5-ribbed, pubescent on ribs, narrowly oblong-elliptic. Pappus 1-seriate, white, c 5 mm long, barbellate.

1. Leaves decurrent, wings on intemodes confluent toothed to deeply pectinate. 2 - Leaves not decurrent usually auriculate at base and sometimes also slightly extending onto the intemode. 4

Margins of evergreen bushland with Grtidia, Brucea, Discopodium, Hagenia, Kiaesa, etc.; 2550 m. KF; Sudan (Imatong Mts ), Kenya R6p. Dem. du Congo, Burundi. Friis et al. 1508.

54. PLUCHEA Cass. (1817) S. Kong-Jones in Englera, 23 (2001; 2001a - electronic supplement). Shrubs or perennial herbs. Leaves alternate, entire, denticulate, serrate or pinnatifid, puberulous to densely woolly, sometimes glandular-viscid, sessile or subsessile, decurrent or not. Capitula heterogamous, campanulate or hemispherical, disciform shortly pedunculate, solitary or a few in clusters terminating branches, or in corymbose cymes. Receptacle epaleate, sometimes honey-combed, concave or convex. Phyllaries 3-10-seriate, imbricate; outer ovate or ovate-lanceolate, usually firm or leathery, appressed, inner lanceolate. Florets white, pink or ma­ genta; outer female, numerous; inner bisexual or sterile. Corolla of female florets filiform, 4-5-lobed, style branches filiform, of bisexual and sterile florets 5-lobed, tubular-campanulate. Anthers tailed at base. Style bifur­ cate, branches papillose-glandular or puberulent. Cyp­ selas cylindric to 4-7-angled, glabrous or pubescent. Pappus of white, 1-seriate, scabrid bristles. A genus with about 80 species common in the warm tropics, 6 species are currently known from the Flora area.

2. Plant up to 30 cm high, leaves 3-5 mm wide 6. P. nogalensis - Plant usually 1-1.5 m high; leaves 10—25(—35) mm wide. 3 3. Leaves pectinate, both surfaces with dense c 1 mm long tuberculate-based hairs and stalked glands; capitula c 8-9 mm long; florets dark-magenta. 4. P. somaliensis - Leaves serrate or serrate-denticulate, glabrous to appressed pubescent, hairs c 0.3 mm long; floreis usually white, rarely pink. 2. P. ovalis 4. Phyllaries with distinctly reflexed tips. 5. P. kelleri - Phyllaries appressed. without reflexed tips. 5 5. Capitula 8-10 mm high at anthesis; leaves woolly tomentose, 10-25(-40) x 5—10(—20) mm. 3 P . sordida - Capitula 3-7 mm high at anthesis; leaves glabrous to sparsely pubescent, 25-50 x 10-35 mm. 1. P. dioscoridis

1. P. dioscoridis (L.) DC. (1836); Baccharis dioscoridis L. (1755); C.onyza dioscoridis (L.) Desf. (1815)- type: “Habitat in Syria Aegypto. D Hasselquist.” Hasselquist s.n in Linnaean Herbarium No. 992.6 [LINN, lecto., UPS isolecto. selected by King-Jones in Willdenowia 29: 211 (1999)| C.onyza baccharoides Sch. Bip. ex A. Rich. (1848); C. dioscoridis (L.) Desf. var. baccharoides (Sch. Bip. ex A. Rich.) Schweinf. & Asch. in. Beitr. FI. Aethiop. 144 (1867); Pluchea dioscoridis (L.) DC. var. baccharoides (Sch. Bip. ex A. Rich.) Cufod. in N. Giorn. Bot Ital., n.s. 50: 105 (1943) - type: TU, near Adwa PI abyss. I, Schimper 146 (P holo., K iso.). Blumea baccharoides Sch. Bip. ex A. Rich. (1848); Conyza modatensis Sch. Bip. ex Schweinf. & Asch. (loc. cit.) - type: EW, near Adegama Modat, PL abyss. II, Schimper 1018 (P holo., K iso ). Pluchea dioscoridis (L.) DC. var. pseudovalis Cufod. in N. Giorn. Bot. Ital., n.s. 50: 105 (1943) type: GG/SD, Corradi 1791,1792 and 1797 (FT syn ); GG/SD, El Dire, Corradi 2025 and 2026 (FT syn.). Shrub, or erect, richly branched perennial herb, 0.75-2 m high; branches striate, terete, pubescent often viscidglandular, glabrescent with age. Leaves ovate-oblong, narrowly elliptic or linear-lanceolate, up to 8.5 x 2.5 cm, sessile or subsessile. with cuneate or auriculate-cordate and sometimes slightly decurrent base, glandular puberulent or pubescent on both sides, rarely glabrous; margins entire to serrulate; apex acute or sub-acute.

145. ASTERACEAE: 54. Pluchea

145

Figure 86 PLUCHEA D IOSCORIDIS 1 - part o f leafy stem; 2 - upper part o f stem with inflorescence; 3 - cypsela with pappus x 8; 4 - cypsela without pappus x 16. (No specimens cited in orig.) Drawn by Ruth Koppel. (Reproduced from FI. Palaestina, vol. 3, PI. 501.)

C ipitula campanulate or ovoid, 4-5 x 3-4 mm, few to s veral in much-branched, paniculate, corymbose cymes: p . iuncle and inflorescence branches densely glandularp ibescent or pubescent. Receptacle narrow. epaJeate. Fhvllaries c 4-5-seriate; outer 1.5-2 x c 1 mm, ovate, s iarsely glandular-pubescent to glabrous, pubescent and f ilose at the outer margin, apex acute; inner c 3.5-4 mm 1 ng. progressively narrower and slightly longer, f mbriate at margins near the acuminate apex, otherwise glabrous, sometimes purple-tinged. Corolla of female flor is c 4 mm long, of bisexual florets c 5 mm long. Anthers c 2.5 mm long. Cypselas brown, c 0.7 nun long. - '-ribbed, ribs whitish, glabrous to sparsely pubescent. ; ppus of 13-15. 1-seriate, white, scabridulous bristles, c nin long, persistent. Fig. 145.86. In moist places bordering lakes, reservoirs, permanent pools, etc.. margins of rivers, streams; 1000-2400 m. EW 1 GD WU SU KF GG HA; Egypt. Sudan. Somalia. E rica, Zambia, Zimbabwe. Mozambique to South Africa

(Natal. Transvaal); widespread from the middle East (Syria. Palestine-Jordan), through S Arabia. Yemen. Mesfin T. 1942; Gilbert 1539; Robertson 1404. King-Jones (2001) cited two specimens for Ethiopia [Corradi 1949 and Pachva 270) under Pluchea bojeri (DC.) Humbert, a species first described from Madagas­ car and now reported to occur as far north as the extreme southern tip of Ethiopia. Pluchea dioscoridis and P. bojeri are similar in gross morphology, including capitular fea­ tures. The major feature used in separating them by King-Jones (2001) was the shape of the capitulum, i.e., campanulate in the former and cylindrical or nearly cylin­ drical in the latter. Re-examination of capitula in P. dioscoridis indicated that they could also be cylindrical particularly before anthesis. Until more material from the extreme southern tip of Ethiopia is examined. P. bojeri is excluded from this account. P. bojeri may also be a syn­ onym of P. dioscoridis. The isotypc of P. bojeri ai Kew (Bojer s.n.) lias all capitula in bud.

146

145. ASTERACEAE: 54. Pluchea

2. P. ovalis (Pers.) DC. (1836); Baccharis ovalis Pers. (1807); Pluchea dioscoridis (L.) DC. var. ovalis (Pers.) Cufod. wArrn. Nat. Hist. Mus. Wien 56: 161 (1948) - type: Senegambia, Sene­ gal River, Rousillon 20 (P holo.). Shrub, c 1-4 m high; upper stem and branches usually pubescent to tomentose, rarely glabrous or glabrescent, terete, striate, densely leafy. Leaves elliptic or ovate, 2.5^L5(-8.5) x l-2(-2.5) cm, pale green to ashy-green, pubescent to tomentose, hairs short (c 0.3 mm), narrowed towards the sessile auriculate-clasping base, decurrent in unequally dentate wings, margins denticulate-serrate; apex obtuse or subacute. Capitula heterogamous, cam­ panulate or hemispheric, 5-6 x 3.5-4 mm, on short un­ equal pedicels in dense terminal paniculate corymbose cymes Phyllaries 5-6-seriate, leathery, firm, puberulous. with sessile glandular hairs, apex acute; outer ovate to ovate-lanceolate, 1.5-2.5 x 1 mm; inner c 4-4.5 mm. Flo­ rets white or pink. Cypselas glabrous or with few short hairs on the angles, brown, c 0.6 mm long. Pappus c 13-65, white, persistent, barbellate. Fig. 145. 87. Mixed scrub thicket along lake margins, flood plain, riverine scrub; 400-1830 m. EE GJ SU GG SD BA HA; Cape Verde, Morocco, Egypt, Sudan, Somalia, E Africa, Rwanda, Burundi, Rep. Dem. du Congo. Togo, Senegal; Saudi Arabia, Bahrain. Oman. Mesfin T. et al. 4095; Carr 689; Mooney 7290. 3. P. sordida (Vatke) Oliv. & Hiern (1877); Laggera sordida Vatke (1875) - type: Tanzania, Zanzibar Island, Hildebrandt 1022 (B holo., de­ stroyed). Bushy, perennial herb, c 0.3-1 m high; upper stem and branches densely hispid. Leaves ovate-elliptic to nar­ rowly obovate, tomentose on both sides, narrowed to the sessile base, 10-25(-40) x 5-10(-20) mm; margins entire to serrate, apex obtuse or acute. Capitula campanulate. 8-10 mm high, in branched globose cymes; peduncle up to 12 mm long. Receptacle concave; naked when cypsela fully mature. Phyllaries 5-6-seriate. lanceolate, outer c 3 x 1 mm, densely pubescent; inner c 7-8 mm long, pubes­ cent along the middle striae, purple, inner surface silvery and shiny when dry. Cypselas c 1 x 0.3 mm, cylindric. pu­ bescent. Pappus white, 3 3-3-8 mm long, barbellate. In seasonal marsh fringing lake; 1300-1500 m. SU GG (only known from around lakes Langano and Chamo [=Lake Maigharita]); Kenya, Tanzania. Ash 3335; Mesfin T. 3033; Vatova 1446. 4. P. somaliensis (Thell.) Thulin (2004); Blumea somaliensis Thell. (1923)-type: Ethiopia. “Webi Suabeli” [= Wabi Shebelle], 1891/92, Keller 9 (Z lecto ). P. sennii Chiov., nom. nud.; Pluchea frutescens Benth. var. parvifolia Chiov., in Flora Somala 2: 261 (1932)-type: Ethiopia [SD/BA], Somalia. “Dolo". 17 Sept. 1929, Senni 773 bis (FT holo.) P. pectinata F.G.Davies & J. P. Lebrun (1981) — type: HA, Ogaden. Mustar hill. Ellis 347 (K holo ).

P. arguta auct., non Boiss. (1856): Cufodontis (1967), quoad Senni 773 bis. Laggera helichrysoides Chiov., nom. nud., quoad Vatova 2 (FT) Profusely branched shrub, 1-3.5 m high, aromatic, branches and upper stem thinly pilose, striate, terete. Leaves obovate in outline, up to 6 x 3 cm, yellowtshgreen, serrate above, pinnatilobed in lower parts, densely to sparsely pilose and sessile-glandular, sessile ind decurrent with deeply divided or laciniate wings on stem; hairs c 1 mm long. Capitula campanulate, 7-8 x 4-5 mm. in loose cymes terminating branches; peduncle up to 10 mm long. Phyllaries 5-6-seriate; outer ovate, 2-2.5 x c 1 mm, with green appendage, sparsely pubescent with both glandular and eglandular hairs; middle series lanceolate, c 4 x 1 mm, sparsely pubescent with both glandular and eglandular liairs. margins ciliate; inner oblanceolate to ob­ long. c 6.5 x 0.8 mm. margins pectinate. Florets purple, fe­ male c 4 mm long; bisexual c 5 mm long, corolla 3-lobed in female, 5-lobed in bisexual florets. Cypselas brown, c 0.8 x 0.3 mm, c 7-ribbed, sparsely pilose, narrowly obovate. Pappus white, c 4.5 mm long, c 10-13, 1-seriate. Fig. 145.88. Amongst dry riverine forest, stream bed; salty mud bordering Wabe-Shebelle river, 200-500 m. SU (middle Awash) SD HA; Somalia. IECAMA BH-45; Gilbert et al. 7510;v4s/j 1160. P. somaliensis was compared with P. arguta Boiss. wluch it resembles in vegetative morphology, e.g., leaf dissection However, the later is distinct in having differ­ ent types of phyllaries, glabrous or nearly glabrous leaves with the base being either sessile-auriculate or only slightly decurrent

3. P. kelleri {Thell.) Thulin (2004); Blumea kelleri Thell. (1923) - type: BA/HA, Bessara. 1891/92, Keller 21 p. p. (Z holo ). B. serrata Chiov. (1932) - type: Ethiopia. Soma­ lia, “Dolo”, 17 Sept. 1929. Senni 773 (FT holo ). Bushy perennial herb or shrublet, forming clumps of up to 3 ft. aromatic, 10-35 cm high; indumentum of 1-2 mm long, white translucent liairs mixed with short glandular hairs or sessile glands. Leaves bright yellow-green, nar­ rowly oblanceolate, 10-45 x 2—10(—20) m m base auricu­ late, not decurrent, margins irregularly dentate to pinnati­ fid, apex acute. Capitula solitary, campanulate, 5-10 x 5-12 mm; peduncle 1.5-4.5 cm long, ebracteate or with a few short bracts Phyllaries 6-7-seriate, lanceolate, gradedmonomorphic, spreading or reflexed from near the tip or half-way up, glandular with short stalked hairs; outer 1-3 x 0.5 mm; inner 5-10 x 0.5-0.8 mm. with long, acumi­ nate, reflexed tips. Florets purple. Cypselas c 1 x 0.3 m m cylindric, pubescent. Pappus white, 4.5-6 mm long, barbellate Acacia-Commiphora bushland on limestone; 250-1180 m; SD HA. Somalia. Gilbert & Gilbert 3986, Ash 1146; Gilbert et al. 7607.

145. ASTERACEAE: 54. Pluchea

147

Figure 87. PLUCHEA OVALIS 1 - up­ per flowering and fruiting branch x W, 2 - caudex x c 5; 3 - outer phyllary x c 5; 4 - inner phyllary x c 5; 5 - female floret x 6 V2 ; 6 - bisexual floret x c 5; 7 - sta­ men x 14; 8 - stigmas x 14; 9 - cypsela x 14. From Lebrun 7602 and 7572. Drawn by A. Cleuter. (Reproduced with permission from Flore des Spermalophyles du Parc National Albert Vol. II, PI XLV.)

A C l e u t e r del.

6.

nogalensis Chiov. (1929) type; Somalia. Bacino del Nogal, 27 June 1924, V. Puccioni & J. Stefanini 937/1031 (FT holo.) P. pteroclada Chiov. (1932) - type: Somalia/Ethicpia [HA] border, Goddere, 1931, Guidoti 47 (FT holo.).

Bi >hy. perennial herb, up to c 30 cm high, aromatic; lower stem bark glabrous, fissured, young branches pube cent to woolly, narrowly winged. Leaves oblanceolate to narrowly lanceolate, 2.5-5.5 x 3—10(—15) mm. nar­ row ed into the sessile decurrenl base with wings on

intemodes. c 1-2 mm wide and denticulate, scabrid-pilose to woolly on both surfaces, margins denticulate to serrate, apex acute. Capitula campanulate, heterogamous, 5-6 x 4-4.5 mm at anthesis, few to several in teminal corym­ bose cymes; peduncle green, white-woolly or pubescent, 5-10 mm long. Receptacle epaleate. narrow. Phyllaries 4-5-seriate; outer ovate, c 2 x 0.7 mm, green in middle, purplish at apex, loosely cottony at base and up to middle with sessile glands; middle lanceolate, 3-3.5 x c 1 mm, purplish and sparsely pilose in upper parts and with sessile glands; inner linear to linear-lanceolate, c 5 x 0.5 mm.

148

145. ASTERACEAE: 54. Pluchea. 55. Pseudoblepharispermum

Figure 88.

PLUCHEA S O M A U E N S IS

1 upper part o f a flowering branch x '/j; 2 - leaf x 2; 3 - capitulum x 3; 4 outer phyllary x 4; 5 - inner phyllary x 4; 6 - pistil and young cypsela with pappus x c 6. (No specimens cited in orig.) Drawn by Flelene Lamourdedieu. (Reproduced with permis­ sion from Bull Mus. Natl. Hist. Nat., B, Adansonia , ser. 2, vol. 3 (1981). p. 318, PI. 1, as P. pectinata.)

purplish toward apex, sparsely pubescent and with sessile glands especially from middle to apex. Florets purple or violet: outer numerous, female, c 4.5 mm long, corolla filiform, 3-lobed, c 3.7 mm long, inner bisexual, few. c 5 mm long; corolla tubular c 4.6 mm long, 5-lobed. Cyp­ selas brown, c 0.6 mm long (only immature seen), sparsely pilose, ribbed. Pappus wliite. 1-seriate, 3-4 mm long, shorter in female florets, scabrid. Rocky stream bed in Acacia-Commiphora bushland; c 400 m. HA: Somalia. IECAMA BH-39.

Shrub. Leaves simple, alternate, densely white-tomentose. Capitula small, densely congested into terminal secondary heads, disciform, pedunculate. Receptacle paleate; paleae hairy . Outer florets female, filiform, cen­ tral florets functionally male. Anthers without tails. Style undivided. Cypselas ellipsoid, with uncinate hairs. Pappus absent. A monotypic genus so far recorded from southern Ethiopia only.

P. brem eri J.-P. Lebrun & Stork (1981) 55 PSEUDOBLEPHARISPERM UM J.-P. Lebrun & Stork (1981) J.-P. Lebrun, A. Stork & J. Wuest in Bull. Mus. Natl. Hist. Nat. Pans, Ser. 4, Sect. B, Adansonia 3(4): 419-424 (1981)

-ty p e: BA. 150 km SW of Kelafo. near Goddere, Boudet 7805 (ALF holo.). Shrub, richly branched, up to 30 cm high, whole plant densely tomentose. Leaves linear-oblong or ovate, up to 40 x 12 mm, margins entire, petiolate; petiole c 4 mm

145. ASTERACEAE: 55. Pseudoblcpharispermum, 56. Sphaeranthus

long, not decurrent. Receptacle flat, paleate. Capitula in glomerules. paired, or many together terminating branches, pedunculate; peduncle up to 1 cm long, nar­ rowly winged. Phyllaries multi-seriate; outer ones linem-spathulate. c 3 mm long, pubescent toward the apex aid also sessile-glandular, inner ones oblong. 4-5 mm long. Female florets (1—)2—3 mm long; ovary triangular, pibescent. some hairs circinnate; corolla filiform, glab dus , 3-lobed. lobes ciliate. glandular. Bisexual florets 2 3, c 5 mm long; ovary sterile, ciliate; corolla narrowly ft ruiel-shaped. ciliate. 5-lobed, lobes hairy, glandular. Stamens shortly exserted; anther base saccate, without tails. Cypselas ellipsoid, pubescent. Pappus absent. Open, deciduous scrub with Boswellia on gypseous . HA. only known from the type. Lebrun et al. (loc.cit.) compared this genus with I. Iepharispermum, from winch it differs in having cyps : las without pappus, and with Sphaeranthus, from which i differs in its shrubby habit and in the leaves being non-decurrent over the stem. They were unable to assign i to any tribe, but Anderberg [in Plant Syst. Evol. 176: 45-177 (1991)1 placed it near Sphaeranthus in the Plucheeae.

again, but as it is a common and widespread species, it is described here from material from Kenya and Tanzania. S. flexuosus O. Hoffm.. which is known from Sudan, might also occur in tlie northwestern part of the Flora area and hence it is included in the key. It is a smali. hairy plant with branch leaves 5-15 x 3-5 mm. with subentirc margins and glomerules of about 7-8 x 6-8 mm. A few specimens with immature glomcmles al FT (i.e., Corradi 1816 and 1818. from GG. between Atana and Murulc) have unwinged stems and might probably represent S. kirkii Oliv. & Hiem. a species know n from Kenya (K l) and beyond. Cufodontis (1967; 1099) citcd the above specimens under .V. amaranthoides Bunn f. an Asian species. S. kirkii. if present in tlic Flora area, could easily be distinguished from S. flexuosus and S. ramosus (Klatt) Mesfin. two species with unwinged stems, on the basis of its long leaves (up to 7.5 x 1.5 cm) and ovoid, con­ ical or elongate-ovoid glomcmles with bright pink sub­ tending bracts.

1. Stem not winged, leaves only slightly dccurrcnt. 2 - Stem distinctly w inged at least in the upper parts. w ings extending between tlic nodes. 3 2.

56 SPHAERANTHUS L. (1753) "■ R oss-C raig in H o o ker’s Icon. PI. 5(6): 1-90 & Tab

JSb 1-3525 (1954). Perennial herbs or shrubs, usually aromatic; stem erect or procumbent and spreading, usually winged, wings often denticulate. Indumentum of 1-seriate or basally multiscriite simple and/or capitate liairs. and sessile glands, .eaves simple, alternate, linear oblong, lanceolate, ellipjjic. sessile, decurrent with short or elongate wings, finely •uiberulous to scabrid-hispid. usually densely glandu­ lar-punctate. margins denticulate, dentate, rarely entire, ipex acute-apiculate or rounded. Glomcmles globose, o oid. conical or cylindrical, pedunculate, peduncle usuilly short. Receptacle round, ovate, oval, narrowly obong; linear flat or convex in longitudinal section. Lower­ most subtending bracts variously linear, lanceolate, obong. ovate, elliptic or rounded, hidden or visible al intJicsis. Capitula sessile or on very short secondary rcceptaclcs. Capitular bracts 2-18. flat or folded lengthwise, ‘ lorcts purple, lilac or violet. Female florets: corolla filiform or narrowed from base upwards, usually gla­ brous. 5-lobcd; ovary pubcsccnt or glabrous, liairs simple >r anchor-shaped; style bifurcate, slightly exserted or in­ cluded. Bisexual florets: corolla tubular-funnel-shaped. Jandular or glabrous. 5-lobcd; ovary pubescent, rarely Llabrous: stamens usually included; style minutely bifid at apex. Cypselas. if developed, with anchor-shapcd or uncinate hairs. Pappus absent. A genus with about 38 species distributed throughout the tropics of the Old World from Africa to Australia, with a few species in Iraq and Iran. So far about 9 species arc known from tlic Flora area. S. hullatus. which is know n from Ethiopia only from the type, lias not been collected

149

Leaves obovate to oblong or oblong-elliptic, ob­ tuse. subentirc. shaggy or densely hairy; recepta­ cle globose; outennost subtending bracts not visible. S. flexuosus Leaves linear to linear-lanceolate, denticulate, apprcsscd-hispid; rcccptaclc convcx: outennost subtending bracts visible. 8. S. ramosus

3. Rcccptaclc globose, ovate, conical, narrowly ob­ long, or linear. - Receptacle flat or convcx.

4

9

4. Corolla of bisexual florets deeply and abruptly constricted just above the broad base and immedi­ ately below the lobes. 7. S. gomphrenoides - Corolla of bisexual florets uniformly tubular or funncl-sliapcd. gradually widcr upwards. 5 5. All bracts ± hidden at anthesis. lower ones not larger than the others: capitular bracts 7-17. flat; female florets 7-18. 6 Lower subtending bracts always visible, usually larger than the others; capitular bracts 3-4; fe­ male florets (3-)4-6. 6. S. steetzii 6. Rcccptaclc linear; glomcmlc narrowly c> lindnc. (15—>25—60 mm long. 2. S. ukambensis Rcccptaclc and glomcmlc round, ovate, conical, or narrow ly oblong. 7 7. Ercct subshmb or slinib. c 1-1.2 m high: leaves 5-12 mm long. 2. S. ukambensis - Ercct or procumbent creeping herb, up to 50 cm or more high: leaves 16-100 nun long. 8 8. Leaf margin dentate to coarsely dentate, surface bullatc. veins impressed above, prominent be­ neath; ovary of female florets with bifid liairs. co­ rolla lobes of both female and bisexual florets with both long setae and sessile glands 1 S. hullatus

150

145. ASTERACEAE: 56. Sphaeranthus

Figure 89.

SPHAERANTHUS BULLATUS 1 part o f flowering plant x 1; 2 - leaf from main stem x 1; 3 - part o f leaf margin, lower surface x 2; 4 -part o f peduncle x 1V2\ 5 - glom erule o f capitula, part cut away to show receptacle x 2; 6 - two o f the outerm ost subtending bracts x 5Vi; 7 - phyllaries x 5'/a; 8 - fem ale floret x 24; 9 & 10 bisexual florets in two stages x 24. (No specim ens cited in orig.) Drawn by Stella Ross-Craig. (Re­ produced from Hooker's leones Plantarum , Vol. 36, Ser. 5, 1954. Tab. 3503B.)

-

9.

Leaf margin senate to denticulate, surface ± smooth, not bullate, ovary- of female florets with anchor-shaped hairs; corolla lobes glabrous or sessile glandular. 3. S. suaveolens Lowermost subtending bracts densely pilose with both capitate glandular and eglandular hairs, apex long apiculate and recurved; ovary of female flo­ rets with anchor-shaped hairs. 5. S. neglectus Lowermost subtending bracts glabrous or sparsely glandular-hairy at base, apex rounded; ovary of female florets with simple hairs. 4. S. zavattarii

spinescent; the others linear-spathulate, hairy near the apex and similar to capitularbracts. Capitula on very short secondary receptacles. Capitular bracts 8-14. linear or linear-spathulate. 2.3-2.8 mm long, scarious, hairy near apex, ciliate on margins near apex, apex with short spine. Florets mauve, purple or violet. Female florets 8-14, c 2 mm long; corolla nanowed from base to apex, glabrous or with a few glands, ovary hairy with minute bifid liairs; style bifurcate, exserted. Bisexual florets 2-3. 2.5-3 mm long, corolla narrowly tubular-funnel-shaped, lobes glan­ dular and usually also with long setae, tube sparsely glan­ dular. stamens exserted; ovaiy sterile;’ style simple, exserted. Fig. 145.89.

1. S. bullatus M attf (1936) Wooded grassland and busliland with permanent to types: Tanzania, Endlich 556. Volkens 1632. senu-pennanent water courses; riverbanks; c 320 m; SD Petholtz 217, Jaeger 22. Winkler 3758. Broun 1914 & [Dolo Odo); Kenya. Tanzania. Sebsebe & Ensermu 2735; 5217. Zimmermann 152. Stuhlmann s.n. (all B syn.. destroyed). E. Brumpt s.n. S. gallensis Sacleux (1938) - type; Ethiopia (re­ gion not knowa probably SD). Doungatta. mission du Bourge de Bozos. Dec. 1907, E. Brumpt s.n. (P holo.) 2. S. ukambensi.s Vatke & Hojfm. (1894) type: Kenya. Ukamba. Adi River. Ilildebrandt Perennial herb; stem erect or decumbent, branches erect 2612 (?B holo . K iso.). or spreading, leafy, winged, wings coarsely incisedS. cvlindraceus Cufod. (1943) - type: SD. Mega. dentate. deciduous or broken up with age. sparsely mi­ Corradi 1865 (FT syn.); SD. Giarcorsa. Ruspoli 0-2100 m. EW SU KF SD; Sudan. Uganda. Gilbert & Thulin 871; Gilbert 3529; Mooney 7586.

4

4 S. zavattarii Cufod. (1943) tvpes: GG. Gondaraba. 28 Mav 1939. Corradi 1839 & 1852 (FT syn.). Erect, annual or short-lived perennial herb, up to c 30 cm luah: stem and branches winged, wings deciduous or bro­ ke n in lower parts, continuous in upper parts, sparsely liairy, hairs both eglandular and capitate glandular, mar­

Figure 92. SPH AERANTU H S SUA VEOLENS var A B YS­ SINICU S 1 part o f upper stem with glomerules x 2 bi­ sexual iloret x 8. var ANG U STIFO LIU S 3 part o f upper stem with aglom erule x 3 4. 4 glom erule, longitudinal section \ \'/2; 5 & 6 - phyllaries x c 4; 7 bisexual floret x 8 var T E T R A P H Y L L U S 8 & 9 parts o f flowering stems x 1 i. 10 bisexual floret x 8. 1 & 2 from Schimper 219; 3 7 from Kotschy 500. 8 from Diimmer 2594; 9 & 10 from Small 1184 Drawn b\ Stella RosS'Craig. (Reproduced from Hooker s leones Planlarum. Vol. 36. Ser 5. 1954, Tab 3509.)

gins dentate or denticulate. Leaves oblancclate. 3-6 x 0.8-1.3 cm. sparsely hairy on both surfaces, with both eglandular white hairs and sessile glands, decurrcnt. mar­ gins widely dentate to denticulate, apex obtuse-apiculate. G lo m e ru les su b g lo b o sc to dcprcssad globose. 5-6 x 7 - 9 mm. shortly pedunculate, pcdunclc 3-6 mm long, not winged sparsely hairy and with sessile glands. Receptacle flat. Subtending bracts large and almost enclosing the capitula. leathery . 4-6 x 2.5-3.5 nun. concave, rounded to oblong-elliptic, glabrous or sparsely liairy or glandular at base (inyoung glomerules). margins hyaline and purplish, apex rounded Capitula sessile. Capitular bracts 4; 2 later­ als folded inwards. 4.5 x c 1.5 nun, scarious to leathery : the outer and inner flat to slightly concave, c 4.5 x 1 nun. oblong, scarious. apex round to truncate. Female florets 1-2. stipitatc. c 4.5 mm long; corolla filiform-tubular.

154

145. ASTERACEAE: 56 Sphaeranthus

Perennial herb; stem repeatedly branched, winged, wings dentate or denticulate to entire, deciduous or broken in older parts, sparsely to densely pubescent, hairs long, both capitate-glandular ;ind eglandular (especially in younger parts). Leaves oblong or narrowly oblong-lanceolate, c 1-4 cm long. 2-7 mm wide, densely pilose on both sur­ faces with both long capitate-glandular and uni- or rnulb'Scr.atc hairs, decurrent, margins denticulate with sharp mucronatc teeth, apex acute- or obtuse-mucronate. Glomerules depressed-hemispherical, 5-6 x 8-10 mm, shortly pcdunculatc: pcduncle 4-10 mm long, not winged, pilose with both capitate-glandular and long eglandular liairs. Rcccptaclc flat Subtending bracts: lowermost ovate to ovate-elliptic. (4-)5-6 x 2.2-2.7 mm, sparsely to densely pilose with both capitate-glandular and eglandular hairs strongly curved at base, the irud-nb thickened, apex long apiculate and recurved; other sub­ tending bracts similar 111 sh ap e. 3.5-4.5 mm long, ciliate and glandular in the upper parts. Capitula sessile. Capitular bracts '-7 . scarious and thin; 2 laterals 3.5-4 nun long, folded lengthwise, puberulous. apex rounded and densely ciliatc; the outer and inner c 3-4 nun long, narrow ly oblong to elliptic, puberulous and ciliate on mar­ gins near apex, apex obtuse or truncate, minutely mucronate. Female florets (3-)4-7(-10). 3-4.5 mm long, co­ rolla filiform, glabrous: ovary with forked and hooked liairs; style bifurcate, exserted. Bisexual florets 1-2. 3.5—5 mm long; corolla tubular but wider near apex, tube sparsely glandular, lobe densely glandular and with a sin­ gle stifTbristly hair, stamens about equal to corolla; ovary hairy: sty le minutely bifid. Fig. 145.94.

Figure 93. SPHAERANTHUSZA VA TTARII 1 part o f flow­ ering plant x V4; 2 detail o f leaf surfacex c 9 . 3 glomcrulc x 2. 4 - glomerulc. longitudinal scction x 2. 5 subtending bracts \ c 3. 6 - 8: phyllaries. lateral, adaxial and abaxial, respectively \ 4 1 / 3; 9 - female floret x 11; 10 - bisexual floret x 11. (No speci­ mens cited in orig.) Drawn by Stella Ross-Craig. (Reproduced from Hooker's leones Plcinlarum. Vol. 36. Ser 5. 1954. Tab

3512.)

slightly wider near the base, sessile-glandular at base, oth­ erwise glabrous: ovary liairy. liairs simple; style deeply bi-furcatc. included or only slightly exserted. Bisexual florets c 2.5 111111 long, shortly stipitate; corolla tubular (but constricted about half-way up), glabrous; stamens in­ cluded: ovary c 2 111111 long, glabrous: sty le slightly longer than stamens Fig 145 93. No habitat data from the Flora area available [in Ken­ yan specimens tlic following liabitat information is pro­ vided for the specics alluvial flats, damp, swampy grassland, open grassland with scattered tal\Acacia seyal war. fistula and occasional. Icacia mellifera; 100-400 m| SD; Kenya. Corradi 1839. 1852. 5 S. neglectus R. /• Pries (1916) - types: EW. Acrour. 1900 m, Schweinfurth & Riva 1056 (UPS syn., K iso.) & ?Zimbabwe. Chihinde. Baum s.n. (B sy n. ?destroyed).

No habitat data from the Flora area: 1900 111 EW, Rep. Dem. du Congo. Zambia. Zimbabwe. Namibia. Schweinfurth & Riva 1056. 6. S. steetzii Oliv. H iem (1877) ty pe: TU. near Schire. Dschogardi. PI. abyss. II. Schimper 525 (K holo ). S. anguxtijoUus auct., non DC. (1836). quoad Schimper 525 Sch. Bip. ex A. Rich. (1848). Oligolepis angustifolius (DC.) Steetz (1863). nom. nud.. quoad Schimper 525. Sphaeranthus nuhicus auct. non Sch. Bip ex Steetz (1864). quoad Terraciano & Pappi 2782, GioJit 2. I atova 1384 Bahlrati 1546 (all in FT). Perennial herb or sub-shnib: stem erect or prostrate, densely branched from near the base upwards, w inged, puberulous to glabrous, the wings inucronulate and punctate Leaves linear to narrowly oblong. 1-5 cm long. 2-8 mm w idc. sparsely puberulous to glabrous, densely giandular-punciaie 0 11 both sides, decurrent, margins denticulate with sharp teeth, apex mucronate to apiculate. Glomerules subglobosc. 8-10 x 7-8 mm, shortly pcdunculatc. pcdiinclc up to 1.2 cm long, sparsely puberulous and glandular, usually notwinged. Receptaclc suborbicular Subtending bracts: lowermost oblong-ovate to ovate. (3—>4—6 \ 2-2 111111. glabrous or glandular, round and apiculatc al apex, margins minutely ciliate: other bracts similar or oblong, rounded and mucronate at apex

145. ASTERACEAE: 56. Sphaeranthus

155

Figure 94.

SPH AERAN TH U S

NEG IJICTUS

1- flowering branches \ 1; 2 large le a f from lower part o f stem x 1.3 de­ tail o f leaf surface x 1. 4 glomcrule x 2; 5 glom crule. longitudinal sec­ tion \ 2; 6 & 7 subtending bracts (out­ erm ost left) x c 6: 8 10: phyllaries. lateral, adaxial and abaxial x c 6: 11 female floret x I I S . 12 bisexual flo­ ret x 1112 . (No specimens cited in orig.) Drawn by Stella Ross-Craig. (Repro­ duced from Hooker's leones Plantarum. Vol. 36. Ser. 5. 1954. fab 35I5A .)

12

CaiMtuJa sessile. Capitular bracts 3(-4): 2 laterals folded lei ithwise. c 4 mm long, scarious. ciliate at the apex and on the midrib, the inner and outer bracts, when present, na nowly elliptic. 3-4 nun long, scarious. pubcmlous at the apex, ciliate on the margins. Florets red-violet or pale pu pie. FemaJe florets 4-6, 3.5-4 mm Jong; corolla nar­ rowed upwardly, glandular; ovary puberulous with forked an: hooked liairs; style bifurcate, exserted Bisexual flo­ ret 1 or 2, 3.5-5 mm long; corolla tubular or cylindrical, gh ndular; stamen c 4-5 mm long; ovary glandular; style mi mtely bifid. Fig. 145. 95. On rocky and sandy ground along lake margins, in me ist depression in dry and bare deforested areas, in tem­ po arilv water-logged soil in dry areas; 500-2400 in. EW Tl GD SU IL SD BA; Sudan. Uganda. Kenya. Ash 748; Ry ling & Ermias 1238; Mesfin T. 7294.

nun long, concave, leathers except at margins, sparsely glandular-hairy, margins ciliate. apex with a 0.5-3 nun long awn; others similar but membranous. Capitula ses­ sile. Capitular bracts 3(-4). 2 laterals folded lengthwise. 3-4 mm long, membranous except for mid-rib and apex. truncate. ciliatc or fimbriate at the apex; inner and. when present, outer oblong-oblanceolate. truncate. 3.5-4 mm long, membranous, hairy and ciliatc at the apex. Female florets 4. 3.5-4 nun long, shortly stipitatc; corolla gradu­ ally narrowed from base to apex, glabrous, glandular; ovary glandular and with anchor-shaped hairs; style bifur­ cate. included. Bisexual floret (l-)2 . 3-4 nun long, co­ rolla broadly cy lindric. deeply and abruptly constnctcd just above the broad base and also below the lobes, ses­ sile-glandular at base, in the constricted areas and on the lobes; stamen exserted; ovaiy glandular and with an­ chor-shaped hairs at base; style bifurcate, exserted Fig. 145.96.

Along stream banks in open grassland with scattered 7. gomphrenoides O. Hoffm. (1894) type: East Africa, sine loco. Fischer 335 (B Acacia. Tamarix. Ficus, etc.; wet roadside ditches; 1550-2745 m. SU SD (Wachile) HA: Kenya. Tanzania. holo., destroyed, K fragment of holo ). IF.CAMA RS-X2XJ. de Wilde 4345; Corradi 1675. Pe ennial herb, up to 60 cm high; stem erect or at first pn jumbenl'spreading with erect or ascending branches, roi ung in the lower nodes, all winged, wings broken or 8. S. ramosus (Klatt) Mesfin (2004); de iduous in older parts, minutely ham . distantly Athanasia ramosa Klatt (1895) - type: HA. nu cronulate, glandular-punctate. Leaves linear to nar­ Ogaden-Steppe, 1891/92. Keller 31 (Z holo.). row Iv oblong, decurrent. 1.5-8.5 x 0.2-2.2 cm. sparsely nu lately pubescent, margins mucronulatc-denticulate. Low spreading perennial herb or shrublct, aerial pans up glandular-punctate, apex acute or rounded and apiculate to 25 cm high. Stems several from the root apex or the or iristate. Glomerules sub-globose or ovoid, 10-13 x short rhizome, terete, densely sessile-glandular and 9 - 2 nun. pedunculate; peduncle 1-5 cm long, narrowly sticky, not winged, brownish with closely spaced interlo >roadly v\ inged. Receptacle conical in longitudinal secnodes. leafy toward the apex. Leaves linear to lineartio i Subtending bracts: lowermost ovate-oblong. 3-4 lanceolate. 10-30 x 1-3 nun. sessile, appressed hispid.

156

145. ASTERACEAE: 56. Sphaeranthus, 57. Litogyne

Figure 95.

SPH AE R A N TH U S S T E E T Z II 1 - part o f flowering stem and branches x 1. 2 large leaf x 1; 3 - detail o f leaf surface x 1; 4 - glom erule with part cut away to show receptacle x c 2; 5 - a lowerm ost subtending bract x 5'/*; 6 - one of the other subtending bracts x 51/*; 7 & 8 phyllaries, lateral and adaxial view x 514; 9 - female floret x 12; 10 - bisexual flo­ ret x 12. (No specim ens cited in orig.) Drawn by Stella Ross-Craig. (Repro­ duced from Hooker s leones Plantarum , Vol. 36, Ser. 5, 1954, Tab. 3516A.)

broadly denticulate, apex acute. Glomerules subglobose to hemispheric, 8-10 x 6-7 mm, pedunculate; peduncle c 5 mm long, sparsely ciliate, bracteate and leafy below, not winged. Receptacle convex or flat 3-4 mm wide. Sub­ tending bracts: lowermost broadly oblong to broadly ob­ long-elliptic with acute apex, sparsely pubescent through­ out, 4-4.3 x 2.8-3.2 mm; inner series narrowly oblong with hairs confined to near the apex, 4.8-5 x 1-1.2 min, apex mucronate. Capitula sessile. Capitular bracts 6: 2 lat­ erals folded lengthwise, stiff, 4-4.2 mm long, scarious at margins near the apex, ciliate at apex and on midrib; 2 in­ ner bracts similar to outer; 2 innermost bracts flat, scarious or membranous, oblong, 4-4.5 nun long. Florets purple. Female florets 4, 4-4.5 mm long; corolla nar­ rowed upwardly, glabrous, ovary glabrous, style bifur­ cate, c 4.5 mm long, equal to or slightly exserted from co­ rolla, branches c 0.5 mm long, stigmatic surfaces mar­ ginal. Bisexual florets 3, 4.5-5 mm long, corolla tubular or cylindric, glabrous, stamens c 2 mm long, ovary gla­ brous, style c 6 mm long, much exserted, bifurcate, papillose, branches c 0.5 mm long. Edge of seasonal pools; altitude not known. HA (Balleh); Somalia. Hemming 1547 (FT K). S. ramosus is morphologically allied to S. flexuosus O. Hoffm. and S. kirkii Oliv. & Hiem. In S. flexuosus, the leaves are obovate to oblong or oblong-elliptic, with ob­ tuse apex, subentire margins and dense, raided shaggy hairs; the receptacle is globose with the outermost sub­ tending bracts not visible. In S. kirkii, the leaves are 30-75 x 10-15 mm long, the glomerules ovoid, conical or elongate-ovoid and subtending bracts visible and bright pink.

57 LITOGYNE Harv. (1865) Erect perennial herbs or shmblets, aromatic. Leaves alter­ nate. ovate or oblong, entire or serrate, decurrent. Capitula heterogamous. disciform, campanulate, in terminal cory mbose cy mes or few in clusters, rarely solitary, pedunculate. Receptacle convex to slightly conical, epaleate, smooth or minutely alveolate. Phyllaries few- to multi-seriate, imbricate, increasing in length inwards. Florets purple, outer female, inner bisexual. Corolla of fe­ male florets filiform, 2-3-lobed, shorter than the style, those of bisexual florets tubular-campanulate, (3-)5lobed. Anther base sagittate, tailed, connective ovate or ovate-oblong, obtuse. Style undivided or minutely notched, thicker above with tuberculate hairs. Cypselas cylindric or narrowed below, 5 - 10-ribbed. Pappus absent in female florets: of 2-3 smooth bristles in bisexual florets or absent. A monotypic tropical African and south African genus commonly encountered from W Africa to Sudan and So­ malia, and south to Namibia and S. Africa (Natal, Cape).

L. gariepina (DC.) A. Anderb. (1991); Ethulia gariepina DC. (1886); Epaltes gariepina (DC.) Steetz (1864) - type: S Africa, Gariep River, Drege 2721 (K iso.). Bushy and much branched perennial herb or shrub, aro­ matic, up to 1 m high; stem narrowly winged especially in young branches and above, wings narrow, entire to sparsely serrate. Leaves lanceolate or linear-oblong, 20-50(-60) x 4-8(-9-12) mm, viscid-glandular and sca­ brous-pubescent to glabrescent, decurrent; margins entire

145. ASTERACEAE: 56. Sphaeranthus, 57. Litogyne

157

Figure 96. SP H AERAN TH U S GOMPHRENOIDES. 1 - part o f flowering stem x I; 2 - leaf from main stem o f young plant x 1; 3 - detail o f leaf surfacc x 16; 4 - glom erule, part cut away to show receptacle x c 2; 5 - an outermost subtending bract x 5 >/3; 6 - one o f the other subtending bracts x 5'/3; 7 & 8 phyllaries, lateral and adaxial x 5 */3; 9 female floret x 12; 10 - bisexual floret x 12. (No specim ens cited in orig.) Drawn by Stella Ross-Craig. (Repro­ duced from H ooker's leones Plantarum. Vol. 36, Ser. 5, 1954, Tab. 3522B.)

or parsely serrulate, apex acute or mucronate. Capitula cai loanulate or sub-turbinate at the base, 4-5 x c 4 mm, usi ally 3-6 on short peduncles at the apex of branches; pei Lncle up to 10 mm long, pubescent, liairs yellow. Ph iiaries c 4-senate. 2-4 x c 1 mm. leathery, pubescent wi p. both simple. 1-seriate and sessile-glandular liairs; ou j r and middle ovate to ovate-lanceolate; innennost lineaj : r oblong, acuminate, often purple-tipped. Florets pur­ ple corolla of female florets c 3 nun long, style exserted. bif ^cate at apex, of bisexual florets 5-6 mm long; anthers ex: erted; style exserted. undivided or minutely notched, pa; illose. Cypselas cylindric, c 1 nun long. 5-ribbed. Pa »pus absent or of 2-3 smooth bristles on those from bise? ual florets, 2-3 nun long. Fig. 145.97.

Tribe 8. G N A P H A L I E A E Rydb. (1917) Inuleae - Gnaphalieae Cass. (1822). Inuleae - Gnaphaliinae Less. (1830).

Shrubs, perennial or annual herbs. Leaves simple, alter­ nate. pubescent to densely tomentose; margins entire, flat or revolute. Indumentum of flexuous intertwined hairs, rarely also of myxogenic hairs. Capitula campanulate. hemispheric or rarely conical or turbinate, heterogamous and radiate or homogamous and discifonn. Receptacle epaleate or sometimes fimbrillate. arcolate or honey­ combed, flat. Phyllaries few to multiseriate. scarious throughout or upper ones brightly colored and scarious and lower (stereome) suberized (sclcrenchymatous). So far no specimen of L. gariepina lias been seen from these fenestrate or not. Female florets Filiform and tubular the Flora area. But since tliis weedy species was recorded, or ligulate: ligulate florets fertile. Bisexual florets bisex­ as f- rhulia gariepina. from Sudan (?Kordofan. Kotschv ual or functionally male; corolla tubular to tubular35 bank of white Nile, Andrews A 37) and from Somalia campanulate. 5-lobed. Anther base sagittate and tailed: (H iran. Region. Bulo Burte district. 4° 05' N ; 45° 15’ E. connective oblong, obtuse. Style bifurcate, branches with K i :nar 17153). it is most likely to occur in the lowlands sw eeping liairs at apex or on outer surfacc; stigmatic areas of S and SW Ethiopia. Tlie name “Epaltes kotschyi Sch. separated, converging (in Athrixia) or confluent (in Bi : in Schweinf. & Asch. (1867:284) and also cited as Phagnalon). Cypselas dark brown, glabrous or with an l icorrcct name under Epaltes alata (Sond.) Steetz by myxogenic or not myxogenic liairs. Pappus of barbellate Ct fodontis (1967: 1098) is perhaps based on “no. 45. bristles, or of scales, or absent. Ki schy 1837. 1838. Aethiopia”. This specimen was col­ About 110 genera and 1600 species with worldwide let icd not from the Flora area but from Kordofan in Sudistribution; 9 genera in the Flora area. da The above description is based on NE African sp c miens. 1. Capitula radiate. 2 Habitat not recorded from the Flora area [in Somalia Capitula discoid or disciform. 3 L gariepina is known from mixed deciduous bushland wi h Acacia and Dobera and from C.ombretum wooded gr ssland; 130-360 m], ?S and SE Ethiopia; Mali in west AJ ica to East Africa, and south to South Africa.

2.

Ray florets white, pink to light purple; pappus usu­ ally 1-seriate and bristles barbellate or sometimes also with shorter outer bristles. 58. Athrixia

158

145. ASTERACEAE: 57. Litogyne

Figure 97. LITO G YN E GARIEPINA 1 part o f plant x Vy, 2 - capituluir x 6; 3 capitulum , cross section x 12 4 outer floret x 14; 5 - inner floret x 14, 6 - style branches o f inner floret x 20, 7 - stam ens x 12. All from Beesley 30. Drawn by Linda Gurr. (Repro­ duced with perm ission from FI. Trop. E. Afr.. C om positae, fig 77.)

I *u*«

-

Ray florets pale yellow, pappus 2-seriate. bristles more or less equal and numerous 59 Macowania

3

Annual cottony herbs, capitula terminal oraxillaiy. in sessile globose cymes. 65. Fiiago Perennial herbs or shrubs, if annual, capitula not globose 4

4.

-

Capitula not discernible, tangled and held together by densely matted indumentum, held flat on one plane. 63. Chiliocephalum Capitula discernible, with long or short peduncles. although densely formed in some taxa. not tan­ gled and held together. 5

Hairs on cypselas setaceous, multicellular; anther base obtuse or minutely tailed 60 Phagnalon Cy pselas glabrous or with duplex my xogenic liairs. or short, erect, usually 3-celled liairs: anther base sagittate with long tail. 6 6

Inner phy llaries opaque and cotton-white; female florets more numerous than the bisexual ones, stereome not fenestrate. 64. Gnaphalium Apices of phyllaries more or less the same colour as the rest; stereome usuallyfenestrate. 7

7

Annual herbs, weeds or cultivated plants.

8

145. ASTERACEAE: 58. Athrixia

8.

9.

-

159

Shrubs or perennial herbs, rarely annual, native plants 9 Capitula in spicate axillary' glomerules, each 2.5—3 5 x 2-2.5 mm; weed of usually wasteland and gardens. 64. Gnaphalium (only G. americanum) Capitula solitary, showy. 20-70 x 7-12 mm; culti­ vated for its attractive capitula. 66. Bracteantha Involucre 5-15 mm long or across, hemispheric or campanulate; phyllaries many-seriate, yellow, white, pink, red or cream. 6 1. Helichrysum Involucre 2-4(-5) mm long or across, conical or campanulate; phyllaries 3-5-seriate, pale yellow, white or pink. 10

1( Involucre campanulate; phyllaries ± equal, pale to bright yellow. 62. Pseudognaphaiium Involucre conical; phyllaries lengthening inwards, bright yellow, white or pink. 6 1. Helichrysum

58. ATHRIXIA Ker Gawl. (1823) G Kroner in Mitt. Bot. Staatssamml. Miinchen 16: 1-268 (I >*80). Si rubs or perennial herbs. Leaves simple, alternate, narrc a margins revolute. Capitula solitaiy or subcorymb< ie, heterogamous, radiate, turbinate-campanulate. Rect ;)tacle epaleate. Phy llaries multiseriate, unequal and si ;:essively longer inwards, apex acute. Florets pink to lif t:i-purple; ray florets female, in one row. fewer than the di >c florets, staininodes often present. Disc florets bisexu< yellow, corolla 5-lobed. Anther base sagittate with tailed auricles. Style branches obtuse, with hairs dorsally. C pselas oblong. 3-6-ribbed, sparsely hairy. Pappus C( imonly of uniseriate, free, barbellate bristles, rarely also with very short outer setae. A genus with about 14 species, mainly found in tropi­ c s and southern Africa and Madagascar, 1 species in the F >ra area. A rosmarinifolia (Sch. Bip. ex Walp.) Oliv. & Hiern 1877); Klenzea rosmarinifolia Sch. Bip. ex Walp. (1843) - type. TU, Mt. Kubbi, near Aksum, PI. abyss. I, Schimper 265, wrongly indicated by Walpers as ‘267’ P holo . G K M iso.). E 2Ct, perennial herb or shrub with several stems from a wijody rootstock; stem 0.5-1.5 m high, terete, striate, glab> :us below, leafy above, floccose to whitish tomentose ai nve or puberulous at the extremities, branches erect or a.1.. ending. Leaves linear or narrowly linear-lanceolate, up tc " 5 x 0.6 cm. leathery, white-tomentose beneath, nearly gi tbrous, shiny and longitudinally canaliculate above, s< isile; margins entire and revolute; apex acute or apicula e Capitula heterogamous, radiate, turbinate or c; mpanulate, 10-12 x 5-6 mm. solitary or a few together on very short pedicels, terminal and subtenninal on branches. Receptacle epaleate, flat, areolate. Phyllaries n my-seriate, lengthening inwards, imbricate and only

Figure 98 A TH R IXIA RO SM ARINIFO LIA 1 - upper stem portion x '/*; 2 - outer phyllary x c 7; 3 - inner phyllary x c 7; 4 ray floret x 5>/3; 5 - style o f ray floret x 5 '/3; 6 - disc floret x 5 */3; 7 - cypsela with pappus. All from Quarre 7294. (Reproduced with permission from Flore d'Afrique Centrale, part 2, 1989, p. 229. PI. 47.)

the brownish tips reflexed; outermost oblanceolate, con­ tinuous with bracts on peduncle, 3-3.5 x 0.5-0.8 mm, woolly at the back, apex acute-apiculate; inner linearoblong, ciliate. 7-7.5 x c 1 mm. Ray florets white or pink to light purple, c 10-12. female and fertile. 12.5-15 mm long, glabrous; tube c 4.5 mm long, lamina oblongelliptic, 7.5-8 mm long, 3-dentate at apex. Disc florets bi­ sexual, c 8.5 mm long, 50-55. Corolla tubular slightly wider in upper half, c 6 mm long, 5-lobed, lobes papillose, tube glabrous. Anthers c 3 mm long, base sagittate with acute base reaching filament-collar joint, connective ob­ tuse. Style bifurcate, c 6.3 mm long, base wider and thick­ ened, branches obtuse, c 1 mm long with sweeping hairs half-way on outside. Cypselas dark brown, c 1.3 x 0.4 mm, c 10-ribbed, glabrous except at base where there is a ring of erect bristles, rarely with a few glands. Pappus 2-seriate, white; outer short, c 0.5 mm long, inner c 5-6 nun long, barbellate. Fig. 145.98. Banks of streams and rivers in open scrub or thicket border, including Erica, Juniperus, Hypericum, Heli­ chrysum, Kniphofa, etc.; open rocky places in

160

145. ASTERACEAE: 58. Athrixia, 59. Macowania, 60. Phagnalon J

Juniperus-Podocarpus forest; 1890-2800 m TU GD GJ

SU AR KF GG SD BA HA; Sudan, E Africa, R6p D6m. du Congo, Mozambique, Malawi, Zambia, Zimbabwe. Gilbert 1104; Mooney 5016; Mesfin T. 6982.

59. MACOWANIA Oliv. (1870) Klenzea Sch. Bip. ex Walp. (1843), p.p. min., excl. lecto. Antithrixia auct, non DC.: Benth. in Be nth & Hook. f. (1873) p.p.; Oliv. & Hiem (1877); O. Hoffm in Engl. & Prantl (1894). B.L. Buitt & J. Grau in Notes Roy. Bot. Gard. Edinburgh 31: 373-376 (1972). Intricately branched dwarf shrub Leaves alternate, crowded, linear or linear-lanceolate with usually promi­ nent midrib below, sessile, margins revolute, apex mucronate. Capitula heterogamous, radiate, campanulate or tuibinate, solitary or a few together, sessile or shortly pedunculate and terminal. Receptacle flat, honey­ combed, epaleate. Phyllaries multiseriate, increasing in length inwards, light brown or hyaline, glabrous to thinly glandular pubescent. Ray florets pale yellow or whitish, 1-seriate, female and fertile, rarely sterile; ovary ribbed, pubescent or rarely glabrous, pappus of white, caducous or persistent scabrid bristles; tube glabrous or haity, lamina narrowly elliptic, 3-fid at apex; style branches sub­ acute. Disc florets bisexual, usually all fertile; ovary ribbed pubescent or rarely glabrous, pappus of white caducous or persistent scabrid bristles. Corolla tubular be­ low, gradually widened above, 5-lobed. Anther base sagittate, smooth or fimbriate. Style branches obtuse at apex. Cypselas cy lindric or narrowly spindle-shaped (el­ liptic), ribbed, glabrous or hairy. Pappus of bristles.

glabrous except for the hairy mid-vein, also minutely glandular, lower surface of old leaves white-woolly, of young leaves appearing glabrous owing to the woolly hair being matted with a viscid exudate, margins of young leaves revolute. Capitula sessile and solitary or a few in shortly pedunculate corymbs, 9-11 x 5-7 mm, usualh su­ perseded by some of the upper leaves. Phyllaries tuibinate, straw-yellow, glabrous except for the minute glands on the outermost; middle and lower ones ovate to ovate-lanceolate; innermost oblong-lanceolate. Florets pale yellow or whitish. Cypselas villous, c 3 mm long, 10-ribbed. Pappus white, c 6 mm long, persistent.

Diy montane slopes; GD/TU WU; Yemen. Polunin 11650; Schimper 1443. 2. M. abyssinica (Sch. Bip. ex Walp.) B.L. Burtt (1972); Klenzea abyssinica Sch. Bip. ex Walp. (1843); Antithrixia abyssinica (Sch. Bip. ex Walp.) Vatke (1875) - type: TU, M t Scholoda, near Adwa, PI. abyss. I, Schimper 227 (P holo., BM K iso.). Similar to M. ericifolia except in the following features: Leaves narrowly elliptic, 10-20 x 1-2.5 mm, flat mid­ vein raised below, both surfaces white-felted when young, upper surface of fully grown leaves finely glandu­ lar-pubescent to glabrescent lower white-woolly, upper usually reaching only to base of capitula. Capitula not su­ perseded by the leaves, 7-10 x 5-8 mm. Phyllaries sparsely glandular-pubescent middle and lower oblong to oblong-lanceolate. Fig. 145.99. Montane slopes; c 2200-2800 m; EW TU SU; Yemen. Pullen 574; Friis et al. 4532; Pappi 1177.

A genus with 11 species, nine of which occur in South Africa and two in the Flora area and Yemen. 1. Leaves linear, subglabrous above, 1-2.8 cm long, upper reaching the top of the capitula or some overtopping it; middle and lower phyllaries ovate to ovate-lanceolate, glabrous. 1. M ericifolia - Leaves narrowly elliptic, thinly woolly above, 1-2 cm long, upper usually reaching only to the base of the capitula; middle and lower phyllaries ob­ long to oblong-lanceolate, sparsely glandular pu­ bescent. 2. M. abyssinica 1. M. ericifolia (Forssk.) Burtt & Grau (1972); Aster ericifolius Forssk. (1775) - type: Yemen, M t Kurma, Forssk&l in Heib. Forssk. 992 (C holo., BM iso.). Antithrixia angustifolia Oliv. & Hiem (1877) type: GD. Gerra Abune Tekle Haimanot, 2440 m, 1863, Schimper 1443 (BM holo., K W iso.). Antithrixia abyssinica auct., non (Sch. Bip.) Vatke (1875), p.p., quoad spec. excl. syn. Dwarf shrub or perennial herb; young branches leafy, lower parts bare with only the conspicuous grey or dark leaf scars. Leaves linear tapering to the acute apex, up to 28 x 1 mm, flat mid-vein raised below, upper surface of young leaves thinly white-woolly, those of older leaves

60. PHAGNALON Cass. (1819) M. QaiserA H W. Lack in Willdenowia 15: 437-450 (1986). Bushy, perennial herbs or small shrubs; stem densely white-tomentose, rarely glabrous. Leaves simple, alter­ nate, undulate or dentate, usually with revolute margins, sessile and caudate or slightly decurrent on stem, light green above, white tomentose beneath. Capitula hetero­ gamous, disciform, usually solitary at the end of branches or a few together in loose paniculate cymes. Receptacle epaleate, flat Phyllaries multi-seriate, imbricate, scanous at margins, usually dark-brown at middle or purplebordered, obtuse to acute-acuminate. Florets yellow, rarely red or purple-tinged; outer florets female, filiform, usually more numerous than the inner bisexual florets; co­ rolla 3-5-lobed, lobes hairy. Bisexual florets narrowly funnel-shaped, corolla 5-lobed, lobes papillose and hairy; anthers ecaudate or caudate at the base. Style bifid, stig­ mas rounded, papillose. Cypselas oblong-elliptic, whitehairy, light brown, indistinctly 8-ribbed. Pappus 1-seriate with 5-10 scabrous setae. A genus with 6 species in tropical north and northeast Africa 5 of them occurring in the Flora area. P. nitidum Fresen. (1833), recorded by Cufodontis (1967: 1105), is

161

145. ASTERACEAE: 59. M acowania, 60. Phagnalon

Figure 99.

M ACOW ANIA A B YSSIN IC A 1 habit x V*; 2 - outer phyllary x 4'/*; 3 inner phyllary x 4 '/2; 4 - floret x 4 Vi. All from Mesfm T. & Tewolde B.G.E. 8805. Drawn by Birhanu Mihretu.

)jnd in N Africa and in the Middle East (cf.. Qaiser & ack 1986: 448). i . Capitula on peduncles up to 1 cm long, arranged in leafy racemes throughout stem branches. 1. Capitula on peduncles up to 15 cm long, arranged in loose paniculate cymes arising near the apex of branches. 2 Leaves very distinctly dentate, lanceolate to obovate; margins of phyllaries not distinctly scarious. 2. P. stenolepis

-

Leaves entire, undulate or very niinutley and re­ motely dentate; phyllarics with distinct broad scarious margins. 3

3.

Outer and middle phyllarics ovate to obovate. ob­ phagnaloides tuse atP.apex, without purple borders, glabrous or glabrescent; pappus setae 9-10. 3. P. quartinianum Outer and middle phy llarics acutc to acuminate. purple- or dark brown-bordered or with a dark brown band from middle to apex; pappus setae 5-7. 4

-

162

4.

-

145. ASTERACEAE: 60. Phagnalon

Margins of phyllaries purple-bordered, apex acute-acuminate; young leaves narrowly elliptic 4. P. schweinfurthii with flat margins. Margins of phy llaries broadly scarious. dark brown in middle or with dark brown band from middle to apex; young leaves linear with strongly revolute margins. 5. P. abyssinicum

1. P. phagnaloides (A. Rich.) Cufod. (1967); Pluchea (Laggera) phagnaloides Sch. Bip. nom. nud.. quoad Schimper 685; Blumea phagnaloides A. Rich. (1848); Phagnalon scoparium Schweinf. (1867), nom. nud.; P. scoparium Sch. Bip. ex Oliv. & Hiem (1877), nom illegit. - type; GD, Simen Mts., Mt. Silke, 9000 f t, 1851, PI. abyss. II, Schimper 685 (P holo., K iso.). Perennial herb or subshrub up to 30 cm high. Stems as­ cending to suberect, branches angled. Leaves simple, al­ ternate. glabrous to arachnoid, 10—25(—30) x 2-6 mm, lan­ ceolate to elliptic, narrowed below into a small winged petiole, margins denticulate, revolute, apex acute. Capitula axillary, solitary, 7-9 x 8-9 mm, arranged in leafy racemes, peduncle 5-15 mm long, glabrous to sparsely glandular. Phyllaries multiseriate, glabrous, graded monomorphic with the outermost smallest and broadest, ovate, 2-3 x 1-1.5 mm, margins undulate, apex scarious, translucent; middle series 3-4 x c 1 mm; inner ones linear or linear-lanceolate, 4.5-7 x 0.5-1 mm, apex acuminate. Outer florets female, 4-6 mm long, filiform. Bisexual florets 5-6 nun long; corolla tube sparsely hairy, limb glabrous; stamens exserted; style bifurcate, truncate and papillose at apex. Cypselas brown, sparsely hairy, round, c 1 mm long. Pappus setae 6-7, scabrous above. 4-5 mm long. Erica arborea bushland; evergreen woodland; 1830-3200 m. GD; not known elsewhere. Schimper 81; Lemma G.S. 649; Mesfin T. & Zerihun W. 2848. This species can easily be distinguished from the oth­ ers by its leafy, racemose inflorescence.

2. P. stenolepis Chiov. (1911) type: TU, near Guendepta, Schimper s.n. (RO holo.). Blumea fruticulosa Hochst., nom. nud., quoad Schimper s.n. (fide Cufod., 1967:1106). Perennial, straggling herb, woody at the base, or subshrub, up to 45 cm high, densely branched with the loosely branched stem usually covered with dense appressed woolly liairs. rarely subglabrous to almost gla­ brous. Leaves alternate, oblanceolate, dark green above, grey-green to almost whitish beneath, 20-45 x 5—10(—15) iiuii, narrowed to a sessile attenuate base, margins denticulate to widely dentate-serrate with apiculate teeth, apex acute. Capitula heterogamous. disciform. 8-9 x 5-6 nun. campanulate to subhemispheric. arranged in pani­ culate cymes at the apex of branches, peduncles white to­ mentose to subglabrous in upper parts, 2.5-10 cm long.

slender, c 0.5 mm wide. Phyllaries multi-seriate, progres­ sively longer inwards, linear-lanceolate, all more or less similar, dark purple in upper parts, apex acute to acuminate, glabrous or sparsely floccose in young ones; outer 1.2-2 x 0.5-0.8 mm; middle 3-4 x 0.8—1 mm; inner­ most 7-8 x 0.5-0.8 mm. Outer florets female, many more than the inner bisexual ones, c 5-6 mm long, filiform or slightly wider at base, ovary hairy. Bisexual florets 35-40, 5.5-6 mm long; corolla narrowly cylindric. ex­ panded in upper parts, 5-lobed, lobes glabrous or sparsely papillose, recurved, purple-tinged; stamens slightly exserted; anthers tailed; style with 2(-3) branches, trun­ cate and papillose at apex. Cypselas 7.5-8 nun long, cov­ ered with white bristles. Pappus setae 5-6, equal to or slightly longer than, the corolla, scabridulous near the apex. Degraded mountain slopes with evergreen bushland or scattered Acacia, Becium and Aloe\ c 2500-2700 m. TU SU; Sudan, Yemen, western Saudi Arabia. Schimper s.n.; Mesfin T. 8507 Senni 71 (FT).

3. P. quartinianum A. Rich. (1848) type: EW/TU, Mareub. Ouartin-Dillon s.n. (P holo.). P. nitidum auct., non Fresen. (1833); Oliv. & Hiem (1877); Engler (1892), quoad Schimper 251 p.p. Subshrub up to 50(-75) cm high Stems erect, densely branched, woolly older branches glabrescent. Leaves simple, alternate, linear, 35-60 x 1-5 mm, densely woolly above, more sparse beneath, sessile-auriculate, margins entire or denticulate toward the apex, revolute. Capitula solitary in open corymbs, 11-13 x 9-10 mm. pedunculate, peduncle 5-10 cm long, glabrescent to tomentose. Phyllaries multiseriate. graded monomorphic, margins broad, scarious. translucent, silvery white; outer obovate, 2.5-3 x 1.5-2 mm, apex obtuse; middle 4-5 x 1.5-2 mm, similar to outer ones; inner ones 7-8 x c 1 mm, linear, apex acuminate. Outer florets female, filiform Bisexual florets 30^t0; corolla 6-7 mm long, tube sparsely hairy, lobes papillose on outside; stamens exserted; style bifur­ cate, branches truncate, papillose. Cypselas brown, round, c 1 mm long. Pappus setae 9-10. white, barbellate. 6-7 mm long. Moist river valleys, montane slopes; 1900-3000 m. EW TU SU WU; not known elsewhere. Schimper 25 lp.p.; Scott 200; GelahunA. & Zerihun W. 283.

4 P. schweinfurthii Sch. Bip. ex Schweinf (1868) type: Sudaa Jebel Scheltal. Schweinfurth 365 (K syn ), Schweinfurth 364 (P syn.).

var. schweinfurthii Perennial herb, up to 30 cm high. Stem erect, sparsely branched, woolly Leaves simple, alternate, linear. 15-65 x 3-5 mm, glabrescent above, densely white woolly be­ neath. sessile, amplexicaul, margins entire, flat, apex

145. ASTERACEAE: 60. Phagnalon, 61. Helichrysum

163

aci ic or acuminate. Capitula solitary1, 14—18 x 10-12 mm, pei uncle densely white woolly, 5-7 cm long. Phyllaries mi lV'.seriate. glabrescent, densely floccose or woolly exter lally, outer ovate to lanceolate, 1.5-2.5 x 1-1.5 mm, mar:.;ins membranous, silvery-wliite. purple streaked, apt > acuminate; middle ones lanceolate, 4.5-6 x 1—1.5 m i !. apex acute to acuminate; inner ones 6-8 mm long, ma rgins membranous, silvery white, apex purplish or brownish. acuminate. Outer florets female, filiform. Bise> ual florets pale yellow. 35^-5. 7-8 mm long; corolla 6-7 mm long, tube sparsely liairy. limb c 1 nun long; sta­ men- exserted; style bifurcate, branches truncate. pa| iilose. Cypselas brown, round, c 1 nun long. Pappus set il 5-7. white. 6-7 mm long. Fig. 145.100. Habitat not known; 2500-2700 m. EW (Kohaito pla­ teau ); Sudan. Egypt. Schweinfurth s.n. (not traced). According to Engler (1910), this species is known fron: the extreme north-western parts of Eritrea. This in­ formation is based on specimens collected in 1892 and 18 - by Schweinfurth, but the vouchers have not been tra.ed. Var. androssovii (B. Fedtsch.) Qaiser & Lack (in Wi idenowia 15; 13 (1985)] occurs in Saudi Arabia, Egvpt, Iran, Pakistan. Afghanistan and Turkmenistan. 5. ’. abyssinicum Sch. Bip. ex A Rich. (1848) - type: SU. ‘Choa’, Quartin-Dillon & Petit s.n. (P ecto., cf Qaiser & Lack p. 446). P. hypoleucum Sch. Bip. ex Oliv. & Hiem (1877), nom illegit. - type: GD. Simen Mts.. Mt. Aber. near Jschenausa, PI. abyss. II, Schimper 854 (K lecto., BM WU isolecto.). P. nitidum auct.. non Fresen (1833): Oliv. & n iem (1877), quoad Schimper 251 p.p Penninial hert>. up to 50 cm liigh. Stem woody. with straggli ig to suberect branches, wliite woolly. Leaves simple, alternate, linear, 15—55(—100) x 2-10 mm, glabrous to arac.inoid above, white woolly beneath, sessile to subampiexicaul. shortly decurrent, margins denticulate, re\olute, apex subacute to mucronate. Capitula terminal, soitary, con mbose or cymose, 9-12 x 8-9 mm, peduncle de tsely wlute woolly. 5-8 cm long. Phyllaries multiseri­ ate graded monomorphic; outer 2-3 x 1.5-2 111111. ovate, glabrous to woolly, apex obtuse, margins scarious. undulat ;o dentate, dark brown to blackish at base; middle ones 4-5 x 0.7-1 mm. glabrous, brownish or blackish at b a ; a p e x subacute to acuminate; inner ones linear. 5-7 x c nm. Outer florets female, filiform. Bisexual florets 35-45, yellow; corolla 4-5 mm long, tube sparsely liairy, limb glabrous except for the papillose lobes; stamens ex: eited; style bifurcate, branches often unequal in width, truncate and minutely papillose at apex. Cypselas brown, round, sparsely hairy, 1-1.2 inm long. Pappus setae 6-7, white, unequal, barbellate. 6-9 mm long. Rocky areas in montane forests, dry montane slopes wi h low shrubs and herbs; 1800-3660 m. EW TU GD GJ W l SU AR BA HA; not known elsewhere. Friis et al. 1366; Ash 1399; Mooney 5029.

Figure 100. PHAGNALON SC H W E IN FU R TH II 1 - upper part o f stem (no magnification given in orig.); 2 outer phyllary x 5. (No specimens cited in orig.) Drawn by Magdi El-Goharv. (Reproduced from Students' Flora o f Egypt, cd. 2. PI. 197A.)

61. HELICHRYSUM Mill. (1754), corr. Pers. (1807), nom. cons. M esfin T adesse & T. Reilly in D .J.N . H in d et al. (eds),

Advances in Compositae System a tics: 379-450 (1995). Erect, or straggling, perennial herbs, subslirubs or shrubs, rarely annual; stem lanatc. tomentose, glandular pubes­ cent to partly glabrcsccm. Leaves simple, alternate, linear lanceolate, elliptic or ovate, petiolate or sessile with the decurrent bases sometimes forming narrow wings on intemodes. woolly or glandular-pubescent, margins entire or undulate, flat or revolute, apex acute to acuminate. Capitula small to large, cylindric, conical, campanulate to depressed globose, heterogamous or homogamous. few to numerous in loose or congested terminal coiymbose cymes or panicles, rarely solitary. Receptacle flat or slightly convex, areolate. honey-combed or fnnbrillate. Phyllaries few' to multiseriate. white, yellow, pink, red. or­ ange or light brownish-yellow, sometimes with varied

145. ASTERACEAE: 61. Helichrysum

164

combinations of colour, translucent to opaque, glabrous or sometimes pubescent at the base or in the middle; stereome usually not fenestrate (in Flora area). Florets yellow or whitish yellow, rarely pinkish, c 4 to more than 900. proportion of female to bisexual florets variable; co­ rolla glabrous, lobes 0.2-0.5 mm long, papillose or hairy on outside; corolla of female florets usually (3—)4(—5)lobed, narrowly tubular to filiform, usually narrowed in upper parts, of bisexual florets 4-5-lobed. tubular below or funnel-shaped in upper parts. Anthers sagittate at base with narrow translucent tails, apical appendage lanceo­ late. Style base dilated, branches truncate and penicellate, sUgmatic surfaces not confluent at apex. Cypselas light brown lo black, oblong to oblong-obovate. glabrous to papillose, rarely hairy. Pappus white or yellow. 1-seriate, setae few to many, bases free or cohering by patent cilia, sometimes falling off together, scabnd or barbellate in middle, tips or upper parts scabrid. baibellate or cells im­ bricate. the uppennost cells often clavate. A genus with more than 600 species found mainly in southern Africa and Madagascar (c 245 species in S. Af­ nca). Also found in Southern Europe. SW Asia S India. Sri Lanka and Australia; 23 species in the Flora area. H. gracilipes Oliv. & Hiem. H. rosulatum Oliv. & Hiem and / / mmmoanum Oliv. & Hiem. all recorded by Cufodontis (1967: 1111) as probably occurring in Eritrea (EE), are re­ stricted to the island of Socotra. The cultivated ornamental plants sometimes referred to as H. bracteatuni (Vent.) Andrews (1805) are now placed in genus no. 65. Bracteantha Anderb. & Haegi. 1. 2.

-

3. -

Plants with spines in leaf or inflorescence axils; leaves less than 1 cm long. 2 Plants without spines, leaves usually longer than 1 cm. 3 Cushion-fonning perennial herbs or shrubs: capitula 10-15 x 5-8 mm; phyllaries 10-14 mm long; florets 122-130. 1. H. citrispinum Densely branched shrub, not cushion-fonning; Capitula 6-6.5 x 4-4.5 mm, phyllaries 4.5-5 mm long: florets 48-50. 2. H. horridum Erect annual herbs, the roots not surviving beyond the life-span of the aerial parts. 23. H. foetidum Perennial herbs with erect, scrambling or scandent branches, or cushion-forming shrubs. 4

4. -

Involucre (6 -)8 -15 x (4 - )5 -15 mm. Involucre 2-6 x 1-5 mm.

5.

Florets very numerous (280-910); involucre 12-15 x 7-15 mm. 6 Florets 19-50, involucre 8-10 x 4-6 mm 8

6.

7.

Leaves 2-5 mm wide; shrub, subshrub or bushy. much branched, perennial herb. 3. IL argyranthum Leaves (5-) 10-45 mm wide; erect perennial herb. 7 Capitula homogamous, florets bisexual; phyllaries white, pink or red; rootstock thickened, woody 5. H. formossisimum

-

Capitula heterogamous, female florets outnumber­ ing bisexual ones; phyllaries bright yellow; roots slender. 23. H. foetidum

8.

Leaves 8-15 mm wide; phyllaries pale brown. 6. H. quartinianum Leaves 2-3.5 mm wide; phyllaries white, yellow­ ish-white or pink 8. H. gofen *e

9.

Erect perennial heibs with woody rootstock; leaves mostly basal, cauline leaves few; capitula in com­ pact or loose terminal clusters. 10 Shrubs or perennial heibs with scandent or strag­ gling branches or cushion-forming; leaves all along the stem and branches, capitula terminal or axillary. l | 13

10. Capitula heterogamous, female florets outnumber­ ing bisexual florets. 11 - Capitula homogamous, all florets bisexual. 12 11. Capitula 2—2.5(—3.5) mm wide; phyllaries pale or brownish-yellow, pink or purple, flat; florets 9-18. 19. H. globosum - Capitula 4-5 mm wide; phyllaries pale golden brown, crisped or crumpled; florets 24-39. 18. H. gerberifoUum 12. Involucre 5-6 x 4-5 mm; phyllaries yellowish or golden brown 18. EL gerberifoUum - Involucre 4-5 x 2.5-4 mm; phyllaries bright cream

or golden yellow.

17. H. nudifolium

13. Phyllaries silveiy-white, white with red or pinkish tinge, pink, light green or rarely pale yellow. 14 - Phyllaries yellow, yellowish brown, pale or creamy yellow, rarely whitish-brown. 18 14. Leaves 1-10(-14) mm wide, oblanceolate, oblanceolate-spathulate, oblong, linear or narrowly linear-lanceolate. 15 - Leaves 12-35 mm wide, ovate, broadly elliptic, or oblong-oblanceolate. 16 15. Capitula conical to cylindric, 2.7-5 x 1-2.3 mm. 14 H. giumaceum - Capitula oblong-campanulate. campanulate or urceolate. 6-8 x 3-4.5 mm. 15. H. jijigaensis 16. Florets 4-5; perennial herb or shrub with straggling or scandent branches. 20. H. schimperi - Florets 20-30; erect perennial heibs or shrubs. 17 17. Leaves narrowly lanceolate, usually 1-veined, densely formed on stem and branches; phyllaries light green. 4. H. elephantinum - Leaves oblong-oblanceolate, strongly 3-5-veined from base of blade, sparsely set on stem and branches; phyllaries white. 10. H. harennensis 18. Shrubs or cushion-forming bushy and woody pe­ rennial heibs. 19 - Perennial heibs with erect or straggling or scandent branches. 20 19. Leaves 7-10 x 0.5-1.5 mm; capitula 2.5-3 x 1-1.5 mm; florets 8-9. 12. H. somalense - Leaves (5-)10-70 x 3-20 mm; capitula 3.8-5 x 2.8-4 mm; florets 2 4 ^ 5 . 9. H. splendidum

145. ASTERACEAE: 6! Helichrysum

2 i Receptacle without fiinbrills; leaves densely woolly-tomentose on both surfaces. 7. Receptacle fimbrillate; leaves variously pubescent, not woolly except in young ones. 21 21

Perennial herbs with erect branches; leaves 1-5 (-8) mm wide. 11. H. forsskahlii Perenial herbs with scandent or straggling branches; leaves (5-) 10-20 mm wide. 22 Leaves linear to narrowly elliptic, strongly decurrent. 1-veined or veins not prominent; capitula 2.5-3 x 1-1.5 mm; phyllaries 1.5-2 x 1 mm. 16. H. stenopterum Leaves ovate-lanceolate, oblanceolate or elliptic, sessile or petiolate, 3-veined; capitula 3.5-4.5 (-5) x (1.5—)2.5—3 mm; phyllaries (2-)2.5-4 x (0,5-) 1-2.2 mm. 23

2

Capitula cylindric; leaves petiolate, petiole up to 1 cm long, ovate-lanceolate or oblanceolate; florets pale yellow. 13. H. ogadensis Capitula tubular-campanulate: leaves sessile or subsessile with cuneate base, rarely petiole up to 5 mm long, blade elliptic; florets bright yellow. 24

24 Female florets more numerous than the bisexual ones: phyllaries 0.5-0.7 mifi wide. 22. H. hedbergianum Female florets fewer tlian the bisexual ones; phyllaries (1.5—)2—2.2 mm wide. 21. H. traversii 1 H. citrispinum Del. (1843); Gnaphalium citrispinum (Del.) Schweinf. & Asch. (1867) - type: GD. Simen Mts., Ferret et ( ialinier s. n. (MPU holo.. not seen). Helichrysum spinosum Sch. Bip. (1843). nom. nud.. quoad Schimper 666 and 1246.; Elichrysum spinosum Hochst. (1844). nom. nud.; Gnaphalium spinosum Sch. Bip. (1845) - types: GD. Simen Mts.. Mt. Silke. PI. abyss. II. Schimper 666 (K isosyn ); Simen Mts.. Mt. Bacliit. Schimper 1246 (BM K isosyn.). Helicrysum new it auct.. non. Oliv. & Hiem (1877): Moeser (1910). quoad Steudner 266 and 350. //. newii Oliv. & Hiem var. gunae Schweinf. ex Engl, in liber die Ilochgebirgsfora des tropischen A f­ rica: 426 (1892) - type: GD. Guna. 3100 m. Steudner 350 (B holo.. destroyed. K iso.). //. citrispinum var. hoehnelii auct.. non (Schweinf.) Hedberg. quoad Mooney 5277 and 5287: Cufod. (1966). Cushion forming perennial herb or shrub, often 50-75 cm high and 0 5—1 m across, in very old individuals the plant r lay be up to 3 m in diameter with death of stem and branches progressing ccntrifugally; branch stems an aricate. terete, palc-tomentose. leafy and spiny. Leaves I ijear-lanceolate or oblong. 4-10 x 1-2 mm. often crowded towards the apex of branches, densely white or grey floccose-tomcntose on both surfaces, sessile, mar­ gins revolute, apex obtuse or subacute; spines in upper I aves up to 1.5 cm long. Capitula homogamous. broadly c ijupanulate or depressed-globose. 10-15 x 5-8 nun. sol­

165

itary at the ends of the short leafy- branches. Receptacle flat, epaleate. honey-combed. Phyllaries multiseriate and H. sclerochlaenum spirally arranged, lanceolate to elliptic, 10-14 x 1.5-2.5 mm, white, leathery with dense black hairs on outside at base, scarious in upper half or at apex, apex sub-acute: stereome mostly not fenestrate. Florets yellow, numerous (122-130), all bisexual. 3.5-3.7 mm long. Corolla tubular to slightly funnel-shaped above, c 3.2-3.5 mm long. 5-lobed, lobes densely black-papillose. Anthers c 2 mm long. Style bifurcate, c 2.8 mm long, stigmatic branches c 0.8.mm long. Cypselas light brown to black, c 0.8 x 0.3 nun. papillose. Pappus setae white, 3.2—3.7 nun long, barbellate. 1.

-

Stem with usually many spines up to 1.5 cm long. leaves 7-10 mm long, spreading or some crowded at points of stem or branch bifurcation. var. citrispinum Stem with very few, or completely without, spines; leaves often 4-6 mm long, erect and overlapping, closely appressed to stem. var. hoehnellii

var citrispinum Rocky mountain slopes and meadows, afro-alpine com­ munities with scattered Erica and Senecio: 3000-4500 m. GD GJ SU AR BA; E Africa. Mooney 5277; O. Hedberg 5558; Mesfin T. 5341. var. hoehnelii (Schweinf) Hedberg in Svmb. Bot. Upsal. 15(1): 206(1957); Helichrysum hoehnelii Schweinf. (1892) - type: Tanzania, Kilimanjaro. Teleki 115 (B holo.. de­ stroyed). Ncotypc: Tanzania. Kilimanjaro. O. Hedberg 1194 (UPS neo.. EA K S isonco.). Similar to var. citrispinum in habit and floral details but differing by absence of spines, and short leaves (only up to 6 mm long), erect and closely appressed to the stem. Open stony mountain slopes on light loamy soil de­ rived from volcanic rock, c 4270 in. BA; Kenya. Tanza­ nia. Mooney 7250. 8331. Mesfin T. 5560 from Tulu Dimtu. Bale Mountains. BA. has a few spines, but is otherwise as var. hoehnelii. 2. H. horridum (Sch. Bip.) A. Rich. (1848); Gnaphalium horridum Sch. Bip. (1845) - typo Ethiopia |region not known], PI. aby ss. II. Schimper 975 (P holo.. BM K iso ). Densely and intricately branched spincsccni shrub. 1-1.5 m high, branches terete, indumentum on young parts floccose-tomemose. older parts glabrous and dark grey, spines 0.5-2.5 cm long, straw-yellow when young, straight, axillary and tenninal. Leaves crowded on young shoots, often sub-fasciculate, linear, light-green above, tomentose beneath. 3-8 x 1-2 nun. furrowed along midvein. sessile; margins entire, rcvolutc. apex acuteapiculate. Capitula campanulate. homogamous. 6-6.5 \ 4-4.5 nun. solitary or in cymose clusters and terminal or axillary , leafy. Receptacle epaleate. flat. Phy llaries 5 (>seriate, erect, pale yellow; outermost ovate-oblong, thinly

166

145. ASTERACEAE: 61 Helichrysum

woolly on outside at base, c 1.5-3 x 1—1.5 mm, innner lin­ ear-oblong, 4.5-5 x 1 mm, obtuse at apex. Florets golden yellow, 44-50, all bisexual, 3.7-4.2 mm long. Corolla tu­ bular below, funnel-shaped above, 3-3.2 mm long, 5-lobed, lobes setose. Anthers 1.7-1.8 nun long. Style c 4 mm long, branches 1-1.2 mm long. Cypselas obscurely angular to subterete, 0.8-1.2 x 0.3-0.4 mm, oblong, gla­ brous to sparsely setulose. Pappus expanded toward the apex, setae white. 3—3.5 mm long, barbellate. In Erica arborea scrub with Lobelia rhynchopetalum above afro-montane forests; (2500-)3300-3600 m. GD SU; not known elsewhere. Mooney 6422; Lemma G.S. 622; Mesfin T. 7182. 3. H. argyranthum O. Hoffm. (1895) - type Tanzania, Kilimanjaro. Volkens 1531 (B holo., destroyed, BM K iso.). Shrub, sub-shrub or bushy, richly branched perennial herb with slender, brittle stems and spreading branches, up to 1 m high; lower parts of stems and branches with dry, shriv­ elled persistent leaves. Capitate glandular liairs on upper parts of stems, branches, leaves and peduncles. Leaves lanceolate to linear-lanceolate, (2-)4-6.5 cm long. 2-5 mm wide, faintly aromatic, upper surface light green, gla­ brous or rarely tliinly araclmoid. lower surface wliitetomentose. base sessile-auriculate. margins entire, revolute, apex acute-apiculate. Capitula homogamous. campanulate to depressed-globose, c 15 x 7 mm at anthesis, in compact or loose cymes. Receptacle epaleate. honey-combed. Phyllaries 4-5-seriate. silvery wliite and pinkish or purplish at base or entirely pink, shiny. 3-7 x 1-3 mm. ovate to oblanceolate. glabrous: stereome fene­ strate. greenish w ith a single branched vein. Florets yel­ low. very numerous (c 440—175). all bisexual. Corolla tu­ bular- campanulate, c 3.5 mm long. 5-lobed. lobes c 0.5 mm long. Anthers c 1.7 mm long, sagittate at base. Styles bifurcate, c 4.2 nun long, branches c 0.7 mm long, trun­ cate at apex. Cypselas (only immature seen) oblong. 0.6-0.7 mm long, glabrous. Pappus setae wliite, c 3 mm long, slightly fused at base, barbellate. Margins of montane forests, open grassy mountain or hill slopes with Erica and Helichrysum. wet valleys, 2230-3060 in EE/?/ SU SD KF GG; Sudan. Uganda. Tanzania. Rwanda. Rep Dem du Congo. Mesfin T. 6918; Gilbert 337; Friis et al. 1456. 4 H. elephantinum Cufod. (1966) - type: GG/SD. 'Elefantcnberg' |= Elephant Mountain], 2950 in. Kuls 114 (FR holo.. not seen). Profusely branched, bushy shrub up to 75 cm lugh and c 2 ni across, rarely erect perennial herb Stems at first spreading and then erect, slightly viscous and aromatic, densely leafy , white-woolly. Leaves alternate, narrowly lanceolate, gradually narrowed to base, sessile to slightly auriculate. up to 9 x 1.2 cm, upper surface green, lower white tomentose, margins entire; apex acuminate. Capitula c 4 x 2 mm. hcterogamous, campanulate-urceolate. arranged in dense corymbose cymes, epedunculate or

peduncle 3-4 nun long, densely tomentose mixed with erect glandular liairs. Receptacle conical, glabrous. Phyllaries subequal, the outer continuous with bracts >n peduncle, light green with scarious tip and margins, densely glandular-pubescent throughout except at base. 3-3.5 x 0.8-1.2 mm. linear to narrowly oblanceolate. stereome not fenestrate, light green. Florets 21-31, golden yellow, 2.5-3 mm long. Female florets 8-12. fun­ nel-shaped; corolla 5-lobed. 2.5-3 x 0.8 mm (in expanded part). Bisexual florets 10-19, tubular and funnel-shaped (both occumng together); corolla 5-lobcd, 2.5-3.5 x 1 mm (in expanded upper part), lobes papillose, ovary gla­ brous. Stamens 2.8-3 mm long; anthers 1.2-1.5 mm long, connective acute-acununate. base tailed. Style c 2.2-2.6 mm long, bifurcate at apex, brandies c 0.5 mm long, trun­ cate and penicillate al apex. Cypselas (only immature seen) glabrous, oblong, c 0.5 mm long. Pappus setae white, c 2.2-2.8 mm long, shorter than the corolla tube, scabrid. free at base Fig. 145.101. Moist Hagen ia abyssinica. Allophyllus abyssinicus, Schefflera abyssinica forest with dense undergrowth of shmbs and lianas: 2470-3320 in; SU GG SD BA. not known elsewhere Mooney 8503; Miehe 1870. Mesfin T. 8293. 5 H. formosissimum Sch. Bip. ex A. Rich. (1848); Gnaphalium formosissimum Sch. Bip (1845); //. elegantissimum DC. var majus Oliv. & Hiem. in FI. Trop. Afr. 3: 350 (1877) - type: GD. Simen Mts.. Mt. Silke. PI. abyss II,Schimper672 (P holo.. BR K iso ). Helichrysum formosissimum Sch. Bip. ex A. Rich, var latifolium A. Rich.. Tent. FI. Abyss. 1: 422 (1848) -ty p e. SU. Choa', Petit s.n. (P holo). 1H. formosum auct.. non Sch. Bip.. quoad Schimper s.n. (26.X. 1850. Bachit. 12000'. Ad rivulos montis): Klatt (1873). ?H. formosissimum Sch. Bip ex A Rich forma nivea Chiov. in L A. Savoia-Aosta. La Fsplorazione dello Cabi-llebi Scebeli ...: 415 (1932) - type: ?BA. Basile 98 (TO. holo., not seen). ?H. fomosissimum forma rosea Cluov. ( loc.cit.) — type: ?BA, Basile 183 (TO holo., not seen). H. elegantissimum auct.. non DC. (1838): Englcr (1906. 1910). Erect, perennial herb, (0.2—)0.5—1.5 in lugh. stem usually solitary , from a thickened rootstock, terete, obscurely stri­ ate. lanate-pubescent in upper parts, glabrescent in lower parts, leafy. Leaves ovate-oblong or oblong-lanccolatc. (1—)3—13.5 x (0.5-) 1-4.5 cm, with one prominent midvein and several longitudinal veins (especially apparent on larger leaves), scabrid with numerous glandular liairs. and sometimes also araclmoid above or w oolly -tomentose beneath and loosely cottony or araclmoid above, ovate oblong or oblong-lanccolatc. sessile and clasping stem and sometimes also dccurrent on stem, margins entire, revolute: apex acute-apiculate. Capitula homogamous, hemispherical. 1.2-12 x 1.2-1 5 cm. in terminal longpedunculate corymbs; peduncles w hitc-tomentosc. up to 4 cm long. Reccptacic epaleate. flat. Phy llaries multisenate. wliite. pink or red. sluny. glabrous, lanceolate to

145. ASTERACEAE: 61. Helichrysum

167

lin :ar-lanceolate. (3—)5—15 x 3—5(—7) nun. spreading, ap :x acute; stereome fenestrate or not. Florets yellow, ve > numerous (c 550-750), all bisexual, c 4 mm long. Corolla tubular-campanulate, c 3.8 mm long. Anthers c 1.1 mm long. Style c 3 mm long, branches c 0.6 nun long. C> pselas grey, c 0.8 x 0.2 mm, papillose, otherwise glabn us. Pappus setae white. 3-3.5 mm long, barbellate. Fig. 145.102. 1. Leaves scabrid w ith numerous stiff and glandular hairs, sometimes also thinly arachnoid. var. formosissimum Leaves thinly arachnoid above, densely woollytomentose beneath, sometimes also with a few glandular liairs above. var. guilelmii va formosissimum Along streams and damp spots in Erica arborea bushlai d or forests, in marshy ground with Kniphofia, along SU ams in degraded woodland with Erythrina. Podoca rus. Ficus, Echinops, etc.; (2050-)2600-4345 m. GD G. SU AR IL GG SD BA; Sudan. E Africa. Rwanda. Rep. D( m. du Congo. Mooney 7186; Friis et al. 3625;Mesfin T. 55 o. Friis et al. 7158 is a straggeling specimen from an unusual habitat, AningeriaSyzygium rain forest with much undergrowth of C.yathea mannii, at 2300 m in IL. va . guilelmii (Engl.) Mesfin in Comp. Newsl. 22: 14 (1992; non rite publ.) & in Hind et d\.^ Advances in 1 ompositae Systematics: 413 (1995). Helichrysum guilelmii Engl. (1892) - type: Tanza­ nia. Kilimanjaro, above Kibosho. Volkens 1556 (K lecto.. selected by Hedberg 1957: 215). In moist or sliaded spots in Juniperus forest. Erica ar*< rea bushland. along streams in valleys and on steep slopes; 2900-3800 m. BA GG: E Africa. Rwanda. Rep. D< m. du Congo. Friis et al. 5689; Moonev 8346; Scott 161 6. li. quartinianum A. Rich. (1848) -type: TU, Cliire, Scholoda. 1862, Quartin-Dillon s.n. (P holo., K iso.). Gnaphalium quartinianum (A. Rich.) Schweinf. & Asch. (1867). Erect, perennial herb, up to 60 cm high, with a thick woody rootstock; stems several from the rootstock, each unbranched except in the inflorescence, densely ar. chnoid-tomentose. Leaves sessile, oblong-elliptic to oblanceolate, 3-veined with often the median vein promi­ ner!:. up to 8 x 1.5 cm. grey arachnoid-tomentose on both surfaces, margins entire, apex acute. Capitula homogaiiious. c 10 x 1.5 mm, campanulate, in dense terminal paniculate corymbs, pedicels c 1 cm long, densely woolly, will; both white and black hairs. Receptacle flat, fimbrillate. Phy llaries 4-5-seriate; outer ovate. 4.5-5.2 x 2-j£ 2 mm; inner 5.6-6.5 x 1.8-2 mm, white, opaque or pal. brown (when dry), scarious, narrowly oblanceolate to I'vate-elliptic. pubescent in lower half, with black

Figure 101. H ELICH RYSU M ELEPHANTINUM : 1 - upper part of*stem x '/2; 2 capitulum x 7; 3 - phyllary x 12: 4 —bisex­ ual floret x 11; 5 - female floret x 11,6 - barbellate pappus bris­ tle x 30. (No specimens cited in orig). Drawn by Damtew Teferra. (Reproduced with permission from Opera Bot. 121, 1993, p. 131, fig. 2.)

1-seriate hairs near base, glabrous above, 1-veined. Flo­ rets yellow. 47-54, all bisexual, 5-5.4 mm long. Corolla 3.5-4.6 111111 long, tubular below, funnel-shaped above. 5-lobed, lobes papillose, c 0.5 mm long. Stamens c 3.5 nun long; anthers c 2.2 nun long. Style 3-4.5 mm long, branches 0.5-1.1 mm long. Cypselas cy lindric-tetragonal.

168

145 ASTERACEAE: 61 Helichrysum

102 HELICH RYSU M FO RM O SSISSIM U M : 1 - upper part Figure

o f plant x c '/j; 2 capitulum x 1 ,3 capitulum , cross section x 60; 41outer floret x 12; 5 - inner floret x 12; 6 - style branches o f inner floret x 50; 7 - stamens x 30. J from F ries & Fries 1293. 2, 7 -8 from Tweedie 4219; 3^6 from Rich 85. Drawn by Linda Gurr (Reproduced with permission from FI. Trop. E. Afr.y C om positae, fig 90.)

L 6 u **

multt-ribbed. 1-1.2 x 0.5 mm. glabrous, whitish or grev Pappus wlute. c 4 mm long, barbellate Wooded grassland on black clay soil, grassy patches along stream banks. Erica arborea bushland. 1400-2500 (-3070) m. TU GD SU WG KF IL BA; Sudan. Uganda. Tanzania. Burundi. Rep. Dem. du Congo. Gabon. Cameroun. Nigeria. Zambia. Malawi. Angola. Gilbert 345; Friis et al. 2404. Ensermu K. A Melaku W. 2959. A record from Bale Mountains. Hillman 453. repre­ sents a subalpine form of this species also known from East Africa.

7 H. sclerochlaenum Moeser (1910); Achvrochne sclerochlaena Sch. Bip. ex Vatke (1875), nom. non rite publ - ty pe: Ethiopia without further locality. IS53, Schimper 1060 (P holo., K iso.) Gnaphalium sclerochlaenum Schweinf. & Asch (1867). nom. nud., quoad Schimper 778. Erect, short-lived perennial herb, up to 60 cm high; stem woody below, glabrescent. branched, branches terete, as­ cending. lanatc-tomentose. Leaves oblong-elliptic to obovatc-oblong, densely woolly-tomentose. 3-7 x 1-2 cm. 3-veined with veins arising about 0.5-1 cm from base

145. ASTERACEAE: 61. Helichrysum

of blade and running almost parallel to inid-rib. sessile and narrowed to the amplexicaul base; margins entire; apex sub-apiculate. Capitula heterogamous, oblong. 4-5 x 1-1 8 mm. sessile, in dense, branched sub-globose termi­ nal cymes. Receptacle epaleate. Phyllaries 3-4-seriate, sub-equal in length 4-4.5 x 0.5-1 mm, shiny and pale yel­ lowish- or whitish-brown, scarious toward apex, leathery and light green below; outermost oblong-lanceolate, thinly arachnoid or w oolly at base, apiculate at apex; inner linear-oblong, glabrous, fimbriate at apex. Florets light yellow, 7-13, c 4.5 mm long; c (6 -)9 -l 1 female, c 1—2(—3) bisexual. Corolla 3.1-3.3 nun long. Stamens 4.1-4.3 mm long; anthers 1.5-1.6 mm long. Style 3.8-4 mm long. Cypselas glabrous, c 0.8 mm long. Pappus white. 3.5-3.8 mm long, scabrid. Vlontane forests and woodlands; c 2450m. ?GD SU; not known elsewhere. Ash 640; Schimper 1060. \chyrocline sclerochlaena Sch. Bip. was not ac­ cepted by Vatke (1875) since he cited it in synonymy onl The name should therefore be dated only from Moeser (1910).

8. H. gofense Cufod. (1966) - type: GG, Dita-berg pDita Mountain’], Kuls 763 |FR holo.). Perennial herb with w oody prostrate stems forming dense clu:nps or mats; branches subterete, arising from almost all the nodes, erect, often (3-) 10-15 cm liigh, leafy throughout with concealed intemodes. Leaves oblonglint ar, linear or linear-lanceolate, sessile, crowded, (0.5-) 1 -4 cm long. 2-3.5 mm wide. appressed-tomentose especia ;. on the upper surface. 6-7-veined, veins parallel, im­ mersed above, slightly raised beneath Capitula heterogarnous, campanulate. 8-10 x 4-6 mm at anthesis, termi­ nal. sessile or on up to 5 mm long, araclmoid. pedunculate, globose or branched cymes. Receptacle flat, epaleate. Ph llaries multiseriate, imbricate, oblanceolate, shiny, white or yellow ish-white, sometimes pink or pink-tinged at base, glabrous throughout or with numerous black hairs at l ase, (3—)5—11 x (1—)2—3 mm. middle series w idest and lor ifest (c 7.5 x 2 mm), innermost series narrow and small, stereome not fenestrate. Florets yellow. 35^40; 8-14 female. 22-30 bisexual. Corolla tubular-campanulate. 3.5-3.7 mm long, glabrous. 5-lobed, lobes with dense black papillae or hairs. Anthers 1.7-1.8 mm long. Style 3.J -4 mni long, branches c 1 mm long. Cypselas grey, obloi g-elliptic. c 1 nun long, glabrous or sparsely papillose. Pa ipus wliite. c 4 mm long, barbellate. Moist and open afro-alpine grassland and heathJand. sa aercd Erica-Helichrysum scrub in open, rocky ground; (3! 75-)3500-4375 m. AR GG BA; not known elsewhere. Mi oney 5195; O. Hedberg 4215; Mesfin T. 5538. The variation in habit, leaf and capitula size in this sp.cies seems to be correlated with differences in alt tudinal zonation. Plants growing in liigh alpine com­ munities, i.e., between 4050 m and 4375 m have 1-3 cm hij:h erect branches. They produce fewer but larger

169

capitula and smaller leaves (usually less than 1 cm long). Those growing at lower altitudes, i.e., between 3275 m and 4050 m, have more numerous but smaller capitula on the 10-30 cm high branches. The leaves in these forms are often between 1 cm and 4.5 cm long. 9. H. splendidum (Thunb.) Less. (1832); Gnaphalium splendidum Thunb. (1800) - type: S Africa, Cape of Good Hope, Thunberg in Herb. Thunberg (sheet 19257, UPS). G. abyssinicum Sch. Bip. (1845); Helichrysum abyssinicum (Sch. Bip.) A. R ich (1848) - type: Mt. Silke, PI. abyss. II, Schimper 663 (BM K UPS syn.); Mt. Kubbi. Schimper 127 (BM K syn). Shrub or bushy perennial herb with woody base. (20-) 50-75 cm liigh. and sometimes up to 1 m in diameter, stem branched and leafy above, branches subvirgate. terete, white-woolly above, grey and with withered leaves below. Leaves linear, linear-oblong or lanceolate, (5-) 10-20 (-25) x 3-6(-8) mm, 3-5-veined. densely grey­ ish-white tomentose on both sides, sessile; margins entire, often revolute; apex sub-obtuse with short cartilaginous mucro. Capitula heterogamous, campanulate. 4-5 x 3 - \ mm, subsessile, crowded in dense, globose or corymbose terminal clusters. Receptacle flat, epaleate. Phyllaries 4-5-seriate, yellow above, translucent in the middle and light brown below, shiny; outermost light brown and loosely woolly at base on outside, 1.5-2 x 1-1.5 mm, ob­ long or ovate, acute at apex; innermost glabrous, 3-4 x 1.5 mm, oblong-oblanceolate or ovate-oblong, acute at apex. Florets yellow, 40-55, 3-4.2 mm long; 5-12 female, 36-43 bisexual. Corolla tubular below, funnel-shaped in upper parts, 2.5-3 mm long, 5-lobed, lobes sessileglandular. Anthers 1.5-1.6 mm long. Style c 3 mm long, branches c 0.6 mm long. Cypselas light-grey. 0.8-1 x 0.3-0.5 mm, minutely papillose or glandular, otherwise glabrous. Pappus white, 3-3.5 mm long, barbellate. Erica arborea bushland, afro-alpine grassland and hcathland; (2500-)2900^300 m. TU GD WU SU AR BA HA; Sudan. Tanzania. Malawi, Zimbabwe, South Africa. O. Hedberg & Mesfin T. 7049; Mooney 6438; Friis et al. 3417. 10. H. harennensis Mesfm (1993) type: BA, Bale Mts., 2-3 km on Fincha Haberra-Simbiro track, c 3500 m. Mesfin T. 7858 (ETH holo., K UPS iso.). Erect, perennial herb, up to 80 cm liigh, with short rhi­ zome; stem solitary', unbranched except in the inflores­ cence. pubescent; liairs often purple. Leaves alternate, sessile, slightly decurrent on or clasping stem, 4.5-8 x 1-2.5 cm, oblong-oblanceolate, strongly 3-5-veined from base of blade, veins prominent below, decreasing acropetally, margins entire, sometimes revolute; surfaces thinly and loosely arachnoid and/or pubescent with purple-tipped hairs. Capitula numerous, heterogamous. 3-4 mm long, campanulate in cory mbose cymes, several held together by woolly indumentum; the ends of the

170

145. ASTERACEAE: 61 Helichrysum

11 H. forsskahlii (J.F. Gmel.) Hilliard & Burtt (1980); Gnaphalium fruticosum calycibus/Iavis, ... fohis linearibus ... Forssk (1775). nom. non rite publ.; G. forskahlii J.F Gmel. (1792); Helichrysum cymosuni (L.) D. Don subsp.fruticosum Hedbcrg in Symb. B907-1908, 2: 420 (1911) - type: GD, Gaffat [near Debre TaborJ, 1863, Schimper 1122 (B holo., BM K iso.). Gnaphalium dominici-saccardoi Fiori (1940) type: KF, Beka, near Gogeb, Saccardo 13 (FT holo.). Ei set, perennial herb, (10—>30—75(—L00> cm high; with a th .:k. woody rootstock, stem simple, shortly pale tomento e above, puberulous and shiny below. Leaves often or y up to middle of stem, usually 3-5-veined starting fr< n the middle of the blade, pale white-tomentose be­ ne -th, squamous-scaberulous and bright green above, na newly to broadly elliptic; margins entire; apex acuteac minate; radical leaves up to 15 x 3 cm, attenuate into th petiole, petiole up to 5 cm long; cauline leaves narro ved towards the sessile, auriculate-amplexicaul base, up to 14 x 4.5 cm, uppermost linear. Capitula heteroga nous, oblong-campanulate, 4-5 x 2-2.5 nun, sessile or su sessile on veiy short peduncles arranged in dense br ached sub-globose terminal clusters or occasionally in lo< sely branched, several to about a dozen smaller clustei Receptacle fimbrillate. Phyllaries 3-4-seriate. pale or br ■•nish-yellow, pink or purplish, shiny; outermost br udly elliptic, 3—3.2 x 1—1.2 mm, obtuse, woolly below: ini ermost oblong-obtuse, 3.5-4.5 x 1-1.4 mm, leathery be ow, glabrous; stereome fenestrate. Florets yellow, 9-18,8-14 female, 1-4 bisexual, 4-4.2 mm long. Corolla 3.. -3.6 mm long. Anthers 1.2-1.3 mm long. Style c 3-3.3 mi i long, branches c 0.3 mm long. Cypselas oblongob '.ate. terete, glabrous, c 1-1.1 x 0.4-0.5 mm. Pappus wl te, c 3.5 mm long. Forest margins, open moist montane and afro-alpine gn island and bushland with Erica, Helichrysum. etc.; 13 m)-3660 m. GD GJ SU AR IL KF GG SD BA HA; Suda l E Africa Cameroun. Nigeria, Bioko, Gabon. R\ anda, Angola, Rep. Dem. du Congo, Malawi, Mada­

175

gascar. O. Hedberg 5668; Evans & Lythgoe 28; Mesfin T. 5367. 20. H. schimperi {Sch. Bip. ex A. Rich.) Moeser (1910); Achyrocline schimperi Sch. Bip. ex A. Rich. (1848); Gnaphalium schimperi (Sch. Bip. ex A. Rich.) Schweinfurth & Asch. (1867) - type: TU. Mt. Scholoda. PI. abyss. I, Schimper 393 (K lecto., BM isolecto.). Achyrocline adoensis Sch. Bip. ex A. Rich. (1848) type: TU, Mt. Scholoda, PI. abyss. I, Schimper 41 (P hoio., BM K iso ). Gnaphalium schimperi v ar. strammeum Schweinf. & Asch. (1867), nom. nud.; Helichrysum schimperi var. straminea (Sch. Bip. ex Schweinf. & Asch.) Chiov. (1935) nom. nud. Intricately branched, perennial herb, or shrub with scram­ bling or semi-scandent branches, up to 2.5 m long; stem whitish- or light yellowish-tomentose. sparsely leafy throughout. Leaves 3-7.5 x 1-3.5 cm. dark olive-green and thinly floccose above, white cottony beneath, strongly 3-veined from petiolate base; petiole up to 1 2 cm long, whitish- or yellowish-tomentose; margins entire, undulate or serrate: apex acute. Capitula homogamous or heterogamous. cylindric. 3-4 x 1-2 mm, slightly fragrant, numerous, in dense paniculate cymes held together by dense white, pinkish or brownish woolly indumentum. Receptacle smooth, flat. Phyllaries 3-4-seriate. scarious, up (o 1.5-3.2 x 0.5-1 mm. concave and enclosing florets, white or pale yellow, often tinged pink, woolly at base; outermost small {c 1.5 x 1 mm), others subequal: stereome narrowly fenestrate or not. median vein clearly visible. Florets pale yellow, 4-5, 0-4 female. 0-4 bisexual, c 3 mm long. Corolla of female florets filiform-tubular, c 2 .2 mm long, 3-lobed. narrowed upwardly but expanded al base; corolla of bisexual florets cylindric, c 2.5 nun long. 5-lobed. lobes c 0.4 mm long. Stamens c 2.5 mm long; an­ thers c 1.5 mm long. Style c 2.5 mm long; brandies c 0.3 nun long. Cy pselas brown, glabrous, c 0.6 x 0.2 mm. ob­ long-tetragonal. Pappus white, c 2.5 mm long, barbellate, caducous. Margins of dry Juniperus-Podocarpus forest, mixed montane bushland, often becoming numerous in cleared areas, bushes along stream banks; 1750-3300 m. EW TU GD GJ WU SU AR GG SD BA HA; Sudan, E Africa, Socotra. Rep. Dem. du Congo, Rwanda, Malawi. Zambia. Zimbabwe. Friis et al. 1168; Mooney 8913; Mesfin T. 6606. 21. H. traversii Chiov. (1911) type: SU, ‘Scioa’, Mt. Entoto, Dec. 1886. Traversi s.n. (FT holo.) Scrambling or semi-scandent shrub with up to 5 m long drooping flowering branches, ora much branched c 75 cm high perennial herb or undershrub, usually growing in small gregarious clumps, aromatic; stem terete, densely appressed pubescent to tomentellous; hairs yellowish brown. Leaves ovate-lanceolate to narrowly elliptic. 10-50 x 5-15 mm, 3-veined from the base, upper surface

176

145. ASTERACEAE: 61. Helichrysum

Steep mountain slopes and gorges with bushes (Helichrysum, Senecio, etc.), and small trees (.Hypericum, Hagenia, Teclea, Olea, etc.); 2100-3350 m. GD SU AR KF GG BA HA; Kenya, Tanzania. Burger 1470; Jackson 708; Mesfin T. 8300. 11 22. H. hedbergianum Mesfin & Reilly (1995) type: SU, Addis Abeba, Mooney 4738A (K

holo.). Helichrysum fruticosum (Forssk.) Vatke var. latifolium Cufod. in Senck Biol. 47(4); 255 & 275 (1966) - type: GJ, Gogeb-Tal, zwischen Nebergedel und Debra Sion, 2600-2800 m; Kuls 76 (FR holo ).

Figure 107. H ELICH RYSU M H EDBERG IANUM 1 - part o f upper stem x 2/s; 2 - basal part o f stem showing larger leaves x 2 /3 ; 3 - capitulum x c 6; 4-6 outer, middle and inner phyllaries, respectively x 8; 7 - pappus bristle x 24; 8 - ray floret, without pappus x 12; 9 - disc floret, without pappus x 12; 10 - stamens x 18; 11 - style x 12. 1 from W. de Wilde & de Wilde 10960; 2-11 from F riiyetal. 1490. Drawn by Pat Halliday. (Reproduced with permission from Advances in Compositae Systematics , RBG, Kew, 1995, p. 437, fig. 22.)

light green, thinly arachnoid to densely tomentose, lower surface densely rusty or brownish tomentose, subsessile, or petiole up to 5 mm long; margins minutely serrulatecrenulate. Capitula campanulate, heterogamous, 3.5-4.5 (-5) x (2-)2.5-3 mm, sessile or subsessile, terminal in dense paniculate, cymose clusters. Receptacle fimbrillate, flat. Phyllaries 4-6-senate, bright yellow, scarious or chartaceous, 2.5-4 x (1.5—)2—2.2 mm, glabrous; outer­ most ovate or rounded, c 3 x 1 mm, light yellow or whit­ ish; innermost oblanceolate, c 4 x 1.5 mm, stereome not fenestrate. Florets brownish-orange to yellow, 7—13; 0-2(-3) female, 6-13 bisexual, c 3 .8 mm long. Corolla of bisexual florets tubular below, funnel-shaped in upper parts, 3-3.3 mm long, 5-lobed, expanded just below the lobes; lobes densely globulose-glandular. Anthers c 1.5 mm long. Style c 2.8 mm long, branches c 0.8 mm long. Cypselas grey, c 0.8 x 0.4 mm, papillose, otherwise gla­ brous, cylindric. Pappus whitish-yellow, c 3 mm long, barbellate.

Lianescent, perennial herb or shnib, aromatic, 0.35-2.5 m high; stem pale grey-green, densely branched, thinly appressed-tomentose. Leaves elliptic, 3-veined, dimor­ phic (those of main stem large, up to 40-52 x 9-12 mm, those of floriferous branches 20-30 x 3-7 mm), tomen­ tose on both surfaces with denser rusty hairs on the lower surface, diminished in size upwards. Capitula tubularcampanulate, heterogamous, 3.8—4 x 1.8-2.2 mm, on short bracteate peduncles held together in terminal corymbose cymes. Receptacle fimbrillate with funbrils overtopping ovaries. Phyllaries c 5-seriate, yellow, scanous, oblong to narrowly oblanceolate, glabrous ex­ cept for the sessile orange glands in middle, outennost 2.2-2.5 x c 0.5 mm, innermost 3.6-3.8 x 0.5-0.7 mm; stereome not fenestrate. Florets yellow, 9—14; 5-9 female, (3-)4-5(-6) bisexual 2.8-3.8(^1) mm long. Corolla of female florets 2 7-3(-3.5)mm long, filiform but narrowed further near the lobes; those of bisexual florets c 2.5—3(—3.5) mm long, tubular with 5 reflexed globose glandular lobes. Anthers c 1.6 mm long. Style c 2.5 mm long, bifid, branches c 0.5 mm long. Cypselas greybrown, c 0.8 x 0.5 mm, minutely papillose, otherwise gla­ brous. Pappus white, c 3 mm long, baibellate. Fig. 145.107. Mixed evergreen woodland with shrub-layer of Erica, Hagenia, Arundinaria, Maesa, Agarista, etc., clearings in open Juniperus forest, shrub-clumps along streams, dry sunny, rocky slopes in valleys, open hill tops; 1625-3050 m. GJ SU; not known elsewhere. PichiSermolli 977; W. de Wilde 5987; Friis et al. 1490. 23. H. foetidum (.L.)Moench. (1794); Gnaphaliumfoetidum L. (1753) - type: “Habitat in Aethiopia.” ?South Africa. Herb. Clifford: 402, “Gnaphalium 13”, fol. 8 [BM, lecto., selected by Hilliard & Burtt in Taxon 47: 361 (1998)]. Erect, sweetly scented, annual herb, 0.5-1.2 m high; stem usually branched only in the inflorescence, rarely a few from the base, glandular-pubescent or thinly tomentose in upper parts or in young plants, terete, faintly striate. Leaves oblong-lanceolate, cordate-amplexicaul, 40-95 x 5-22 mm, flat, both surfaces sometimes scabrous and glandular-pubescent, sessile, margins entire; apex acuteapiculate. Capitula heterogamous, depressed-globose, 5-10x5-15 mm, sessile or shortly pedunculate, several to numerous in crowded, corymbose, cymes. Receptacle

145. ASTERACEAE: 61. Hclichrysum

177

fla, smooth or areolate. Phyllaries 5-9-seriate, pale lemon-yellow to golden yellow or cream, shiny, leathery and rigid below, scarious and reflexed above; apex acute; outermost obovate, thinly woolly at base on outside, 4-7 x 1 5-2.3 mm; innermost 5.8-9 x 1.8-2.5 mm. spathulate, e reeding the florets and often spreading; stereome not ft nestrate. Florets pale yellow, c 300-1100; c 250-930 fe­ ll ale, c 20-190 bisexual, c 4 mm long. Corolla tubular but narrrowed above, 3.3-3.8 mm long. Anthers c 1.5 mm long. Style c 3 mm long, branches 0.5-0.7 mm long. Cypselas oblong, 0.5-0.7 mm long glabrous. Pappus white, 3 p-3.8 mm long, barbellate. In contrast to an earlier narrow circumscription of H. f : tidum, e.g., Hilliard and Burtt [in Bot. J. Linn. Soc. 82: IS - (1981 this species is found to be widespread in tn>aical Africa, occurring chiefly in disturbed parts in g r ssland, hill and mountain slopes, and forest and wood1; nd margins. It is composed of populations with large ( .. jeptacle 7 - 1 0 mm wide), intermediate (receptacle 4 - 6 n m wide) and small (receptacle c 2-4 mm wide) capitula wnh usually bright yellow phyllaries. Individuals or capitula with pale or yellowish white phyllarics liave also been known from the same general habitat of the species. H foetidum. while displaying many of the characteristic features of Helichrysum s.L differs from the rest of the species in the genus by possessing a liigh number of fe­ rn :le rather than bisexual florets. Hilliard and Burtt (1981 1 noted that in the eleven specimens they examined fn m South Africa, although the bisexual florets outnuinb^ red the female ones, there was a specimen in which the f male florets exceeded the bisexual by fifty. In spccin . ns with large capitula studied from Yemen. Ethiopia. Cameroun. Nigeria and Bioko, the range of the female flor Ls is c 8 0 0 -9 3 0 and that of the bisexual ones 4 1-c 160. I specimens with small capitula from Ethiopia. Burundi and Rep. Dem. du Congo, the range of the female florets is c 1 6 0 -3 8 0 and that of the bisexual ones 22-c 190. Fig 145.108.

I

v

Capitula 12-15 mm wide across the radiating phyllaries; leaves often glandular scabridpubescent on both surfaces: phyllaries 6.8-9 mm long. var foetidum Capitula 5—7(—10) nun wide across the radiating phyllaries: leaves often whitc-tomentosc beneath and glandular-pubescent on the upper surface: phyllaries 4-6 mm long. var. niicrocephalum

foetidum Helichrysum foetidum (L.) Mocnch var. macrocepholum A. Rich.. Tent. FI. Abyss. 1: 426 (1848) type: 'TU . Ambactcha, Quartin-Dillon s.n. (P syn ); TU. Mt. Scleuda. near Adwa. Ouartin-Dillon s.n. (P svn). H. foetidum (L.) Moench var. intermedia Chiov. in Ann. R. 1st. Bot. Roma 8: 177 (1904) - type: EW. Assaorta, Lake Minat to Urug Valley. Pappi 3328 (FT syn.); EW. Mcnsa, Mt. Ira. 2400 in.. Terracciano cCPappi 1440 (FT syn.).

Figure 108 H ELICH RYSU M FOETIDUM Habit Repro­ duced with permission from M anual o f the Flowering Plants o f Hawaii, vol. 1, 1990. PI. 28 p.p.)

Middle cauline leaves 90-95 x 15-22 mm. densely glan­ dular and scabrid pubescent on both surfaces. Capitula usually less than 50 per plant, c 12-15 mm across the radi­ ating phyllaries. Florets c 800—900; c 800-860 female, c 40-50 bisexual. Corolla c 3.7-3.9 mm long Phyllaries c 6.8-9 x 2.3-2.5 mm. Margins of cultivations, partly cultivated hill and mountain slopes, roadsides: 200-4000 m. EW TU GD WU SU KF BA HA; Sudan. Uganda. Kenya. Bioko. Ghana. Nigeria. Cameroun. Bprundi. South Africa. Ma­ deira. Tenerife; Saudi Arabia. Yemen: Spain. Mooney 8592; Schimper 332; Mesfin T. 8438.

var microcephalum .1. Rich.. Tent. FI. Abyss. 1: 426 (1848) type: TU. Mt. Soloda. near Adwa. PI. ab\ss. II. Schimper 927 (P holo.. K iso.). Helichrysum glutinosum A. Braun (1841). non (Hook, f.) Benth (1866); Gnaphalium foetidum I. var. glutinosum (A. Braun) Sch. Bip in Hot. Zen. 3 173 (1845) - type: Plant cultivated in Hort Carlsruhc in 1840 from seeds sent by Schimper from Ethiopia (locality unknown) (B holo.. K iso.). Helichrysum foetidum var. pallidum Less. (1832). nom. nud.. quoad Schimper 1508.

145. ASTERACEAE: 61. Helichrysur.i, 62. Pseudognaphalium

178

Gnaphalium buchingeri Schweinf. & (1867), nom. nud.. quoad Schweinfurth 2453.

Asch.

Middle cauline leaves 40-75 x 5-20 mm, glandularpubescent above, thinly white woolly beneath, rarely both surfaces glabrous. Capitula usually over 100 per plant, in dense, paniculate, corymbose cymes, 5—7(—10) mm across the radiating phyllaries. Florets c 280-300; c 250-275 female, c 20-30 bisexual. Corolla 3.3—3.5 mm long. Phyllaries 4-5 8 x 1.5-1.8 mm. Wooded grassland, margins of Erica arborea bush­ land, open Acacia scrub, mixed evergreen bushland. Eu­ calyptus plantations, margins of cultivation, disturbed places; 1500-3900 nv. TU GD WU SU AR IL KF ?GG SD BA HA; Cameroun Bioko, Sudan, Somalia, E Africa. Rwanda Burundi. Malawi, Zambia; Yemen Mesfin T. & Kagnew 1997; Mooney 5295; Friis et al. 1446

Imperfectly known and excluded taxa Of the names listed by Cufodontis (1966: 1109-1115) un­ der Helichrysum as occurring in the Flora area the fol­ lowing were found to liave been either transferred to other genera, or not occurring in the area, or names tliat cannot be identified from the protologue and for which no voucher specimens have been traced so far. Valid names that have not been reduced to synonvmv are shown in bold. H. bracteatum (Vent.) Andr. = B racteantha bracteata (Vent.) A. Andcrb. & Haegi (1991) H. conglobatum (Viv.) Steudel. (1840) This name, which is the correct name for a species from the Eastern Mediterranean and the Middle East, was re­ corded from the Flora area by Cufodontis (1966: 1109) following the determination in 1962 at Kew of a specimen from Kofele, Ethiopia. Mooney 8542. This specimen has not been traced. The present author lias examined speci­ mens of this species from North Africa but, so far. lias not found matching specimens from the Flora area.

As it is closely related to H. foetidum var. foetidum, the published account (Brenan 1954; Lisowski 1989) might result from an error in determination.

62. PSEUDOGNAPHALIUM Kirp. (1950) O.M. Hilliard in Bot. J. Linn. Soc. 82: 267-292 (1981). j Annual or perennial herbs. Leaves simple, alternate, lan­ ceolate. linear-lanceolate, oblanceolate or spathulate, ses­ sile and auriculate to semi-amplexicaul. or widely decurrent on stem, margins flat, crenulate. undulate orentire. woolly or glandular pubescent, often discolorous. Capitula urceolate or campanulate, heterogamous, in small clusters, 2.8-4 x 2-3.3 mm. Receptacle fimbriate or smooth and honey-combed. Phyllaries 3-5(-7)-seriate. equal to or slightly exceeding the florets, pale white, grey, light brownish yellow, light to bright yellow, stereome usually fenestrate, rarely undivided. Florets pale to bright yellow or yellowish green I6~cl30; 15-c 120 female; 1-10 bisexual. Corolla of female florets filiform and nar­ rowed upwards, of bisexual more or less uniformly tubu­ lar; lobes glandular hairy on outside. Anther-base sagittate, tailed. Style bifurcate at apex with truncate branches. Cypselas light brown, usually glabrous, rarely with duplex liairs or papillose. Pappus setae scabrid. api­ cal cells slightly inflated, base cohering by patent cilia A genus with over 40 species distributed in Africa Asia and America; 5 species in the Flora area. P. marranum (Philipson) Hilliard, which is known from Su­ dan. may also occur in the Flora area and hence it is in­ cluded in the key. 1

Small herbs, usually at most reaching 10-25 cm above ground; capitula in dense globose clusters: peduncles indistinct. 2 Large herbs, usually longer than 25 cm; capitula in loosely branched paniculate corymbose cymes peduncles evident. 4

2

Phyllaries dark-brown or grey at least in lower half capitula 2 8-3 x 2.3—2.5 mm. 4. P. melanosphacrum Phyllaries yellow, straw-yellow, or light brown ai base in some, capitula (3.5-)4-4.2 x (2.5—)3—3.3 mm 3

H. decorum au c t. non DC. (1838): Rochet d'Hericourt (1846), quoad Rochet d 'Hericourt 29 (?P). t

The specimen lias rio( been traced The species is other­ wise known from Southern Africa, but not from the Flora area.

3.

Leaves densely white-woolly throughout, silvery, linear-lanceolate; cypselas glabrous; phyllaries white or pale brownish-white. P. marranum Leaves sometimes greenish and thinly floccose above, not silvery, usually broadest near apex; cypselas papillose: phyllaries pale to bright yellow. ij I 5. P. luteo-album

4

Plant without glandular liairs or. if glands present plant not sticky ; leaves oblanceolate. broadest near the apex, sessile but not auriculate or decurrent, phy llaries and florets usually bnghi yellow. 5. P. luteo-album

H. rochetii Del. in Rochet d Hericourt (1846). nom. nud.. quoad Rochet d'Hericourt 25 (?P). The specimen lias not been traced. H. setosu m I1ar\ (1895) Cufodontis (loc.cit.) listed tliis name for Ethiopia follow­ ing published literature (Brenan 1954). A voucher speci­ men was not cited This species is known to occur in East. South-Central and South Africa but not in the Flora area.

145. ASTERACEAE: 62. Pseudognaphalium

-

Plant sticky from glandular hairs; leaves linear lan­ ceolate or elliptic, broadest at base or in middle, sessile and auriculate or decurient on stem; phyllaries and florets pale to straw-yellow. 5

5.

Erect bushy herbs, rarely more than 40 cm high; leaves erect-patent, linear often only up to 5 mm wide; female florets c 100-120; receptacle smooth. 3. P. oligandrum Tall herbs, up to 1.5 m high with main stem or branches often climbing, hanging or ascending through shrub-clumps; leaves spreading, lanceo­ late to elliptic, broadest at base or at middle, often 10-20 mm wide; female florets 15-25; receptacle fimbrillate. 6

-

6.

Leaves widely decurrent on stem, linearlanceolate, stem erect, stamen c 3 mm long.

-

Leaves sessile-auriculate, not decurrent; stem and/or branches often climbing, grow ing through shrub-clumps; stamens c 1.5 mm long. 2. P. richardianum

1. P. petitianum

1 P. petitianum (A. R ic h .)M e sfin (1992); G naphalium petitianum A. Rich. (1848) - type: GD. Ouodgerate, P etit s.n. (P syn.); SU. between Mayegouagoua & Debre Sina, P etit s.n. (P syn.). Ei ct, annual or short-lived perennial herb, c 1.2 m high; st< m branched in upper parts, sub-terete to sub-tetragonal, th :ily white-woolly, leafy throughout. Leaves linearlai ceolate, up to 9 x 1.5 cm, thinly white-tomentose be­ neath, green and thinly floccose-scabrid above, sessile an i widely decurrent at base, giving the upper part of the stt m and branches a winged appearance, margins entire, sli ntly revolute, apex acute. Capitula heterogamous. tubu ar 3.5^1 x 2.2-2.5 mm. sessile, or on short loosely cotto \ pedicels, many held together by loose white or cotto \ indumentum, in dense corymbose and often paniculate cymes. Receptacle fimbrillate. Phyllaries 4-5se ate, light yellowish-green, scarious in upper parts, le; r.hery below, 2.2-4 x 1-1.5 mm. apex obtuse; outer and m idle series ovate to oblong-ovate, woolly at base; inne most linear-oblanceolate. Florets yellow, 16~c 24; 15-22 female, 1-3 bisexual. 3.3-3.6 mm long. Corolla 3- i.2 mm long. Anthers c 1 mm long. Style c 2.2-3 mm lo ig. branches c 1 mm long. Cypselas obovate, grey, c 0.5 x .4 mm, papillose otherwise glabrous. Pappus white, 3- •• 3 mm long, barbellate. Degraded mountain grassland with Aloe, Vem onia, B t ctum, Rumex, etc.; 2800-2875 m. TU SU, not known elsewhere. P etit s.n; Sue E dw ards et al. 10\M esfin T. 8436. B urtt ( 1981); G napalium richardianum Cufod. (1966); G. auriculatum A. Rich. (1848), non Lam. (1788), nom. llegit. - type: SU, ‘Choa’, P etit s.n. (K iso.).

2 P.

richardianum {Cufod.) H illiard

A nual or short-lived perennial herb; stem erect or often gr iwing through and over shrub-clumps. up to 1.5 mlong. Lcaves oblanceolate, up to 8 x 2.5 cm, green above, softly grev -tomentose beneath, sticky, odourous, sessile-auricu­

179

late, margins entire to shallowly crenate. apex acuminate. Capitula heterogamous, tubular-urceolate. 3.5-4 x 1.8-2 mm. sessile or shortly pedicellate, in branched but com­ pact cymose clusters. Phyllaries ovate to oblanceolate, 3^4 x 1-1.5 mm, shiny, silvery-green, grey or brownish; outermost hairy all along through the middle: innermost hairy only at the middle. Florets pale to bright yellow, 26-30; 23-25 female, 3-5 bisexual, 1.8-3 mm long. Co­ rolla 1.8-2.2 mm long. Stamens 1.3-1.5 mm long; anthers c 1.2 mm long. Style 1.3-1.5 mm long, branches c 0.5 mm long. Cypselas obovate, light brown, c 0.6 x 0.3 mm. gla­ brous. Pappus white, c 2 mm long, baibellate. Margins of montane evergreen bushland with Erica, Hagenia, Arundinaria, Maesa, etc.; 2100-3200 m. EW SU AR KF BA; Sudan, Rep. Dem. du Congo. Friis et al. 1442; W. de Wilde 8975 and 8957. 3. P. oligandrum (DC.) Hilliard & Burtt (1981); Anaphalis oligandra DC. (1838); Gnaphalium oligandrum (DC.) Hilliard & Burtt (1976) - type: Madagascar, Bojer 13 (G-DC). Helichrysum steudelii Sch. Bip. ex A. Rich. (1848); Gnaphalium steudelii (Sch. Bip. ex A. Rich.) Schweinf. & Asch. (1867) - type: TU. Adwa, PI. abyss. I. Schimper 231 (P holo., BM K iso.). G. undulatum auct., non L. (1753): Cufodontis (1967). Erect, bushy annual, or short-lived perennial herb, 30-50 cm high; stem densely branched from the base, grey or whitish tomentose. Leaves linear-lanceolate, erect-patent, bright green above, grey or white-tomentose beneath, stick>', 30-50 x 3—5(—7) mm, lower leaves often withering or turning light brown upon drying, sessile, margins en­ tire, revolute, apex acute. Capitula heterogamous, cam­ panulate, 3-3.5 s c 3 mm, in dense, terminal, corymbose clusters. Receptacle epaleate, flat. Phyllaries 6-7-seriate, diminishing in size inwards, firm; outer and middle series narrowly elliptic with acute-acuminate apex, c 3.5 x 0.8 mm. incurved at base, slightly spreading or up-turned at apex, pale-white, loosely cottony up to middle; innermost membranous, c 2.5 x 0.5 mm, oblong-elliptic. Florets pale brownish-yellow, c 110-130; c 105-120 female. 7-10 bisexual, c 2.2 mm long. Corolla tubular, 2-2.5 mm long. Anthers c 1 mm long. Style c 2 mm long, branches c 0.3 mm long. Cypselas grey, c 0.4 x 0.2 mm, minutely papillose, glabrous. Pappus white, c 2 mm long, barbellate. Grassland, banks and mounds around cultivations, open bushland along mountain slopes; 2100-3300 m EW TU GJ WU SU SD, E Africa. Rep. Dem. du Congo, Rwanda. Angola. Malawi, Zambia, Zimbabwe. Mozam­ bique, Botswana, Namibia. South Africa, Madagascar. Mesfin T. 6401; Ash 2607; W. de Wilde 8234. 4. P. m elanosphaerum (Sch. Bip. ex A. Rich.) Hilliard (1981); Gnaphalium melanosphaerum Sch. Bip. ex A. Rich. (1848) - type: GD, Simen Mts., near Enschedcap. PI. abyss. II, Schimper 1360 (P holo., BM K iso.).

180

145. ASTERACEAE: 62. Pseudognaphalium, 63. Chiliocephalum

Cypselas light brown, c 0.6 x 0.1 mm, densely papillose. Pappus white, 1.7-1.8 mm long barbellate. Open grassland, grassy bank of water course, bare soil beside rivers, degraded montane forest with remnant/lcflcia, Olea and planted Eucalyptus, etc.; 1500-3000 m. EW GD SU BA; not known elsewhere. Sue Edwards 2144; Mooney 6414; Mesfm T. 8312. 5. P. luteo-album (L.) Hilliard & Burtt (1981); Gnaphalium luteo-album L. (1753) - ty pe. “Habi­ tat in Helvetia, G. Narbonensi, Hispania, Lusitania ” Heib. A. van Royen, No. 900.286-294 [L lecto., se­ lected by Hilliard & Burtt in Bot. J. Linn. Soc. 82: 206, 244(1981)1. subsp hiteo-a!bwn

Figure 109 PSEUDOGNAPHALIUM LUTEO-ALBUM 1 habit x Vi, 2-4: outer, middle and inner phyllaries x 8 ,5 - female floret x 12, pappus in outline; 6 - bisexual floret x 12, pappus in outline; 7 - stamens with tailed anthers x 24; 8 - style x 24; 9 pappus bristle x 12; 10 - cypsela x 24; 11 - part o f a capitulum. longitudinal section. All UomJulien in M AU 15484. Drawn by Pat Halliday. (Reproduced from FI. Mascareignes. 109. Com poshes, PI. 25.)

Annual or short-lived perennial heib, stems solitary or several, prostrate or ascending, rarely erect and up to 10 cm high; glabrous or glabrescent below, slightly woolly in upper parts, obscurely striate. Leaves linear to spathulate, sessile-amplexicaul with mid-vein prominent below, 10-25 x 2-3.5 mm, tomentose on both sides, margins en­ tire to undulate, apex obtuse to subacute. Capitula hetero­ gamous, campanulate. 2.8-3 x 2.3-2.5 mm. sessile or shortly pedicellate, in dense, globose, terminal and subterminal clusters, sometimes some of the lower leaves overtopping the clusters of capitula. Receptacle epaleate, flat. Phyllaries 3(-4)-seriate. subequal, 2.5-3 x 1—1.5 mm, with a translucent band between the lowermost green part and the upper dark-brown or grey part; outermost ovate-oblong, dark-brown and woolly near base, c 2.5 x 1 mm; middle series oblong-oblanceolate, c 3 x 1 mm, dark-brown in middle and greenish below; innermost nar­ rowly oblanceolate. c 2.5 x 1 mm, daric-brown near apex with a green band from middle to base. Florets green­ ish-yellow, c 35^45; c 30-35 female, 4-10 bisexual, 2.4-2.5 mm long. Corolla 1.7-1.8 mm long. Anthers c 0.8 mm long. Style 1.7-1.8 mm long, branches c 0.3 mm long.

Erect annual or short-lived perennial herb, up to 60 cm high, usually with several ascending branches from the base, or partly trailing on the ground before turning up­ wards: stem white-woolly throughout with denser wool near inflorescence, striate, subterete. Leaves oblongoblanceolate to spathulate, sometimes linear. 10-60 x 5-12 mm. diminishing in size upwards, sessile and semiamplexicaul, densely woolly tomentose on both surfaces, especially on lower side, sometimes thinly floccose above or even glabrescent. margins entire or indistinctly erenulate, apex obtuse or acute with sharp mucro. Capitulaurceolate. heterogamous. 4-4.2 x 3-3.3 mm, ses­ sile or shortly pedicellate, in crowded corymbose or cymose clusters at the ends of the stems and brandies, and from the upper axils, often held clear of the foliage. Re­ ceptacle epaleate, flat. Phyllaries 6 -7 -seriate, diminishing in size inwards, 1.5-3.5 x 0.5-0.9 mm, bright golden yel­ low to pale yellow with sometimes brownish blotches at middle or at base, glabrous except at base in the outer­ most; outermost c 3.5 x 0.9 mm, ovate-lanceolate, green­ ish below, scarious in upper parts, innermost c 1.5 x 0.5 mm. translucent. Florets yellow, c 90-120; c 85-110 fe­ male; 6-10 bisexual c 3.5 mm long. Corolla c 3 mm long. Anthers c 1.3 mm long. Style c 2.5 mm long, branches c 0.5 mm long. Cypselas light-brown, papillose, c 0.5 x 0.2 mm. Pappus white, c 2 mm long. Fig. 145.109. Roadsides, wasteland, margins of cultivations, dis­ turbed banks of rivers and water courses; 1500-2650 m. EW TU GD GJ SU AR XL KF SD BA; a cosmopolitan weed, in Africa found from west Africa to Somalia, and south to South Africa Mesfin T. 7951; Burger 541; Moo­ ney 8933. Subsp. affine (D. Don) Hilliard & Burtt occurs in Ne­ pal and adjacent areas.

63. CHILIOCEPHALUM Benth. (1873) Mesfin Tadesse in Kew Bull. 47: 131-135 (1992). Perennial herbs; stem erect or decumbent, sparsely to densely woolly, rarely glabrescent Leaves simple, alter­ nate, erect-patent to spreading, slightly to strongly

181

145. ASTERACEAE: 63. Chilioccphalum

110. CHILIOCEPHALUM SC H IM PERI 1 - part o f plant x 1 / 3 ; 2

Figure

T

\fr. \\

ED

*

r-

- corolla o f female floret x 26; 3 - style o f female floret x 26; 4 - bisexual floret x 26; 5 - style o f bisexual floret x 26; 6 stamen x 26; 7 - cypsela o f female floret, ventral face left, dorsal face right x 26; 8 - cypsela-wall in cross-section x 170; CU = cuticle, EP = epidermis, EM = embryo. ED = endosperm. T = testa. VB = vascular bundle; 9 capitulum with protruding florets x 13; 10 phyllaries, outer above, inner below x B . 1-9 from Schimper 890; 10 trom Anderberg 1629. Drawn bv A. Anderberg. (M odified and reproduced with permission from Bot. Jahrb. Sysl. 110(1). 1988. p. 4. fig. 1.)

isexual. Corolla of female florets filiform. 3-lobcd. those bisexual florets 5-lobed. tubular. Anther base sagittate, ■ityle bifurcate, branches obtuse. Cypselas terete, oblong. ;iabrous. Pappus absent. A genus with only 2 species, both confined to highands in the Flora area. Erect herbs; leaves, especially lower ones, long decunent on stem, blade often up to 50 x 15 mm; phyllaries thickish or firm (at base), oblong, often with dark brown glandular liairs at middle, apex obtuse. 1. C. schimperi Stem procumbent or usually mat-forming; leaves sessile, not decurrent, blade of middle leaves up to 35 x 7 mm: phyllaries thin throughout, ob­ lanceolate to oblong-elliptic, whithout glandular liairs, apex acute. 2. C. tegetum

1. C. schimperi Henth. (1873); Helichrysum chiliocephalum Hilliard & Burtt (1973) - type: GD. Siincn Mts.. 1853. Schimper 890 (K holo., E S iso., not seen). Kralikio mvriocephala Sch. Bip. ex Schweinf. & Asch. (1867). nom. nud. K. schultzii Sch. Bip ex Schweinf. & Asch. (1867). nom. nud. Erect annual or short-lived perennial herb. 50-75 cm high: stem unbranched except in the inflorcsccnces. sub­ glabrous in lower parts, woolly to floccosc 111 upper parts. Leaves alternate, erect-patent to spreading, up to 5 x 1.5 cm. oblong-lanceolate, with 3 veins (prominent above), green and sparsely pilose above, pubescent to greyishwhite tomentose beneath, upper leaves clasping stem to shortly decurrcnt. lower leaves distinctly long-decurrent 011 stem; margins entire; apex obtuse, mucronulate. Capitula numerous. 3—3.5 nun long, all held together by woollv indumentum, forming flat tenninal corymbose cymes: cyme branches also matted together. Phyllaries subequal, shiny and translucent, pale yellowish to brownish, indis­ tinctly 2-seriate. 1.5-2 x 0.5-1 111111. glabrous above, glan­ dular-pubescent below: stereome undivided; margins

182

145. ASTERACEAE: 63. Chiliocephalum, 64. Gnaphalium

2. C. tegetum Mesfin (1992) - type: BA, Harenna forest, Kecha, 6° 42.5 ■N; 39° 44’ E, Mesfin T. 5649 (ETH holo., K iso ). Perennial herb with spreading or matforming stems; stem up to 30 cm long, floccose or woolly, rarely glabrescent. Leaves alternate, 3-veined from near the middle of blade, erect-patent to spreading, 15-35 x 4-7 nun, narrowh el­ liptic to oblanceolate; green and glabrous to thinly arach­ noid above, densely tomentose to woolly beneath, sessile with slightly decurrent base but not forming distinct wing on stem, margins entire, apex acute-mucronate. Capitula numerous. 2.5-3 mm long, all held together by woolly indumentum forming flat terminal corymbose cymes; cyme branches also matted together. Phyllaries 9-12 per capitulum, subequal, indistinctly 2-3-seriate. 1.8-2.6 x 0.5-0.8 nun. pale yellow' to greenish, membranous, trans­ lucent with numerous indistinct greenish striae, ob­ long-elliptic to oblanceolate with loosely matted hairs from base to middle, rarely glabrous; margins denticulate in upper parts Florets 8-11, yellow; outennost female, bi­ sexual at center Female florets 7-9. some strongly un­ equal in size. 2.2-2.5 mm long; corolla filiform or tubular, same width as ovary. 1—1.2 mm long, translucent. 3—(-lobed with gland tipped lobes; style 0.8-1 nun long, terete, bifurcate at apex, branches c 0.3 nun long, truncate; ovary 0.2-0.4 mm long, glabrous, indistinctly striate. Bi­ sexual florets 1-2; corolla tubular but wider than ovary. 2 1-2.3 mm long, often shorter than female florets; sta­ mens c 1 mm long, filament c 0.2 mm long; anthers c 0.8 mm long, base tailed, connective acute. Cypselas c 0.4 mm long, elliptic. 4-5-veined, glabrous. Pappus absent. Fig 14 5.111.1-10.

Figure 111 CH IU O CEPH ALU M TEGETUM 1 - habit x 2 leaf node x I 3 - capitulum x 9; 4-6: outer, middle and in­ ner phyllaries (m agnification not provided in original) x 9; 7 female floret x 9. 8 - bisexual floret x 9; 9 - anther x 18. 10 style tip x 18. C SCH IM PERI. 11 - outer phyllary x 13. 1 & 2 from Mesfin 5649,3-10; from J. de Wilde mes. cottom Receptacle epaleate. flat or conical. Phyllanes multiseriate: outer ovate-lanceolate, loosely cottom up to th e middle, inner elliptic-ovate, apex apic j late. margins membranous. Outer florets numerous, 12 male. corolla filiform. Central florets few, bisexual, fer­ tile or stenle; corolla tubular, anther base sagittate; st> le bifurcate with truncate apices. Cypselas obovate to subterete. Pappus of equal bristles. A genus with about 50 species in Europe. Asia and North Africa. A few species, including F. vulgaris Lam., are widespread weeds Cufodontis (1967:1104) recorded it as occurring in Etluopia. So far no specimens have been found from the Flora area that would match the desenption of this species. But as it is a widespread weed it is briefly described and a key to distinguish it from the na­ tive species is provided. I

Figure 112 GNAPHALIUM RUBRIFLORUM 1 - habit x Vi; 2 - capitulum x 8; 3 - bisexual floret x 20; 4 - cypsela x 40. 1-3 from Auquier 3556.4 from Lebrun 9303 (Reproduced with per­ mission from Flore d'Afrique Central, part 2. p. 63, PI. 11.)

yellowish base; innermost about as long as the florets, pale-brown or buff. Female florets c 80-100, bisexual 2-3, whitish with red-purple tips. Cypselas c 0.5 mm long, papillose, mucilaginous when wet. Pappus setae copious, scabnd, fused at the base into a smooth ring, caducous. Roadside ditches, weed in gardens and farms; 9601370(-?2440) m. EW EE TU ?SU, native in the warmer parts of N America, ranging down the east side of S Amer­ ica to Argentina. Widespread as a weed in Australia, New Zealand, Iraq. India. Sri Lanka, China. Hawaii, Mauritius & S Africa (Natal. Transvaal, Transkei). Mesfin T. & Sebsebe D. 3984, Ryding & Seleshi N. 1694; Mesfin T. 8691. Material of this taxon is sometimes also named G. pennsylvanicum Willd. (1809), the name adopted by Hilliard for FI. S. Afr. 33: 28 (1983). The species has sometimes been refeued to as Gamochaeta purpurea auct., non (L.) Cabrera (1961). or Gnaphalium purpureum auct, non L. (1753), e.g., by Oliv. & Hiem, FI. Trop. Afr. 3: 345 (1877).

Leaves narrowly spathulate to round with the wid­ est part toward the tip. upper ones exceeding the clusters of capitula. 1. F. abyssinica Leaves elongate-lanceolate with the broadest pan toward the base, upper ones not exceeding the clusters of capitula. 2. F. vulgaris

1 F. abyssinica Sch. Bip. ex A. Rich. (1848); F. germanica L. var. abyssinica (Sch. Bip. ex A. Rich.) Penzig in Atti Congr. Bot. Genoa 1892: 345 (1893); F. vulgaris Lam. var. abyssinica (Sch. Bip. ex A. Rich.) Cufod. in Bull. Jard. Bot. Etat Brux. 36, SuppL. 1104 (1966) - type: TU, near Adwa. PI. abyss. III. Schimper 1913 (P holo.. K iso.. G GOET TUB). F. pyramidata auct.. non L. (1753): Schweinf. & Asch. (1876). F. evacoides Chiov. (1940) - type: SU, Addis Abeba. Senni 1732 (FT holo.). Erect, annual herb, up to 15 cm high; stem densely whitewoolly, 1-2 mm wide, dichotomously branched. Leaves spathulate to oblanceolate, sometimes oblong-linear 5-25 x 1-5 mm. tomentose on both surfaces, hairs denser on lower surface, sessile; margins entire, apex obtuse to acute-apiculate Capitula heterogamous. campanulate. sessile, in dense globose axillary or tenninal cymes. Phyllaries light-green, with scarious margins, sometimes pink-tinged at apex and deep red at base, c 3 x 1.5 nun, al­ ternate, apices long acuminate, loosely cottony up to mid­ dle. Florets pale yellow, c 2.5 mm long. Cypselas grey-green, densely white papillose, c 0.5 x 0.2 mm. Pappus white, c 2.5 mm long, barbellate. Fig. 145.113 Cultivated fields on coarse sandy soil; open gravelly montane plateau; 500-3300 m. EW TU GD SU, Sudan. Sue Edwards et al. 3848; Mesfin T. 8596; Friis et al. 6687. 2. F. vulgaris Lam. (1778) - type: from France.

J45. ASTERACEAE: 65. Filago.66. Bracteantha. 67. Osteospermum

F. germanica L. (1763), nom. illeg., non Huds. 1762) - type: “Habitat in Europa.” “Gnaphalium” in Fuchsius. Hist. Stirp., 222, 1542, lecto.. selected by Wagenitz in Willdenowia 4: 48 (1965). Erect or decumbent, annual or short-lived perennial he:b, up to 30 cm high; whole plant densely woolly orcottoi v stem terete. Leaves linear or linear-spathulate, ere jf-patent or closely appressed to stem, especially at ba e, sessile, sub-amplexicaul. 4-12 x 2-3 mm. margins en :re; apex acute. Capitula campanulate, sessile, numerou m dense globose cymes. No specimen was examined from the Flora area. (See nc c under the genus.)

66. BRACTEANTHA* Anderb. & Haegi (1991) A .Anderberg & Haegi in Opera Bot. 104: 102-105 (1991). B i;nnial or perennial herbs, forming ectomycorrhiza. L aves sessile, glandular-hairy. Capitula disciform, soli­ tary or few together. Receptacle epaleate. flat. Phyllaries yellow, wlute, reddish or pinkish, opaque. Outer florets yellow, female, filiform, fewer than the central florets. C :rural florets bisexual. Cypselas more or less quadran­ gular with 2-3 vascular bundles, glabrous. Pappus of free, b irbellate bristles.

or sessile. Capitula radiate, heterogamous, solitary or corymbose, subsessile or shortly pedunculate to scapose. Phyllaries 1-2-seriate, subequal, imbricate. Receptacle epaleate. Ray florets female or rarely neuter, yellow, white, pink, mauve, purple, or blue. Disc florets bisexual or functionally male, yellow, orange or reddish. Corolla actinomorphic, funnel-shaped, 5-lobed. Anthers ecalcarate, caudate, connective ovate-triangular, obtuse or acute-acuminate. Style sterile or fertile, unbranched or shallowly bifid with a short apical collar of hairs, or deeply bifurcate with liairs extending down below the branches. Cypselas mono- or polymorphic: ray cypselas terete or triangular to 3-winged, smooth tuberculate, aculeate or beaked, straight or curved; disc cypselas, when present, laterally flattened with thickened margins, obovate. orbicular or elliptic. Pappus absent. A tribe with 8 genera and about 110 species distrib­ uted mainly in S Africa tropical and N Africa Atlantic Is­ lands, southern and central Europe, south-west and West Asia, east to Afghanistan. 2 genera (one indigenous) with 3 species in the Flora area. 1.

-

A genus with 5 species native of Australia, now natural zed elsewhere. One species is cultivated as an omamental plant in the Flora area

E reel. biennial herb, up to 1.2 m high; stem branched in ipper parts, scabrid. Leaves oblong-lanceolate. 5-12 x t 6-1.5 cm. shortly petiolate or narrowed at base into a i tnged petiole, glabrous and densely resinous-punctate, i upitula solitary and terminating main stem and branches, i r a few in open cymes. 2-7 x 0.7-1.2 cm; peduncle up to 2 cm long, pubescent. Receptacle flat, honey-combed, liy llaries multiseriate, leathery below, scarious. stiff and Winy above, golden yellow to red. lengthening inwards; uiermost 6-10 x 5-7 mm. sub-orbicular, obtuse at apex; nnermost up to 15x5 mm. lanceolate, acute at apex. Flo•ris yellow, numerous. Cypselas light-brown, glabrous. ! 3-2.5 x 0.7-0.9 mm. slightly tetragonal, longitudinally iireaked. Pappus setae yellow, c 7 mm long, sub-plumose w ith longer hairs near apex. Culm ated ornamental; 2000-2400 m; SU HA; native :>f Australia. Mesfin T. 8343; IECAMA 95; Friis et al. 9310.

Tribe 9. C A L E N D U L E A E Cass. (1819) Herbs or subshrubs. Leaves simple, alternate, glabrous, dandular or pubescent, margins entire or serrate, petiolate

Cypselas straight, glabrous or verrucose, 3-winged; sty le bifurcate with distinct stigmatic areas; indigenous plants. 67. Osteospermum Cypselas incurved to various degrees and tubercu­ late on the back, inner distinctly curved; style un­ branched or slightly bilobed; ornamental plants. 68. Calendula

67. OSTEOSPERM UM L. (1753)

I .bracteata (Vent.) Anderb. & Haegi (1981); Xeranthemum bracteatum Vent. (1804); Heli­ chrysum bracteatum (Vent.) Andr. (1805) - type: from Australia.

185

T. Norlindh in Studies in the Calenduleae, I: Monograph of the genera Dim orpholheca , Castalis , Osteosperm um , Gibbaria and Chrysanthem oides , Gleerup, Lund (1943); & in Svensk Bot. Tidsskr. 48: 148-154 (1954) & in Bot. Not. 113: 385-399 (1960). Herbs, subshrubs or shrubs. Leaves alternate or basal, cauline ones nearly opposite, entire or pinnatifid. Capitula radiate, heterogamous. solitary or corymbose, pedun­ culate. Receptacle epaleate. Phyllaries greenish with scarious margins. 1-3-seriate. Ray florets yellow or vio­ let. female and fertile. 1-seriatc. style bifurcate, branches filiform with distinct stigmatic surfaces. Disc florets yel­ low. functionally male or sterile. Corolla tubular. 5-lobed. Anthers sagittate, not tailed at base. Style, when present, shortly 2-lobcd. Cypselas terete or 3-angled, smooth or tu­ be rculatc A genus with about 70 species distributed from S Af­ nca to Arabia; 2 species in the Flora area. 1.

-

Plants annual; capitula in open, branched corymbs; lower and middle stem leaves pscudopetiolate. ovatc-lanccolatc in outline, margins grossly sinuate-dentate to pinnatifid. teeth or slender lobes up to 10 mm long. 1. O. vailliantii Plants perennial; capitula solitary: lower and mid­ dle stem leaves sessile, oblong-oblanceolate. margins denticulate to entire. 2. O. monocephalum

186

145. ASTERACEAE. 67. Osteospermum

Figure 113.

OSTEOSPERM UM V A ILLIA N TII 1 - upper part o f plant x '/2; 2 - phyllary x 5!/*; 3 - cypsela x c 2'/j. O. MONOCEPHALUM : 4 -9 : leaves x '/a , 4 and 5 normal forms, 6 and 7 luxuriant forms from damp grassland near a river, 8 and 9 depauperate forms from burnt grassland, 4, 6 and 8 low er and 5, 7 and 9 median stem leaves; 10 - phyllary x 11 cypsela x c 2V2. 2 and 3 from Norlind>i s.n. (c u lt); 4, 5, 10, 11 from Fries. Morlindh and Weimarck 3300a, 6 ,7 from Fries et al. 3737, 8, 9 from Fries et al. 3418. 1 draw n by E.M. Tweedie, 2-11 by S. Ekblom. [ 1 reproduced by permis­ sion o f the author from A gnew, A.D.Q., Upland Kenya Wild Flowers, 1974, p. 487; 2-11 reproduced from Norlindh. T.. Studies in the Calenduleae, I: Mono­

graph o f the genera Dimorpholheca. Caslalis, Osteospermum. Gibbaria and Chrysanthemoides (1943), figs 28 & 30

/\ V ■; W 5w

1. O. vailliantii (Decne) T. Norl. (1943); Tnpterts vailliantii Decnc. (1834) - type: Egypt. Mt Sinai. Bove s.n. (P holo ). T. cheiranthifolia Sch Bip (1841). nom. nud . quoad Schimper 1480. T. cuneifolia Sch. Bip. ex Schweinf. & Asch. (1867). nom. nud. T. rueppellii Sch. Bip. ex Schweinf & Asch (1867). nom. nud. Erect, annual herb. ( 10-)25-75 cm lugli. Stem terete, stiff, scabrous, leafy and often branched below the infloresccncc. woody at base, up to 15 nun in diameter, strialesulcate. Leaves alternate. or those al base opposite, simple to pinnatifid. subglabrous to densely glanduJar-pubcscent. membranous, lower ones obovate. oblanceolate to elliptic, blade narrowed below into a w inged petiole. 3-15 x 1-4 cm. upper oblong-lanceolate to narrow ly lanccolatc. sessile with seini-amplcxicaul or auriculate base, margins dentate or grossly sinuate- dentate to pinnatifid. teeth or slender lobes up to 10 mm long, rarely subentire. Capitula radiate, urceolate or campanulate. 15-25 x 5-7 mm at anthesis. in open, terminal branched cory mbs, nodding in fruit Peduncle densely glandular-pubescent. 2-10 cm long, bracteate. Receptacle flat, c 3 mm wide. Involucre 3-5 x 3-4 mm. Phyllanes inegularly 2-seriate. 8—13, linearlanceolate. 4-5 x 1—1.5 nun, green with scanous margins, apex acuminate, sparsely glandular-pubescent on outside Ray florets yellow, 8-13,10-20 mm long. Disc florets yel­

P Pi

fv\ \vvl T 6 \i

low. Corolla c 4 nun long, tubular below, limb campanu­ late. 5-lobcd Anthers c 1 mm long. Style bifurcate, stig­ matic areas marginal Cypselas yellowish brow n. glabrous or vcrrucose. fusifonn (excl. wings), 7-10 x 1-1.5 mm (excl. wings). 3-wingcd. wings pellucid-scarious. 1.5-2.5 mm wide, extending and overarching the apex. Fig

145.113.1-4. Open places in Juniperus forest. Acacia-Fuclea scrub on dark clay soil. Acacia-Commiphora deciduous bushland, degraded sites in Acacia bushland or succulent bushland; 1300-2800(-3400) m. EW TU WU SU AR SD BA HA; Egypt. Sudan. Somalia. Djibouti. Uganda. Kenya and Tanzania. Friis et al. 3746.6727; Mesfin et al. 7186 2 O. monoccphalum (Oliv. c£ H iem ) T. Norl. (1943); Tripteris monocephala Oliv. & Hiem (1877) type: Mozambique. Manganja Hills. 1861. Metier s.n. (K. holo ). Perennial herb with one to several erect stems from a subterranean, strongly lignified rot apex. 25-35 cm high. Stem solitary or rarely branched, striate-sulcatc. sparsely hairs Lower leaves narrowly elliptic, opposite, sessile or subsessile. entire or dentatc-dcnticulatc. 20-40 x 4-10 mm. surfaces sparsely pilose with denser hair on mid-vein and margins, upper leaves alternate, slightly amplexicaul. linear or linear-elliptic Capitula radiate, solitary at apices of stems and branches, campanulaate.

187

145. ASTERACEAE: 67. Osteospermum, 68. Calendula

2 -50 x 5-7 mm at anthesis, fading quickly, pi aunculate. Peduncle 1-3 cm long, bracteate, densely g indular-pubescent. Receptacle c 5 mm wide. Involucre 8- 10 x 5-6 mm. Phyllaries sub-biseriate, linear-elliptic w ith narrow scarious margins, 5-6 x 7-10 mm at ai inesis, sessile-glandular on outer surface. Ray florets c 1 . yellow. 20-25 x 3-4 mm, margins of tube densely white pubescent. Disc floret corolla yellow, glabrous, 5 lobed. Anthers c 1 mm long. Style bifurcate, stigmatic aj ;as marginal. Cypselas light brown, glabrous or sparsely ciliate toward apex, oblanceolate (excl. wings), li -11.5 x 2-2.5 mm (excl. wings), 3-winged; wings danc-margined, 3-3.5 mm wide, extending and overarching the apex. Fig. 145.113.5-12. Open grassland with scattered Combretum molle and oiittr trees and occasional thickets with sparse grass c c . er: c 1850 m. SD; Tanzania, Zam bia Zimbabwe, Mo­ zambique, Malawi,-/Rep. Dem. du Congo and Angola. GiIbert & Ermias 8431 (ETH); Gillett 14615 (FT). In Africa this species has earlier been known from southern Tanzania and south to Zimbabwe. The recent find fron) southern Ethiopia i.e. “Sidamo. 10 km SW of K i:re Mengist on road to Negelle”, extends its range iii rhwards and the species probably occurs also in similar h; bitats in Kenya. According to Norlindh (1943), Osteospermum monoct -halum is distinguished from O. vailliantii by its “large c} rselas, w hich are 13-17 mm long excluding the wings, ain: by the lowermost and often also the median leaves bein^ opposite.” Also the “stems are simple or sparingly br inched, and the heads are solitary (hence the specific ep ihet monocephalum) at apices of the stem and of the branches The cypselas in the single specimen exam­ ined from the Flora area are slightly smaller, but all other feidures of the specimen fit the description of the species g i . en by Norlindh (loc. cit.).

68. CALENDULA* L. (1753) Annual, or perennial, herbs or subshrubs, often glandular oi aromatic. Leaves simple, alternate, sessile or shortly p< isolate, entire, shallowly toothed or lobed. Capitula radi lie, heterogamous, solitary, terminal, pedunculate. Invc lucre campanulate or hemispheric. Receptacle flat, ej ueate. Phyllaries 2-seriate, herbaceous and partly sci Jious-margined. Ray florets female, fertile, yellow or oi mge; style bifurcate, branches linear. Ray cypselas pi yinorpluc, incurved to various degrees, tuberculate on th back; outer cymbiform, rostrate or triquetrous; inner di unctly curved, sometimes winged; innermost nearly ar nular and transversely rugose. Pappus absent. Disc flore .i bisexual, functionally male, yellow, orange, yellov. -red or brownish Corolla tubular, 5-lobed. Anthers ec; idate. Style unbranched or slightly bilobed, apex acme, conical, surrounded by a ring of short liairs. A genus of about 15 species distributed chiefly in the Mediterranean region. North Africa, Macaronesia, south­ ern and central Europe, Turkey, Iraq, Iran and Yemen.

The following 2 species occur in Ethiopia; one widely cul­ tivated as an ornamental plant, and the other only recently recorded as a weed. 1.

Capitula 2-4 cm in diam., nodding at maturity; fruit strongly incurved; leaves lanceolate. 2.

Capitula 5-10 cm in diam., erect at maturity; fruit crescent-shaped; leaves oblong to oblongobovate. 1. C. officinalis

1. C. officinalis L. (1753) - ty pe: “Habitat in Europae arvis.” Linnaean Her­ barium No. 1035.4 [LINN lecto., selected by Alavi in Jafri & El-Gadi (ed.), FI. Libya 107: 195 (1983)j. Annual herb, 30-75 cm high. Stem erect or procumbent, pubescent. Leaves oblong to oblong-obovate, 2-6 x 1-2 cm. margins entire or minutely and remotely denticulate, sessile with clasping base. Capitula showy, solitary, on stout peduncle, 3-6 cm across the rays. Phyllaries 2-seriate. linear-lanceolate, graded monomorphic, mar­ gins scarious, apex acuminate. Ray florets female, fertile, yellow or deep orange, closing at night. Disc florets func­ tionally male, yellow or orange. Cypselas heteromorphic, outer narrowly beaked, inner smaller, strongly curved, transversely rugose-tuberculate on outer surface. Pappus absent. Fig. 145.114. Cultivated in gardens in the larger towns of the Flora area; 2300-2600 m. EW (Asmara) SU (Addis Ababa) HA (Harar). Cultivated almost throughout the world. Anonymous (Students o f Asmara University) 24; Mesfin T. 5837. 2. C. arvensis L. (1763) - type: “In Europae arvis.” Loefling s.n in Linnaean Herbarium No. 1035.1 [LINN lecto., se­ lected by Heyn& al. in IsraelJ. Bot. 23: 182 (1974)]. Similar to C. officinalis except for the small size of the florets and the following characters: stem slender, up to 20 cm high, finely pubescent; leaves lanceolate; capitula 2-4 cm wide, nodding at maturity; fruits strongly incurved, almost forming a circle, with many prickles on back; inner prickles blunt; outer prickles sharp. Weed in farmland; 2450 m; TU; widespread in central Europe, the Mediterranean area eastwards to Afghanistan and India also in Australia, S Africa and N and S Amer­ ica. Friis et al. 10544.

T ribe 10. A S T E R E A E Cass (1819) Perennial heibs or shrubs, rarely annual herbs. Stems and/or branches erect, ascending, scandent or straggling. Leaves alternate, simple to pinnatisect. Capitula hetero­ gamous, radiate or disciform, campanulate to hemi­ spheric. Phyllaries one- to several-seriate, herbaceous or chartaceous. imbricate, usually lengthening inwards, mar­

188

145. ASTERACEAE: 68. Calendula

Figure 114.

CALENDULA O F FIC IN A LIS 1 -

upper part of stem x Vi; 2 - phyllary x 5'/4; 3 - ray floret corolla and style x 2 Vi; 4 - style of ray floret x 10V4; 5 disc floret, without ovary x 8l/ r 6 stamens x 9; 7 - style of disc floret x 9 , 8 - outer cypsela, side view x p i ; 9 - inner cypsela x 2Vi; 10 - inner cyp­ sela, side view, 11 - longitudinal sec­ tion of capitulum x 2 Vi. 1-7, 11 from Lecordier in MAU 20868; 8 -1 0 from Wyatt 019. Drawn by Pat H all iday. (Reproduced from FI. Mascareignes, 109. Compostes, PI. 55.)

gins and often also apex scarious. Receptacle flat or con­ vex, rarely conical, epaleate. rarely paleate (Ceruana). Ray florets one- to several-seriate, female, fertile, tubular below, strap-shaped above, or filiform, pale yellow, cream, white, blue-violet, or pink. Disc florets herma­ phrodite, tubular below, campanulate in the upper half, yellow, yellowish or white. Anther base obtuse, short. Style bifurcate at apex, branches linear or linearlanceolate, papillose all round toward the apex. Cypsela obovate, distinctly to slightly compressed, 2-4(-5)-ribbed or costate, usually pilose. Pappus of 1-seriate or rarely 2-seriate baibellate bristles, sometimes of scales, rarely absent. A tribe with c 100 genera with worldwide distribution and occurring mainly in temperate Europe and montane areas in the tropics and subtropics; 10 genera in the Flora area.

1.

Receptacle with coriaceous paleae. 69. C eruana Receptacle without paleae, smooth or rarely with short fimbrils. 2

2.

Pappus 1-seriate to 2-seriate of 20-30 equal or subequal capillary 3-5 mm long bristles. 3 Pappus absent, or, if present, of subpaleaceous scales, or 2 weak bristles, up to 2 mm long . 11

3.

Shrubs or subshrubs with erect, scandent, stragg­ ling or climbing branches, 1—3(—6) m high. 4 Perennial or annual herbs, stem unbranched except in the inflorescence, or densely branched and bushy, 0.1-1 m high. 6

4.

Shrub with scandent, straggling or climbing branches; inflorescence pendant. 73. Microglossa Shrub with erect or ascending branches; inflores­ cence erect. 5

-

145. ASTERACEAE: 69. Ceruana, 70. Grangea

5. 6.

7. -

Oise florets bisexual and fertile; cypsela strongly compressed, asymmetric. 77. Conyza Disc florets bisexual but sterile; cypsela slightly compressed, not asymmetric. 72. Psiadia

C. pratensis Forssk. (1775); Buphtalmum pratense (Forssk.) Vahl (1790) type. Egypt, ‘Kahirae’ [Cairo], Forsskdl in Herb. Forssk. 1084 (C holo., BM LD iso.).

Ray florests conspicuous, corolla lobes spreading or distinctly overtopping disc florets, (1—)2—10 mm long. 7 Ray florets with insconspicuous, 0.7-2(-3) mm long corolla limbs, or limbs absent. 9 Ray florets 2-3-seriate, filiform, rays usually inrolled lengthwise upon drying. 76. Erigeron Ray florets 1-seriate, rays spreading, coiled back­ wards or folded irregularly upon drying. 8

Erect, annual herb, 40-60 cm high; stem richly branched from near the base, resilient, hirsute or pubescent, stri­ ate-terete, light yellowish green; branches ascending. Leaves stiff and held erect, light yellowish green, scabrid hispid on both surfaces, ovate, narrowed to a clasping, sessile, often auricled base, margins grossly dentateserrate, rounded at apex; lower leaves 5-7 x 1-2.5 cm. up­ per shorter, oblong, sometimes lyrate or sub-entire. Capitula campanulate or hemispherical, erect, 4-5 x 5-6 mm, disciform, subsessile to pedunculate; peduncle terete, sparsely hairy ; in open indeterminate cymes or ter­ minal and leaf-opposed along the branches of dichotomous cymes, surmounted at or near the base by 2 or more bracts equalling or over-topping the capitula. Phyllaries 2-3-seriate. unequal; outennost herbaceous, exceeding the florets, 3.5-5 x 0.5-1 mm. erect, pilose, oblonglanceolate, acute at apex; inner coriaceous, about as long as the florets, 3.3-4 x 0.6-1 nun, linear-oblong pilose, apex acute. Paleae coriaceous, margins fimbriatc-ciliate. oblong-elliptic. Florets orange-yellow. Ray florets fe­ male. fertile, c 3-seriate; corolla narrowly tubular. 1.6-2 mm long, 3-5-lobed. sessile-glandular; style bifid, c 2 nun long, branches lanceolate. Disc florets bisexual, fertile; corolla tubular below, campanulate above, capitateglandular, 2-2.2 mm long. 4(-5)-lobed; anther c 0.6 mm long, base obtuse; style bifid, c 2 mm long, branches lan­ ceolate with acute papillose appendices. Cypsela light brown, obovate, 2-2.2. x 0.5-0.7 mm. compressed or ob­ scurely angled, appressed hairy. Pappus coroniform with a row of short, whitish scales. Fig. 145.115.

8. Leaves ovate-lanceolate to lanceolate, glabrous, glaucous, shortly auriculate at base; capitula in a long panicle; cultivated plant. 75. Symphyotrichum - Leaves linear, elliptic or oblanceolate, sessile, or shorth petiolate, without auricles; capitula soli­ tary, usually held on long peduncle clear of the fo­ liage; indigenous plant. 74. Felicia 9 -

]89

Marginal florets eligulate; corolla filiform; style much exserted. 77. Conyza Marginal florets ligulate; corolla tubular below, limb or ligule spreading; style included or only slightly exserted. 10

11 Corolla of ray florets unilabiate or bilabiate, bright yellow or orange; florets conspicuous, not hidden by the pappus. 78. Nidorella - Corolla of ray florets tubular below and with spreading, 2-3-fid limb, white or pale yellow, flo­ rets hidden by the pappus. 77. Conyza 1 Pappus coroniform with minute, scarious scales, apex of cypsela flat or flattish. circular. 70. Grangea Pappus absent, or. if present, with minute bristles, usually 2; cypsela compressed, including apex. 71. Dicrocephala

Weed of disturbed sites in sandy river beds; 1600-2430 m; EW GJ; Egypt. Sudan. Chad. Nigeria Ghana. Mali. Senegal. Gambia. Ash 867; BartolommeiGioli 36 (FT).

69. CERUANA* Forssk (1775); Fayed in Mitt. Bot. Staatssarnml. Munchen 15. 425-576 11979).

70. GRANGEA* Adans. (1763)

rect. annual herbs. Leaves simple, alternate, hispid, ilose. oblong-obovate to oblong-linear, margins irreguarly serrate to pinnatifid. Capitula heterogamous, hemipherical, erect, disciform, in open indeterminate cymes; edunculate. Phyllaries 2-3-seriate. herbaceous, a few utemiosi leafy. Receptacle flat, tuberculate. paleaceous, 'aleae coriaceous, oblong-elliptic, fimbriate at margins id apex, exceeding the ovaries. Florets orange-yellow, lay florets female, fertile, c 3-seriate; corolla narrowly ubular, 3-5-lobcd, sessile-glandular; style slightly . vserted. bifid at apex. Disc florets bisexual, fertile; Drolla tubular below, campanulate above, capitateilandular, 4-5-lobcd; anther base obtuse; style bifid, ranches with lanceolate, acute, papillose appendices, ypsela compressed Or obscurely angled, with an annular ipex crowned with short, whitish bristles. A monotypic genus.

A. Fayed in Mitt. Bot. Staatsscunml. M unchen 15: 425-r576 (1979). Annual herbs. Stem solitary or several from the root apex, erect or prostrate, usually pilose. Leaves alternate, piiuiatilobed to bipinnatisect, sessile or petiolate. Capitula heterogamous. disciform, medium, broadly campanulate or hemispheric, pedunculate, solitary and terminal or in groups of 2 or 3 from leaf axils. Receptacle epaleate. coni­ cal or globular. Phyllaries 2-3-seriate. membranous, subequal. scarious at margins and apex. Florets light or golden yellow. Ray florets female, fertile, several-seriate; rays tubular below, widened above into apically 2-4lobcd limb; style bifid, often exserted. Disc florets bisex­ ual. fertile; corolla !ubular below, campanulate in upper parts. 4-5-lobcd; anthers with obtuse tail; style bifid, usu­ ally exserted. branches narrow, apices deltoid, papillose. Cypsela elliptic or oblanceolate. compressed. Pappus

190

145. ASTERACEAE: 69. Ceruana. 70. Grangea, 7) Dicrocephala

most acute at apex; inner obtuse. Florets golden yellow. Ray florets female, fertile, numerous; corolla tubular be­ low, rounded-campanulate above, with 2-4-lobed limb. 1-1.5 mm long, sessile-glandular Disc florets bisexual, fertile; corolla tubular and sessile-glandular below; tuoe c 1 mm long, funnel-shaped above, 4-5-lobed. 1.4-1.8 mm long; anthers c 0.5 mm long, obtuse at base; style c 1 5-2 mm long. Cypsela pale whitish, elliptic to oblanceolate. compressed except for the expanded apex, c 2.5 x 0.7 mm, sparsely sessile-glandular and ciliate toward the base. Pappus coroniform. w ith laciniate. 0.5-1 mm long scales. Fig. 145.116. Disturbed areas along lake margins; secondary ever­ green bushland on alluvial soil; 1500-1700 m. SU AR; a widespread weed northwards into Egypt, south to S Af­ rica, Madagascar, also in Asia. Gilbert & Tewolde B.G.E. 2456,A sh 266; Getachew A. 2910.

71. DICROCEPHALA L Herit. ex DC. (1833) A. Fayed in Mitt Bot. Staatssamml. Mtinchen 15 425-576 (1979)

lig u re i 15. CERUANA PR-X TENSIS'. Flowering branch x Vi.

(No specimen cited in orig.) Drawn by Magdi El-Gohary. (Re­ produced from The Weed Flora o f Egypt, 1994, p. 57).

coromfonn; corona crowncd with laciniate, short and nar­ row scalcs. often caducous. A genus with about 10 species: nine of which are eonfined to subsaharan Africa and Madagascar. 1 widespread and weedy species in the Flora area. G. m aderaspatana (/_.) Poiret (1812): Artemisia maderaspatanus L. (1753) - type: "Habitat in India.” Linnaean Herbarium No. 988.47 (LINN lecto., selected by Fayed in Mitt. Bot. Staatssamml. Mtinchen 15: 452 (1979)|. Annual licrb. 30-50 cm high: stem erect or prostrate (lower branches often prostrate), angular, sparsely to dcnsch pilose, hairs denser towards terminal parts, whit­ ish. Leaves alternate, pinnatilobed to bipinnatisect. dcnsch pilose with long silky hairs on both surfaces, up to II \ 4.5 cm: lower cauline leaves much longer with wider segments, margins coarsely serrate, petiolate. petiole up to 2.5 cm long, narrowly winged. Capitula globu­ lar-conical. 6-10 x 8-12 nun. solitary and terminal or in groups of 2 or 3 per branch, peduncle pilose, up to 3 cm long. Receptacle conical, epaleate. Phyllaries 2-3-senate, subequal, oblong-elliptic, membranous. 5-6 x 1.5—2 mm. margins scarious, ciliate. sparsely liairy on outside; outer­

Annual or perennial herbs. Leaves alternate, simple, sessile or petiolate. serrate or crenate-serrate. lobed to pinnatipartite. Capitula heterogamous, disciform, hemi­ spherical to oblong, solitary and terminal or in paniculate racemes, pedunculate. Receptacle convex, columnar or conical, epaleate. Phyllaries 2-seriate, membranous, mar­ gins scarious. fimbriate. Florets greea whitish yellow or reddish purple. Ray florets numerous, female, fertile, co­ rolla narrowly tubular below, roundcd-campanulate above, 3-4-lobed or dentate at apex; ovary glabrous or glandular, oblique at apex; pappus absent; style bifid at apex. Disc florets bisexual, few; corolla tubular below, campanulate above, glabrous or sessile-glandular. 4-5lobed; anthers orange-yellow, obtuse at base, connective triangular, ovary obovoid, glabrous or glandular; sty le bifid, papillose Cypsela light brownish, obovate. com­ pressed. glabrous Pappus absent or reduced into an annu­ lar disc or 1-3, short, caducous bristles. A paleotropical genus with four species; 3 species in the Flora area 1 Leaves petiolate, petiole without auricles; capitula short-pedunculate. 1. D. integrifolia - Leaves sessile or subsessile, auriculate at base; capitula long-pedunculate 2 2. Plant 30-100(-150) cm lugh; capitula 6-8 mm across; plant of mid-altitudes. 2. D. chrysanthcm ifolia Plant 2-10(-20) cm liigh; capitula 4-5(-6) mm across; plant of afroalpine habitats. 3. D. alpina I. D. integrifolia ( / ../ ) O. Kuntze (1891); Hippia integrifolia L. f. (1781) - type: From India Linnaean Herbarium No 1039.1 [LINN lecto., se­ lected by Fayed in Mitt. Bot. Staatssamml. Mtinchen 15: 492 (1979)1

145. ASTERACEAE: 70. Grangea, 71. Dicrocephala

191

Figure 115. GRANGEA M AD ERASP A T A N A. 1 - habit x 3/4; 2 - floret x 7. (No specimens cited in orig.) Drawn by B hola Ram. (Reproduced with permis­ sion from FI. India , vol. 12. fig. 37, 1995).

Dicrocephala latifolia (Lam.) DC. var. schimper ana Sch. Bip., nom. nud.. quoad Schimper 1537; D. atifolia (Lam.) DC. var. sonchifclia auct.. non (M.B ) 3C. (1836): Asch. in Schweinfurth & Asch. (1867). quoad Schimper 1537. D. abyssinica Sch. Bip., nom. nud., quoad Schimper 949 and 176. D. latifolia auct., non (Lam.) DC. (1836): A. Rich. 1848); Sell Bip. in Schweinfurth & Asch. (1867); Vatke (1875). sut ;p. integrifolia En-;: or ascending annual herb. 30-75 cm high; stem sul :ate, sparsely pilose, branches sometimes prostrate. Le ves alternate, dark grey-green above, pale green be­ ne; fih, lyrate pinnatifid to pinnatipartite, membranous, (2- i3.5-11.5 x 1.5-6.5 cm, segments ovate or ovatelar .eolate, with serrate or dentate margins, the terminal sej ment being the largest, glabrous to sparsely pilose; leaf petiole up to 4.5 cm long. Capitula heterogamous, sul globose. 3-4 x 3-4 mm, in open, tenninal. divaricate

panicles, short-pedunculate. Receptacle hemispherical, epaleate. Phyllaries c 2-seriatc. subequal, oblong-elliptic 0.9-1.3 x 04-0.5 nun. glabrous, membranous, light green, with scarious fimbriate margins and apex. Florets greenish white, pale yellow or purplish. Ray florets many-seriate, female, fertile; corolla tubular below, wid­ ened and 2-3-fid at apex. Disc florets bisexual, fertile; co­ rolla c 1-1.2 mm long, sparsely sessile-glandular, tubular below, funnel-shaped above. 4(-5)-lobed. lobes glabrous or glandular. Cvpscla light brown, narrowly obovate. compressed, margins thickened. 1-1.2 x 04-0.5 mm. apex with a mass of globose, sluny, sessile glands. Pappus absent or with 2 weak or caducous bristles. Fig. 145.117 Weed of arable, fallow and waste-land, normally growing in wet spots and depressions, swampy ground in forests by stream banks and along watercourses, rarely in open rockv places; 1750—3500 m. EW TU GD GJ SU IL KF SD BA HA; widespread in Africa. Arabia and Asia. Mooney 6409; E.F. Gilbert 428; Friis et al. 2226. Subsp. gracilis (DC.) Fayed occurs in Nepal, India and Madagascar.

192

145. ASTERACEAE: 71. Dicrocephala

117. DICHROCEPHALA IN TE G R IF O U A .. 1 - habit x %\ 2 Figure

capitulum x 714; 3 - phyllaries x 15; 4 - ray floret x 28; 5 - disc floret x 28, 6 - stamens x 28; 7 - style x 28; 8 cypsela x 28. 1 from Johnston 840, 2 -7 from Govinden in MAU 20505, 8 from BN 41. Drawn by Eleanor Catherine. (Reproduced from FI. Mascareignes, 109. C om posees, PI. 26.)

2. D. chrysanthemifolia (BI.) DC. (1833); Cotula chrysanthemifola BI. (1826) - type: Java, Cheribon, Blume s.n. (L holo,). Dicrocephala abyssinica Hochst. (1841), nom. nud., quoad Schimper 87; D. abyssinica Sch. Bip. ex A. Rich. (1848); D. chrysanthemifolia (BI.) DC. forma abyssinica (Sch. Bip. ex A. Rich.) Asch. in Schwein­ furth & Asch., Beitr. FI. Aethiop.. 145 (1867); D. chrysanthemifolia (BI.) DC. var. abyssinica (Sch. Bip. ex A. Rich.) Fayed in Mitt. Bot. Staatssaml. Mtinchen 15: 506 (1979) - types: TU, Mt. Scholoda, QuartinDillon s.n. & PI. abyss. I, Schimper 87 (P syn., BM K isosyn.) & 176 (P syn., BM K L M isosyn.). D. chrysanthemifolia (BI.) DC. forma macrocephala Asch in Schweinf. & Asch., Beitr. FI. Aethiop. . 145 (1867) - type: GD, Simen Mts.. Debre Eski, 1850, Schimper 77 (P holo., K iso.). Erect, stiff, widely branching annual, or short-lived, pe­ rennial heib, 30-75 cm high; stem solitary or several

from the base, ascending, scabrid-pubescent to gla­ brescent, striate. Leaves pale yellowish green, glandu­ lar-viscid, alternate, oblong to oblanceolate, 1.5—8(—12) x 0.5-2.5 (-3.5) cm, scabrid-pubescent with denser hair on veins beneath, margins grossly dentate to pinnati­ lobed. sessile or subsessile, with auriculate-amplexicaul base. Capitula heterogamous, disciform, globose to spherical (in fruit), 6-8 x 5-7 mm, solitary or in open racemes, pedunculate, peduncle sparsely pilose, bracteate. Receptacle epaleate, columnar. Phyllaries c 2-seriate, elliptic, green with white scarious, fimbriate margins and apex, c 1.5-2 x 0.7 mm, pilose up to middle. Florets greenish yellow, white, or dark red-purple. Ray florets 3-4-seriate, numerous, female, fertile, c 1 mm long (excl. ovaiy); rays 3-4-lobed at apex, tube densely sessile-glandular. Disc florets bisexual, fertile, 1.5—2 mm long; corolla tubular below, funnel-shaped above, 4(-5)-lobed, densely capitate-glandular. Cypsela light brown, obovate, compressed, with thickened margins,

113

145. ASTERACEAE: 38. Haplocarpha, 39. Berkheya

Figure

66.

HAPLOCARPHA RU EPPELLII 1 habit x 2/ 3 ; 2 -3 - leaf x 2/ 3 , 4 ray floret x 3; 5 - disc floret, cypsela removed x 4; 6 - disc floret, opened out x 6 ; 7 an­ ther o f disc floret x 10; 8 9 - style and stigma o f ray floret x 6 ; 1 0 style and stigma o f disc floret x 6 : 1 1 multicellular leaf hair x 40; 12 cypsela (im ­ mature, with pappus) o f ray floret x 16. 13-14 cypsela (with pappus) o f disc floret x 8 . 1. 4 -8 , 10-11, 14 from Twee die 1813; 2 from Fries 1279, 3 from Galpin 7918. 9 from Greenway & Kanuri 13631; 12 from Liebenberg 1578; 13 from Volkens 1551. Drawn by C.M. W ilmot-Dear. (Reproduced with

permission from FI. Trop. E. AJr.. C om positae, fig 57.)

• mate. Style thickened below the fork, branches linear. Cypselas obconical. ribbed, appressed or villous pubes­ cent. Pappus 2-seriate, of numerous, entire or deeply iaciniate scales. A genus of 74 species in southern and tropical Africa; 2 species in the Flora area. Pappus of oblong, entire scales, 1.2-1.5 111111 long, apex dent iculate; leaf margins wavy-dentate with large (1.5-7 mm) spinescent teeth; peduncle densely glandular hispid. 1. B. spekeana

Pappus of deeply lacerate, fimbrillate scalcs. 2.5-3.2 mm long; leaf margins entire with short (1-2 mm long) spines; peduncle densely white woolly. 2. B. chicsiana 1. B. spekeana Oliv. (1873) - type: Uganda. Unyoro, Grant s.n. (K holo.). B. spekeana var. abyssinica Fiori in .V. Giorn. Bot. Ital. n.s. 47: 46 (1940) - types: KF. Jimma [Gimma|. 10.1937. Saccardo s.n. (FT syn.); KF. Abba Kella. 8.1938. Mariottini s.n. (FT syn.).

114

145. ASTERACEAE: 39. Berkheya. 40. Gazania

Erect perennial herb. 0.5-1.5(-2.5) m high, with a woody rhizome. Aerial stem branched in upper parts, striatesulcate, densely glandular liispid. terete, often brownish or purplish brown. Leaves alternate, simple, sessile to semi-amplexicaul. narrowly elliptic, linear-lanceolate or lanceolate, 4-20 x 1-5 cm, with the largest leaves at the base of the stem, upper surface dark green with pale green midrib, densely glandular hispid, lower surface densely white-tomentose except for major veins, margins wavydentate with large (1.5-7 nun long) spinescent teeth and smaller (1-4 nun long) spines between the teeth, spines pale yellowish brown. Capitula lie mi spheric, 40-60 x 10-15 nun at anthesis, solitary, terminating stem and branches. Peduncle absent or up to 25 mm long, densely glandular-hispid. Phyllaries graded monomorphic. 3-4senate, linear-lanceolate, densely glandular liispid to to­ mentose on outer surface, margins with several upwardly pointed spines, apex acuminate, spinescent, outermost 5-15 x 1—2(—3) nun, middle 10-25 x 1-2 nun. innermost sometimes glabrous. Ray florets yellow, neuter, tube 6-7 mm long, densely pubescent, rays narrowly oblanceolate. 20-25 x 3-4 mm. apex 3-4-fid. Disc florets yellow, bisex­ ual. Corolla tubular-campanulate, 7-8 mm long, lower pan densely pubescent hairs rigid, slightly hooked at apex, 5-lobed. lobes 2.5-3.3 mm long. Anthers not exserted. 3-3.2 nun long, yellowish brown, connective lanceolate, c 0.5 nun long, base sagittate. Style bifurcate, 8.5-9.5 mm long, branches 1.7-1.9 nun long. Cypselas black, tuit>inate, tetra- to pentagonal near apex, ndged. 1.5-2.5 x 0.7-1.0 mm. densely appressed pubescent, liairs wliite. Pappus of 18-20, entire, oblong, yellowish scales, 1.2—1.5 mm long, sub-2-seriate, margins pectinate, apex denticulate. Fig. 145.67. Grassland with scattered slinibs. evergreen montane bushland, forest edges, roadside slopes; 1400-2800 m. GJ SU AR IL WG KF GG SD; Sudan. Rep. Dem. du Congo. Rwanda. E Africa. Nigeria. Mesfin T. & Kagnew 2468; Mooney 8185; Friis et al. 104. 2. B. chicsiana Chiov. (1912) ty pe; SU, Uata Daleccia near Holetta. 2650 m. Negri 583 (FT holo.). Berkheya spec, nova affinis B. zeyheri (Han & Sond.) Oliv. & Hiem Cufod.: Cufodontis (1967: 1168), quoad A'. Hildebrandt 120. Berkheya aethiopica Roessl. (1970) - type: GJ. Ainewetsch Mt, c 10 km SSW of Bahir Dar. Sehald 2348 (STU holo.. WU iso.). Erect, perennial herb. 30-80 cm high. Stem unbranched except in the inflorescences, reddish at base, pale green higher up. stnate-sulcate, terete, sparsely puberulous. Leaves alternate, simple, sessile-auriculate. oblong to ob­ long-elliptic, middle cauline ones 6-14 x 1.5-3.5 cm. me­ dium to dark green and scabnd above, with small white bnstly liairs. white tomentose beneath except for veins, margins entire or denticulate, with short (1-2 nun long) spines all along, apex acuminate, spinescent Capitula hemispheric, 45-65 x 10-15 nun at anthesis, solitary, ter­ minating stem or branches. Peduncle densely white

woolly or wliite tomentose. 1-15 cm long. Phyllanes graded monomorphic, 3-4-seriate, linear-lanceolate, densely glandular hispid to tomentose on outer surfaces, margins with several upwardly pointed spines, apex acuminate, spinescent, outermost 4-10 x 1-2 mm, middle 8-12 x 1-2.5 mm, innermost sometimes glabrous. Ray florets yellow, neuter, tube 4-6 mm long, densely pubes­ cent rays narrowly oblong-lanceolate, 15-20 x 3-4 mm, apex 3^4-fid. Disc florets yellow, bisexual. Corolla 5 5-6.5 mm long, tubular-campanulate. densely pubes­ ccnl on tube, liairs rigid, slightly hooked at apex, 5-lobed, lobes 2-3 nun long Anthers not exserted, 2.6-3 mm long, yellowish brown, connective lanceolate, c 0.5 mm long, base sagittate, 0 .3 -0 4 mm long. Style bifurcate, 7-8 mm long, branches 1.4—1.6 mm long. Cypselas black, oblanceolate, tetra- to pentagonal near the apex, ridged. 15-2 x 0.5-0.7 mm, densely appressed pubescent, hairs white Pappus of 10-16, yellowish, deeply lacerate. 2 5-3.2 mm long, sub-2-seriate, fimbrillate scales. Hilly grassland with scattered trees and shmbs, grass­ land along stream margins; 1300-2000 m. GJ SU KF IL WG SD HA; not known elsewhere. W. de Wilde 6835; Mooney 6819; F.nsermu K. & Tamrat B. 491. 40 GAZANIA* Gaert. (1791), nom. cons. 11 Roessler in Mitt. Bot. Staatssaml. Mtinchen 3: 71-500 (1959). Perennial herbs with woody rootstocks. Plant scapose, or with suberect to decumbent leafy stems. Leaves basal and crowded or cauline and alternate, simple; blade nar­ rowly elliptic to dentate-pinnatifid, with linear-lanceo­ late segments, usually glabrous on the upper surface, white-felted tomentose beneath. Capitula radiate, heterogamous, large, showy, pedunculate or scapose, solitary Involucre campanulate. cylindric or obconicc>lindric. Receptacle alveolate with alveolae c 0.5 mm lugh Phyllaries connate, forming a campanulatec\lindrical cup, free parts 2-3-seriate, inserted on the rim of the connate part, with none or few scattered on the fused portion. Ray florets 1-seriate, frequently orange and with a dark basal purple spot and a dark dorsal stripe, sterile, closing at night, 2-4-fid at the apex. Disk florets yellow, bisexual. Corolla funnel-shaped. 5-lobed An­ ther base minutely sagittate, shortly mucronate-acuminate Style slender below, thickened above the mi­ nutely hairy node, bifurcate, branches very short, linear. Cypselas narrow l> turbinate or obovoid-obconical. with silky liairs, 3-5 mm long. Pappus of linear-subulate, subequal scales in two rows. A genus with about 17 species confined to South Af­ rica and Namibia. One species extends to tropical East Af­ rica A number of species, e.g.. G. rigens (L.) Gaertn.. G. anearis (Thunb.) Druce. etc.. are cultivated ornamentals, each with several varieties and/or hybrids. The two spe­ cies inenuoncd arc also known to hybridize in nature. G. rigens (syn. G. splendens) is a glabrous, scapigerous hetb, w ith large orange ray florets. No vouchers of the latter species has been seen from the Flora area.

145. ASTERACEAE'. 39. Berkheya

115

Figure 67. BERKHEYA SPEKEANA: 1 flowering branch; 2 ray floret. 3 style o f ray floret; 4 disc floret; 5 stamens; 6 -style; 7 cypsela with pappus; 8 - scale o f pappus. No magnifications given and no specimen cited in the original publication. Drawn by W. H Fitch. (Reproduced from Trans. Linn. Soc. 29. tab. 66.)

116

145. ASTERACEAE: 40. Gazania, 41. Hirpicium

G. linearis (Thunb.) Druce (1917); Gorteria linearis Thunb. (1798) - type: from South Africa. Thun berg in Herb. Thunberg (UPS). Rhizoinatous perennial herb. 40-75 cm high. Stem suberect and often growing through and over shrubs, or spreading along ground and mat-forming, branching from base, striate-sulcate. leafy in lower parts. Cauline leaves alternate, most simple, some pinnatifid. pseudo-petiolate to sessile, blade with long, petiole-like winged base, oblanceolate to elliptic, leathery, glabrous or nearly so above, wlute woolly beneath except for the prominent midrib, 7-17 x 3—7(—10) mm. margins entire, revolute, apex acute. Capitula radiate, heterogamous. 40-70 x 10-12 mm at anthesis, solitary, scapose; scape up to 35 cm long, glabrous to sparsely pilose, narrowed toward the capitula. Involucre glabrous or hairy, campanulate or con­ nate in lower half forming a cup, pouched at base Phyllaries light green, 2-3-seriale. 8-12 x 6—12(—15) mm. outer ones linear 7—10(—15) x 1 mm, acuminate, margins bristly -ciliate especially in upper half inner ones 8-15 x 1.5—2 mm. margins entire, often purplish, apex narrowly acuminate. Rav florets yellow or orange. 20-22. rays 40-50 x 6-8 mm. usually with dark spots at base, apex 2-4-fid Disk florets bisexual, reddish orange. Corolla c 8 mm long. Immature cypselas 1-2 mm long. Pappus scales 7-8. 3-4 mm long, usually hidden by tlx: longer liairs of the cy psela. J

Leaves or segments linear or linear-lanceolate. 1—5(—10) mm wide. var. linearis Leaves or segments elliptic or broadly lanceolate. (10-) 12-25 mm wide. var. ovali.s

var. ovalis (Harv.) Roessel. in Mitt. Bot. Staatssaml. Munchen 3: 380(1959); G. longiscape DC. var. ova/is Harv. in Harv . & Sonder. FI. Cap. 3: 474 (1864/65) - type: South Af­ rica. Somerset East, Bowker (Mrs. Berber) s.n. (K holo.). Perennial herb. 30-40 cm high. Stem repeatedly branched, ercct. spreading or ascending. Leaves or segments elliptic or broadly lanceolate. (10—)12—25 mm wide A garden ornamental widely cultivated in Addis Ababa and other larger towns; 2350-2400 in. So far only known from a few collections from EW and SU; native of South Africa and cultivated in many other countries. Anon (Students o f Asmara Univ.) 18; Sebsebe D. 63. Var. linearis lias not been substantiated with a voucher specimen. 41. HIRPICIUM Cass. (1820) Berkheyopsis O. Hoffm. (1892). H. R oessler in Mitt. Bot. Staatssaml. Munchen 3: 7 1 -5 0 0 (1959).

Annual or perennial herbs. Stem solitary or several from the root apex, each branched from middle to apex, densely hispid with coarsc. erect or suberect hairs. Leaves alter­ nate, sessile or subsessile. linear oblong or narrowly

oblong-elliptic, simple to pinnatifid. densely hispidsetose on the upper surface, white tomentose on the lower surface except for the mid-veins. Capitula radiate, hetero­ gamous. solitary , terminating main stem and branches. Peduncle densely hispid-setose. Receptacle shallowly alveolate Involucre campanulate or turbinate. Phyllanes fused except at apex and forming a cup-like involucre, 3—l-seriate, linear subulate or triangular-lanceolate, densely hispid-setose on surfaces, margins ciliate toward the acuminate, spinescent apex. Ray florets yellow, ster­ ile. rays 5-nerved. margins becoming inrolled, apex (2-) 3-4-fid. Disc florets bisexual, fertile. Corolla tubular gla­ brous. 5-lobed, lobes oblong-lanceolate, often divided to the middle of the limb, not papillose. Anthers yellowish brown, shortly sagittate with blunt or acute base, connec­ tive whitish, not striated. Style bifurcate, slightly thicker below fork, branches oblong, short, apex obtuse. Cypselas turbinate, densely clothed with long (2-5 mm), 2-seriate bristles, light to dark brown. Pappus scales 2-seriate, outer series 10, much longer than the inner ones, spirally ar­ ranged and overlapping, oblong with setose-acuminate apex, margins ciliate toward the apex, strongly medially veined, inner sencs 10, oblong-lanceolate. A genus with about 12 species found mostly in S, SW, C ?nd E Africa: 2 in the Flora area. 1

Perennial, leaves mostly pinnatifid; disk floret co­ rolla 7-8 nun long; anthers 3—3.5 mm long; style 8.5-9.5 mm long. I. H. beguinotii Annual, leaves mostly simple; disk floret corolla 4.5-5 nun long; anthers 1.5-1.8 mm long; style 5—5.5 mm long. 2. H. diffusi m

1 H. beguinotii (Lanza) Cufod. (1 9 6 7 ); Gazania beguinotii Lanza (1939) - type: SD, Mega, Cufodontis 631 (FT holo.). Perennial herb. 20-35 cm high. Stem with several ascend­ ing branches from near the root apex, densely hispid with coarse, ercct or suberect hairs, striate, terete. Leaves alter­ nate, pinnatifid, discolorous, scabrid-hispid above, densely wlute tomentose beneath except for the nud-vem most 5-9 cm long. 1-2 cm wide between leaf teeth, seg­ ments 5-10 x 3-5 nun, apex acute, margins revolute. Capitula 30-35 x 8-10 mm at anthesis, solitary terminat­ ing main stem and branches. Peduncle densely hispid, 5-8 cm long, narrowed toward the base of the capitula. Phy Uaries 3-4-seriate. fused for most part with only the apices free, outer surface densely hispid-setose, margins ciliate near apex 6-20 mm long, apex ending in sharp, spiny, spreading or reflexed bristles. Ray florets bright yellow, sterile, 13-15, rays oblong to oblong-elliptic, 15-25 x 3—I irun, apex 2-4-fid. Disk florets yellow, bi­ sexual. Corolla tubular limb slightly wide, 7-8 mm long, glabrous. 5-lobed. lobes 3-3.5 mm long, reaching middle of limb. Anthers brownish yellow, 3—3.5 mm long, base sagittate, obtuse, connective oblong-lanceolate, c 0.5 mm long, not striate Style bifurcate, 8.5-9.5 mm long, branches c 1 mm long, obtuse at apex. Immature cypselas light brown, densely villous, c 3 x 1 mm, cuneate-obovate.

117

145. ASTERACEAE: 41. Hirpicium

Figure 68.

H IRPICIU M DIFFU SU M

1 habit x Vy, 2 & 3 - leaves x Vy, 4 capitulum x 114; 5 -c a p itu lu m , cross section x 6 - phyllary x 4; 7disc floret x 3; 8 - style x 4; 9 - style arms x 10; 10 - anthers x 4; 11 - ray floret x 3; 12 - cypsela with pappus x 3. 1 from Johnston s.n. (type); 2 from Thomas 3414; 3 from Paulo 415; 4-12 from Tweedie 2644. Drawn by Cam illa Speight. (Repro­ duced with permission from FI. Trop. E. Afr., C om positae, fig. 59.)

P a 'pus scales 2-seriate, outer series 10. oblong lanceoIat< inner series 10, lanceolate. Dense, low, seasonally water-logged or marshy grass­ land on black soil; 1920-2130 m. SD; probably not known elsewhere. Mooney 7319; Gilbert & Sebsebe D. 88 2; Gillett 14202.

2. 1 diffusum (O. Hoffm.) Roessler (1959); Berkheyopsis diffusa O. Hoffm. (1892) - type: Taninia, Kilimanjaro, Johnston s.n (B holo., BM K iso.). Ere cl, bushy annual herb with dense, spreading branches, 20- 35 cm high Stems several from the root apex, densely

hispid-setose, greenish yellow, striate-sulcate, terete. Leaves simple, oblong-lanceolate to narrowly oblongelliptic, 15-70 x 3-10 mm, densely hispid-setose on the upper surface, white tomentose on the lower surface ex­ cept for the mid-vein, narrowed toward the base to sessile-auriculate, apex acute, margins revolute. Capitula 20-30 x 6-9 mm at anthesis, solitary, terminating stem and branches. Peduncle densely hispid-setose, 1-6 cm long. Phyllaries 3-4-seriate, fused for most part with only the apices free, surfaces densely hispid-setose, margins ciliate near apex, 4-20 mm long, apex bristly or spinescent especially in young inflorescences. Ray flo­ rets yellow, sterile, rays oblong, 10-20 x 3-4 mm, apex 3-4-fid. Disk florets yellow, bisexual. Corolla tubular gla­

118

145. ASTERACEAE 41. Hirpicium

brous, 4.5-5 mm long, 5-lobed to near middle of limb, lobes 1.5—1.8 mm long, epappillose. Anthers brownish yellow, 1.5-1.8 mm long, base sagittate, acute, connec­ tive oblong-lanceolate, not striated. Style bifurcate, 5-5.5 mm long, branches c 0.5 mm long, obtuse. Cypselas brown, cuneate-obovate, 3.5-4 x 1.3-1.6 mm (0.4-0.5 mm wide near base), densely villous with long 2-senaie hairs. Pappus scales 2-seriate, outer series 10, stipitate at base, subspirally arranged, oblong- or ovate-lanceolate, medially thickened, denticulate at margins toward the apex, mostly aristate-acuminate. 3-6 x 0.8-1.2 mm, inner series 10, lanceolate, c 1.5—1.8 mm long. Fig. 145.68

Sclerostephane Chiov. (1929), which is quite conunon in Somalia, is included in the key since it might occur in the far southeastern comer of Ethiopia. Wagenitz and GamalEldin (in Bot. Jahr. Syst. 104; 91-113 (1983)] recognized 5 species from Somalia in Sclerostephane. Two of the spe­ cies were then described as new . The genus is related to Pulicaria with the main difference being in the morphol­ ogy of the cypsela. In Sclerostephane, the upper margin of the cypsela is extended to form a woody tube and the 2-seriate pappus bristles are inserted at the apex of the tube. 1. -

Receptacle with paleae. Receptacle without paleae.

Acacia-Lannaea woodland, open and degraded Acacia-Commiphora bushland, 1130-1700 m. SD, E Africa. Mesfin T. & Vollesen 4115; Friis et al. 2686; Ensermu K. 1133.

2.

Capitula sessile or shortly pedunculate, solitary or often congregated and concealed in the leaf axils or forks of branches; ray florets pale yellow, c 4-5 mm long. 43. Geigeria Capitula on up to 10 cm long pedicels, solitary at the apex of branches or corymbosely arranged; ray florets bright yellow, 12-14 mm long. 42. Anisopappus

-

Tribe 6. IN U L E A E Cass. (1819) A Anderberg in Canad. J. Bot. 67: 2277-2296 (1989). Perennial herbs or shrubs, rarely annual herbs. Leaves simple to pinnatifid, alternate, variously pubescent to woolly-tomentose, rarely glabrous, margins serrate, serrulate or denticulate, rarely entire. Indumentum of 1-seriate or basally multiseriate, erect patent or flexuous liairs and capitate or sessile glands Capitula hetero­ gamous. radiate or disciform or homogamous. Receptacle paleate or epaleate. Phyllaries several- to multi-seriate, herbaceous to cartilaginous, or apically and marginally scanous. Ray florets (when present) 1-to 2-seriate, female, yellow. Bisexual florets yellow, corolla tubular or fun­ nel-shaped. Anther base sagittate with long or short tail, connective acute-triangular. Style bifurcate, with mostly acute or rarely obtuse (in Anisopappus) sweeping hairs above the point of bifurcation Stigmatic areas distinct at base, confluent at apex. Cypselas oblong-cylindric, about 10-ribbed. sparsely to densely pubescent. Pappus of barbellate or plumose (in Pegolettia), slender bristles, in one to several rows, or of cup-forming scales on outside and bnstles on inside, or separate scales only. A tribe with about 40 genera distributed in Europe, Asia and Africa; 9 genera in the Flora area. In a recent investigation of the characters that define the Inuleae, taken in the wide sense, Anderberg (1989) wrote that it consists of “...three separate monophyletic taxa...” and proposed that Inuleae Cass, (sensu Merxmuller, H„ Leins, P., and Roessler, H. 1977 in; Heywood, V. H„ Harbome, J.B. and Turner, B.L. (eds ), The Biology and Chemistry o f the Compositae, 1. pp. 577-602 with about 200 genera and over 2000 species should be di­ vided into the tribes Gnaphalieae Rydb. (comprising the Inuleae - Gnaphaliinae and the Inuleae - Athrixinae sensu Merxmuller et al., loc. cit.), the Inuleae Cass, sensu stricto, and the Plucheeae (Benth.) A. Anderb. (= Inuleae - Plucheinae Benth.). This reclassification of the Inuleae is adopted here as the taxa recognized under each tribe seem to be more related to each other than across the tribal boundaries.

3. Capitula radiate. - Capitula discoid or disciform.

2 3

4 6

4.

Pappus 1-2-seriate, all setaceous, without a short outer row 44. Inula Pappus 2-seriate, outer series short and usually cup-forming or of separate scales, the inner senes of long bristles. 5 5. Cypselas with a cartilaginous annular tube at apex; pappus bristles arising from the apex of the tube. Sclerostephane - Cypselas without a cartilaginous annular tube; outer pappus usually cup-forming or scale-like and separate, inner pappus of long capillary bns­ tles. 48. Pulicaria 6. Pappus 1-seriate, of more or less equal capillary, barbellate bristles. 7 - Pappus bi- to several-seriate, equal or unequal, of barbellate or plumose bristles or incised scales 8 7. Capitula discoid; all florets bisexual, corolla 5-lobed, lobes with dense orange glands. 45. Pentanem a - Capitula disciform, outer florets female, corolla 2-4-lobed, glabrous, inner florets bisexual or fe­ male, corolla 5-lobed, lobes papillose. 46 Blumea 8. Outer pappus of deeply incised scales, inner of more or less equal plumose bristles; cvpseias 12-20-ribbed or costate; corolla zygomorphic with one large lobe with bristles. 47. Pegolettia - Pappus of several-seriate, equal or unequal, barbellate and spreading bristles; cypselas 4-angled or 10-ribbed; corolla equally 5-lobed, lobes glandular or glabrous. 9 9. Pappus bristles irregular in size; cypselas 10ribbed. ribs conspicuous, not hidden by the indumentum; leaves simple and linear or deeply pinnatifid 49. Iphiona - Pappus bristles more or less equal; cypselas 4angled or 4-ribbed. ribs hidden by dense indu­ mentum. leaves spathulate to round. 50. Iphionopsis

145. ASTERACEAE: 42. Anisopappus

42. ANISOPAPPUS Hook. & Arn. (1 8 3 7 )

H. m & mKirkia*. 45-73 (1964); S. Ortiz & J.A.R- Paiva in Boi J. Linn. Soc. 117: 3 9 ^ 6 (1995); S. Ortiz et al. in Anales Jard Bot. Madrid 54: 378-391(1996). Erect, perennial or annual herbs. Leaves simple, crenate or dentate, pubescent. Capitula radiate, heterogamous, larie and solitary, or clustered and few. Receptacle pa sate, paleae folded lengthwise. Ray florets female, yel­ low Disc florets yellow, bisexual. Anthers minutely calcaj :tte, with long or very short tails. Style with obtuse style ha rs. Cypselas ellipsoid or with indistinct edges. Pappus of ;hort scales, or sometimes absent. A genus with about 17 species found mostly in south­ ern and tropical Africa and extending into Madagascar an I. China. Two taxa: one species and one subspecies in the Flora area.

119

mm long, rays narrowly oblong, 3-fid at apex; tube c 2.5 mm long, sparsely glandular. Paleae c 5-7 mm long, nar­ rowly oblong, pubescent near the apex. Disc florets yel­ low, 3.1-4.2 mm long; corolla 5-lobed, tubular below, slightly expanded in upper parts, sparsely glandular in lower parts, lobes glandular hairy. Cypselas cylindric, 1.5-2.5 x 0.5 mm, tetragonal, multi-ribbed, pubescent. Pappus scales 0.5-1 mm long, squamulate, unequal, many short and broad, some long and bristle-like. Fig. 145.69. Dry evergreen forest, Combretum-Terminalia wood­ land wooded grassland with Comb return and Cussonia, bushland with Erica arborea, Hypericum quartinianum, Protea gaugedi, short grassland; 1400-2250 m. WG IL KF BA, West Africa, Sudan, Rep. Dem. du Congo, East Africa to Mozambique, Malawi, Zambia. Mesfin T. & Kagnew 2438; Friis et al. 506; Mooney 8440.

Recently Ortiz & Paiva [in Bot. J. Linn. Soc. 117: 39-46 (1995)] and Ortiz., Paiva & Rodr.Oubina (in Anal. Jard. Bot. Madrid 54: 383 (1996)] made thorough com­ parative morphological and nomenclatural studies on A. africanus (Hook, f.) Oliv. & Hiem (1877) and A. chinensis (L.) Hook. & Am. (1837) and concluded that they should be combined into one species, i.e., A. 1 4. chinensis (L.) Hook. & Arn. (1837); chinensis. While the present author accepts their taxo­ Verbesina chinensis L. (1753) - type; “Habitat in nomic studies, their nomenclature as regards the wide­ China. Osbeck.” Osbeck s.n. in Linnaean Herbarium spread taxon in Africa is not accepted, since it does not 021.3 [LINN, lecto., designated by Wild in Kirkia 4: follow the rules of priority within ranks. Ortiz., Paiva & 50 (1964)]. Rodr.Oubina recognized 3 subspecies in A. chinensis of which only one is found in the Flora area, subsp. su :sp. africanus (Hook./.) Ortiz & Paiva in Bot. J. Linn. africanus. The epithet subsp. africanus (1995) lias prior­ Soc. 117: 44(1995); ity at that rank over subsp. buchwaldii (1996). A. Telekia ajricana Hook. f. (1864); Anisopappus chinensis subsp. chinensis occurs in Madagascar and in africanus (Hook, f.) Oliv. & Hiem (1877); - type: Asia to China, while a third subspecies, A. chinensis Cameroun Mt. Cameroun, Mann s.n. (K holo ). subsp. lobatus (Wild) Ortiz & Paiva occurs in Zambia and A. chinensis (L.) Hook. & Am. subsp. buchwaldii Zimbabwe. (O. Hoffm.) Ortiz., Paiva & Rodr.Oubina in Anal, •ard. Bot. Madrid 54: 383 (1996); Sphacophyllum buchwaldii O. Hoffm. (1898); Anisopappus buch­ 2. A. holstii (O. Hoffm.) Wild (1964); waldii (O. Hoffm.) Wild (1964) - type. Tanzania, Sphacophyllum holstii O. Hoffm. (1895) - type: Usambara Mts., Buchwald 221 (B holo., destroyed; Tanzania Usambara. Holst 3915 (B holo., destroyed); BM C OIK iso ). Usambara. Kwa Mshuza. Holst 8898 (K neo., P Anisopappus inuloides Hutch. & B.L. Burtt (1932) isoneo.. selected by Beentje 2002). - type: Congo, Quarre 2847 (BR holo., K iso.). A. suborbicularis Hutch. & B.L. Burtt (1932) - Ercct amiual herb, 15-25 cm liigh. Stem slender, c 1 nun type: Congo, de Witte 482 (BR holo., K iso.). wide, sparsely branched in upper parts, striate-sulcate, Telekia niamniamensis Schweinf., nom. nud. pilose. Leaves simple, alternate, petiolate, blade ovate, 1.5-4 x 0.6-2 cm, membranous, hairy on veins of particu­ E ect, stiff perennial herb, c 1 m high; stem glabrcscent to larly the lower surface, glabrous or glabrcscent above, d( nsely crisped-pubescent. Leaves alternate, up to 6.5 x 3 margins serrate, petiole up to 10 mm long, densely ciliate. era, deltate to ovoid-oblong, light-green, 3-5-veined from Capitula campanulate, 3-5 x 3-4 mm at anthesis, not ti e base, with only the midvein being prominent, sparsely much altered in fmit, in open terminal or axillary cymes, s< tbrid above, densely pubescent below with denser hair peduncle up to 2 cm long, pubescent. Phyllaries whit­ ir the prominent reticulate veins, base cordate-hastate, ish-green with dark green mid-rib. folded inwards. 1-1.5 n irgins crenate except near tip; apex obtuse or acute; x 0.3-0.5 mm, pilose with liairs mostly on mid-rib and to­ P uolate, petiole 1-1.5 cm long, crispid-pubescent. ward the apex, apex apiculate. Paleae scarious, folded flat C Lpitula solitary at the end of branches, or several corym­ or folded. 1.5-2 mm long, apex apiculate, similar to bs sely arranged on up to 10 cm long, densely scabridphyllaries. Ray florets yellow, female, rays 1-3 mm long. p lose, peduncles; involucre depressed-hemispheric. Disc florets yellow. Corolla tubular-campanulate, c 1.5 P ivllaries 3-4-seriate, up to 10 x 1 min. light green, mm long, glandular in lower parts, 5-lobed, lobes c 0.3 si bequal, linear-oblanceolate. tapering to an acute apex; mm long. Style bifurcate, c 1.4 mm long. Anthers yellow, d nsely pubescent on outside. Ray florets yellow, 12-14 I.

Capitula 10-25 mm in diameter, ray florets 12-14 nun long; perennial herbs, c 1 m high. 1. A. chinensis subsp. africanus Capitula 3-5 mm in diameter; ray florets 1-3 mm long; annual herbs, c 15-25 cm high. 2. A. holstii

120

145. ASTERACEAE: 42. Anisopappus, 43. Geigeria

Figure 69.

ANISOPAPPUS C H IN E N SIS subsp AFRICANUS. 1 - flowering branch. 2 - outer phyllary; 3 ray floret, 4 disc floret and palea. 5 inner phyllary ; 6 two anthers; 7 - style arms. No magnifi­ cations given and no specimens cited in original publication. Drawn by W.E. Trevithick. (Reproduced with permis­ sion from FI. W Trop. Afr., vol. 2, 1963, fig. 255. as A. africanus.)

c 0.6 mm long. Cypselas black, columnar, c 10-ribbed. c 1 x 0.3 mm. glabrous. Pappus absent. Woodland with Acacia, Combretum, Entada, Sclerocarva and Zanthoxylum on steep slopes with abandoned cultivations; 1550 m. GG: E Africa. Gilbert et al. 8971. Until recently this species has only been known from East Africa; the specimen cited was collected from an area between Jinka and Konso (GG). It is not known whether it represents a natural population or a recent introduction But the relatively recent collections of other species, such as I an ilia roscheri Rchb.f. [Orchidaceae. Bidgood & Cribb in Kew Bull. 54: 378 (1999)1 from the same general area with similar distributional gaps, might indicate that this is a natural component of the vegetation. Further col­ lections in Gamo Gofa might also bring forth other re­ cords showing similar vicariance.

43. GEIGERIA Griess. (1830) Diplostemma Hochst. et Steud. ex DC. (1838). Araschcoolia Sch. Bip. ex Benth. & Hook. f. (1873). U. M crxm uller in Milt. Bot. Staatssamml. Mtinchen 1, 7

239-316 (1953). Bushy herbs, up to 50 cm high, with prostrate or erect branches, or rosulate acaulescent herbs; stem winged or wingless. Leaves simple, opposite or alternate, linearlanceolate to linear-elliptic, glabrous to woolly. Capitula sessile to shortly pedunculate, often concealed amongst the leaves, terminal and solitary or aggregated, or in the leaf axils or forks o f the branches, heterogamous, radiate Involucre ovoid or campanulate. Phyllaries multiseriate, imbricate, outermost leafy, inner scarious or leathery Re­ ceptacle flat or convex, paleate. Ray florets uni-seriate, fe­ male, tubular below; rays oblong to linear-oblong, 3-fidor 3-lobed at apex. Paleae conduplicate. fimbriate. Disc florets yellow, tubular, fertile, 5-lobed. Anthers sagittateauriculate at the base, with long caudate-acuminate tails;

style branches linear-lanceolate, apices obtuse to sub­ acute. Cypselas indistinctly 3^-angled. hairy. Pappus of 2-seriate. similar scales, or the inner series longer and bristly. A genus w ith about 25 species mainly found in south­ ern Africa; 2 species in the Flora area. 1.

Leaves glabrous or glabrescent. glandular, shinv; stem up to 50 cm high or. if acaulescent, leaves glabrous and shiny. 2. G. alata Leaves densely whitish-araneose or cottony: acau­ lescent herbs. 1. G. acaulis

1 G. acaulis Benth & Hook. f. ex Oliv. & Hiem (1877); type: Sudan. Cordofan, Arasch-Cool, Kotschy 104 (K holo.. M P Z iso). Araschcoolia acaulis Sch. Bip., nom. nud. . Diplo­ stemma acaulis Sch. Bip., nom. nud. Acaulescent perennial herb, with slender, up to 15 cm long taproot, rarely annual, stem rarely up to 2 cm long, araneose. Leaves crowded at base, sessile or subsessile, up to 8.5 cm long. 5-9 mm wide, narrowly oblanceolate, tapering to the base, densely grev-white araneose or cot­ tony. margins entire, apex subobtuse, mucronate. Capitula solitary, sessile in the crowded leaf axils, villose, c 10 mm in diameter, ovoid. Phyllaries 2-3-senate, subequal, 5-7 mm long, lanceolate, tapering to an acuminate apex, ciliate at margins near apex, densely cot­ tony at base. Paleae c 2.5 mm long. Ray florets female, pale yellow, rays up to 5 mm long, oblanceolate. apex 3-fid Disc florets pale yellow, c 2-3 mm long, cylindriccampanulate, with short tube, corolla 5-lobed with long narrow lobes, papillose. Cypselas c 1 mm long, narrowly cylindric, hairy. Pappus bi-aristate: outer series palea­ ceous. c 1 mm long, oblong, obtuse at apex; inner series aristate. longer than the outer series. Fig. 145.70. Alluvial flats in open Acacia bushland with numerous water courses; 325-650(-1900) m. EW SD GG; Sudan,

121

145. ASTERACEAE: 43. Geigeria

Figure 70. GEIGERIA ACAU LIS 1 habit x 2/3; 2 - capitulum , cross section (simplified) x 6; 3 - inner phyllary x 6; 4 - palea x 12; 5 - outer floret x 10; 6 style o f outer floret x 18; 7 —inner floret x 10; 8 -s ta m e n s x 18; 9 -s ty le o f inner floret x 24; 10 - cypsela x 12. 1 from Gilbert 4731; 2 -1 0 from Polhill & Greenway 452. Drawn by Pat Halliday. (Reproduced with permission from FI. Trop. E. Afr., C om positae, Fig. 68.)

Ug; ida. Kenya, Tanzania, Angola. Zimbabwe. Namibia. Sou li Africa (Transvaal). Mooney 9535; Gilbert et al. 814 I . Sebsebe D. & E nserm u K. 2758. 2. G. alata (DC.) Benth. & Hook. f. ex Oliv. d- Hiem 1877); Cichorium alaium Hochst. & Steud . nom. nud.; iplostemma alaium DC. (1838) - type: Yemen. Mt. • :dder. Schimper 853 (K M). Am jaJ herb, up to 60 cm high; stem dichotomously brai'.hed. glabrous or sparsely pubescent, often 3wimged. Leaves oblong-lanceolate, bright yellow-green. ses< le. decurrent. up to 5 x 1.2 cm. glabrous to sparsely pub scent, densely glandular-punctate, lateral veins fused

and running parallel to margins, margins entire; apex acute-mucronate. Capitula sessile, solitary or several at th e fo rk s o f th e b r a n c h e s , u r c c o la te .

1 0 - 1 2 - f lo w c r e d .

Phyllaries imbricate. 5-6-seriate. c 4.5-5.5 mm long, pu­ bescent. glandular-punctate, ovate and terminated by a spinous, reflexed awn. Ray florets bright yellow, female. c 4 nun long, tube c 2.5 mm long, glandular-punctate. Disc florets yellow, campanulate, densely glandularpunctate. c 3.5 mm long: corolla 5-lobed. Cypselas c 1.5 mm long, densely sericeous. Pappus c 1.8 mm long. 2-seriate. outer series paleaceous, c 1 mm long, inner sc­ ries similar but with elongate aristae. River-bed in dry bushland with Boswellia. Commi­ phora. and Acacia, open dry' bushland. subdesert with lava;

122

145. ASTERACEAE: 43. Geigeria, 44 Inula

60-1500 m. EE SD HA; Egy p t Sudan. Yemen, Saudi Ara­ bia. Mali, Niger, Chad. Somalia. Djibouti, Kenya Namibia. Ellis 221; Gilbert et al. 8189; Gillett 15092.

44. INULA L. (1753) Petrollina Chiov. (1911) Perennial herbs or shrubs. Leaves simple, alternate, entire or serrate, sessile to shortly petiolate, scabrid-pubescent or tomentose. Capitula usually in terminal corymbose cymes, occasionally on short lateral branches, hetero­ gamous. radiate; involucre hemi-spherical to turbinate. Receptacle plane or slightly convex, areolate or foveolate. Phyllaries multiseriate, imbricate, herbaceous or scarious. Florets yellow; ray florets female or absent, one- to multi-seriate. Female florets few or many. 1-2-seriate, tu­ bular 3-fid at apex. Disc florets bisexual, tubular, corolla 5-lobed; anthers sagittate and tailed at base. Cypselas cy­ lindric. slightly constricted at apex, angular or 4-10ribbed. Pappus one- to several-seriate, persistent, un­ equal. scabrid.

late, peduncle 0 5 - 1.5 cm long, villous. Receptacle naked, flat c 8-10 mm across. Phyllaries multisenate, imbncate, more or less equal, 9-10 x 1.5-2.5 mm. apex acute, cili­ ate; outermost series foliaceous, densely villous, middle lanceolate, villous, successively wider; inner linear shortly hairy Florets yellow , outermost with strap-shaped rays. Cypselas oblong, 1.5-2.0 mm long, costate, with erect patent short setae near the apex. Pappus brow msh. I-seriate, barbellate. Erica arborea scrub; 3200-3600 m. GD; not known elsewhere. Schimper 1061; Lemma G.S. 690; PichiSermolli 2621. Inula arbuscula is very closely related to the more widespread I. confertiflora, the differences being mainly in the size and shape of the phyllaries and the ty pe and de­ gree of indumentum on the leaves and phyllaries. More collections from the type locality o fI. arbuscula and from Wello and Gojjam might further reveal the affinities be­ tween them

A genus with about 100 species found in Europe Asia and Africa; 4 species in the Flora area.

2. I. confertiflora A. Rich. (1848) - type: SU. 'Choa', Petit s.n. (P holo not seen).

1

Shrub. 1.5-2.5 m lugh; stem much branched above, branches terete, striate, white tomentellous and leafy at extremities. Leaves densely fonned towards apex of branches, submembranous. elongately lanceolate, 3—13.5 x 1-4.5 cm. wlute tomentellous beneath, nearly glabrous above, reticulately veined, narrowed at base, shortly petiolate. margins denticulate, teeth gland-tipped, apex acute. Involucre sub-hemispheric, 10-18 mm across Capitula many-flowered, in dense corymbose or rounded terminal cymes, pedunculate, pcdunclc with foliaceous arbuscula bracts, up to 2.5 cm long. Receptacle flat, naked. 3-5 mm across Phy llanes multiseriate. unequal, imbricate, ciliate; outer successively shorter, appendaged. more obtuse lan­ ceolate and hairy at the back, inner linear, acute. Florets yellow, outermost with strap-shaped rays. Cy pselas ob­ long. 1.5-2.2 mm long, terete, c 10-nbbed. with short, erect-patent setae near the apex. Pappus 1-seriate, c 3 nun long, pale brownish, minutely scabrid. Fig. 145.71.

-

2

3.

Leaves sessile to shortly petiolate. cuneate at the base; inflorescence closely surrounded by leaves: capitula hemispheric, usually 10-20 mm wide at anthesis. Cauline leaves cordatc-auriculate. amplexicaul; inflorescencc carried above leaves: capitula obconic to turbinate, usually 4-7 mm wide at anthesis. Phy llaries nearly equal in length, densely shaggy leaves dark-green, slightly paler beneath. 1 .1, Phy llaries unequal, outermost c half as long as the innermost, becoming glabrous towards center, leaves light yellowish green, silvery beneath 2. I. confertiflora Infloreccncc orange-yellow; phyllaries uniformly light-green, lanceolate to linear-lanceolate, taper­ ing to an acute apex, plants of usually dry w ood­ land 3. L paniculata Inflorescence purple or yellow: phy llaries darkbrown at apex, obtuse; plants of usually forest margins. 4. I. mannii

1 I. arbuscula Del. (1843) type: Ethiopia (region not known|. Ferret el Galinier s.n. (P holo ). I fruticosa Sch. Bip . nom. nud quoad Schimper 664. Shrub, up to 3 m high; branches terete, striate and puberulous below, villous. leafy and brownish above. Leaves ovate or lanceolate. 5-15 x 2.5-6 cm wide, nar­ rowed at the base into a short petiole, pale tomentose beneath, less so above, margins denticulate, apex acute or subacute, mucronulate; upper leaves much smaller In­ volucre hemispheric, c 2.5-3 cm wide Capitula manyflowered. in dense corymbose, terminal cymes, peduncu­

Margins of and clearings in Juniperus Podocarpus forest. Erica arborea scrub, along stream banks in Euca­ lyptus plantations, montane grassland on slopes with scat­ tered Juniperus, Hagenia. Hypericum, etc.; 2500-3730 m. WU SU AR BA HA; not known elsewhere. Mesfin T. 5828; Friis et al 5705; Mooney 6449.

3 I. paniculata (Klatt) Burtt-Da\y (1935); Monactinocephalus paniculatus Klatt (1896) type: South Africa. Transvaal. Rehman 6068 (K iso ). Inulaster macrophyllus Sch. Bip ex A. Rich. (1848); Inula macrophylla (Sch. Bip. ex A. Rich.) Schweinf & Asch. (1867). non Kar. & Kir (1841). nom. illegit.. I. decipiens E. A. Bruce (1947) - type: TU. Mt. Scholoda. PI. abyss. II, Schimper 912 (K syn.. P isosvn.); ibid., PI. abyss. II, Schimper 938 (K syn.. P isosyn.).

145. ASTERACEAE: 44. Inula, 45. Pentanema

123

Erect, perennial herb, up to 75 cm high; rootstock thick, pale-brown villous; stem sub-tetragonal orcylindric, striat pubescent. Leaves both radical and cauline; radical leaves petiolate. up to 60 cm long, petiole up to 10 cm long, clasping at the base, semi-circular and more or less canaliculate or flattened above, sparsely hirsute or gl ibrescent blade up to 45 x 15 cm, ovate-elliptic to obLinceolate. base cuneate and shortly decurrent on the pi uole, forming narrow wings, margins coarsely and irrc siiilarly dentate, scabrous with tuberculate hairs above, p bescent to browny-tomentose beneath, reticulate venati in prominent especially on lower surface; cauline leaves smaller, gradually diminisliing upwards, at first sessile with cuneate base but higher up becoming cordate, a'.ncled and amplexicaul, and passing into the bracts of tl e inflorescence. Inflorescence a dense, terminal, corym­ bose cyme Capitula obconic to turbinate, 4-6 mm wide; pciiuncle up to 1 mm long, scabrous-pubescent. Phyllaries 3-4-seriate. outer lanceolate, 2-3 inm long, inner progressively longer, c 5 mm long, linear-lanceolate, tapering to an acute apex, ciliate on the margins, pubescent outside, s imetimes purple-tipped. Florets all bisexual, ora iLie-yellow. Corolla narrowly tubular below , w idening and funnel-shaped in the upper parts, 4.5-5.5 mm long, 5-lobed; lobes glabrous. Cypselas cylindric, puberulous. 1 2 x 0.2-0.4 mm. Pappus 1-seriate, 3.5-4 mm long, s.:abrid. Pennisetum schimperi grassland, broad-leaved decid­ uous woodland with Combretum, Terminalia, Faurea, et; , grass', glade in moist, semi-evergreen forest; 14002500(-2600) m. TU GD SU AR WG KF GG SD BA HA; Sudan, E Africa. Zambia, Zimbabwe, Malawi, South Africa (Transvaal). Mesfin T. & Kagnew 2439; Ash 673; \lt>oney 5021. 4. L mannii (Hook. f .) Oliv. & Hiern (1877); Vernonia ?mannii Hook. f. (1864) - type: Cameroun, Mann 1933 (upper part of sheet] (K syn.); ibid., Mann 1314 (radical leaf only] (K syn.). Laggera heteromalla Vatke (1875); Petrollinia heteromalla (Vatke) Chiov. (1911) - type: GD, near Debra Tabor, 26. Nov. & 15. Dec., 1863, Schimper 1528 (B holo. destroyed, K lecto., WU iso.). 1 rect. robust, perennial herb, up to c 2.5 m high; stem and 1 'inches dark purplish red, angular ribbed, greyi imentose Leaves both radical and cauline; radical leaves i p to 60 x 20 cm elliptic, attenuate at the base into a short I etiole, sinuate or entire and denticulate, glandular-pilose i i both surfaces, daik-green above, paler beneath. | ! iuline leaves up to 50 x 15 cm, decreasing in size acrol tally, lanceolate-elliptic, sessile, with cordate-auricuue base, margins denticulate; apex acute, mucronate; irface light green often with reddish midrib above, liose, grey-green tomentose beneath. Inflorescence of lense corymbose cymes on both stem and branch apices, apitula cylindric to turbinate, c 5-7 mm wide at anthesis; t duncle up to 1 cm long, densely hirsute. Receptacle reolate or smooth. Phyllaries 4-5-seriate, imbricate, light ;reen with dark brown apex on upper half, linear-

Figure 71 INULA CONFERTIFLORA . 1 - upper part o f stem; 2 - capitulum in longitudinal section x 2x/i\ 3 - ray floret x 5. (No specimens cited in orig.) Drawn by W. Burger. (Reproduced with permission from Burger: Families o f Flowering Plants in Ethiopia , 1967, fig. 68:4.)

lanceolate, grey-hairy; the inner linear and tapering to an acute or acuminate apex, densely hirsute. Florets yellow or reddish purple. Corolla 5-6 mm long, cy lindric but slightly wider in upper parts. 5-lobed. Cypselas 1.2—1.5 mm long, oblong-cylindric, 8-10-ribbed, glabrous. Pappus 3.5—4 mm long, scabrid. Evergreen forest margins, montane slopes with de­ rived evergreen bushland; 2100-2600 m. TU/GD GD SU KF WG; Cameroun, Congo (Kinshsa). Rwanda. E Africa. Malawi. Zimbabwe. W. de Wilde 9533; Friis et al. 498; Benedetto 238. Mooney 6132 from 'marshy ground in grassy valley’ and Anderberg 1638 from Marge bog’ may belong to this species.

45. PENTANEMA* Cass. (1818) Vicoa Cass. (1829). Inula Sect. Vicoa O. Hoffm. (1890). Mesfin Tadesse in Kew Bull. 50: 401-408 (1995). Annual or perennial herbs. Leaves simple, alternate, rarely opposite. Inflorescence a lax cymose panicle. Capitula small, hemispheric, heterogamous or homo-

124

145. ASTERACEAE: 45. Pentanema, 46. Blumea

Erect, annual herb, 0.4-1 m high. Stem simple or branched above, branches sparsely pubescent. Leaves thin, alternate, sessile, 15-80 x 5-15 mm, linearlanceolate to narrowly lanceolate or narrowly elliptic, ta­ pering to an acute apex, auriculate and amplexicaiL at base, puberulous. margins entire to serrate-crenate. mid­ vein prominent below. Capitula homogamous, discoid. 10-15 x 4-5 mm at anthesis, not much altered in height in fruit, arranged in lax cymose panicles; peduncle puberulous. up to 4 cm long, with small (3-10 mm long) linear auriculate bracts which increase in size dow nw ards and pass into the upper leaves. Receptacle epaleate, pitted Phyllaries multiseriate. unequal, 2-6 mm long, increasing inwards, with or without orange glands, outer green, linear-acuminate, sparsely wlute-hairy, spreading or re­ curved; inner linear, tapering to an acute apex, 1-veined, margins hyaline. Florets 40-85, bisexual, yellow, 4-6 mm long. Corolla 3.8-4 9 mm long, narrowly cylindrical, gla­ brous. 5-lobed, lobes densely orange-glandular, c 0.5 mm long. Stamens 4.5-5.2 mm long; anthers c 3 mm long (incl. tailed connective); tail c 0.5 mm long, slightly ex­ ceeding filament-collar joint, connective c 0.3 mm long, ovate. Style c 4.8-5 2 mm long, bulbous at base, bifurcate at apex, branches c 1 mm long, stigmatic surfaces mar­ ginal. confluent at apex. Cypselas light green, c 1-1.3 mm long, narrowly oblong-elliptic, pubescent, base hyaline, annular shirty. Pappus of 3—3.2 mm long scabridulous bristles. Fig. 145.72. Figure 72. PENTANEM A INDICUM'. Habit x ‘/j. (No specimen cited in orig.) Drawn by Bhola Ram. (Reproduced with permis­ sion from FI. India , vol. 13, fig. 9.)

Grassland with patches of bare, pale brown silty soil; 500-800 m. EW IL (Gambella); West Africa, Sudan. Uganda Tanzania, Mozambique, Malawi, Zam bia Zim­ babwe, Angola; Pakistan, India, Sri Lanka. Gilbert & Friis 8365; Pappi 6280; Terracciano & Pappi 2785.

gamous. Phyllaries multiseriate, imbricate, unequal, nar­ rowly linear, tapering to an acute or acuminate apex, mar­ gins scarious. Receptacle epaleate. Florets numerous, ra­ diate or discoid (in the Flora area). Corolla narrowly tubu­ lar 5-lobed; anthers sagittate and tailed at base; style bifur­ cate, stigmatic branches wide, truncate or obtuse at apex. Cypselas narrowly oblong-elliptic, smooth. Pappus of 1-seriate, barbellate bristles.

Cufodontis (1967: 1117) cites Beccari s.n.. from EW, Mai mentai. as this species, but the specimen has not been seen by the present author.

A genus with about 12 species centered in the Middle East and Asia. The following cosmopolitan species is found in the Flora area.

P. indicum (L.) Y. Ling (1965); Inula indica L. (1763) - type: “Habitat in India orientali.” Linnaean Herbarium No. 999.16 [LINN, lecto., selected by Anderberg in Taxon 32:652 (1983)). Vicoa auriculata Cass. (1829). - type: Sri Lanka. Merat s.n (P not traced). Inula leptoclada Webb. (1849); Vicoa leptoclada (Webb.) Dandy (1956) - type: Cape Verde Ils, Webb s.n. (P holo.). Inulaster kotschyi Hochst. (1841), nom. nud. quoad Kotschy 108 (Sudan); Varthermia kotschyi Schweinf. & Asch. (1867), nom. nud., quoad CienIcowsky s.n (Sudan); Conyza kotschyi Schweinf. & Asch. (1867), nom. nud.

46 BLUMEA DC. (1833) A J. Randena in Blumea 10: 176-317 (1960); A.A. Anderberg in PI. Syst. & Evol.. 176: 105 (1991) & Willdenowia 25 19-24 (1995). Annual or perennial herbs. Indumentum of 1-seriate, multiseriate and capitate glandular hairs. Stems simple or branched, usually erect, sometimes ascending, terete, gla­ brous, pubescent or tomentose. Leaves simple, alternate. Unear-elliptic to obovate-lanceolate, margins serrate, dentate, denticulate or pinnatifid, sessile with rounded semi-clasping base or petiolate with tapering base, pubes­ cent to villous, apex acute-apiculate. Capitula hetero­ gamous, campanulate, disciform, pedunculate or sessile, in lax or compact terminal and axillary panicles or cory mbs. Receptacle epaleate, flat, convex, areolate or alveolate. Phyllanes 4-5-seriate, imbricate, increasing in length inwards, herbaceous, linear to linear-oblong, densely pubescent to villous. Florets purple, pink or vio­ let, numerous, outermost female, innermost bisexual or female. Corolla of female florets filiform, 2-4-lobed or

145. ASTERACEAE: 46. Blumea

bilabiate, glabrous; of bisexual florets tubular-funnelsha; ed with the tube expanding from the base upwards, sometimes with sessile glands in upper parts, 5-lobed, lobts papillose. Stamens 5; anthers sagittate with tailed b a s.. connective triangular with obtuse apex. Styles exserted, bifurcate. Cypselas brown, angular ribbed, pu­ bescent. Pappus white, 1-seriate, barbellate. A tropical and subtropical genus with about 100 spe­ cie: many occurring in SW Asia. A few species are native to Africa, 3 species in the Flora area. ■ a note on the relationships between Blumea and oth-:i genera. Randeria [in Blumea 10; 191 (I960)] wrote tha "Laggera and Blumeopsis perhaps have the closest aff: uties ...” with Blumea and also provided the differen­ tia] characters. Wild (1969), in the Compositae of the Flo' ^ of Zambesiaca area, argued that Laggera can not be maintained as a distinct genus and thus reduced it to synom my under Blumea. Although Laggera and Blumea sh2 ret some characters the author here follows the traditioi al definition of these genera and keeps them apart. Blumea aurita (L. f.) DC., transferred to Laggera by v Randeria (1960: 298) is now Pseudoconyza viscosa (Mill.) D’Arcy [in tribe Plucheeae (Benth.) A. Anderb.]. The anther tails in Pseudoconyza viscosa (syn. B. aurita) do rut reach up to the joint of the filament-collar, but are dist inctly longer than in members of Laggera. However, antiier tails with unequal lengths were observ ed in some spe ::imens of P. viscosa. The female florets are bilabiate in P viscosa, whereas in Laggera thev are distinctly 4 (-1 -lobed. Recently, Anderberg (in Willdenowia 25: 19-24, 19'- 5) ”resuscitated” the genus Doellia Sch. Bip. ex Walp. ( 1M 3) and included in it two of the species that were kept in i'umea by Randeria and others, i.e., B. bovei (DC.) Vatke and B. caffra (DC.) O. Hoffm. The major reason for thi s the presence of “conspicuous resiniferous ducts appeanng as longitudinal, red strips in the mature cypselas." According to this and and also according to an earlier Study, Doellia was placed in the tribe Plucheeae, close to the Laggera clade”. (Uumea appears to be a polyphyletic genus but monoph ly, based on a single or a few characters and particu­ lar;;, taking only a tiny segment of the genus, must not be employed for recognition of genera. In relating Blumea to other genera in Inuleae s.l., Randeria wrote '‘further divi­ sion of the Plucheinae into genera has been acliieved on :hc basis of differences in the pappus and other individual characters of the capitula, florets and phyllaries. The same ge itral trend for delimiting genera by single or very few ch .racters seems to chracterize the entire Compositae and other families like Gramineae, Cruciferae and Umbellife r ie .” Randeria studied 49 of the species in the genus rather we i i. but left about 49 others as “taxa and names of uncer­ tain status”, an unusually large number of names to be untreaed in a work that purports to be a revision of a genus. Anderberg (1991:105) reiterated this and in his latter work (Anderberg 1995:19), where he transferred two of

125

the species to Doellia, also wrote ’’more work remains to be done before the taxonomic situation in Blumea is un­ derstood in detail,...” The name Doellia is not typified, as is clearly stated by Anderberg (1995 :20), but is being proposed for use w ith an amended description because of its association with Blumea abyssinica Sch. Bip. ex A. Rich. (=B. bovei). This may be sufficient to warrant the recognition of Doellia as a genus but due to the reservations expressed above, and also due to the large number of names that still need to be addressed in Blumea, including the widespread B. axillaris that occurs in the Flora area, the author prefers to maintain the three African species in Blumea until a more comprehensive treatment is obtained. 1.

2.

Plant up to 75 cm liigh; leaves linear or ob­ lanceolate, rounded or clasping at base; capitula pedunculate, peduncle up to 4.5 cm long. 2 Plant up to 1.5 cm liigh; leaves obovate to elliptic, often narrowed or tapered to the sessile or petiolate base; capitula sessile or peduncle only up to 1 cm long. 3. B. axillaris Capitula 3-8 mm wide; corolla lobes of female flo­ rets glabrous. 1. B. bovei Capitula 8-15 mm wide; corolla lobes of female florets with 1-4 erect bristles. 2. B. caffra

1. B. bovei (DC.) Vatke (1875); Convza bovei DC. (1834) - type; Egypt, Sinai, Bove 118 (G-DC holo., K iso.). Blumea abyssinica Sch. Bip. ex A. Rich. (1848) type. EW, near Aguar IAgerJ, Medat [ModatJ Valley, PI. abvss. II, Schimper 1006 (P syn.. K isosyn.) & 1032 (P syn.). Perennial herb, up to 50 cm high, with a woody rootstock, rarely annual with a long taproot. Stems solitary or sev­ eral. each ascending from the apex of the rootstock, branched or unbranched, terete, light green to dark purple, sparsely pubescent. Leaves thick, linear-oblong to ob­ long-lanceolate, 15-80 x 2-8 mm, sparsely pubescent on both sides to glabrous, sessile-auriculate with rounded base, margins sharply denticulate, apex acute-apiculate. Capitula heterogamous, campanulate, 5-6 x 3-4 mm at anthesis, solitary or in open, branched coiymbs. pedunculate, peduncle up to 4.5 cm long, bracteate. sparsely to densely pubescent. Receptacle flat, alveolate. Phyllaries c 4-seriate, purple-tinged, linear, densely pu­ bescent with both glandular and eglandular liairs; outer c 4 xQ.5 ram; inner c 5-6 x 0.5 nun, margins scarious. Florets purple. Corolla of female florets filiform, c 2.7—3.1 mm long, 2-3-lobed; of bisexual florets 4.0-4.8 nun long, 5-lobed. lobes papillose and with stiff bristles, funnelshaped. Cypselas c 0.7 x 0.3 mm. sparsely pubescent, subtetragonal. Pappus white, 2.5-3 nun long, barbellate. Fig. 145.73. Wet montane grassland w ith seasonal streams; river banks; 1200-2200 in. EW TU GJ SU HA; Egypt. Sudan. Somalia; Yemen. Saudi Arabia, Oman. Palestine. Afganistan. Schweinfurth & Riva 1097; Schimper 111; Mesfin T. & Sebsebe D. 3764.

126

145. ASTERACEAE: 46. Blumea

monomorphic. linear to linear-lanceolate with narrow scarious margins, densely pubescent on outside mainly along the grey-green (dark purple when diy) median striae. 3-6 x 0.5-1 mm, apex long acuminate. Florets pur­ ple. Corolla o f female florets filiform. 3-4-lobed. c 3 mm long, glabrous: style c 4 mm long; corolla of bisexual flo­ rets tubular, funnel shaped above, c 3.5 mm long, 5-lobed; style c 4.5 mm long. Cypselas black, densely white pubes­ cent. c 1 x 0.5 mm. Pappus white, pubesccnt. 3—4 mm long Riverine scrub with Acacia, Tamarix. etc.; saline hab­ itats and hot springs; 14(H)-1600 m. SU; Kenya, Tanzania Mozambique. Zimbabwe. Botswana, Namibia. South Af­ rica and Madagascar. Friis et al. 713; Gilbert & Gilbert 3965.

Iigurc 73. BLUMEA BOl E l 1 upper stem portion. 2 female floret x 4 1'3'.3 - bisexual floret x 4 1/3; 4 -cypsela with pappus x c 7. (No specimens cited m orig.)D raw n by Ruth Koppel (Repro­ duced from FI. Palaestma. vol 3. PI. 500.)

2 B. caffra (DC.) (). Hoffm. (1889); Conyza caffra DC. ( J836) - type: S Afnca. Cafra. Key River. Drege s.n. (G-DC hoio.. K fragment). Erect perennial herb, c 50-75 cm high, stems several from the woody rootstock, terete, glabrous to sparsely ciliatc. dark-purplish or reddish-brown at base. Leaves sessile, yellowish-green, oblong to oblong-lanceolate alternate, diminished in size upwards. 3-6 x 0.5-1 cm. wider at base, sparsely hairy on veins beneath Capitula solitary at leaf axils or several iti open corymbose cymes, pedunculate. 8-15 x 6-7 mm. peduncle up to 4 cm long, pubescent, bracteate. Phyllaries multi-seriate, graded

3. B. axillaris (Lam.) DC. (1836); Conyza axillaris Lam. (1786) - type. Maurice (Mauritius), C'ommerson s.n. (P-LA holo.. P -JU 8615. P iso ). B. lacera (Burm. f.) DC. (1836); Conyza lacera Bunn f ( 1768) - type: Java. Burmann s.n. (G-DC. not seen). B. lacera forma elata robusta Vatke in Linnaea 39; 485 (1875). nom. non rite publ.. quoad Schimper 810. Blumea drageanoides Sch. Bip. ex A Rich (1848) - types TU/DG. near Ferrokoba Schimper 633 (P syn.. K isosyn.), GD, Wogcra. Schimper 1297 (P syn , S isosyn ); TU. Cliire, Ouartin-Dillon s.n. (P syn ); Choa. Petit s.n (P syn., K isosyn). B. lacera forma humilisflaccida Vatke in Linnaea 39: 485 (1875). nom. non rite publ.. quoad Schimper (1862) 804 & (1862) 182 B. lacera form elata robustior Vatke in Linnaea 39: 485 (1875). nom. non rite publ.. quoad Schimper 810 B. lacera m c\.. non (Bunn, f.) DC. (1834). quoad Schimper 810 B. solidagmoides (Poir.) DC. (1836). B. hieracifolia auct non DC (1836). quoad Negri 519 (FT). B. lacera (Bunn f.) DC. var. glabrescens Chiov.. nom. nud.. quoad Vatovn 397 (FT). Annual or shon-livcd perennial herb, up to 1.5 m high, w ith a long taproot Stem usually solitary, sometimes sev­ eral from the base, smaller and narrower than the central one and ascending, stnatc. terete to angular often densely tomentose to veiutinous with silky pale brownish hairs, sometimes also with glandular liairs. Leaves obosatc to elliptic-obovatc. up to 12 x 3.5 cm, progressively dimin­ ishing in size upwards, narrowed to the sessile base or shortly petiolate. petiole up to 3 cm long, tomentose to veiutinous on the upper surface, tomentose to woolly on the lower surface, margins serrate, apex acute or obtuse Capitula heterogamous. campanulate. 4-6 x 3-4 nun] ses­ sile to pcdunculatc. in axillary and terminal dense pani­ cles; peduncle up to I cm long, densely tomentose or veiutinous; luurs dark purple. Receptacle flat or convex, alveolate. Phy llancst 4-scriatc, linear densely veiutinous. also with glandular hairs; outer 2-2.5 x 0.3-0.5 nun, pur-

127

145. ASTERACEAE: 46. B lu m ea.47. Pegolettia

Figure 74. BLUM EA A XILLA IUS 1 part o f flowering plant x : r. 2 capitulum . longitudinal section x 9; 3 4 phyllaries x 9: 5 outer floret (pappus removed) x 18:6 inner flo­ ret (pappus removed) x 18: 7 sta­ mens x 18. 8 sty le o f inner floret x 18. 9 cypsela x 36: 10 pappus bristles x 18. All from Gueho in MAIJ 15199. Drawn by Pat Halliday. (Reproduced from FI Mascareignes . 109. C om posees. PI. 19.)

pie ringed; inner 5-6 x 0.3-0.5 mm. margins scarious. Flor:ts pale purple, whitish at base of corolla. Corolla of fen lie florets filiform, c 3.4 mm long, glabrous, those of bisi >.ual florets funnel-shaped, c 4.6 mm long. 5-lobed. lobes papillose. Cy pselas light yellowish-brown, somewha angular, sparsely pubescent on the nbs. oblong, shin, c 8.0 x 0.3 mm Pappus white. 3.0-3.5 mm long. Fig 45.74. disturbed places in rocky hills with remnants of tall Albizta woodland, dense bushland of ('arissa and PteroIbbI ilw, Acacia bushland. grassland with Acacia on allu­ vial soil, stream or river banks, near waterfall, waste places; 850-2575 m. EW TU GD SU KF GG SD HA; widespread in Africa and SE Asia. Burger 3710; Gilbert 46(> Mesfin T. & Sebsebe D. 3773.

47. PEGOLETTIA Cass. (1825) Slmibs. subshrubs or herbs with erect or spreading branches; old stems glabrous, smooth, brown, reddish, yellowish brown, or greyish; young branches green, stri­ ate. glandular-pubescent. Leaves alternate, filiform, lan­ ceolate. oblong or spathulate in outline, simple to pinnatifid. glandular-pubescent. Capitula discoid, homo­ gamous. solitary to loosely corymbose, tenninal. Phy Maries leathery. lanceolate to narrow ly oblong or nar­ rowly obovate. glandular margins ciliate. apex acute to acuminate. Receptacle epaleate. flat. Florets bisexual, yellow, turning purplish vvith age; corolla glandularliairy 5-lobcd. lobes narrowly triangular, glandular base widened into a ring-shaped foot: anther base sagittate, tails branched: style bifid al apex and swollen at base.

145. ASTERACEAE: 47. Pegolcttia. 48. Pulicaria

128

Erect, annual herb, 8-60 cm high, stem sparsely to densely branched, green, glandular-pubescent. Leaves simple, 5-40 x 1-10 mm. narrowly linear to lanceolate or spathulate. glandular-pubescent, sessile, margins serrate, apex obtuse with a small mucro. Capitula 5-10 x c 5 nun, in loose corymbs, rarely solitary. Phyllaries c 3-senate, spreading at anthesis. apex acute, inner longest. Florets yellow or purplish (with age), 14-30. Corolla 6-8 x 1 1.5 mm. zygomorphic with one large lobe with bristly hairs at apex and the rest minute. Anthers c 2 mm long, base sagittate with long tails. Style 5-6 mm long, branches c 1.5 mm long. Cypselas 3-4.5 x 1-1.5 mm. thinly pubes­ cent. 12-16-ribbed. ribs with protrusions toward the apex. Pappus 1-seriate, with more or less equal plumose bristles and deeply inctsed scales externally. Fig. 145.75 Grassland with scattered Acacia, grassland on loamy soil, rocky hills with Acacia and Balanites. subdesert: 260-2000 m. EE SU GG SD HA: Senegal. N igena Chad. Algeria. Egypt. Sudan. Somalia Kenya. Tanzania. Zim­ babwe. Angola. Zambia. Mozambique. Botswana Swazi­ land. Namibia. S Africa. Cape Verde Island: Yemen. Saudi Arabia. Oman. Pakistan. India. Mesfin T. & O Hed­ berg 1116: Gilbert et al. 230: Sue Edwards et al. 2763.

48. PULICARIA Gaertn. (1791) E Gamal-Eldin in Phanerogam. Monogr 14: 1-311 (1981).

r t Figure 75 PEGOLETTIA SE N E G A L E SSIS 1 habit \ * j. 2 4 phyllaries (outer, middle and inner) x 3. 5 - cypsela with pappus x 6; 6 outer pappus x 10; 7 corolla with style branches x 6; 8 stvle branches x 10. 9 stamen with tailed anthers x 10. (No specimens cited in orig.) Drawn by M J Vesque & D Godot de Mauroy. (Rearranged and reproduced from Flore de Mada­ gascar et des Comores. C om posces. Tome II. tig. 110.)

Cypselas narrowly cylindrical, tapered to base. 12-20ribbed. ribs with knob-like protrusions toward apex, glan­ dular-pubescent Pappus of one to several-seriate barbellatc or plumose bnstles and small or large, entire or in­ cised bnstle-Iike scales. A genus with 9 species mainly occurring in S Afnca; one species in the Flora area. P. senegalensis Cass. (1825) - type: Senegal. Gay 86 (P holo.. K iso ). Kuhnia arabica DC. (1838) - type: Yemen. Gesser Mts. Schimper 863 (G lecto.. FT K M NY iso ).

Perennial or rarely annual herbs, or shrubs: whole plant usually sparsely to densely hairy . Leaves simple, alter­ nate. auriculate or cordate, sessile, rarely petiolate. mar­ gins denticulate, entire or serrate. Capitula hcterogamous. radiate, hemispheric or rarely campanulate (in P. soma­ lensis). solitary or in paniculate cymes. Receptacle epaleate. flat or convex, arcolate. Phyllarics few- to many-seriate, sparsely to densely pubcsccnt and glandular to tomentose: outer usually herbaceous and shorter Flo­ rets yellow. Ray florets female, with shon or long ray s. glabrous, pubesccnl or glandular Disc florets bisexual Corolla tubular 5-lobcd. lobes acutc. Anther base sagittate, tailed Style bifurcate, branches obtuse. Cypsclas c 10-ribbed. cylindric to slightly angular, sparsely setose, light to dark brown Pappus dimorphic and 2-scnatc: outer cup-shaped or scale-likc. 0.1—0.7 mm long: inner setaceous or flat and wider toward apex in P. somalensis). 2.5-5 mm long, barbellate. or usually smooth toward the base. A genus with about 54 species in Europe. Asia, Mid­ dle East. North, tropical and South Afnca. 11 species in the Flora area 1

2.

Leaves petiolate: petiole narrowly w inged, up to 1.5 cm long: blade ovate to lanceolate. 8. P. petiolaris Leaves sessile with cordate or auriculate-amplexicaul base or narrow ed from about middle to base, blade obovate-spathulalc. oblanceolate. oblong or linear. 2 Outer pappus scale-like, all or most separate. 3

145. ASTERACEAE: 77. Conyza

Moist bank in low scrub on denuded hillsides, among bushes in woodland; 2200-3000 m. EW TU SU AR KF SD BA HA; Sudan, E Africa, Cameroon, Rep. Dem. du Congo, Rwanda, Burundi, Malawi, Zambia; Yemen. Mooney 5005; Mesfin T. 8430, 7534. 1< C. schimperi Sch. Bip. ex A. Rich. (1848); Erigeron schimperi (Sch. Bip. ex A. Rich.) Schweinf. & Asch. (1867) - type: TU. Adwa, PI. abyss. I, Schimper 121 (P holo., FT K iso.). C. schimperi var. adoensis Vatke in Linnaea 39: 482 (1875) - type: TU. Mt. Scholoda, near Adua, 15 Aug. 1862, Schimper 29 (B holo., BM iso.). C. schimperi var. bipontini Vatke (loc. cit.) - type: TU, Hedscha [East of Adua], 5 Oct. 1862, Schimper 354 (B holo., BM iso.). C. schimperi subsp. longepapposa R.E. Fries in Acta Hort. Berg. 9(6): 127 (1928). Ei :ct, perennial herb, with deep-seated taproot or su )shrub, 50-100 cm high; stem woody, terete, green or so netiines purple-tinged, striate, sparsely to densely pubc scent with often both glandular and eglandular liairs. ra ely with only densely capitate-glandular liairs, young bi nches tomentose. Leaves pale green, pinnatisect or pi natilobed, sessile, 1-5 cm long, sparsely to densely pubt ;:ent, often with dense sessile glands, lobes triangular w ih sharply drawn-out teeth. Capitula campanulate, 3-4 x 3 mm, in dense, corymbose cymes; peduncle absent or uf to 3 mm long, sparsely to densely pubescent, sometii ;cs tomentose or densely capitate-glandular. Recepta­ cle convex, epaleate. Phyllaries 3-4-seriate; outermost lii ..ar-oblong. densely pubescent and sessile-glandular, gr *nish in middle, rarely purple-tinged at margins and apex, 1-2 x 0.5-0.8 mm; innermost glabrous or sparsely se sile-glandular in middle, c 3 x 0.5 mm. Florets yellow . E: temal florets female. 2.5-3 mm long, ligulate; ligule slightly wider than the tube, c 0.8 mm long. 2-3-fid at ap :x; tube c 1.5 mm long, pubescent; pistil slightly e.\ erted. Central florets bisexual, c 3 nun long; corolla pa iillose-liairy at middle and below, c 2.5 mm long, 5- obed, lobes sessile-glandular. Cypsela light brown, ot >vate. 0.8-1 x c 0.5 mm, densely pubescent. Pappus se ,ie 1-seriate, 2.5-3 mm long, white, barbellate. Dry, scrubby grassland, dry, grassy and stoney m ntane slopes with/tosa, Carissa, etc.. presumably derri ed from Juniperus forest; 1500-3300 m. TU GD GJ W SU KF SD BA HA; Sudan, E Africa, Rep. Dem. du C< ngo. Rwanda Burundi. F m setal. 3063; Moonev 5838. 7088. 1” C. stricta Willd. (1803) type. India Vellore. W of Madras, Klein s.n. in Herb. WILLD (B holo.). Ei et annual or short-lived perennial herb, woody below, 4( '5 cm high, with long taproot; stem and branches le; , sparsely pubescent and sessile-glandular, terete, su ate. Leaves simple and linear-spathulate or spathulate to unnatifid with linear segments, usually held erect, light yellowish green, sessile, sparsely to densely hispid-

209

pubescent and sessile-glandular, up to 7 x 2 cm. margins entire to crenulate. Capitula campanulate 2.5-3 x 2-2.5 mm, in dense coiymbose cymes, peduncle up to 6 mm long, densely pilose and sessile-glandular, rarely sparsely hispid. Receptacle convex, pitted. Phyllaries 3-4-seriate. linear-lanceolate, acute at apex; outermost hispidpubescent and densely sessile-glandular, greenish in mid­ dle, 2-2.5 x c 0.3 mm; innermost glabrous. 2.5—3 x c 0.4 mm. yellowish. Florets yellow. External florets female. 2.5-3 mm long, ligulate or minutely bilabiate and usually 2-3-fid, ligule same width as tube, pubescent, tube filiform, sparsely hispid. Central florets bisexual, c 3.5 mm long; corolla campanulate. sparsely hispid and ses­ sile-glandular below. 5-lobcd. Cypsela light brown, obovate, c I x 0.5 111111. pubescent and sessile-glandular. Pappus setae 1-seriate. 2.5-3 mm long, w hite, barbellate. 1. -

Leaves distinctly pinnatilobed to pinnatisect. var. pinnatifid a Leaves not lobed. margins dentate or serrate. var. stricta

var. stricta C. macrorrhiza Sch. Bip. ex A. Rich. (1848); Erigeron macrorrhizus (Sch. Bip. ex A. Rich.) Schweinf. & Asch. (1867) - type: T1J. near Adwa PIabyss. II, Schimper 1104 (P holo., BM K WU iso.). Conyza macrorhiza forma silvestris Vatke in Linnaea 39: 483 (1875) - type: TU, Schahagenna (west of Adua], Schimper 238 (B holo.). C. vatkeana Oliv. & Hiem . Bot. Agr. Indus­ try. Abissinia sett.: 33 (1912); A arborescens L. forma rehan (Chiov.) Chiov. in Savoia-Aosta (ed.). La Esplorazione dello Uabi-Scebeli ...: 417 (1932) lype: SU. Entoto, Tra\>ersi s.n. (FT holo.). A. arborescens L. var. cupamana Chiov. in Savoia-Aosta (ed.). loc.cit. 417 (1932). based on the

invalid name A. arborescens var. /loribus abortivus Guss. in Florae Siculae Syn. 2(1): 456 (1844). A. arborescens auct., non L. (1763): Chiov. (1937). Cufod. (1947). Erect, perennial herb, 30-60 cm high. Older stems grey, glabrous, younger ones densely appressed white tomen­ tose. Leaves bipinnatisect. whitish or silvery tomentose. basal ones up to 35 cm long, upper cauline leaves 3-5 cm long, segments 5-20 x 1-2 mm. Capitula subglobose. 3-4 x 3-4 mm at anthesis. Phyllaries grey -green with scanous margins. 2-3 x I mm. ovate-lanceolate. Florets yellow.

224

145 ASTERACEAE: 81. Artemisia. 82. Dendranthema

Wormwood (common English name for A. absinth­ ium) was widely introduced in temperate countries as a medicinal, aromatic and ornamental hert>. About a hun­ dred years ago, the alcoholic drink called absinthe and prepared from the essential oils of this species was widely used in the temperate world. C. Kowalchik & W H. Hylton (eds.) w rote (in Rodale 's Illustrated Encyclopedia o f Herbs, 1987. Rodale Press. Emmaus. Pennsylvania) ”if you want to sec what that beverage [absinthe) did to peo­ ple. you need look no farther than the nearest reproduction of lidgar Degas's painting The Absinthe Drinkers (...) A whole history could be written about the rise and fall of absinthe as an addicting and deteriorating drink that led to serious mental disturbance, to seizures, and sometimes to death. It is no wonder then tliat absinthe is illegal in most countries of the world, including the United States and Canada. France, which was the leading producer of ab­ sinthe. was also one of the last countries to ban it (in 1915). Wormwood, however, is still used to flavor alco­ holic beverages, including vermouth and Campari r Al­ though there is no documented study known to the author on the industrial use of A. absinthium in Ethiopia, its use in local ARAKI should be discouraged

82. DENDRANTHEMA* (DC.) D esM oul (I860) Figure 133 ARTEM ISIA A B SIN TH IU M 1 - habit x 1/ 3; 2 capitulum, longitudinal section x S'/a. (No specimens cited in orig.) Drawn by John H. Rumelv. (Reproduced with permission from Intermountain Flora, Vascular plants o f the Intermountain West, USA, vol. 5, 1994, p. 153 p.p.)

C y p selas m in u tely striate , d ark b ro w n , n arro w ly o b o v ate,

0.8 x 0.2 mm. Fig. 145.133. Widely cultivated especially in the northern and cen­ tral parts of the Flora area for its aroma and widely applied in rituals called 'adbar” and in flavoring a locally distilled alcoholic drink called AREK1 (in amharic); 1700-2350 (-2440) m. EW TU GD SU WG HA; native to the Medi­ terranean region but widely cultivated in the temperate world for the active ingredient used in the alcoholic drink absinthe (see below). Griaule 89; Sebsebe D. 3037; Sebsebe D. & Nigist 2399. Artemisia absinthium is similar to A. abrotanum L. with which the Ethiopian specimens should be compared. Although the type of A. rehart is from Entoto (SU] it was described by Traversi as “pianta odorosa cultivata” . Cur­ rently the species is only known from cultivations in pri­ vate gardens. In his initial description and diagnosis, based on hand-written notes on a sheet attached to Chiovenda 2746/2766, another cultivated specimen obtained from Gonder (GD) on 31 Oct. 1909, Chiovenda wrote that his species, i.e., A rehan, is affiliated to A absinthium, thus offering another clue for the true affinity of this taxon A. rehan is, therefore, considered synonymous with the widespread Mediterranean species.

Perennial herbs or subshrubs. Leaves alternate, pinnatisect, lobed, serrate or rarely entire. Capitula radiate, heterogamous. arranged in open corymbs (rarely soli­ tary), or all radiate except for the few central ones, pedunculate. Receptacle epaleate, convex to conical. Phyllaries ashy-greea generally with dark brown mar­ gins. Ray florets female, fertile, white, pink, or yellow. Disc florets yellow, bisexual. Corolla tubular, often with sessile glands Anthers translucent ecalcarate, ecaudate, base obtuse, connective ovate-triangular. Style bifurcate, branches truncate and penicellate at apex. Cypselas obovate, faintly 5-8-ribbed, often with myxogenic cells in rows. Pappus absent. A genus of about 37 species distributed in Asia, mainly in China and Japaa The widely cultivated garden chrysanthemums’, a complex hybrid group, are believed to have been raised in China from D. indicum (L.) Des Moul.. a species native to India. This group of hybrids as well as others derived from it are widely cultivated in Ethiopia. All of them are here treated as belonging to D. x grandiflorum (Ramat.) Kitam D x grandiflorum (Ramat.) Kitamura (1978); Anthemis grandiflorum Ramat. (1792) - type: Probably from China or Japaa not traced. Erect or spreading perennial herb, 50-75 cm liigh. Stem solitary or often branched, grey pubescent or tomentose, stoloniferous and with short creeping rhizomes. Median cauline leaves chartaceous, ovate-oblong, usually thick, shallowly lobed to pinnately cleft or lobed, dissections of lower segments sometimes reaching mid-rib, those of up­ per segments one-third to one-half the depth of the blade,

145. ASTERACEAE: 82. Dendranthema. 83. Santolina

225

petiolate. 3-12 x 2.5-7.0 cm across the segments, upper su ace ashy green with usually sparse white hairs, lower su ace sparsely lo densely grey-pubescent or tomentose. pe le 1-3 cm long, clasping the stem. Capitula hemi­ spheric, 35-60 \ 8-12 mm at anthesis. solitary or 1-3 per br itch, peduncle 1-4 cm long, grey tomentose. Phyllaries he tjaceous. 3-4-seriate. densely pubescent to tomentose, espocially at base, grey-green with dark brown scarious mi rgins. 3-7 x 1-2.5 nun: outer lanceolate. 3-4 x c 1 mm. gn en in middle, grey-brown at margins, inner 6-7 x 2-2.5 nu i. oblanceolate. green in middle and with wide, sc; nous, often lacerate or lobed margins. Ray florets w lute, yellow or pink-purple, one to several-seriate, often qu lied, lincar-lanceoalte, female, fertile or female-sterile, 10 5 x 3-5 mm. apex acute or minutely 2-3-fid. Disc flc rets yellow, bisexual. Corolla tubular, slightly bulbous at ase. 3.5-4 nun long, glabrous. 5-lobed. lobesc 0.7 nun loi g Anthers translucent, base obtuse, not reaching filami nt-collar joint, c 1.5 nun long, connective triangular lai .eolate. Style bifurcate, 2-2.8 mm long, branches trunca eand pcniccllate at apex. c. 0.7 nun long. Cypselas light bn \s n. c 1.8 nun long, slightly curved. 5-grooved. Fig. 145.134. ’ * (Gardenornamental: 1770-2440 in. EW SU IL SD HA. ( it achew A. 1352; Afeworki & Alehr el 16; Dem el T. 326. his is a widely cultivated plant, often known in commi : _c as 'the chrysanthemum'. It is known in many cultiva ad forms, displaying different colour types of the rays (y How. pink, lilac, shades of red. white, etc.). and differ­ ent si/cs of capitula. The capitula in the most hybridized forms consist of quilled or rolled in* ray florets. It is assuncd to have been derived from D. indicum (L.) Des VI ml. (Chrysanthemum indicum L.). In some floras the name Chrysanthemum morifolium Ramat. (1792) or Dendranthema mori folium (Ramat.) Tzcvlev. (1961) is utilized for certain cultivars of this hybrid.

83. SANTOLINA*

Figure 134. DENDRANTIIICMA G R A N D /E L O R I A/ Upper part o f stem \ 2■? From Afeworki (i. MeliretA. 16. Drawn b\ Birhanu Mihrctu.

(1753)

Sn ;;ll shmbs or densely branched and bushy perennial he bs. aromatic. Stems erect, branched, brittle. Leaves al­ ter late, dentate to pinnatifid or pinnatisect. Capitula ho uogamous. solitary , discoid, pedunculate. Receptacle co vex. palcatc. Phyllaries light green. 2-3-seriate. sc; nous at margins, keeled in the middle. Paleae with a median resin canal. Florets bisexual, yellow. Corolla basaiiv saccate around the cypsela. especially adaxially. \\i h a ratlier long. bent, greenish tube and a distinct. 5- 'bed limb. Anthers translucent, yellow, bases shortly saj iitatc. connective lanceolate. Style bifid, amis with ma rginal stigmatic areas, apex truncate, densely pa ' Hose. Cypselas oblong. 3-5-angled, sometimes with nr \ogcmc cells. Pappus absent. A genus of about 8 species distributed in southern Eu­ ro: h. particularly in Spain and Croatia, and in North Afric i (Morocco and Algeria): 1 species introduced as a ga den ornamental in the Flora area.

S. chamaecyparissus L. (1753) type: “Habitat in Europa australi. " Linnaean Herbarium No. 985.1 |LINN lecto.. selected by Arrigoni in Webbia 34: 260 (1979)). Shnib or erect, bushy, richly branched, perennial herb. 20-60 cm high. Stem loosely tomentose. silvery-grey to white. Leaves alternate, pinnatifid to pinnatisect. 2 0 - 4 0 x 4-5 nun. loosely to densely tomentose. segments nar­ rowly oblong or linear. Capitula solitary , hemispheric, discoid. 8-15 x 6-8 mm. peduncle striatc-sulcate. loosely tomentose. Phyllaries light green with scarious margins. 3-4.2 x 0.5-1 9 nun at anthesis. Paleae carinatc and thick­ ened in the middle. 7.5-8.5 x 0.8-1.1 nun. ciliatc toward the apex. Florets homomorphic, yellow. Corolla with greenish tube and yellowish, glandular limb, c 3.5-4.0 nun long. 5-lobed. lobes c 0.8 111111 long. Anthers translu­ cent. sagittate at base, not reaching filamcnt-collar joint, c 1.0 nun long, connective c 0.5 mm long. Style bifurcate.

226

145. ASTERACEAE: 83 Santolma. 84. M atricaria. 85. Cotula

84 MATRICARIA* L. (1753) Annual herbs. Leaves alternate, pinnatisect. Capitula soli­ tary. shortly pedunculate, radiate and hclerogamou.s or discoid and homogamous. Receptacle conical to subulate at maturity, hollow, epaleate. Phyllaries 2-3-seriate. im­ bricate. margins and apices scarious. Ray florets (w iien present) female, fertile, white Disc florets bisexual, vellow. Corolla 4- lo 5-lobed. tube swollen in fruit, lobes rarel\ with resin canals. Cypselas slightly dorsiventrally compressed, with 5 mainly adaxially arranged thin nbs, sometimes with resin canals, abaxially and on the ribs covered with myxogenic cells. Pappus absent or a small corona or sometimes, especially in ray cypselas, an adaxial auricle. A genus of about 7 species widespread in Europe, the Middle East, temperate Asia. North Africa and North America. 1 species. M. recunin L.. lias been under cultiva­ tion in the Institute of Agricultural Research (IAR). Ambo M. recutita L (1753) type: "Habitat in Europa." No original material Type cons prop.: Podpcra in FI. Exsicc. Reip. Boh.-Slov. No. 946. II (K), proposed b> Jcffre> in Jarvis (ed ), Taxon 41: 566 (1992). under consider­ ation. KI. chaniomilla L. (1753). - type: "Habitat in Europae agris. cultis/’ Herb Clifford: 415. “Matricaria T ’ [BM lecto.. selected by Grierson in Notes Roy. Hot. Gard. Edinh. 33: 252 (1974)|. Ercct annual herb. 10-50 cm high. Stem glabrous, solnan or sometimes branched from near the base Leaves pinnatisect. glabrous, lower ones 2-5 cm long, oblong or oblanceolate in outline, segments c 0.5 mm w ide Capitula solitary or 3-10 in subcorvmbose cymes. 12-17 x 1-4 mm at anthesis. radiate, heterogamous; peduncle stnate. glabrous, slightly wider at base of reccpatacle. Phyllaries oblanceolate. 2 5—3.5 x 0.7-1.2 mm. glabrous, obtuse al apex. Ray florets 12-15, female, fertile, patent at antliesis. reflexed in fruit. 4-8 x 2-3 mm. apex minutely 2-3-fid Disc florets yellow, bisexual. Corolla tubular. 1.1-15 nun long. 5-lobcd. lobes c 0.2 mm long. Anthers c 0.5 nun long, obtuse at base, conncctivc lanceolate Style bifur­ cate. c 1—1.5 nun long, branches c 0.4 mm long. Cypselas brow n, c 0.7 x 0 3 nun. w ith 5 w hitish ribs. Fig 145 135 Figure 135 M ATRICARIA RECUTITA I - habit; 2 capitulum, longitudinal section x 20; 3 cypsela x 3.(No speci­ men cited in orig.) Drawn by Ruth Koppel (Reproduced from FI. Palaestina. vol. 3, PI. 581.)

No habitat data available; SU (Ambo). Benedetto 34 (FT).

85. COTULA L. (1753) 3.8-4.2 mm long, slightly thicker below the fork, green­ ish, narrow ed at base, branches truncate and pcnicellate at apex, c 1 mm long, stigmatic surface c 0.9 mm long. Cyp­ selas oblong, 3-5-angled, with myxogenic cells. 0.8-1.2 x 0.5-1.0 mm. Ornamental plant or cultivated as a low hedge; 2000-2440 m. EW SU HA; native of southern Europe, cultivated elsewhere. Amare G. (IECAMA) 206.

Annual or perennial herbs. Leaves alternate orsomcitmcs opposite or rosulate. pinnatisect. lobed or occasional!) en­ tire. Capitula solitary, pcdunculatc. generally disciform or discoid: peduncles sometimes swollen below the rcccpta­ clc. Receptacle flat to conical, epaleate. rarely with short pedicels. Phyllaries 2-3-scriate. monomorphic. Ray or marginal florets female, fertile, one- lo several-seriate, generally stalked, tube short or absent, lamina absent.

145. ASTERACEAE: 85. Cotula

227

Central florets corolla (3-)4-lobed, abaxially sometimes saccatc. lobes generally with central resin canals, one lobe occasionally expanded to a radiate lamina. Ray cypselas domventrally flattened, often laterally w inged, with 2 lat­ eral (occasionally 3 or 4) vascular strands, and sometimes with 2 lateral resin canals, generally with blunt. 1-seriate liairs sometimes with myxogenic cells. Pappus absent. Di; c cypselas oblong, without wings. A genus of about 55 species found mainly in South Af­ rica. several species occur in Australia. South America and tlie South Oceanic Islands; a few species are also found in East and North East Africa and in Mexico; 3 spe­ cies in the Flora area. 1.

Leaves opposite, petiole-base with a long sheath: receptacle epaleate: capitula over-topped by leaves. 1. C. cryptocephala Leaves alternate, petiole-base with a short sheath: receptacle with short pedicels; capitula equal to or projected well beyond leaf tips. 2

2.

Pedicles of marginal florets longer than those of the central florets; peduncles much longer than leaves 2.C. abyssinica Pedicels of marginal florets about the same as those of the central florets; peduncles not much longer than leaves. 3. C. anthemoides Figure 136. COTUL-X CRYPTOCEPHALA: 1 - habit x V a.

1. C. cryptocephala Sch. Bip. ex A. Rich (1848) type: GD. Simcn Mts., Mt. Buahit. near Enchetcab. 7 July 1838. PI. abyss. II. Schimper 1171 (P holo.. BM K UPS iso.). Perannial herb. 5-10 cm high. Stems procumbent or rhizo ttatous, often several from the caudex. glabrous. Le.i\es opposite, bipinnatipartitc. oblong or oval in outliijrjj 3-8.5 cm long including the sheathing connate petio k . segments shortly linear, with acute apex, glabrous. Capitula solitary , terminal. 6-8 x 3—4 mm at anthesis. sh inly pedunculate, overtopped by the leaves: peduncles 1-2.5 cm long, pilose-pubescent in uppcrparts. Phyllarics 2( 3)-seriate. ovate. 2.5-3.2 x 1.5-2 mm at anthesis. glabnms. margins membranous, dark brown towards the apex Marginal florets 1-seriate, female. Central florets bistaoial, yellow. Corolla tubular, glandular, c 1.1 mm lonii. 4-lobed. lobes c 0.1 nun long. Anthers c 0.2 mm long Style c 1.2 mm long, bifurcate, branches 0.2 nun lo: l Cvpselas monomorphic. flat, winged, c 1.7 x 1.1 yellowish green. Fig. 145.136.1.

COTULA ABYSSINICA: 2

old receptacle with phyllaries and but without florets x VA. COTUL/\ ANTHEM OIDES: 3 part o f stem with capitula x '/2; 4 - old re­ ceptacle with phyllaries and marginal tubercles but without flo­ rets x 3 12 . 1 from Hedberg 996; 2 and 4 from Hedberg 945; 3 specimen not given in original. [1-2, 4 drawn by M. Induss. combined and reproduced with permission from Sytnb. Bol. Upsal.. vol 15 (1957). figs 15A & B & 16. 3 drayvn by Magdi El-Goharv. reproduced from The Weed Flora o f Egypt ( i 994), p. 6 4 1. marginal

tubercles

Demerki. PI. abyss. II. Schimper 1381 (P holo., K iso.). (\ pusilla auct., non Thunb. (1800): Sch. Bip. ex Schyvcinf. & Asch. (1867).

Erect, annual or short-lived perennial herb. 0.3-30 cm high. Stems pilose, usually richly branched near the base, branches ascending, procumbent at base. Leaves alter­ nate. bipinnatipartitc. 1-3.5 cm long, shortly petiolate. segments pilose, with 1-2 nun long, erect or spreading yvhite hairs, linear. 5-7 x 1-1.5 mm. apex acute, often pur­ On moist ground in short grass, along patlis. roadside plish (in highland forms), petiole half-clasping or shortly rnjj rgins. river banks, etc.; 2800-3750 m. GD SU BA; sheathing, short, or as long as the blade of the leaves. Uganda. Keny a. Getachew A. 3005; Hillman 38; Negri 372. Capitula disciform, 4-5 x 2-3 111111 at anthesis. solitary , terminating stem and branches: peduncle sparsely pilose. 1-13 cm long. Rcccptacle flat or slightly convex, tubercuI!. abyssinica Sch. Bip. ex A. Rich. (1848) late or pedicellate, with unequal tubercles or pedicels types: TU. Adyva. PI. abyss. I. Schimper 137 and clearly seen after the fall of the fmits. marginal tubercles .875 (P syn ). Simen Mts.. Demerki. Schimper 1381 1-seriate, much longer than the inner ones. Phy llaries iP syn ); TU. Kuavcta. Mcmsah. Quartin Dillon s.n. 2-3-scriatc. ovate-oblong, obtuse with broyviiish scarious (P syn.). C. abyssinica var. nana Sch. Bip ex A. Rich.. margins, glabrous or sparsely pilose. Marginal florets fe­ Tent. FI. Abyss. 1:419 (1848) - type: DG. Simen Mts.. male. 1-seriate, yvhitish. pedicellate, tubular; cypselas

228

145. ASTERACEAE: 85. Cotula

Figure 137. COTULA A B Y SS I M C A 1 - habit x 1; 2 - leaf x c 2: 3 capitulum , longitudinal section \ 7; 4 phyllary x 10; 5 - pedicellate mar­ ginal floret x 20; 6 - central floret x 20; 7 - stamens x 60; 8 style \ 40; 9 - cypsela x 2. 1 from Bogdan A3166; 2 9 from Greenway 14995 Drawn by Helen Ireland. (Reproduced with permission from FI. Trop E. Afr., C om positae, fig. 110.)

winged, arising from the pedicels. Central florets bisexual, yellow; corolla tubular, slightly constricted in middle and in lower parts, c 1-1.2 nun long, glabrous. 4-lobed, lobes c 0.2 mm long, papillose; anthers translu­ cent. c 0.2 mm long, arms c 0.1 nun long; cypselas oblong, smooth, c 1.2 x 1.0 nun. glabrous, terete to obscurely an­ gular. Figs 145.136.2; 145.137. Afromontane and afroalpine grassland, often in damp sites, and heathland with Helichrysum. Lobelia. Agrostis. etc.. roadside margins, weed between paving stones and among annual crops; (1600-)2000-4220(-4600) m. EW TU GD GJ SU BA; Sudan. E Africa. Rep. Dem. du Congo, Saudi Arabia. Mesfin T. 4810; Miehe & Miehe 1181; O. Iledberg & Getachew A. 5380b.

3. C. anthemoide.s L. (1753) type; Habitat in Insula Helenae, Hispania.' Herb Clifford. 417, “Cotula J ” [BM lecto., selected by Humphries in Jarv is & Turland (ed ). Taxon 47(2): 3*59 (1998)1. C. dichrocephala Sch. Bip. ex A. Rich (1848) type; TU, near Adwa. PI. abyss. II. Schimper 1325 (P holo.. K iso ). Erect annual herb. 5-10 cm high. Stem simple or branched, hirsute or pubescent, branches spreading or prostrate, ranging up to 15 cm. Leaves alternate, deeply bipinnatifid. obovate in outline, hirsute, 1-2.5 cm long, half-clasping at base, segments lanceolate or ovate, apex acute-apiculate. Capitula solitary, terminal, numerous per

145. ASTERACEAE: 85. Cotula. 86. Anthemis

229

p ant. 4-5 x 2-3 nun at anthesis. shortly pedunculate, p duncles not exceeding the leaves. Receptacle flat or convex, tuberculate after the fall of the fruits, with more or It ss equal tubercles or pedicels. Phyllaries 2-seriate, ob­ it ng, with thin scarious margins, apex obtuse, c 1.7 x 1.0 n n at anthesis. Marginal florets female, several-seriate. C itral florets bisexual, yellowish;.corolla tubular, c 1 n m long, densely glandular, 4-lobed. lobes c 0.1 mm long; anthers translucent c 1 nun long, style bifurcate, c 1 mm long, branches c 0.2 mm long; cypselas mononh. rphic. narrowly winged, densely glandular, c 1.5 x 0.7 n m, oblanceolate or obovate. Fig. 145.136.3-4. Weed in agricultural fields, roadsides, grazed grass1; nd, etc.; (800-) 1480-2850 m. EE EW TU GD SU SD; widespread in tropical Africa and Asia. Pulschen 228; z rihun W. 289; Mooney 6466. The low altitudinal record was obtained from Onovenda (1904:187) who based it on the specimen. / ippi 3432. Assaorta. Majo Illalia [EE],

86. ANTHEMIS L. (1753) Perennial herbs. Stem usually dccunibent or creeping. L caves alternate, variously lobed to pinnatisect. occasiona l> entire. Capitula solitary or loosely corymbose, p . dunculate, radiate. Receptacle convex to narrowly conii al, palcate. rarely basally or totally epaleate. Phyllaries 3 4-seriate. margins scarious or dark purple. Paleae s.arious. oblong or subacutc. occasionally with a median n sin canal, apex truncate, acute or acuminate. Ray florets female, fertile, or neuter, white, rarely reddish on the 1 wer surface. Disc florets bisexual, yellow. Corolla 5 iobed. tube basally much swollen in fruit, rarely pilose. / others brown, base obtuse or shortly sagittate. Style bif rcatc. brandies truncate, penicellate. Cypselas turbinate, smooth, prismatic. 10-ribbed. sometimes tuberc ilatc. rarely with myxogenic cells. Pappus a shallow, oft n adaxially more developed corona, or an adaxial auric e. or absent. A genus of about 210 species found mostly in southern 1 rope. SW Asia and N Africa; lspccies in the Flora area. A. tigreensis J. Gay ex A. Rich. (1848); A. abyssinica J. Gay ex R.E. Fries var. tigreensis (J. Gay ex A. Rich.) Chiov. in Ann. R. 1st. Roma 8: 187 (1904) - types; TU. Cliire. November 1839. QuartinDillon (P syn.); Adua, July 1839. Quartin-Dillon s.n. (P syn ): Adwa. PI. abyss. Ill, Schimper 1848 (P K syn). Maruia cotula auct. non DC. (18.37), quoad ?Schimper 539: Schweinf. (1862) Anthemis cotula (Schweinf.) Oliv. & Hiem (1877) non L. (1753), nom. illegit. A. abyssinica J. Gay. nom. nud quoad Schimper 539; A. abyssinca Schweinf. & Asch. (1867). nom. nud ; A abyssinica J. Gay ex R.E. Fries (1928)- type; GD. Sirnien. foot of Mt. Buahit. Schimper 539 (K P lecto.).

Figure 138. A N T H E M IS T IG R E E N S IS 1 flowering branch \ 2 capitulum. longitudinal scction x c 3; 3 phyllary x 5V2; 4 ray floret x 4; 5 - disc floret x 4; 6 - stamens and style x 20; 7 stamens x 30; 8 -s ty le x 10. All from Mabberley 348. Drawn by Helen Ireland. (Reproduced with permission from El Trop. E. Afr.. C om positae. fig. 108.)

A. cotula L. var. atromarginata Vatke in Linnaea 39; 501 (1875) - type: GD. near Mt. Guna, 3700 m.. 1863. Schimper 1422 (B holo., destroyed). A. semiensis Pic.Semi. (1950) - types: GD. Simen Mts., Mt. Dcdscha, Schimper s.n. (P syn); GD. Dedsclia. 14000 ft. Schimper 220 (P syn.). Anthemis pvgmaea Schweinf & Asch. (1867). nom. nud.; Anthemis pygmaea Sch. Bip. ex Oliv. & Hiem (1877), nom. illegit.. non Bertolini (1843). Pvrethrum rarum Steud., nom. nud.. quoad Schimper s.n. (P). Anthemis aulacospermum Hochst.. nom. nud.. quoad Schimper 2069 (P). Erect, perennial herb. 5-60 cm high. Stem single or sev­ eral from the caudcx or taproot, usually decumbent, loosely branched with branches terminating in solitary capitula. Leaves alternate. 2-3-pinnatisect 2-7 cm long, sparsely to densely hispid-pilose with short crect liairs. sometimes wolly tomentose in high-altitudc forms, ob­ long or elliptic in outline, petiolate. or middle and upper

230

145. ASTERACEAE: 86 Anthcmis, 87. Argyranthcmum

ones sessile, segments narrowly linear, apex acuteapiculate. Capitula radiate, heterogamous, solitary, termi­ nal. 35-45 x 6-8 mm at anthesis, peduncle wider near the base of the receptacle, striate, sparsely to densely pubes­ cent. Phyllaries oblong, pubescent, 2-3-seriate, 4-5 x 1-1.5 mm, greenish in the middle and usually dark-grey or brown on the crisped margins. Ray florets white, neu­ ter. oblong-elliptic, 15-25 x 4-6 mm, apex minutely 2-3-fid: sometimes ray florests much reduced in size. Pa­ leae narrowly lanceolate, c 3.5 x 0.5 mm, apex dark brown, setaceous, equalling or exceeding florets. Disc flo­ rets yellow, bisexual. Corolla tubular, glandular, c 2.5 mm long, 5-lobed, lobes c 0.7 mm long. Anthers brown, c 1 mm long, connective lanceolate. Style bifurcate, c 2 mm long, branches c 0.5 mm long, truncate, penicellate. Cyp­ selas light brown, oblanceolate. curved at apex, c 1.7 x 0.7 mm, faintly longitudinally grooved, glandular, slightly keeled abaxially, convex adaxiallv. Pappus absent. Fig. 145.138. Afro montane and afroalpine grassland, commonly found with Senecio, Helichrysum, Artemisia, Ursinia, etc., in Erica arborea bushland; (1800-)2300-4620 m. EW TU GJ GD WU SU AR BA HA; E Africa. Mesfin T. 7777; Ryding 2050; O. Hedberg & G. Aweke 5363.

87. ARGYRANTHEMUM* Webb ex Sch. Bip. (1844) C. J. Humphries in Bull. Brit. Mus. Nat. Hist., Bot. 5 147-240(1976). Shrublets or shrubs. Leaves alternate, variously dissected. Capitula radiate, pedunculate, solitary or 2-50 per branch, in open corymbs, peduncle erect. Involucre hemispheric. Receptacle convex, epaleate. Phyllaries 3-4-seriate, imbri­ cate. many-veined, greenish part thickened, margins and apex scarious. Ray florets female, fertile, wliite, rarely yel­ low or pink, many-veined. Ray cypselas 3-angled, gener­ ally laterally and adaxiallv strongly 1-3-winged, some­ times coalesced into groups, wings sometimes reduced, of­ ten apically projected into a pappus-like corona; true pappus absent. Disc florets bisexual, yellow or deep pur­ ple-red. Corolla tubular-campanulate. sparsely glandular. 5-lobed. lobes rarely reddish purple. Anthers with short tails, apex lanceolate Style bifurcate, the bases swollen, apices truncate-penicellate. Disc cypselas obconic. 4-angled or laterally compressed, with 1-2 wings, or pris­ matic to terete and wingless, sometimes coalesced with the ray cypselas. Pappus coronifonn or absent. A genus with about 24 species, all endemic lo Macaronesia (Canary Islands. Madeira and Salvage Is­ lands); 2 widely cultivated species in the Flora area. 1

Leaves glaucous, crowded around the base of the peduncles, the lower caducous, primary segments 0.5-5 x 0.5 mm, secondary segments 2-15 x c 2 mm. 1. A. foeniculaccum Leaves green, evenly spread along the stem, the lower persistent, primary segments 2-30 x 1-2 mm, secondary segments 1-10 x 0.5-4 mm. 2. A. frutescens

1. A. foeniculaceum (Willd.) Webb ex Sch. Bip. (1844); Pyrethrum foeniculaceum Willd. (1809); Chry­ santhemum foeniculaceum (Willd.) Desf. (1829) type: Tenerife. Broussonet in herb. Willdenow (B-WILLDNo. 16200, holo , RNG, photo, not seen). Perennial herb, 60-100 cm high. Stem procumbent to iscending, bushy, with branches throughout, glabrous, intemodes short Leaves obovate in outline, 2-3pinnatisect, 4-10 x 1-6 cm, glabrous, glaucous, petiolate, crowded around the base of the peduncles; pnmaiy seg­ ments commonly 7-8, 0.5-5 x 0.5-1.0 nun, subopposite, acuminate; secondary segments 2-10, alternate to subopposite, 2-16 x 1.5-2 mm, apex acute. Capitula soli­ tary or 2-5 in open corymbs, peduncle up to 20 cm long, bracteate or ebracteate. Involucre 10-18 mm in diameter. Phyllaries 3-seriate, the outer ones triangular to obspathulate, scarious with a fleshy midrib; inner ones with an expanded, scarious, hyaline apex. Ray florets fer­ tile, lamina 13-23 x 3.5-5 nun, wliite. female, apex ob­ tuse to 2-3-fid Ray cypselas trigonous, arcuate, 2-3winged. with 2 w ide, lateral wings and 1 small, cunei­ form. ventral w ing, (3-)5-6 x 3-5 mm. Pappus coronifonn. dimidiate on the dorsal edges, dentate. Disc florets yellow, bisexual. Corolla campanulate in upper parts, tu­ bular below, 3-3.5 mm long, sessile-glandular on tube, 5-lobed. lobes c 0.5 mm long. Anthers light brown, c 1 mm long. Style bifurcate, c 3.5 nun long, branches c 1 nun long, truncate, penicellate. Disc cypselas obconical. later­ ally compressed to more or less quadrangular, usually with one small ventral wing, 2-3 x 1-2 mm. arcuate in outer series. Pappus coronifomi. dentate. Cultivated ornamental; c 2000 m. ?SU(probably culti­ vated in Addis Ababa) HA; native of Tenerife, but now w idely grown as an ornamental in many parts of the trop­ ics. Demel T. 345. 424. This species is easily distinguished from the next one b> its candelabra branching pattern, the deeply pinnatisect glaucous leaves, and the short intemodes which give the new leaves a crowded appearance at the base of the peduncles. Usually the lower leaves are deciduous, thus leaving the stem bases quite bare. J. Francisco-Ortcga (pers. comm., 1997). who works on the evolution of some Argvranthemum species in the Canary Islands, commented that the specimens from Ethi­ opia might also belong to the A. haouarvtheum Humphries & B ramwell group, or they might also repre­ sent liybrids between A. frutescens (L.) Sch. Bip and A. broussonetu (Pers.) C.J. Humphries. The specimens were compared with the published descriptions and illustra­ tions of the above named species (cf. Humphries 1976) but they are different in a number of respects. In both.4. haouarytheum and A. broussonetii the leaves are devel­ oped tliroughout the stem and are not crowded on short intemodes. a feature which together with the candelabra type of branching pattern distinguishes A. foeniculaceum from other related species. It is therefore recommended that specimens of cultivated Argyranthemum in the Flora area should also be compared with either authentically

145. ASTERACEAE: 87. Argyranthcmum. 88 Chrysanthemum

231

n imed species or with reference to the above cited litera­ ti ire.

cate. arms truncate, apices penicellate. Disc cypselas pris­ matic w ith a narrow wing, or terete vvitli a thick undulat­ ing wall, thus apparently ribbed Pappus absent.

2 A. frutescens (L .) Sch. Bip. (1844); Chrysanthemum frutescens L. (1753); Pyrethrum frutescens (L.) Gaertn. (1791) - type: "Habitat in Canariis insulis.” Herb. Clifford: 417, "('hrysanthemum 5” |BM lecto.. selected by Humphries in Bull. Brit. Mus. (Nat. Hist.), Bot. 5: 181 (1976)).

A genus currently believed to consist of only 2 spe­ cies Both species arc cultivated in the Flora area. Many of tlic species previously described within this genus have now been assigned to Tanacetum.

S ibshrubs. 40-60 cm liigh, glabrous, or hispidulous only o i the leaf midribs. Stem procumbent to erect, branching tl iroughout or only at the base. Leaves pinnatisect to b pinnatisect, glabrous or sparsely hispidulous on the r .idvcins. coriaceous, sub-succulent, 2-8 cm long, prir an' segments 2-8, opposite. 2-30 x 0.5-2 mm. lanceoI; tc to linear-lanccolate; secondary segments 2-6. suba:>positc to alternate. 1-10 x c 1 mm. linear-lanceolate; l mm. Ray florets c 16. female, fertile. 8-15 x 2-5 mm. \ lute, apex minutely 1—3-fid. Ray cy pselas 3-angled. 3-5 > 2-4.5 mm. arcuate. 3-winged, the 2 lateral wings e.x\ tnded. coriaceous, with convex dorsal surface; the vent il wing oblanceolate; pappus coroniform, margins 1 iciniatc. Disc florets bisexual, yellow. Corolla tubular. ‘ -3.5 mm long. 5-lobcd. lobes c 0.5 mm long. Anthers 1 ^ht brown, c 1.2 mm long. Style bifurcate, c 3.5 mm 1 ng. branches c 0.8 mm long, truncate, peniccllatc. Disc c vpselas obconic. terete to laterally compressed, cne-winged, irregularly ribbed, 2.5-4 x 1-2 mm. Pappus coroniform, margins laciniatc. Ornamental; rarely found as an escape in "irrigated, cm. partly bare ruderal ground”; 2350 m. EW ?SU; ' idely cultivated ornamental plant in many parts of the \ orld. Ryding 1234.

88 CHRYSANTHEMUM* M l 753) I. Bremer & C .1 I Iumphries, in Bull. Nat. Hist. Mas. Loud. o tj. 23(.1) 135 (1993). nnual herbs. Stems erect, leafy, branched. Leaves alteri :iic. toothed, deeply serrate-dentate and pinnatifid to f ectinatc. sometimes amplexicaul. Capitula radiate, i ^terogamous. solitary at branch apices, or in open (Irom left to right). 4 from Knox 6X1. 5 from Mesfin 721 (left) and Hoitzsch 7 (right). 6 from Eriis ct a| M I2 and ffedberg 5534b (from left to right). 7 Irom Mesjbi -10 nun long, slender, densely bracteatc. glabrous to sp irscly ciliatc. Calyculus (2-)5-6. glabrous, lanceolate. 2 2 5(—4) xO. 1-0.8 mm. purplish at apex. Involucrc 4-5 x 2 A 3.5 mm at anthesis. up to 8 x 4 mm in fruit. Pin llaries (13 ) 13(—14). light-green with pale purple tips. (5—)6—7 ( 8) x (0 5-)0.6-1.8 nun. glabrous. Rav florets (5-) 7- lit-9). female, rarely neuter, bright yellow, tube glabr..vis. lamina (4.3—)7—10(—13.5) x (l-)2.5-4 nun. apex 2 ;-fid. obtuse or rarely acute. Cypselas light brown, cc ;tatc. appressed sctulose in grooves. 1.7-3 x 0.2-0.7 m n. cylindric. Pappus bristles (3—)5—5.6 nun long. Fig. 145.143.

Figure. 143 SENECIO M YRIO CEPIIALU S: t upper pari o f stem; 2 disc floret, longitudinal section \ 15. (No specimens citcd in orig.) Drawn hv W. Burger. (Reproduced with permis­ sion from Families o f Flowering Plants in Ethiopia 1967. tig 6 8 :1 p.p.)

Senecio sp. sensu Cufod. (1967). quoad Mooney 8299.

Ercct. perennial herb. 50—75(—120) cm high. Stem simple or densely branched, woody at base, floccose-tomentosc or woolly on upper intemodes. Leaves simple, narrow 1\ linear-lanceolate or elliptic, often bright-green above and densely wliitc-tomcntosc beneath sometimes upper sur­ face arachnoid and low er surface woolh or both surfaces glabrous (especially in forms growing along river or stream banks). (1.7—)3—8(—12) cm long. (2—)3—1()(—20) nun wide; margins closely serrulate. Capitula radiate, campanulatc. few to numerous in usually dense cors mbose or paniculate cymes. 18-40 x 7-13 mm at anthesis. Peduncle up to 3 cm long, floccosc-lomcntosc lo wooll\ Calyculus (5—)8—11. dark purplish-brown in ilie upper Margins of Ilagenia ScheffJera forest. Podocarpus half. 3—8.1 x 0.4—1(—1.3) mm. arachnoid w ith intcr-mixcd forest, usually in forest clcanngs. and in Arundinana purplish-black hairs. Involucrc (4-)6-8 x 4-6 mm at bamboo thickets. Erica arborea bushland. sometimes anthesis. up to II x 6 111111 in fruit. Phyllaries (13-) al ;o used as a hedge plant at high altitudes |c.g.. Dinsliu (E A)]: 2250-3300(-3900) m. TU/GD GD WU SU AR 21 (—22). 6-1 I x 0.5—1(—2.2) mm. dark green, purplishk SD BA HA. not know n elsewhere. Thulin et al. 3691; brown at apex, arachnoid with intermixed purplish-black Mesfin 7 . 723; Friis et al. 1507. liairs. Ra> florets (7—) 13(—20). lemon-yellow. female, tube glabrous or sparsely pilose, lamina 10.5—22.5(—32) x 2 .2 - 5 .6111111. apex 2-3-fid. rarch obtuse. Cypselas brow 11. 2 S. frescnii Sch. Bip. ex Oliv. & I Hern (1877) type: TU. between Halci and Tcmbicn. Riippell glabrous, costalc. cylindrical. 2.2-3 \ 0.4-0.8 111111. Pappus bristles 4-8.3 111111 long. Fig. 145.142.2. s.n. (FR holo.).

238

145. ASTERACEAE: 90. Scnecio

Juniperus forest at high altitudes. Erica arborea bush­ land, well drained grassy slopes in afro-alpine meadow; 1520-4300 in GD GJ SU AR BA; not known elsewhere Mesfin T. 5540; W. de Wilde 9147; Mooney 8299. 3. S. stcudelii Sch. Bip. ex.4. Rich. (1848) - type: GD, Simen Mts.. Mt. Buahit. PI. abyss II. Schimper 750 (P syn.. BM K isosyn ), Quartin-Dillon s.n. (P syn.. K isosy n ). S. steudelii forma latifolia Vatke in Linnaea 39 506 (1875) - type: GD. Siincn Mts.. Dsclian-Meda. 1863. Schimper 1523 (?B holo . BM K iso ). S. steudelii forma angustifolia Vatke in Linnaea 39: 506 (1875) - type: GD, Simen Mts . Mt. Bachit. above Demcrki. Schimper 1525 (?B syn.. K isosy n ). 1853. Schimper 31 1 (7B syn., K isosyn.). S steudelii Sch. Bip cx A. Rich, forma angusti­ folia canescens Vatke (1875). nom non rite publ Erect, semi-aquatic, perennial herb. 25-150 cm high Stem usually unbranchcd except in tlie inflorescence, leafy in upper parts, floccose-tomentose. Leaves simple, linear-lanceolate to oblanceolate. sessile, with auriculate base; middle leaves 12-20 3 \ 1.5-2.1 cm; upper leaves 3-6 x 0.3-1.5 cm. cordatc-auriculate. thinly floccose to glabrescent. margins denticulate, sometimes subcntirc Capitula radiate, campanulate. (1.9—)3—3.5 x 0.5-13 mm at anthesis. in corymbose cynics. Peduncle 0.6-10 cm long, sparsely pubescent to densely anichnoid-tomcntose. bracteate. Caly culus (2—)5— 6(—9). arachnoid at base, gla­ brous and purplish in upper parts. (3-)3.8-8.2 x (0.3— ) 0.6—I(—I 5) mm. linear-lanceolate, apex ciliolate. some­ times bristly or occasionally scarious and minutely lacer­ ate. margins ciliolate. sometimes denticulate. Involucre 6-12 x 5-6 nun at anthesis. 8-10 x 7-8 mm in fruit Phyllaries (13—)2(>—22. linear-lanceolate. 5.8-9 x 1-2 mm. arachnoid with intermixed purplish black liairs. apex bristly, purplish-black Ray florets (1 0 -)13. female. \cllou. tube glabrous or pilose, lamina 9.2-17(—26) x I 8-4.2 mm. apex 2-4-fid. often w ith deep clefts, rarely acute, obtuse or cmargmatc Cypselas brow n, costatc. ob­ long. glabrous, sometimes scabnd. terete. 3.3-4 x 0 5-0 9 mm. ovate-lanceolate to oblong. Pappus bnstles 3 8-5—15 x 2-10 cm. glab 'us and shiny or puberulous on the midrib beneath, p relate; petiole up to 4 cm long, blade cuneate at base a- d acute at apex, margins regularly dentate. Capitula o? ong-cylindric. 6-9 x 2-3 mm at anthesis. on up to 1 cm 1« ug. slender, puberulous pedicels in dense panicled c nes, terminal. Calyculus filiform. 3-4. Involucre 7-8 x 2 -2.5 nun. Phyllaries 5-8. linear-oblong, glabrous, brown n fruit. Florets 6-12. yellow. Cypselas light brown, mi­ tt itely hairy, longitudinally ribbed, columnar. 2—2.5 x 0.5

Figure 149 SO L/INECIO GIGAS 1 - portion o f leaf and in­ florescence x 12 . 2 - disc floret x 6. (No specimens cited in orig.) Drawn by W. Burger. (Reproduced with permission from Fam­ ilies o f Flowering Plants in Ethiopia , 1967, PI. 68:2. as Senecio

gigas).

nun. Pappus white, 7-8 nun long, barbellate. Fig. 145.150. Montane evergreen forest with Aningeria-Syzygium and Aningeria-Olea, especially along margins and in clearings, derived evergreen bushland; 1300-2450 m. SU AR WG IL KF SD BA; Sudan. E Africa. Rep. Dem. du Congo, Cameroun, Bioko, Nigeria, Rwanda. Burundi, Mozambique. Malawi. Zambia. Zimbabwe. Angola. Friis et al. 2174; Mesfin T. 5028; Burger 2551. 3. S. angulatus (Vahl) C. Jeffrey (1986); Cacalia sonchifolia auct., non L. (1753): ForsskAl (1775); Cacalia angulata Vahl (1794) - types: Ye­ men, Hadie, Forsskal in Herb. Forssk. 1026 (C syn., not seen). 1027 (C svn.. not seen), 1028 (C syn.. not seen) & 1029 (C syn., not seen). Senecio subscandens Hochst. ex A. Rich. (1848); Crassocephalum subscandens (Hochst. ex A. Rich.) S. Moore (1912) - types: SU. Quartin-Dillon & Petit

250

145 ASTERACEAE 91 Solanecio

5-7. oblong-linear, green at anthesis. brown in fruit, gla­ brous except for tufts of liairs at margins near the apex. Florets white with orange-yellow stigmas. 10-12. Cyp­ selas black, cylindnc. ribbed with white hairs in grooves. 2-2.5 x 0.7-1.0 nun Pappus white, 7-7.5 mm long, barbellate. Margins of Juniperus forest, moist scrub in grassland with streams and clumps of Acacia, Ficus, etc.. ditches, margins of cultivations, rarelv cultivated along fences. 1250-2500 in. EW TU WU SU KF GG SD HA; wide­ spread in E and C Afnca to South Africa, westwards to Cameroun Madagascar; Yemen. Mesfin T. & Kagnew 2045; Ash 1287; Mesfin T. 7266. 4 S. nan dens is ( \ Moore) ( \ Jeffrey (1986); Senecio nandensis S. Moore (1902) - type: Kenya. Nandi. Scott Flliot 6987 (BM holo.. K iso ). Senecio caranianus Chiov (1912) - type: SU. Lago Zuai. Negri 812 and 1420 (FT sy n). Subscandcnt. subsucculent, richly branched herbaceous climber, up to 4 m long, scrambling over bushes and over boulders Stem often rooting al lower nodes. Leaves pinnately lobed often from base to middle with upper parts sliarply and shallowly lobulate-dcntate. rarely some ly rately -pinnatifid 5-17 x 2.5-5 5 cm. pale green above floccosc-tomcntosc cspccially on low er surfacc. petiolate petiole narrowly winged, up to 3 cm long. Capitula ob­ long. 5-8 x 2-3 mm at antlicsis. on up to 7 nun long, slen­ der. bractcate pcdiccls in dense paniculatc cynics Calyculus filiform. 3-5. Involucre 5-7 x 2—2.5 mm Phy llancs 6-8. light green glabrous or thinly pubcmlous brown in fruit. Florets 15-20. pale yellowish white. Cyp­ selas brown, columnar, costatc. hairy in grooves. 2-2.5 \ o 5 mm Pappus wlute. 5-6 mm long, barbellate figure 150. S O L l.X IX '/O M A N M I 1 part ofplant: 2 invo­ lucre: 3 lloret. 4 anthers. 5 style arms Drawn by W I l'revithick No magnifications given and no specimen cited m original publication. (Rcproduccd with permission from /•’/ II Trap Afr.. fig. 252. as ( 'rassocepluilmn mannii )

Hillsidc-scmb with Croton. Dodonaea. Rumex and Fterolohium. rock-crcviccs in bushland. along fences 1550-2500 m SU AR KF SD HA (perhaps introduced in KF); Kenya. Tanzania. Rep Dem du Congo. Mesfin 7 & Kagnew 1560; GiUett 14888; If. de Wilde 8133

s.n (P syn.); Ethiopia, without futhcr locality. PI abyss. III. Schimper 1926 (P syn.. not seen) Senecio hojeri DC (1838): Crassocephalum hojen (DC ) Rob> ns (1947)

5 S. tuhcrosus (.Sc li Hip. e x . I Rich.) ( Jeffrey (1986): Senecio tuherosus Sch Bip cx A Rich. (1848) types TU/GD. Djcladjcrannc (in theTacazzc Valley | Schimper 1610 (P sy n . not seen; K isosy n ). OuarnnDiUon s.n. (P syn. not seen); GD. Simen Mts Schoata. Schimper 1361 (P sy n . K isosy n ) Cacaha tuherosa Del. (1848). non Nutt. (1818). nom illegit. I Senecio solanoides Sell Bip cx Asch (1867) type Ethiopia, without further loc . Schimper 1361 p.p. (B holo ) C ahyssinuh Walp (1852). non R Br in Salt (1814). nom nud

Climbing, creeping or straggling perennial herb, up to 3 m tall Stems bright yellow or green, finch stnate-sulcate. glabrous, slightly succulent. Leaves succulent, obovate in outline, up to 20 \ 10 cm. pinnatipartite with (3—)5—11 suboppositc to nearly alternate segments, aunculate at base, glabrous, shiny; petiole narrow ly w ingcd. up to 4 cm long: lateral segments 1-5 x 0.8-3 cm. margins grossly dentate, oblong-ovate or obovate; terminal segments of­ ten the largest, deltoid, rcpand or toothed. 1-5 \ 3-8 cm Capitula narrow ly campanulate. in round dense cy mes. 6-8 x 3-4 mm. pcdunculatc; pcdunclc short, slender bractcate. glabrous to sparsely ciliatc. Calyculus 2-3. lan­ ceolate. lino lucre 6-8 x 2-4 nun al anthesis Phyllaries

Ercct. perennial herb 3 0 -6 0 cm high, with irregular, flat­ tened. globose tuber up to 7 cm long or wide, often with sex cral separate shoots from near the center Stem y ellow ish green in upper parts, deep purple at base, glabrous, sol­ itary . or several from the apex of the tuber, leafy al the

251

145. ASTERACEAE: 91. Solanecio

Figure 151.

SOI^ANECIO H A R E N N E N SIS 1 habit x Vi; 2 - lower portion o f stem with roots x '/2; 3 capitulum. after anthesis x 23/4; 4 - calyculus with m ar­ ginal hairs x 9; 5 - phyllary x 9 : 6 - flo­ ret x 5'/3; 7 - floret without pappus and ovary x 5'/3. 8 - apical part o f stamen (1 o f 5 shown in detail) x 16. 9 upper part o f style with bifid branches x 24; 10 style branches, much enlarged to show pcnicillate apex and linear stig­ matic surface. All from Friis, Gilbert & Vollesen 3546. Drawn by Fat Hallidav. (Reprodced with permission from Kew Bull. 49. 1994, p 138. fig. 1.)

t se. Tuber brown on outside, whitish inside, with a pep1 ry smell. Basal leaves elliptic, ovate-elliptic or spathu-

var. tuberosus

je, 9-30 x 4-14 cm, fleshy, glabrous, slightly glossy, i ight or greyish green, petiole with decurrent leaf base. ~6 cm long. Upper leaves narrowly elliptic, margins arsely to sinuately toothed or lobed, shortly petiolate or :>sile. Capitula oblong, 6-8 x 2-3 nun at anthesis, on up 5 mm long, slender, pilose pedicels, in dense submbelliform cymes. Calyculus 2-3, filiform. Involucre -7 x 1.5-2.5 mm, phyllaries 5-6, linear-oblong, gla­ brous, brown in fruit. Florets 10-20. bright yellow or oruige. Cypselas light browa costate, oblong-elliptic, 4-4.5 \ 1—1.5 mm, densely strigose, liairs silky-white. Pappus • lite, 4-5 mm long, barbellate.

Moist montane grassland with black clay soil, steep slopes with outcrops of marble and evergieen bushland domi­ nated by Euclea schimperi. Rhus natalensis. etc.. open Combretum-Terminalia woodland; 1600-2500 in. EW TU GD GJ SU WG KF; Sudan. Uganda. Kenva. Gilbert 2015; W. de Wilde 6698; Gilbert et al. 8042.

Leaves glabrous or glabrescent; florets bright yel­ low. var. tuberosus Leaves densely liairy; florets bright orange. var. pubescens

Open short grassland with volcanic boulders, deciduous woodland with grass, steep mountain slopes; 1600-2300 m. SU; not known elsewhere. Gilbert et al. 8043; Ash 2968; Hildebrandt 110.

var. pubescens Mesfin in Kew Bull. 49: 140 (1994); Senecio tuberosus Sch. Bip. ex A. Rich. var. pubescens (Mesfin) Rowley in Bradleya 14: 83 (1996) -ty p e; SU. Woliso, Ash 1948 (K holo.).

252

145. ASTERACEAE: 91. Solanecio, 92. Cineraria

6. S. harennensis A/es/m (1994) - type: BA, 31 km on Dello Menna - Goba road. Friis, Gilbert & Vollesen 3546 (ETH holo., C K iso ). Epiphyte with trailing branches, up to 30 cm long. Stem rooting in lower parts. Leaves simple, petiolate, 4-8 x 1.5—3.5 cm, glabrous, purplish beneath, membranous, margins grossly serrate-denate. petiole 1.5-4.5 cm long, slender. Capitula oblong-campanulate. (8-) 10-11 x 2-4 mm at anthesis, up to 12.5 x 5 mm in fruit, in open subumbellate cymes. Peduncle up to 2.5 cm long, glabrous. Calyculus filiform. 5-6, continuous with bracts on peduncle. 1-2.5 x 0.5 mm. ciliate on margins, dark purple. Phyllaries 1-seriate. 8-12, glabrous except for the pur­ plish cilia at apices, 10-11 x 1 mm, linear-oblong, puiple at base, light green in the remaining parts. Florets 10-18. c 10.5—11.5 mm long. Corolla white, c 8-8.2 x 1 nun. gla­ brous. 5-lobed. lobes papillose, c 0.7-1 mm long. Anthers 0.7.1.2 mm long, base obtuse, connective 0.3-0.6 mm long, collar 0.5-0.6 mm long, broader near joint with fila­ ment. Style c 9 mm long, exserted at anthesis. swollen at base and slightly widened below the branches, bifid at apex, stigmatic surfaces along margins, not confluent at apex. Ovary glabrous, light to dark brown, cylindric. Cyp­ selas (only immature ones seen) dark brown, glabrous, costate, c 2 x 0.5 mm. Pappus of 1-seriate. wliite. 7 8-8 nun long, uniformly barbellate setae. Fig. 145.151. Middle to high level epiphyte on Svzygium in moist mixed forest with ScheJJlera, Lepidotrichilla, Bersamo and Ocotea; 1850 m. BA; not known elsewhere. Friis et al. 3546. 92. CINERARIA L. (1753) Annual or perennial herbs. Leaves alternate, simple to pinnatisectly lobed, petiolate and auriculate. palinatcly veined. Capitula radiate, rarely discoid, in loose corymbs. Involucre calyculate, campanulate, rarely cy lindric. Re­ ceptacle epaleate. Calyculus present. Phyllaries 1-senate, connate. Florets yellow'. Corolla tubular-campanulate. 5-lobed. Anthers ecalcarate. Style bifurcate, branches truncate to obtuse. Cypselas obovate. laterally com­ pressed with thickened or winged margins. Pappus of many slender bnstles or sometimes deciduous. A genus of about 30 species confined to Africa and Madagascar. 3 in the Flora area. 1.

Ray florets 6-16 x 3-5 mm. always present; cyp­ selas 2.5-3.7 x 1.0-1.5 mm. 2

Ray florets 3-5 x 1-1.3 mm. sometimes absent: cypselas 1.8-2.2 x 0.8-1.9 nun. 2. C. sehaldii 2. Ray cypselas 3—3.7 nun long; involucre 8-10 x 6-8 mm at anthesis; phyllaries 7-9 mm long in fruit. 3. C. deltoidea - Ray cypselas 2.5-2.8 mm long, involucre 6-7 x 5-6 nun at anthesis; phyllaries 4.5-6 mm long in fruit. I . C. abyssinica l.C. abyssinica Sch. Bip. ex A Rich. (1848) - type: TU. Mt. Scholoda. PI. abyss. 1. Schimper 335 (P syn ); SU. Ouodgerate. Petit s.n. (P syn.).

C. abyssinica Sch. Bip. ex A. Rich. var. minor Sch. Bip. ex Engl (1892). nom. nud. Erect or scrambling, annual or perennial herb. 20-100 cm high. Stem repeatedly branched with straggly branches, slightly woody at base, terete, green, red-violet or purple tinged, striate-sulcate. glabrous or glabrescent. Leaves al­ ternate. simple to lyrately pinnatifid, petiolate. or the up­ per ones sessile and amplexicaul. glabrous to covered with loose floccosc tomentum especially on petioles and veins of developing leaves, thin, light green. 1.5-7 x 1.5-7 cm, margins serrate to regularly lobulate with Geramum-type dissection, base of simple leaves tnincate to cordate, petiole glabrous or loosely floccose. 1—9.5 cm long, auriculate. Capitula radiate, campanulate. 6-7 x 6-8 mm at anthesis. in dense corymbose cymes. Peduncle 5-30 mm long, slender, bracteate. Calyculus 6-9. lineartriangular with attenuate, black-tipped apex, glabrous or rarely floccose at base. 2-3 x 0.54).8 mm. Phyllanes 12-14. oblong, with attenuate and purplish or blackish apex, glabrous. >-6 x 1—1.5 nun. margins scarious. Ray florets bnght yellow. 10-13. female, fertile, tube slender. c 2-3 mm long, lamina oblong-elliptic. 5-6 x 2 8-3.2 mm. 4-stnate. apex shallowly 3-fid. Disc florets y ellow, inner­ most functionally female. Corolla tubular-campanulate. 5-6 nun long, glabrous. 5-lobcd. lobes 0.6-0.8 mm long, shortly papillose Anthers yellow, c 1.5 mm long, ecalcarate, connective c 0.3 mm long. Style bifurcate. 4.8-5.4 nun long, branches 0.6-0.8 mm long, apex trun­ cate. papillose, stigmatic surface cO.5-0.6 mm long. Cyp­ selas black, glabrous, obovatc-oblanceolate. narrowly yellow-margined becoming grey or black at maturity. 2-2.2 x 0.8-1 nun. convex on outer surfacc. flat on inner surface, slightly constnctcd at apex fonning a cup of bnstles Pappus w hite, barbellate. 3-4 nun long. 1-senate. Schefflera Hagenia forest, open montane forests, of­ ten in shelter of boulders, evergreen woodland w ith Aca­ cia, Dombeva. etc.. shon grassland, river banks, margins of cultivations; 2100-3900 in. TU ?GD WU SU AR SD BA; not known elsewhere. Mesfin T. 8482; Ash 1667; Mooney 8594. | 2 C. sebaldii Cujbd. (1968) - tvpe: GD. Simen Mts.. "Buahit-Pass”. 4180 m. Sebald 1108 (STU holo ). Erect annual licit). 10-60 cm luglu lower pan of stem red­ dish-brown. unbranched or little branched, sometimes branches from near the caudex Leaves in lower pans of the stem simple, petiolate. blade orbicular to deltoidtnangular. 1-3 \ 1-5 cm. glabrous on upper surfacc. sparsely to densely appressed pubescent below, margins doubly serrate (Geranium-type), petiole up to 6 cm long, often with a few. small, lateral lobes towards the base of the blade, loosely floccose. Upper leaves pinnahlobed. lyrately pinnatipartite to pinnatisect. sessile to subsessile. ovate-lanccolatc in outline, upper surfacc light green, gls brous or glabrescent. lower surface loosely to densely floccose-tomcntosc Involucre cylindric-campanulatc. 4-5 \ 4 - 5 mm at anthesis. Capitula radiate or occasionally eradiate, in corymbose cymes. 10-12 x 4-5 mm at

145. ASTERACEAE: 92. Cineraria

253

anthesis. pedunculate; peduncle glabrous or loosely floe .;ose. 0.5-3 cm long, slender, bracteate. Calyculus lin­ ear 5-7. c 1-1.3 mm long. Phyllaries 1-seriate, 12-14. ligl t green, linear, with attenuate purplish apex, glabrous. 4-4 5 x 0.5-0.8 mm. Ray florets yellow. 8 (sometimes ab­ sent) female, pistil c half as long as the rays, tube c 2.5 inni long, glabrous, rays 2.5-3 x 1-1.3 mm. often 4-stnate, 3-fid at apex. Disc florets lobed; lobes 0.5 mm lonj; truncate, penicellate. Anthers c 1 nun long, ccakarate. connective c 0.3 nun long, collar c 0.3 nun lon§; Cypselas black, oblanceolate. glabrous. 1.8-2.2 x 0.8- mm, convex on outside, fiat on inner surface. Pappus of white, numerous, c 4 mm long bristles. Evergreen bushland with Rumex, etc.. Eucalyptus plantations, montane grassland. Erica arborea bushland; 23 ' -4200 m. GD WU SU BA; not known elsewhere. Fn ' et al. 5526; Mesfm T. & Kagnew 1886; Mesfin T. & Berhanu 8095. his species can be distinguished from the other two sp< c es by its smaller ray florets and the variously dis­ sected and loosely to densely floccose upper leaves. A ra\ -less form is sometimes also encountered in popula­ tion.- inhabiting drier and rocky areas. The plants from the up >tr altitudes have often been confused with Senecio far,' aceus from which it can easily be distinguished by characteristics of its ray florets and cypselas. Figure 152. CINERARIA DELTOIDEA 1 - part o f flowering branch x '/2; 2 - capitulum x 1sh\ 3 - ray floret, half pappus cut 3. ( deltoideaSond. (1850) away x 3 ,/i; 4 disc floret, half pappus cut away x 3'/2; 5 - part o f type: South Africa. Natal. Gueinzius 343 (MEL stamen x I 1; 6 - style branches x 1 1 ,7 - cypsela x 11 (No speci­ mens cited in orig.) Drawn by Blythe Pascoe. (Modified and re­ holo.. W iso.). produced from Compositae in Natal, 1977, fig. 14 p.p.)

Cineraria gran diflora Vatke (1875) - type: GD, Dschan-Meda [near Debra Tabor). 2850 m. 1863, Schimper 1517 (B holo.. BM K iso ). C. gracilis O. Hoffm. (1906) - type; SD. Gallaiocliland (Jem Jem in Sidamo). Ellenbeck s.n. (B holo. destroyed). ?C. abyssinica Sch. Bip. ex A. Rich, forma longiradiata Oliv. & Hiem (1877), quad Schimper s.n. C. abyssinica forma rothii Oliv. & Hiem (1877) type: ?SU, Roth s.n. (K holo.).

PerL nnial lieib with woody, ascending or trailing rhizome, up lo 1 m lugh. Leaves simple to lvrato-pinnately lobed. gkibrous to sparsely pubescent on veins especially on 10 /er surfaces, petiolate; blade orbicular, deltoid to o\ 5: c-lanceolate. 2-7 x 2-8 cm, margins doubly serrate; pen ile up to 9 cm long, often with tiny leaf lobes near the bast of blade, auriculate at base. Involucre campanulate. 8 -1 ) x 6-8 nun at anthesis. Capitula radiate, terminal, cor mbose. 30-35 x 6-8 mm at anthesis, pedunculate, pc i incle up to 5 cm long, bracteate, sparsely pilose. C, cuius 7-12, linear, glabrous or ciliate at margins, rarely floccose at base. 2.5-3.5 x 0.5-0.8 nun. Phyllaries 11 15, linear-oblong with attenuate and purplish or bl i^kish apex, glabrous. 7-8 x 1.2-1.5 nun, up to 9 mm long in fruit, margins scarious. Ray florets yellow. 8-13. fc nale, fertile, tube slender. 2-4 nun long, rays oblong. 8- i ) x 3-4.5 nun. 4-6-striatcd. apex shallowly 3-fid. Disc florets yellow. Corolla tubular. 5-7 111111 long, glabrous. 5-lobcd: lobes 0.5-0.8 111111 long, shortly papillose. An-

thcrs yellow. 1.5-1.9 mm long, ecalcarate, connective c 0.5 mm long. Style bifurcate, 5-6 mm long, stigmatic sur­ face 0.6-0.8 mm long. Cypselas black, glabrous, obovate-oblanceolate, narrowly yellow margined, 3-3.7 x 1.1—1.5 mm, convex on outside, flat on inner surface, slightly constricted at apex forming a cup. Pappus of nu­ merous, white, 4-5 mm long bristles. Fig. 145.152. In sliade in Erica arborea scrub and overhanging rock on mountain slopes, in protected sites between rocks; afroalpine meadow with Helichrysum, Senecio and Erica bushes, etc.; (2800-)3200-4150 m. ?GD GJ SU BA; Su­ dan. E Africa. Rep. Dem. du Congo, Rwanda, Malawi, Zambia, Mozambique. Zimbabwe, South Africa. Mooney 8328; Mesfin T. et al. 7202, Evans & Hiller 94. Cineraria deltoidea can be distinguished from C. sebaldii and C. abyssinica by a number of combined capitular features as provided in the key. i.e., larger capitula. phyllaries. cypselas. etc. The leaves, which sometimes have been employed to differentiate it from particularly ( \ abyssinica arc variable, displaying llie same type and amount of variation as in the latter species. Populations with lyrato-pinnately lobed leaves, e.g.. Ed­ wards 3584 and Flenly & Leaky 612. and others with sim­ ple cordate rather than deltoid leaves, e.g.. Mooney 5191, Miehe 1781. arc encountered quite frequently.

254

145. ASTERACEAE: 93 Emilia

93. EMILIA (C a ss .) C a ss . (1 8 2 5 )

6.

C. Jeffrey in Kew Bull. 41: 908-920 (1986) & Kew Bull. 52 205-212 (1997). Annual or perennial herbs, stems and leaves mostly gla­ brous Leaves alternate, simple to variously lobed. sessilc-amplexicaul and auriculate to petiolate. green or often also purple tinged. Capitula usually cy mose. discoid or sometimes radiate. Involucre ecalyculate. cylindric to narrowly campanulate. Receptacle epaleate. Calyculus absent Phyllaries 1 -seriate. connate. Florets pink, purple, lavender, violet, orange, red, yellow, or rarely white. Co­ rolla tubular-campanulate. 5-lobed. Anthers ecalcarate. Style branches truncate to obtuse, sometimes with ap­ pendages of fused papillae. Cy pselas oblong, mostly 5-ribbed. glabrous or pubescent. Pappus of numerous slender bnstles. A genus of about 100 species found in tropical Afnca and Asia, similar to Senecio and Cineraria. and distin­ guished by the absence of caly cuius and the 5-ribbed cvpselas of both the ray and disc florets. Some species are widely distributed as weeds |E sonchifolia (L.) DC ex Wight] or as a garden escape \E. coccinea (Sims) G. Don] in many tropical countries. E. sonchifolia (rig. 145.153) is included in the key to enable its identification, should it occur in the Flora area. About 12 species in the Flora area. Cufodontis (Bull. Jard Bot. Etat Brux. 36. Suppl.. 1154. 1967) cited Emilia emiloides (Sch Bip.) C. Jeffrev. as Senecio emiloides Sch. Bip.. from GG. Gardula pla­ teau. quoating Engler (1906). The cited species is so far known only from Sudan (cf. Jeffrey. 1997). and there is no specimen from the Flora area tliat matches the desenption of this species. 1 Capitula radiate: florets bnghi yellow. Capitula discifonn or discoid: florets pink, violet. lavender, purple, wlutc. pale y ellow or orange 4 2

Involucre narrow. 1-2 mm wide at anthesis; rays 5-8. 1-2.2 mm long. 1. E. abyssinica Involucre 2-5 mm wide at anthesis; rays 8-13. 3 -9 111111 long 3

3. Leaves oblong-elliptic: peduncle bracteate throughout: phy llanes w itli 3 broad, purple stnae

2 Leaves orbicular, spathulate to obovatc-spathulatc. peduncle cbractcatc throughout: phyllaries with 3-8 narrow, greenish striae. 3. E. discifolia 4

Plant of swampy grassland: leaves linear-lanceo­ late, sessile. 5-11 x().3-().8 cm. margins rcvolutc. corolla pale yellow: capitula c 10 .x 3 mm in fnnt: achenes glabrous. 3-3.5 111111 long M E. leptocephala Plants not with the above combined features

5. Phy llaries 6 - 1 2 111111 long 111 fnnt. achcncs 2.5-4 111m long. 4 . E. cacspitosa Phyllaries 3-8 mm long in fruit: achcncs 1 .2-3 mm long. 6

-

Florets bnght yellow, orange or orange-red; style apex with a central group of papillae or with a subulate appendage of fused papillae; achenes (2-)2.6-4.7 mm long. 5. E. jeffreyana Florets pale to bright violet, purple or pink, mauve. scarlet, rarely white; style truncate with unifonn papillae, or a few long ones in the middle or at margins achenes 1-2.5 nun long. 7

7.

Leaves linear-lanceolate. 2-5 mm wide, sessile. margins rcvolute; achenes 2-2.5 111111 long; short­ lived perennial with leaves crowded at the base of young branches. 6. E. adama-gibaensis Leaves round, ovate, elliptic or linear-lanceolate. 8-35 111m wide, sessile to pseudo-petiolate. mar­ gins flat, achenes 1-2 mm long, plants annual, or. if perennial, leaves not crowded at the base. 8

8.

Involucre 4.5-5.5 x 2.5-3 mm; phyllaries 4-5.5 111111 long, glabrous; blade of most or basal leaves rounded or orbicular, often purplish 011 lower sur­ face. 7. E. herbacea Involucre (4—)5—12 x (2.5-)3-4.5 mm; phyilancs (4—)5—7.5 nun long, pubescent or glabrous; basal leaf blade deltoid, ovate, tnangular or elliptic. 9

9

Leaves elliptic or the upper ones lanceolate, sessile or narrowed at base into a slender petiole; capitula on usually long peduncles. 8 E. neghellensis Leaves ovate to triangular, pseudo-petiolate with broad wings, wings broadest at base, upper ones often clasping the stem; capitula on usually short peduncles 10

-

10. Involucre (7.5-)8.5-12 111111 long, up to 12mnilong in fruit, achcncs 3-3.7 111111 long. E sonchifoha - Involucre 4 5-7.5 inm long: achcncs 1.0-2.5 111111 long. ]1 11 Plant annual, stem lierbaceous. 1-4 111111 in diam . erect to decumbent: leaves mostly basal, persis­ tent or deciduous, tnangular-ovatc or deltoidovate. 12 - Plant perennial, stem wiry in lower parts. 2.5 -6 111111 in d iam . trailing or creeping: lower leaves deciduous or. if persistent, blade deltoid-ovate with irregularly serrate-dentate margins. 9 E. scrpcntinus 13. Phy llanes with scattered, stout liairs especially on the E. upper 4-5 mm wide; corolla somalialf; lens involucre i.s (4 7-)6-9 nun long. 10. E. arvensis Phyilancs glabrous or with dense tiny, cilia-like hairs: involucre 2-3 5 111111 wide; corolla 4-5 5 111111 long 14 14 Capitulat^"7 5 x 3-3.5 111111: phyilancs 12. densely luury 12. Emilia s\}.=Sebsebe I). 3327,^0350 - Capitula 4-4 5(—5) x 2.5—3 111111; phyllaries (6-) 8(—9). glabrous to sparsely hairy 7 E. herbacea

1 E. abyssinica (Sch. Bip. ex. I. Rich.) ('. Jeffrey ( 1986); Senecio abyssinicus Sell Bip. ex A. Rich ( 184X) tvpe TU. Adowa. PI. abvss. I. Schimper 67 (P holo.. K M LE UPS iso ).

145. ASTERACEAE: 93. Emilia

255

Figure 153. E M IL IA S O N C IIIF O L IA . 1 habit x Vi; 2 capitulum x 3 ,3 - floret x 7 */3; 4 - part o f stamen x 27; 5 - style branches x 27; 6 - cyp­ sela x 41 / 3. All from Rochecouste 180. Drawn by Eleanor Catherine. (Repro­ duced from FI. Mascareigties. 109. C om posees. PI. 53.)

S. bellidifolius A. Rich. (1848). nom illegit.. non K r;th in H.B.K. (1818); S. quarlinianus Asch. in Schweinf. & Asch. (1867) - type: TU. Shire. Q, .nin-Dillon s.n. (P holo ). var. a lyssinica Erect annual heit>, 5-40 cm high. Stem slender, simple or few-branched, striate, angular, often w ith scattered crispe I hairs. Leaves simple, alternate, blade of lower ones r arrowed below' into a petiole-like base, often up to 2 cm lort^, middle and upper leaves sessile with cordate base, 'Dathulate or spathulate-orbicular, thin, 1-5 x 0.5-2 ;; cm. surfaces glabrous, rarely ciliate toward the margi if. margins denticulate, ciliatc. apex obtuse. Capiti radiate, narrowly oblong-campanulate. 5-6 x 6-8 n 11 at anthesis, 3-12 in open cymes. Peduncle slen­ der. gi ibtous, bracteate, 1-5 cm long. Receptaclc flat. 3-4 mm w de. Phyllaries 8-10, light green, linear to narrowly lancec late, unequal in width on the same capitula. striate, glabrous, with narrow scarious margins. 5-6 x 0.7-1.3

nun. Ray florets yellow, 1-seriate, 5-8. female, style not much exserted. tube c 3 mm long, glabrous, lamina ob­ long, 1.5-2.2 x 0.5-0.8 mm, (3-)4-striate, apex 3-fid, papillose. Disc florets yellow. Corolla 3.8-4 nun long, glabrous, tubular, slightly wider near apex, 5-lobed. lobes papillose. 0.2-0.3 nun long. Anthers ecalcarate, 1.5-1.8 mm long, connective c 0.3 mm long. Style bifurcate, c 3.2 mm long, branches c 1 nun long. Cypselas black, oblong or columnar. 1.2-1.3 x 0.4-0.5 nun, costate, setulose, setae white. Pappus of numerous white bristles. 3-4 nun long. 1-seriate. Degraded Terminalia-Erythrina woodland on sandy gravelly soil, Acacia-C 'ommiphora bushland. Cyperus— Xerophyto turf in Acacia busliland, grassland, rarely weed in cultivated fields; 1300-2200 111. EW TU SU GG SD BA HA; E Africa, Nigeria, Cameroun. Sudan, Central African Republic, Rep. Dem. du Congo, Burundi, Rwanda, Ma­ lawi. Mozambique. Zambia, Zimbabwe. Gilbert & Sebsebe D. 8732; Burger 1146; Tewolde B.G.E. 932.

256

145 ASTERACEAE: 93. Emilia

Var. macroglossa C. Jeffrey (1986), which differs by having larger ray florets (2.2—2.5 x 1.0—1.3 mm), occurs in Tanzania.

long. 5-lobed, lobes 0.2-0.3 mm long. Cypselas grey ob­ long, 2-2.3 x 0.5 mm, ribbed, setulose on the ribs, hairs short and appressed. Pappus hairs white, 5-6 mm long, numerous, barbellate.

2. E. somalensis (S. Moore) C. Jeffrey ( 1986); Euryops somalensis S. Moore (1900) - type: Kenya, Marsabit, Delamere s.n. (BM holo.). Senecio megamontanus Cufod. (1943) - type: SD. Mega, Corradi 2077, 2079 (FT syn.).

Acacia-Commiphora bushland; short, heavily grazed grassland, degraded woodland with Ficus, Ozoroa, Commiphora, Combretum, etc.; roadsides, wasteland, margins of cultivated fields, 1350-1920 m. KF SD; Su­ dan. E Africa. Rwanda, Burundi, Rep. Deni du Congo, Zambia. Zimbabwe. W. de Wilde 9432; Mesfin T. 8073; Mesfin T. & kagnew 2475.

Erect, bushy perennial herb, 30-40 cm high. Stem usually irregularly branched, striate-sulcate or angular, glabrous or sparsely floccose in upper parts. Leaves alternate, sim­ ple, lower ones oblong-elliptic with blade narrowed into a 4. E. caespitosa Oliv. (1873) pseudo-petiolate base to narrowly oblanceolate, upper - ty pe: Tanzania, Karague, Grant s.n. (K holo ). ones sessile, narrowly elliptic, sparsely pubescent espe­ Erect, annual herb. 20-75 cm high. Stems usually several cially on veins and margins. 30-50 x 5-10 nun. Capitula from the root apex, rarely solitary, branched near the base radiate, hemispheric. 15-20 x 6-7 mm at anthesis. soli­ and also near the inflorescences, glabrous or sparsely tary. terminating branches, or 1-3 in open cymes. hairy on lower intemodes. Leaves simple, lanceolate, Peduncle sparsely pubescent. 2-4 cm long, bracteate. mostly basal and purple beneath. (3—)5—11 x (0.5—) 1—2 Calyculi absent. Phyllanes green, 12-14, orange striate, cm, glabrous or glabrescent (liairs mostly on petioles and oblong-lanceolate, more or less equal in width. 6-7 x young leaves), base sagittate, margins denticulate-serrate; 1-1.5 mm. glabrous, margins scarious. apex acutecauline leaves sparse, linear-lanceolate, much nanowed apiculate. ciliate. pinkish. Ray florets c 13. orange, fe­ from the middle to the apex. 6-10 x 1—1.5 cm Capitula male. tube c 3.5 mm long, glabrous, lamina oblong, c 9 x discoid, campanulate. 3-4 in open corymbose cymes, 3.5 mm long, 4-striate, apex 3-fid. Disc florets yellow; co­ 10-12 x 5-7 mm at anthesis. up to 17x9 nun in fruit (incl. rolla c 6.5 mm long, glabrous, tubular-campanulate. florets). Peduncle 1.5-12.5 cm long, slightly widened be­ 5-lobed. lobes c 0.7 nun long, papillose. Anthers yellow, c low the receptacle, ebracteate. glabrous. Receptacle flat 2 mm long, ecalcarate. connective c 0.3 mm long. Style 5-7 mm wide in fruit. Phy llanes c 8. green, oblong with bifurcate. 5.8-6 mm long, branches c 0.8 nun long, trun­ narrow, scarious margins, unequal in width on the same cate and papillose at apex. Cypselas brown, oblongcapitula. muhi-stnate. sparsely pilose at anthesis. gla­ elliptic, c 2.6 x 0.6 nun. nbbcd. pubescent on ribs, grooves brous in fruit. 6-6.5 x 1—1.5 mm al anthesis. up to 9-12 x glabrous. Pappus liairs white, numerous, barbellate. 4-5.2 2.5 mm in fruit. Florets pale mauve, 30-50. Corolla 5-6 mm long, uniseriate. nun long at anthesis, 8-9 nun long in fruit, tubular, gla­ Acacia drepanolohium woodland on black clay soil. brous. 5-lobed. lobes c 1 nun long. Cypselas black with Acacia-Commiphora Launaea bushland on dry sandy witish ribs, columnar. 3.5—4 x 0.7-1 nun. glabrous. soil; 1250-1920 m. SD; Kenya. Somalia Mesfin T & Pappus of numerous white bnstles. 5-6 mm long. I ollesen 4260; Ensermu K. 1149. Habitat in die Flora area unknown, elsewhere it is re­ corded from grassland and cultivated fields; 850 m IL; 3. E. discifolia (Oliv.) C Jeffrey (1986); Sudan, Central African Republic. Lisanework \ in Senecio discifolius Oliv. (1873) - type: Uganda. Mesfin T. 6243 Grant s.n (K holo ). Erect, bushy annual herb. 10-40 cm liigh and often 10-20 cm wide. Stem with ascending branches arising in lower parts, light green with scattered wliite hairs, stnatesulcate. leafy especially at base. Leaves simple, alternate, slightly glaucous, orbicular to obovate-spathulate. nar­ rowed below into a winged petiolar base, puberulous. ashy or light green, 2-7.5 x [-4 cm, margins denticulate, apex obtuse. Capitula radiate, campanulate. 7-8 x 8-12 nun at anthesis. solitary terminating branches, or 2-3 in open cymes. Peduncle glabrous, yellowish-green. 5-25 cm long. Reccptacle convex 5-8 mm wide. Involucre callose-thickened. yellowish brown at base. Phyllarics light green. 12-14. linear, striate, glabrous, margins scarious. 6-7 x 0.5-1.8 nun. reflexed after flowering. Ray florets 18-20. orange or golden yellow. 1-senate, female, tube slender. 4.5-5.5 mm long, ciliate near the apex, lamina oblong. 2.5—3 x 1—1.5 nun. 3-fid at apex, reflexed or curled outwardly Disc florets yellow; corolla 5-6 nun

5 E. jeffreyana Lisowski (1991) - type: Zaire. Mt Kaliuzi. Lisowski 85628 (POZG holo). Erect, annual herb. 10-35 cm high. Stem solitary and zigzaging or branched near the base, glabrous to sparsely pilose in lower intemodes. liairs white. Lower leaves ovate-lanceolalc. narrowed into a narrowly winged peti­ ole. 6-8 x 2-3 cm. petiole up to 2 cm long, glabrous, mar­ gins denticulate, apex acute; upper leaves oblonglanccolate. aunculate and semi-amplcxicaul al base. Capitula discoid. 2-4 on long peduncles, held away from the leaves. 8-10 x 3-4 min al antliesis. Pcdunclcs glabrous or glabrescent. up to 6 cm long. Reccptacle conv ex. 3-4.5 mm wide. Phy llanes 10-12. green, glabrous. 6-7 x (0.5-) 1-2 mm. Florets bright orange. Corolla tubular, wider in the limbs. 7.-9.2 nun long. 5-lobed. lobes 1.8-2 nun long, glabrous Anthers yellow . 1.8-2 nun long, conncc-

145. ASTERACEAE. 93. Emilia

ti' : c0.4 mm long, collar c 0.4 mm long. Style bifurcate. 7 -8.6 mm long, exserted, brandies 1.4-1.6 nun long. aflEN with an appendage of fused papillae. Cypselas purphsh-brown (only slightly immature seen), 5-ribbed. white pubescent on the ribs, c 2.6 x 0.6 mm Pappus white. c 4- nun long. Open bushland with Acacia drepanolobium on grey brown or black soil: 1100-1150 in. SD; Kenya. Rep. Dem. du Congo. Central African Republic. Burundi. Gilbert & Sebsebe D. 8784; Ensermu K. 1176: Friis ct al. 8126 .

6 E. adama-gil)aensis.\/^y?/7 & Beentje (2004) type: SD. Adama Giba. 10 km SW of Kibrc Mengist. 1850 in. Gilbert & Ermias 8423 (K holo.. ETH UPS iso.) E :z l ?short-lived perennial herb. 12-25 cm high. Stem b inched and woody near tlic base, younger parts pubescc nt, green, greyish-brown and glabrous in older parts. 2 : 111111 wide. Leaves simple, sessile, linear-lanccolatc. s| iiifcsely to moderately densely pubescent especially on mid-vein and margins, 20-50 x 2-5 nun. margins entire, revolute, apex acute-acuminate. Capitula solitary or 2-3 in cy mes. discoid. 6-8 x 3 .5-4 mm at anthesis. up to 9 nun hjfeh in fruit. Peduncle glabrous, bracteatc or cbractcate. 3 • cm long. 0.2-0.5 mm wide. Rcccptaclc flat. 1.5-1.8 mm wide after anthesis. Phyllaries green. 8. 4.5-5 x 0 fc 1.2 mm at anthesis. up to 6.5 mm long in fruit. glab : j s or sparsely pubescent toward apex, liairs 1-senate, minute. Florets pale pink to almost white: corolla tubular with slighlty expanded limb. 6-7 mm long, lobes c 1.5 n il long, glabrous. Anthers pale yellow to white, c 1.5 mn long, conncctive subulate. Style 6-7 mm long, bifurc; te. apex truncate with short papillae. Achcnes (imma­ ture) brownish. 2-2.5 x 0.2-0.5 nun, densely pubcsccnt. 01 long or oblong-elliptic. Pappus 4-5 nun long. Grassland or bushland with scattered trees, mostly O • nbretun mo lie, on basement complex with very thin sop: alt. 1850 in; SD. Gilbert & Ermias 8423: I 'atova 2* >8 .

In the key to the Tropical East African spccics of E