Flora of Ethiopia and Eritrea (Volume 5 Gentinaceae to Cyclocheilaceae) [5] 9197128562, 9789197128568

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FLORA OF ETHIOPIA AND ERITREA VOLUME 5

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GENTIANACEAE TO CYCLOCHEILACEAE

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Editors: Inga Hedberg Ensermu Kelbessa, Sue Edwards, Sebsebe Demissew & Eva Persson

Gift from the National Herbarium(ETH) Department of Plant Biology and

Biodiversity Management College of Natural Sciences

Addis Ababa University

flo ra of

ETHIOPIA AND ERITREA VOLUME 5

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 - Eritrea East, below and to the east o f the 1000 m contour EW - Eritrea West, above and to the west of the 1000 m contour AF - Afar region, below and to the east of the 1000 m contour to the Eritrean border in the east and the Harerge border in the south TU - Tigray region, above and to the west of the 1000 m contour GD - Gonder region GJ - Gojam region WU -W elo region, above and to the west of the 1000 m contour

SU - Shewa region, above and to the west of the 1000 m contour AR - Arsi region WG Wclega region 1L -

llubabor region

KI

Kefa region

GG - Gamo Gofa region SD

Sidamo region

BA

Bale region

I LA

Harerge region

MAP OF ETHIOPIA AND ERITREA SHOWING MAJOR PHYSIOGRAPHIC FEATURES

35

40

FLORA OF ETHIOPIA AND ERITREA VOLUME 5 GENTIANACEAE

TO

CYCLOCHEILACEAE

Editors: Inga Hedberg, Ensertnu Kelbessa, Sue Edwards, Sebsebe Demissew & Eva Persson

Addis Ababa, Ethiopia Uppsala, Sweden 2006

Published by The National Herbarium, Biology Department, Science Faculty, Addis Ababa University, Ethiopia

'i

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 Agency. Project Leader - Dr. Sebsebe Demissew, and European Coordinator - Dr. Inga Hedberg

V 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 , except some text sections which were prepared in W ord 97. Printed in Times New Roman. Printed by the Educational Materials Production and Distribution Agency (EM PDA), Textbook Production Department, Addis Ababa. Ethiopia. © The National Herbarium, Addis Ababa University', 2006 ISBN 91-971285-6-2

D istributed by: In Ethiopia The National Herbarium Addis Ababa University P.O. Box 3434, Addis Ababa, ETHIOPIA T e l. +251-1-11 43 23 F ax :+ 2 5 1 -1 -5 5 23 50

Outside Ethiopia The Swedish Science Press. P.O. Box 118 SE-751 04 Uppsala SWEDEN Fax: +46-18-36 52 77 e-mail [email protected]

e-mail: [email protected]

Cover illustrations: Front: Convolvulus bidremis drawn by Damtew Teferra. Back: Justicia schimperiana drawn by Sara Howard. Spine: Infructescence o f Solanum memphiticum drawn by Margaret Tebbs

FLORA OF ETHIOPIA AND ERITREA VOLUME 5

CONTENTS Map of the floristic regions of Ethiopia and Eritrea Map of Ethiopia and Eritrea showing major physiographic features Editorial Board Editorial Team Contributors Foreword Acknowledgements Key to families in volumes 2 to 7

ii iii viii ix x xi xii xiii

F lo ra p a r t

146. Gentianaceae by Sileshi Nemomissa 147. Primulaceae by P. Bizzarri 148. Plumbaginaceae by J.R. Edmondson 149. Plantaginaceae by O. Ryding 150. Sphenocleaceae by M. Thulin 151. Lobeliaceae by M. Thulin 152. Campanulaceae by M. Thulin 153. Goodeniaceae by Sebsebe Demissew & Sue Edwards 154. Menyanthaceae by Sebsebe Demissew & Sue Edwards 155. Polemoniaceae by Sebsebe Demissew & Sue Edwards 156. Hydrophyilaceae by Sebsebe Demissew & Sue Edwards 157. Boraginaceae by H. Riedl & Sue Edwards 158. Solanaceae by I. Friis 159A. Convolvulaceae by Sebsebe Demissew 159B. Cuscutaceae by Getachew Aweke & Sue Edwards 160. Scrophulariaceae by E. Fischer 161. Orobanchaceae by C. Parker 162. Lentibulariaceae by I. Hedberg & O. Hedberg 163. Gesneriacae by B.L. Burtt 164. Bignoniaceae by Saliy Bidgood 165. Pedaliaceae by H.-D. Ihlenfeldt 166. Acanthaceae by Ensermu Kelbessa 167A. Globuiariaceae by Ensermu Kelbessa 167B. Myoporaceae by Ensermu Kelbessa 168. Verbenaceae by Sebsebe Demissew 169. Avicenniaceae by Sebsebe Demissew 170. Lamiaceae by O. Ryding 171. Cyclocheilaceae by Sebsebe Demissew

1 18 29 34 37 39 49 58 59 61 63 64 103 161 232 235 309 315 320 322 335 345 496 498 499 515 516 605

A Glossary of Botanical Terms Abbreviations Index to Scientific Names Vernacular Names for Species in Volume 5

609 627 631 663

FLORA OF ETHIOPIA AND ERITREA VOLUME 5 Editorial Team Dr. Inga H edberg, D epartm ent o f System atic B otany, U ppsala U niversity, Sw eden Prof. 01 ov H edberg, D epartm ent o f System atic Botany, U ppsala U niversity, Sw eden M s Eva Persson D epartm ent o f System atic B otany, U ppsala U niversity, Sw eden Prof. lb Friis, B otanical M useum and Library, U niversity o f C openhagen, D enm ark Prof. Sebsebe D em issew , K eeper, The N ational H erbarium , B iology D epartm ent, F aculty o f Science, A ddis A baba U niversity, E thiopia Dr. E nserm u K elbessa, T he N ational H erbarium , B iology D epartm ent, Faculty o f Science, A ddis A baba U niversity, E thiopia M s Sue E dw ards, T he N ational H erbarium , B iology D epartm ent, Faculty o f Science, A ddis A baba U niversity, E thiopia M r M elaku W ondafrash, T he N ational H erbarium , B iology D epartm ent, F aculty o f Science, A ddis A baba U niversity, E thiopia M r Sileshi N em om issa, T he N ational H erbarium , B iology D epartm ent, Faculty o f Science, A ddis A baba U niversity, E thiopia

FLORA OF ETHIOPIA AND ERITREA VOLUME 5 Editorial Board Dr. T ew olde B erhan G ebre E gziabher, G eneral m anager, environm ental Protection A uthority, A ddis A baba, E thiopia E m eritus Prof. 01 ov H edberg, D epartm ent o f S ystem atic B otany, U ppsala U niversity, Sw eden Prof. Sebsebe D em issew , K eeper, The N ational H erbarium , B iology D epartm ent, Faculty o f Science, A ddis A baba U niversity, E thiopia D r M esfin T adesse, (alternate m em ber) T h e N ational H erbarium , B iology D epartm ent, F aculty o f Science, A ddis A baba U niversity, E thiopia Dr. E nserm u K elbessa, (alternate m em ber) C urator, T he N ational H erbarium , B iology D epartm ent, F acu lty o f Science, A ddis A baba U niversity, E thiopia Prof. Ib Friis, B otanical M useum and Library, U niversity o f C openhagen, D enm ark Dr. Inga H edberg, D epartm ent o f S ystem atic B otany, U ppsala U niversity, Sw eden Ms Sue E dw ards, T h e N ational H erbarium , B iology D epartm ent, Faculty o f Science, A ddis A baba U niversity, E thiopia

FLORA OF ETHIOPIA AND ERITREA VOLUME 5 CONTRIBUTORS Or. Sally Bidgood, The Herbarium, Royal Botanic Gardens. Kew, Richmond, Surrey TW9 3DS, England Prof. Paola Bizzarri, Erbario Tropicale di Firenze, via G La Pira 4, 50121 Firenze, Italy Mr Brian L. Burtt, Royal Botanic Gardens, Edinburgh, EH3 5LR, UK Dr. Jennifer M. Edmonds, University o f Leeds, School of Biology, Leeds LS2 9JT, UK Dr. John Edmondson, National Museums Liverpool, William Brown St, Liverpool L3 8EN, UK Dr. Ensermu Kelbessa, The National Herbarium, Addis Ababa University, P.O Box 3434, Addis Ababa, Ethiopia Ms Sue Edwards, The National Herbarium, Addis Ababa University, P.O. Box 3434, Addis Ababa, Ethiopia Dr Eberhard Fischer, Institute o f Integrated Natural Sciences, Dept. Biology-Botany, Uni versitat Koblenz-Landau, Universitatstr. 1, D-56070 Koblenz, Germany Prof. Ib Friis, Botanical Museum, University of Copenhagen, Gothersgade 130, D K -1123 Copenhagen K, Denmark Mr Gebrehiwet Medhanie, University o f Asmara, P.O. Box 1220, Asmara, Eritrea Dr. Getachew Aweke, The National Herbarium, Addis Ababa University, P.O. Box 3434, Addis Ababa, Ethiopia Dr. Inga Hedberg, Department o f Systematic Botany, EBC, Uppsala University, Norbyv. 18D, SE-752 36 Uppsala, Sweden Emer. Prof. Olov Hedberg, Department o f Systematic Botany, EBC, Uppsala University, Norbyv. 18D, SE-752 36 Uppsala, Sweden Dr. Mats Hjertson, Botany Section (Fytoteket), Museum o f Evolution, EBC, Uppsala University, Norbyv. 16, SE-752 36 Uppsala, Sweden MSc. Hans Peter Hofmann, Institute o f Integrated Natural Sciences, Dept. Biology-Botany, Universitat Koblenz-Landau, Universitatstr. 1, D-56070 Koblenz, Germany Prof Hans-Dietrich Ihlenfeldt, Ostseestrasse 40, D-24369 Waabs/Langholz, Germany MSc. Mattias Iwarsson, Swedish Biodiversity Centre, Box 7007, SE-750 07 Uppsala, Sweden Prof. John. K. Morton, Dept, o f Biology, University o f Waterloo, Waterloo, Ontario, Canada N2L 3G1 Dr. Chris Parker, 5 Royal York Crescent, Flat 4, Bristol BS8 4JZ, UK MSc. Eva Persson, Department of Systematic Botany, EBC. Uppsala University, Norbyv. 18D, SE-752 36 Uppsala, Sweden Prof. Harald Riedl, Herbarium, Dept, o f Botany, Naturhistorisches Museum Wien. Burgring 7, A-1014 Wien, Austria Dr. Olof Ryding, Botanical Museum, University o f Copenhagen, Gothersgade 130, D K -1123 Copenhagen K, Denmark Dr. Oskar Sebald, now retired, former address: State Museum of Natural History, Rosenstein 1, D-70191 Stuttgart, Germany Dr. Sebsebe Demissew, The National Herbarium, Addis Ababa University, P.O. Box 3434, Addis Ababa, Ethiopia Prof. Siegmund Seybold, now retired, former address: State Museum of Natural History, Rosenstein I, D-70191 Stuttgart, Germany Dr. Sileshi Nemomissa, The National Herbarium, Addis Ababa University, P.O. Box 3434, Addis Ababa, Ethiopia Prof. Mats Thulin, Department o f Systematic Botany, EBC, Uppsala University, Norbyvagen 18D, SE-752 36 Uppsala, Sweden

FOREWORD The present volum e is the last part o f F lora o f Ethiopia and Eritrea dealing w ith the angiosperm s. The volum e treats 28 fam ilies, Gentianaceae - Cyclocheilaceae and includes the very difficult fam ilies Solanaceae (78 taxa), Acanthaceae (215 taxa) and Lamiaceae (221 taxa). Some o f the fam ilies treated contain im portant medicinal plants o r stim ulants (Solanaceae), food plants ( Convolvulaceae, Solanaceae ), spices (.Lamiaceae , Solanaceae) and ornam ental trees and lianas (Bignoniaceae). The editing o f the contributions w as started in Addis A beba but w as, for various reasons, later transferred to Uppsala. The Flora Project w as initiated in 1980 and w hat remains is Vol. 1 , w hich will contain the ferns, fern allies and G ym nosperm s, general inform ation on the project and a general index to all volum es. V olum e 3 provides background inform ation about the w riting o f a m odem Flora, notes on the collecting and identification o f plants, and inform ation on the form at used for presenting fam ilies, genera and species. The reader is referred to that volum e for inform ation dealing w ith these m atters. A rrangem ents to m ake the Flora available on CD rom are under way, but for obvious reasons it is essential that the w ork will also remain available in printed form. I.H., E.K., S.E., S.D., E.P. O ctober 2006

ACKNOWLEDGEMENTS The production o f Flora o f Ethiopia and Eritrea enjoys the support o f m any institutions and individuals, both within and outside Ethiopia. The editors particularly wish to em phasize the contributions o f the following, all o f whom are gratefully acknow ledged for the adm inistrative support they have rendered to the project: The Ethiopian G overnm ent through the Ethiopian Science and Technology A gency (previously known as Science and Technology C om m ission), in particular the previous C om m issioner, Ato A srat B ulbula and the current D irector General, Dr. Zerihun Kebede, and the Swedish G overnm ent through the Sw edish International Development Co-operation Agency (Sida/SA REC), which has provided the financial support for the preparation and publication of all volum es o f the Flora; the past and present officers o f A ddis Ababa University, especially the office o f the President and its current President, Prof. A ndreas Eshete, the current Vice Presidents, Ato M oham m ed Habib (VP for Business and D evelopm ent). Prof. Tsige G ebre M ariam, Associate VP for Research and G raduate Program m e, the Research D irector, Dr Daniel Fekade. the past and present Deans o f the Faculty o f Science, especially the present Dean. Dr A raya Asfaw, and the past and present Heads o f the Departm ent o f Biology, especially the present Head. D r D awit Abate. This volum e could not have been prepared without the support and assistance o f the curators, keepers and directors of the following herbaria: A ddis A beba (ETH). Copenhagen (C), Firenze (FT and FI), London (BM. K), Nairobi (EA ), Paris (P), Stockholm (S), U ppsala (UPS), W ageningen (W A G ), W ien (W , W U ) and Zurich (Z). Thanks are especially due to the Royal Botanic Gardens. Kew, in England, particularly the past and present Directors, Prot. Gillean Prance and Prof. Peter Crane, and the past and present K eepers o f the Kewr Herbarium , Prof. Gren Lucas and Prof. Simon Owens, and their scientific and technical staff Thanks are also due to the Botanical M useum and Library o f the University o f C openhagen, to the Carlsberg Foundation for supporting extensive field work, which has contributed much to im proving the coverage o f species and their distribution, and to the staff o f the Biology Library, U ppsala U niversity, for their indefatigable assistance in tracing relevant literature and titles. Thanks to the courtesy o f various authorities, authors, editors, artists and Editorial B oards, it has been possible to use illustrations already published in a num ber o f botanical publications. The follow ing authorities and sources are gratefully acknowledged. AETFAT Zomba P lenary Meeting. A revised handbook to the Flora of Ceylon; A ssociation de Botanique tropicale (Flore de M adagascar et des Comores, Flore du Gabon): Families o f flowering plants (J. Hutchm son 1959); Fam ilies o f flo w e rin g plants in Ethiopia (W .C. Burger 1967); Flora Italica: Flora o f Egypt, Flora oj West Tropical A frica ; Flora Z am besiaca M anaging Com m ittee (Flora Zambesiaca); National Botanic Garden o f Belgium {Flore d'A frique Centrale, Flore du Congo. Rwanda, Burundi, Flore des Spermatophytes du P arc N ational Albert): Iconographia Florae Italicae: National M useums o f K enya {Kenya trees a n d shrubs): N ordic Journal o f Boiany: Royal B otanic G ardens, Kew (Flora o f Tropical East Africa , Hooker's leones Plant arum. Kew B ulletin); Students Flora o f Egypt: Stuttgarter Beitr. Naturk.\ Symbolae Botanicae Upsalienses: The genus Utriadaria (P. T aylor 1989); The Israel Academy o f Sciences and Humanities (Flora Palaestina): The Linnean Society o f London (Botanical

Journal o f the Linnaean Society); Webbia. The editors gratefully acknow ledge the contribution o f P ro f Ib Friis in checking and bringing uptodate the nom enclatural issues in the volum e. I.H ., E.K., S.E., S.D., E.P O ctober 2006

K E Y T O F A M IL IE S 1 2 by Sue Edwards, Ensermu Kelbessa & Sebsebe Demissew"

PRIMARY KEY TO 20 ARTIFICIAL GROUPS (Each artificial group contains keys that will lead to the correct family name.) 1. Plants floating on the surface of water or sub­ merged beneath the surface or lying flat on the mud during periods of low' water level. Group 1 - Plants growing upright out of the water (above the water level) or growing on soil, on rocks, or on other plants; neither floating nor submerged. 2 2. Plants without green pigment (chlorophyll), parasites or saprophytes; the leaves absent or reduced to scale-like or bract-like structures. Group 2 - Plants with green pigment (chlorophyll); leaves usually present, at least during the growing season. 3 3. Plants with leaves more than one metre long or trees and shrubs with grass-like or palm-like leaves. Group 3 - Plants with leaves less than one metre long, if trees or shrubs then leaves not grass-like nor palm-like. 4 4. Grass-like herbs: leaves alternate or basal, blades narrow and with parallel veins. Group 4 - Plants not grass-like: leaves simple, deeply divided, or compound, usually with pinnate or palmate venation and usually with a petiole. 5 5. Flowers densely crowded into compact inflores­ cences (heads), individual flowers closely pressed against each other or against their subtending bracts and often difficult to separate from each other. Group 5 - Flowers not densely crowded into compact inflorescences (heads), individual flowers easy to separate from each other if in a dense inflorescence. 6 6. Stamens and pistils naked, without bracts or perianth parts, or on 1 or 2 special bracts forming a cone, but not within a cup-like or tubular structure. Group 6 - Stamens and pistils covered, surrounded by three or more bracts or perianth parts (sepals and/or petals), or within a cup-like or tubular structure, but not forming a cone. 7

7. Flowers with one or more spurs or with a sac-like pouch at the base of the flower. Group 7 - Flowers without spurs and a sac-like pouch at the base of the flower.

8

8. Flow'ers apparently only female; stamens absent, or lacking anthers and modified into staminodes, or stamens united with each other and/or with other parts of the flower and difficult to identify. Group 8 - Flowers either bisexual or only male, with free and distinct anthers. 9 9. Whole plant or some flowers on the plant functionally only male, flowers or reproductive structures with distinct anthers. Group 9 - Whole plant and flowers bisexual, with stamens and a central pistil or group of pistils of which only the style may be visible in the centre of the flower. t 10 10. Each flower with two or more pistils and styles present and arising from separate ovaries, or if only one ovary is present then with two or more styles arising from separate areas on it. Group 10 - Each flower with only one pistil and only one style, or if two or more styles are present then the styles all arising from one point on the top of the ovary. 11 11. Ovary on a stalk, perianth parts attached below the base of the ovary; fruit often on a stalk with a joint or articulation where the perianth parts and stamens were attached. Group 11 - Ov ary on the receptacle, perianth parts attached at base of ovaiy, or on a disc or raised receptacle, or ovary below the level of perianth and stamen attachment. 12 12. Flowers with an inferior or half-inferior ovary, at least some part of the ovary below the level of sepals, petals, or perianth-tube attachment; fruit with scars or persistent perianth parts on top and opposite the fruit stalk. Group 12 - Flowers with a superior ovary, ovary found above the level of sepals, petals, or perianth-tube attachment; fruit with scars or persistent perianth parts where the stalk is attached. 13

1Based on W.C. Burger (1967). Families o f Flowering Plants in Ethiopia, from Experiment Station Bulletin No. 45, Oklahoma Agricultural Experiment Station 2 "Tie National Herbarium, Addis Ababa University, P.O. Box 3434, Addis Ababa, Ethiopia

xiv

KEY TO FAMILIES

13. Flowers with 13 or more than 13 stamens, or with 13 or more than 13 anthers when the filaments are united or absent. Group 13 - Flowers with 12 or fewer than 12 stamens, or with 12 or fewer than 12 anthers when the filaments are united or absent. 14 14. Filaments united, forming a short or long tube. Group 14 - Filaments free, never forming a tube, or united only at the base, or absent. 15 15. Flowers bilaterally symmetrical (zygomorphic). petals always united to form a tube. Group 15 - Flowers radially symmetrical (actinomorphic), petals all similar in size and shape, free or united, or absent. 16 16. Petals and/or sepals united and coming off as a single unit, never as separate petals. Group 16 - Petals free, usually come off separately and not together as a single unit, or absent, sepals free or united, but not forming a colourful tube. 17 17. Flowers with stamens more or fewer in number than petals, or sepals if petals absent. Group 17 - Flowers with the stamens equal in number to the petals, or sepals if petals absent. 18 18. Flowers with either sepals or petals (but not both) and with stamens attached opposite, at the base of, or on the sepals or petals. Group 18 - Flow ers with both sepals and petals, or if only one whorl is present then the stamens alter­ nating between the perianth parts. 19 19. Stamens alternating with the petals and opposite the sepals, if petals absent or fallen off. Group 19 - Stamens opposite the petals and alternate with the sepals if petals absent or have fallen off. Group 20 GROUP l 1 Herbaceous plants without woody stems growing on or under water, freely floating or if rooted in the soil then not growing more than a few (up to 5 cm) centimetres above the water level, lying flat upon the mud during dry periods and not self-supporting. (1-7) = number of the volume where the family account is found.J 1. Plants of ocean water or of salty water close to the ocean shore. - Plants of fresh or of only slightly salty water.

2 5

2. Flowers found on the lower part of the leaf-base and at first enclosed between the edges of the sheathing leaf-base. [6] 177. Ruppiaceae 1 For lily-like and other monocotyledons, see the key at the start of Volume 6.

- Flowers on stalks or in the axils of leaves.

3

3. Flowers enclosed in 2 opposite bracts, each ovary' with 2 to 12 styles. |6J 172. H vdrocharitaceae - Flowers not enclosed in 2 opposite bracts; each ovary with usually only 1 style. 4 4. Flowers in leaf-axils; perianth of 3 scales or absent; stamens with thread-like pollen. 16] 178. Zannichelliaceac - Flowers in terminal spikes; perianth absent; stamens with powdery pollen. [6] 176. Potamogetonaceae 5. Plants without a stem or stem-like portion, flat and circular or oval in outline, each little plant less than 1 cm long (sometimes attached to each other and forming a group longer than 1 cm), floating fiat upon the water surface and not attached to rocks or soil. [6] 182. Lemnaceae - Plants with a stem or stem-like portion, or if the stem is absent, then attached to rocks and plant more than 1 cm long; if less than 1 cm long then with small overlapping leaves. 6 6 . Plants cabbage-like and freely floating, with leaf rosettes above the water level; roots not attached to the soil; /lowers in a single tubular bract, male reduced to several anthers on a stalk above the reduced female flower. (Pistia) [6] 181. Araceae - Plants not cabbage-like and freely floating, without leaf rosettes; leaves usually flat on the surface or beneath the surface; flowers not in tubular bracts, or plants without flowers. 7 7. Leaves floating with 4 fan- or wedge-shaped leaflets attached to a central petiole; darkcoloured nut-like bodies near the base of the petiole under the water. (Marsilea) 11J M arsileaceae - Leaves not floating, and without 4 fan- or wedgeshaped leaflets; nut-like bodies not found at the base of the petiole. 8 8 Plants attached to rocks in fast moving water or in the spray of moving water, superficially looking like a liverwort or moss; leaves alternate or absent, flowers very small with usually 1 or 2 stamens and an ovary with 2 or 3 styles [2,1 J 26. Podostemonaceae - Plants floating or submerged. 9 9. Underwater leaves with small (2 to 5 mm) round bladders that catch very small aquatic organisms. [5J 163. Lentibulariaceae - Underwater leaves without small bladders. 10 10. Styles 2 or more, coming from separate ovaries or from separate areas on the same ovaiy. or styles absent and stigmas 2 or more on separate ovaries, or on separate areas of a solitary ovary. 11

KEY TO FAMILIES

- Style 1, or if 2 or more, then coming from one point on the top of the ovary, or pistil absent in male flowers. 20 11. Leaves large and floating, over 20 cm wide, with petiole attached at or near the centre of the blade; flowers usually over 2 cm across. 12 - Leaves usually less than 20 cm wide, with petiole at or near the edge of the blade; flowers usually less than 2 cm across. 13 12. Petals 3, pistils 3 or more, separate from each other and from the receptacle of the flower. [2.1] 7. Cabombaceae - Petals many, pistil 1 and made up of many parts, united below with the receptacle and separate only near the top. [2,1] 9. Nymphaeaceae 13. Pistils usually 4 or a single pistil with 4 styles and 4 locules; leaves along the stem. - Pistils 3, 6, or more; leaves usually near the base of the plant.

14 17

14. Flowers in terminal spikes; sepals 4, narrow at the base and bract-like; stamens 2 or 4; leaves broad. [6] 176. Potamogetonaceae - Flowers solitary, in leaf-axils; leaves narrow. 15 15. Leaves in whorls, much divided with thread-like leaflets; pistil 1 with ovary 4-lobed. (Myriophyllum) [2,1] 51. Haloragaceae - Leaves in opposite pairs or alternate, simple, narrow; pistils 4 or more. 16 16. Plant growing erect out of the water, often in dense masses; sepals 4, petals present. (Crassula) [3] 88. Crassulaceae - Plant growing submerged; sepals 3, petals absent. (Zannichellia) [6] 178. Zannichelliaceae 17 Flowers in 2-branched spike-like inflorescences. [6] 173. Aponogetonaceae - Flowers in umbels or racemes, or solitary in leafaxils and in whorls around the stem. 18 18. Flowers usually solitary and on a long stalk, with many pistils and stamens in each flower; petals 5 or more; leaves lobed or much divided. [2.1] 5. Ranunculaceae - Flowers usually in umbels or whorls along the stem; petals 3 or absent; leaves entire. 19 19. Stamens 3 or 6, or more; fruit an achene. [6] 174. Alismataceae - Stamens usually 9; fruit a follicle. Butomaceae1 20. Leaves alternate and bipinnate; flowers in heads, each flower with 5 or 10 stamens or the asexual flowers with 10 petal-like staminodes. (.Neptunia) [3] 95. (Leguminosae) Fabaceae

1 Recorded from East Africa but not in the Flora area.

- Leave^ not alternate and bipinnate and the flowers not in heads.

xv

21

21. Flowers unisexual, with anthers, without a pistil or pistil-like structure. 22 - At least some flowers bisexual, all with a pistil or pistil-like structure and some also with anthers. 28 22. Stamen 1 in each flower. - Stamens 2 or more in each flower.

23 26

23. Flowers in dense spikes, each spike on a stalk. Hydrostachyaceae1 - Flowers in the axils of opposite or whorled leaves, each flower sessile or on a short stalk. 24 24. Filament absent, anther sessile, enclosed in a tubular bract. [6] 180. Najacaceae - Filament present, anther on a long and slender stalk, not enclosed in a tubular bract. 25 25. Anthers with 1 theca and opening by 1 slit. [2,1] 53. Callitrichaceae - Anthers with 2 tliecae and opening by 2 slits. [6] 178. Zannichelliaceae 26. Stamens 10 to 20 in each flower; leaves with many slender divisions that usually have serrulate edges. [2,1] 8. Ceratophyllaceae - Stamens 3 to 9 (rarely 10) in each flower; leaves broad, with entire edges. 27 27. Stamens 3 or 6 in each flower (if more than 6 stamens present the plant should have a few bisexual flowers), flowers in a whorl around the stem or sometimes in umbels; all leaves from the base of the plant. [6] 174. Alismataceae - Stamens 8 in each flower; leaves and flowers in whorls along the stem with usually 4 leaves and 4 flowers at each node, the leaves with many slender divisions with entire edges. (Myriophyllum) [2,1] 51. Haloragaceae 28. Ovary-like structure minute (less than 3 mm long) with a wall with parallel spiral grooves. (Leaf-like structures borne in whorls along the stem - rough to the touch in Chara and smooth in Nitella). Characeae ( A lg a e ) - Ovary over 3 mm long, wall without parallel spiral grooves. 29 29. Flowers naked, without a perianth or bracts, not enclosed by a tubular bract. - Flowers with a perianth or bracts, or within a tubular bract.

32

30. Flowers in dense spikes; ovary with 1 locule and 2 styles. Hydrostachyaceae1 - Flowers in leaf-axils; ovary with 2 or 4 styles.

31

30

31. Pistil without subtending bracts, ovary with 2 or 4 styles and 1 locule with 1 basal ovule; fruit not breaking up. [6] 180. Najadaceae

xvi

KEY TO FAMILIES

- Pistil subtended by 2 bracts, ovary with 2 styles and 4 locules, each locule with 1 pendulous ovule; fruit breaking up into 4 parts. [2,1] 53. Callitrichaceae 32. Flowers enclosed within 2 opposite bracts or within a tubular bract; ovary 1-locular and with 3 to 6 styles, styles often 2-branched. [6] 172. Hydrocharitaceae - Flowers not enclosed within 2 opposite bracts nor enclosed within a tubular bract, reproductive parts may be enclosed within a tubular perianth but the flower itself (with its perianth) is not enclosed by a bract. 33 3 3. Leaves opposite or whorled along thestem. - Leaves alternate along the stem or all from the base.

34 37

34. Flowers with a 2-lipped corolla-tube or an in­ ferior ovary; leaves usually opposite and not in whorls. 35 - Flowers lacking a 2-lipped corolla-tube and the ovary not inferior; leaves often in whorls, usually very narrow or withnarrow divisions. 36 35. Petals united to form a 2-lipped corolla-tube; stamens 2 or 4; ovary superior. [5] 161. Scrophulariaceae - Petals free from each other, not forming a tube; stamens usually 4, 5, 8, or 10; ovary inferior. [2.1] 49. Onagraceae 36. Leaves with a folded or lens-shaped structure near the base that traps minute animals. (.Aldrovanda) [2,1] 25. Droseraceae - Leaves without a folded or lens-like structure near the base. 37 37. Leaves simple (not divided) and narrow; sepals 2

to 5, petals 2 to 5; ovary 3- to 5-locular. [2.1] 27. Elatinaceae - Leaves deeply divided into narrow segments with serrulate margins; sepals 8 to 12, petals absent; ovary 1-locular. [2,1] 8. Ceratophyllaceae 38. Floating leaves petiole attached at the centre of the blade. (Nymphoides) [5] 154. Menyanthaceae - Floating leaves with petiole attached at the edge. 39 39. Petioles of floating leaves with a thicker spongy portion at their midpoint. - Petioles without a swollen portion near their midpoint.

40 41

40. Leaves more or less triangular in outline; sepals 4. petals 4; ovary 2-locular and half-inferior. [2,1] 50. Trapaceae - Leaves rounded in outline; sepals 3, petals 3; ovary 1- or 3-locular and superior. [6] 201. Pontederiaceae

41. Ovary inferior; fruit often with a dry perianth or scars where the perianth has fallen off at the top.

42

- Ovary superior; fruit often with a dry perianth or scars where the perianth has fallen off at the bottom. 43 42. Style.' 2, ovary with 2 locules, 1 ovule in each locule; flowers in umbels. [4,1] 132. (UmbeHiferae) Apiaceae - Style 1 or none, ovary with usually 4 locules, 1 to many ovules in each locule; flowers solitary' in leaf-axils or in racemes or panicles. [2,lj 49. O nagraceae 43. Leaves very narrow, without a distinct petiole; ovary with 3-6 locules, fruit breaking up into 3-6 parts. (Triglochin) [6 ] 175. Juncaginaceae - Leaves narrowly elliptic to rounded or arrowshaped. a distinct petiole usually present; ovary with 1-3 locules, fruit not breaking up. 44 44. Style 1; ovary with 1 or 3 locules and usually many ovules; leaves often arrow-shaped with almost parallel, venation. [6] 201. Pontederiaceae - Styles (or stigmas) 2 or 3; ovary with 1 locule and 1 basal ovule; leaves usually elliptic in outline, with pinnate venation. [2, 11 38. Polygonaceae G ROUP 2 Plants without green pigment (chlorophyll); the leaves absent or reduced to scale-like or bract-like structures. Mostly parasites or saprophytes taking their nutrients from the root systems or stems of other plants. 1. Plant climbing and twining about the host, attached to the host plant by root-like structures along the length of the slender stem. 2 - Plant not climbing or twining, attached to stem of the host at usually only one point, or several points below the ground, on the roots of the host. 3 2. Stamens 12 in each flower, anthers opening by flaps; petals separate, arising from a cup-shaped ^ receptacle, fruit with 1 seed. (Cassytha) [2,1] 3. Lauraceae - Stamens 4 or 5 in each flower, anthers opening by slits; petals united with stamens on the short corolla-tube; fruit with 1 to 4 seeds. [5] 160. Cuscutaceae 3. Plants growing upon stems or above-ground parts of their hosts. 4 - Plants growing from below the ground, attached to the roots of their hosts if parasitic. 5

xvi 1

KEY TO FAMILIES

4 Flowers borne on the stem of the parasite and not directly on the stem of the host; leaves usually present. (3J 115. Viscaceae - Flowers or reproductive structures growing directly from the stem of the host, the parasite without a stem or w ith only a short stem and without leaves. [2,1] 14. Rafflesiaceae 5. Flowers few to very many, if solitary on a stalk 1 cm wide or a broad base.

6 10

6. Flowers very numerous (more than 50) and densely crowded together. [3 & 2,2 Appendix] 117. Balanophoraccae l - Flowers fewer than 50, usually loosely arranged in spikes or racemes. 7 7. Flowers unisexual; ovary inferior w ith 1 loculc and 8 to 14 parietal placentas; anthers 8 to 20 on a single column; perianth of 1 whorl forming a short tube with 2 perianth bracts below. (Cytinus) [2,1 ] 14. Rafflesiaceae - Flower bisexual and not asabove. 8 8. Ovary inferior; stamens 3 or 1, or completely united to the sty le and not easily seen. [6] 200. Orchidaceae - Ch ary superior; stamens 4 or 2 on the irregular corolla-tube. 9 9. Chary 1-locular with 4 parietal placentas; plants dirty white or pale yellow, with dull red markings. [5] 162. O robanchaceae - Ovary 2-locular with axile placentas; plants bright yellow. [5] 161. Scrophulariaceae 10. Fiowers small and crowded on a thick central column, enclosed at first in a single large, sometimes petal-like, bract. [6] 181. Araceae - Flowers or fruiting body larger and single, not enclosed in a single petal-like bract. 11 11. Plant represented by a single flower with only the 3 or 4 perianth parts above the soil surface. [2.1] 13. Hydnoraceae - Plant with the flower or reproductive structure coming to the surface or above soil level, not remaining partly sunken into the ground. 12 12. Flowers unisexual, rarely bisexual, with 3 to 10 sepals, sepals sometimes absent in female flowers; anthers 8 to many, on separate fila­ ments or on a single column. [2.1] 14. Rafflesiaceae - Flowers absent; plants variously shaped but often umbrella-shaped and the broad upper part with many small pores or slits beneath that produce minute dust-like spores. Fungi

GROUP 3 Plants with leaves over one metre long, or trees and shrubs with palm-like leaves or grass-like leaves (the leaves or the leaf-segments with parallel veins or the veins arising all at one point and spreading like a fan). 1. Mature leaves with nearly all the secondary veins parallel with the primary vein or midrib of the leaf. 2 - Mature leaves with secondary' veins arising at an angle from the primary vein or midrib of the leaf, or if arising together from the base of the blade then the leaf fan-shaped or round in outline. 12 2. Plants with a trunk or thick stem; leavesin rosettes at the top or along the branches. - Plants without a trunk or thick stem; leaves coming from the base of the plant.

3 5

3. Leaves with sheathing bases; flowers very rcduccd. enclosed in opposite dry' bracts and without sepals or petals. [7] 215. (G ram ineae) Poaceae - Leaves without sheaths; flowers with sepals or petals. 4 4. Flower-stalk long, flowers solitary, >2 cm across; ovary' inferior. [6] 202A. Velloziaceae - Flower-stalk short, flowers numerous. < 2 cm across, usually in large branched inflorescences; ovary superior. (Dracaena) [6] 187. D racaenaceae 5. Leaves up to 3 m long and less than 10 cm wide. with air-chambers within; growing along stream edges and lake shores; inflorescence a brow'n cylinder of thousands of very slender tightly packed flowers. [6] 210. Typhaceae - Leaves and inflorescences not as above. 6 6. Plants with white or brightly coloured flowers, stamens and ovary' not enclosed in dry bracts. 7 - Plants lacking coloured flowers, stamens and ovary enclosed in small dry' bracts. 11 7. Ovary superior; fruit with dry perianth or scars where the perianth has fallen off at the base near the attachment of the fruit stalk. 8 - Ch an' inferior; fruit with dry' perianth or scars w here the perianth has fallen off at the top of the fruit, i.e. opposite the fruit stalk. 9 8. Leaves tough and fibrous, very' difficult to cut across; flowers usually in many-flowered, often branched, inflorescences. (Sansevieria) [6] 187. D racacnaceae - Leaves slender or thick and succulent but lacking strong fibres, usually easy to cut across the long axis of the leaf; flowers mostly in spikes or racemes. [6] ‘L iliaceae’1 1 See key to monocotyledons in front of Volume 6.

xviii

KEY TO FAMILIES

9. Leaves thick and fibrous, very difficult to cut across; inflorescence up to 4 m tall. (Agave) (6J 188. Agavaceae - Leaves not thick, without strong fibres, easier to cut across; inflorescence less than 1 m tall. 10 10. Stamens 6; perianth parts usually all alike. 6 in number; inflorescencc an umbel; at first enclos­ ed in 1 (or more) dry bract. (6] 195. Alliaceae - Stamens 3; perianth parts usually of 2 different kinds. 3+3; flowers solitary or in a cyme, first enclosed in 2 dry bracts. |6j 197. Iridaceae 11. Leaves in 2 vertical rows (when seen from above), leaf-base enclosing the stem by a usually split sheath; stamens and pistil enclosed in two opposite bracts, style branches and stigmas 2, usually feathery. [7J 215. (Gram ineae) Poaceae - Leaves in 3 vertical ranks or row's (when seen from above), leaf-base enclosing the stem by a usually closed sheath; stamens and pistil enclosed in a single bract or subtended by a whorl of 3 or 6 bract-like stmctures; style branches and stigmas 2 or 3, not feathery. |6] 212. Cyperaceae 12. Trunk made up of the large spongy leaf bases; inflorescences over 2 m long. [6] 203A. Musaceae - Trunk absent, or if a trunk present, the leaf-bases not broad and spongy; inflorescences less than 2 m long. 13 13. Leaves palm-like, tough, often with a spiny stalk. leaf-segments with parallel venation, usually spine-tipped. 14 - Leaves not palm-like, without a spiny stalk; leafblade not divided or if divided then the seg­ ments with pinnate venation, not spine-tipped 15 14. Midrib of leaf more or less flattened in crosssection, ieaf curled up in bud from apex to base like a fern leaf; leaflets very stiff, with veins that branch into 2; reproductive stmctures in cones or on a leaf-like part, young cones not enclosed in a woody boat-shaped structure or in a stiff bract. (1) Cycadaceae1 -j Midrib of leaf triangular in cross-section or a midrib absent and the leaf fan- or V-shaped, leaf folded as a V or O in bud. leaflets or blade stiff but flexible with mostly parallel veins: flowers usually on branched inflorescences, young inflorescence enclosed in a woody boat­ shaped structure or a stiff bract. [6] 214. (Palmae) Arecaceae

1Encephalartos and C.ycas are found in East Africa and fossil pollen of Encephalartos has been found in southern Ethiopia.

15. Flowers never present, reproductive structures usual 1> brown patches or lines on the underside of leaves producing minute powdery spores; petiole usually with scale-like hairs; young leaves curled up with the tip innermost (ferns). 111 T h e F erns - Flowers and/or fruits and seeds usually present; petiole without scalc-like hairs; leaves never curled up with the tip innermost when young. 16 16 Plants growing in wet places, the petiole attached near the centre of the blade, leaves round in outline, or if otherwise then flowers and fruits absent 17 - Plants growing in dry or moderately wet places, if growing in wet places (swamps and lake or pond edges) then the petiole attached on the edge of the blade and not near the centre. 19 17. Leaves usually arrow-shaped, with a deep notch where the petiole is attached to the blade; flowers (if present) enclosed in a tubular bract. (6] 181. Araceae - Leaves round in outline, flowers not enclosed in a tubular bract. 18 18. Leaves completely round, without a notch at one side and with an entire edge; flowers >4 cm across, on long narrow stalks at the surface of the water fNetumbo) [2,1 j 9. Nymphaeaeeae - Leaves with a notch on one side or with a round and lobed edge; flowers 2 cm long; herbs or shrubs. 23 - Flowers with 2 or more anthers or pollen sacs, or if each flower has only 1 anther, then the flowers very small (2 cm long, not tightly crowded together; each ovary with many ovules. [2,2] 67. Caricaceae - Inflorescence a fig; flowers >13&14

37

150. SPHENOCLEACEAE by M. Thulin* Airy Shaw, S phenocleaceae in FI. Trop. E. Afr. 3 pp. (1968); Thulin, S phenocleaceae in FI. Zamb. 7(1): 114-117 ' 1983) & S phenocleaceae in FI. Somalia 3: 258-259 (2006). \nnual. glabrous marsh herbs. Leaves alternate, simple, entire, without stipules. Inflorescences term inal, dense, spicate. Flowers bisexual, regular, subtended by a bract and 2 bracteoles. Calyx-lobes 5, shortly joined, persistent. Corolla bell- to um-shaped, soon falling; lobes 5. Stamens 5, inserted on corolla-tube, al­ ternating with the petals; filaments very short; anthers rounded, dehiscing longitudinally. Ovary half-inferior, 2 -locular; ovules numerous on large stipitate axile placentas; style 1 , short, glabrous, with a capitate, slightly 2-lobed stigma. Fruit a circumscissile capsule, obovate-obconic, 2 -locular, membranous. Seeds numerous, minute; testa irregularly ribbed. A single almost pantropical genus, represented by 1 species in the Flora area. The family has often been associated with Campanulaceae, but molecular data indicate a position in the Solanales.

tudinally to the rhachis by a linear base. Calyx-lobes broadly triangular, rounded at apex, 1-1.5 mm long. Co­ Characters as for the family. rolla white to greenish-white or pink, 2-3 mm long; Genus of 2 species, the second one, S. dalzielii N.E. lobes ovate-triangular, united about half-way. Stamens: Br., is confined to West Africa. filaments slightly dilated at base; anthers c 0.5 mm long. Capsule 4-5 mm across, dehiscing below the calyxS. zeylanica Gaertn. (1788) lobes, which fall with the lid, leaving the scarious base type: from Sri Lanka, collector unknown (L persistent on the rhachis. Seeds yellowish-brown, ob­ holo.). long, c 0.5 mm long. Fig. 150.1. Herb, up to 1.5 m tall; stem hollow, often muchWet places, such as in and near pools or in temporar­ branched. Leaf-blade lanceolate to narrowly ovate or el­ ily flooded areas; 500-600 m. IL; widespread in tropical liptic, up to 12 x 5 cm, base attenuate, apex acute or Africa and also in tropical Asia and America (probably subacute; petiole up to c 1.5 cm long. Spikes cylindricintroduced). Friis et al. 7196; Gilbert & Friis 8356; Mesconical, up to 12 cm long, but usually much shorter, c 1 fin T. 6649. cm across, with only a few flowers open at a time; First collected from the Flora area in 1986, this plant peduncle up to 8 cm long; bracts and bracteoles obis quite common in the swampy lowlands bordering the lanceolate-spathulate, the tips arched over the flowers Sudan. except at anthesis. Flowers sessile, attached longi­ SPHENOCLEA Gaertn. (1788), nom. cons.

* Departm ent o f System atic Botany, Evolutionary Biology Centre (EBC), Uppsala University. Norbyvagen 18D. SE-752 36 Uppsala, Sweden

38

150. SPHENOCLEACEAE. Sphenoclea

Figure 150 I SPH ENO CLEA Z E \L s\N IC A . 1 - habit x %; 2 - part o f flowering stem x ens which also occurs in Soma­ lia, Kenya, Tanzania, Zimbabwe, Mozambique. Comoro Is., Madagascar and Reunion. Subsp. recurvata (E. Wimm.) Thulin, with calyx-lobes recurved in fruit, is known from E Africa. Dem. Rep. Congo, Burundi. Zam­ bia, Zimbabwe, Mozambique and Angola. IL L . chireensis A. Rich. (1850) - type: TU, Chire, Quartin-Dillon & Petit s.n. (P holo.). Annual, more or less erect herb. 8-30 cm tall: stem tri­ angular and broadly winged, glabrous or sparsely hairy. Leaves suborbicular to elliptic at the base of the plant, gradually narrowing upwards, 7-16 x 2-5 m m denticulate, glabrous or with a few short hairs. Flowers in lax leafy racemes; pedicels 4-7 mm long, shorter than or equalling the bracts. Calyx-lobes 1.4-2.8 mm long, entire. Corolla 3.5-4 5 mm long, blue. Stamens with an­ ther-tube c 0.8 mm long, sparsely and shortly pubescent on the back; 2 lower anthers each with a tuft of hairs and a hyaline appendage at the apex. Capsule with lower part 2-2.5 mm long, and valves c 1 mm long. Seeds somewhat compressed, c 0.4 nun long; testa almost smooth. Moist places at intermediate altitude. TU; Tanzania Burundi, Zambia, Zimbabwe. In the Flora area only known from the type. 12. L. heyneana Roem. & Schultes (1819) - type: from “ India orientalis” Heyne s.n. (B holo.). L. umbrosa Hochst. ex Hemsl. (1877); L. trialata D. Don var. umbrosa (Hochst. ex Hemsl.) Chiov., in Result. Sci. Miss. Stef.-Paoli, Coll. Bot.: 109 (1916) type. TU, Schimper 11.779 (K holo., B BM FI G MO P S W iso.). Annual, erect or ascending herb, 5-25 cm tall; stem tri­ angular and broadly winged, glabrous. Leaves ovate to suborbicular, gradually narrowing upwards, up to 13-35 x 8-20 mm, usually rather bluntly and shallowly dentate-crenate, glabrous. Flowers in lax leafy racemes; pedicels up to 10-20 mm long, shorter or longer than the bracts. Calyx-lobes 1.2-3.2 mm long, entire. Corolla

3.5-5.2 mm long, pink, pale mauve or pale blue. Sta­ mens with anther-tube 0.6-1 mm long, pubescent or i glabrous on the back; 2 lower anthers each w ith a tuft of liairs and a hyaline appendage at the apex. Capsule with lower part 2.5-5 mm long and valves c 1 mm long. Seeds compressed, 0 4 -0 .5 mm long, testa almost smooth. Grassland, moist places; 500-1650 m. EE EW TU GD; Tanzania, Deni Rep. Congo. Zambia; Oman and from India east to China. Pappi 4478; Fiori 1726; J. De Wilde & Gilbert 292

y

13 L. inconspicua Rich. (1850); L. heyneana Roem. & Schultes var. inconspicua (A R ich) E. Wimm., in Engl. Pflanzenr. IV.276b: 475 (1953)-ty p e: TU, Ouartin-Dillon & Petit s.n. (P holo.). / Annual, erect or ascending herb, 3—15(—35) cm tall; j stem triangular and broadly winged, sparsely to densely hirsute. Leaves broadly ovate to elliptic, gradually nar­ rowing upwards, up to 7-20 x 6-12 mm. serrate or den­ tate with usually acute teeth, sparsely to densely pubescent. Flowers in lax leafy racemes; pedicels up to 5—10(—13) mm long, equalling or shorter than the bracts. Calyx lobes 1.6-3.2 mm long, with 1-2 pairs of teeth al the margin. Corolla 2.8-4 mm long, pale blue and/or white Stamens with anther-tube 0.6-0.8 mm long, glab­ rous or with a few short hairs on the back; 2 lower an­ thers each with a tuft o f hairs and a hyaline appendage at the apex. Capsule with lower part 3-5 mm long and valves c 1 mm long. Seeds compressed, 0.4-0 5 mm long: testa almost smooth. Upland grassland and woodland, roadsides, ditches, old cultivations; 1300-2500 m. TU GD GJ SU AR WG GG SD; S Sudan, E Africa, Nigeria, Cameroun. Dem Rep Congo, Burundi, Malawi; India. Friis et al. 7754; Thulin 1290; Gilbert & Thulin 833. 14. L. exilis Hochst. e \ A. Rich. (1850) ty pes TU, Quartin-Dillon s.n. (P syn.): Schim­ per II: 1182 (P syn., FI FI-W G K MO S W isosyn.). L. tenerrima Chiov. (1911) - type: GD. Asoso, Chiovenda 2012 (FT holo.). L. exilis Hochst ex A. Rich. var. pusilla E. Wimm., in Annal. Naturhist. Mus. Wien 56: 347 (1948) - types: EW, Pappi 764 and GD, Chiovenda 1613, 1655, 1687. 1857 (all FT syn.). L. exilis Hochst. ex A. Rich. var. major E. Wimm., in Annal. Naturhist. Mus. Wien 56: 347 (1948) types: GD, Dschan-Meda. Schimper 1382 (B syn., BM K isosyn.); GD. Gaffat, Schimper s.n. (B syn ). Annual herb, 2-20 cm tall; stem angular, glabrous. Leaves lanceolate to elliptic or broadly ovate. 1.5-20 x 0.5—7 mm. serrate to subentire, glabrous. Flowers few to solitary in upper leaf-axils; pedicels 5-30 mm long, much longer than the bracts. Calyx-lobes 0.5-1 mm long, entire, glabrous or ciliate. Corolla 2.8-9 mm long, blue, mauve or white w ith dark markings. Stamens with anther-tube 0.6-1.6 mm long, papillose to shortly pu­ bescent on the back; 2 lower anthers each with a tuft of hairs and a hyaline appendage at the apex. Capsule with

151. LOBELIACEAE: 1. Lobelia 45

Figure 151.3 LO B E LIA IIOLST11: 1 - habit x 2/3; 2 - flower with bract and bracteoles x 4; 3 - corolla opened out x 4; 4 - stamens with anther-tube x 8; 5 - anther-tube from beiow x 12; 6 anther-tube opened out x 12; 7 - style and longitudinal section through the ovary from a young flower x 8; 8 style-apex with immature stigmas from young flower x 16; 9 -sty le with open stigmas from an older flower x 8; 10 - style-apex with receptive stig­ mas from an older flower x 16; 11 capsule x 4; 12 - seed x 20; 13 - trans­ verse section through a seed x 20. 1 from Thulin 315; 2-10 from Drum­ mond & Hemsley 1512; 11-13 from Thulin & Mhoro 1108. Drawn by Maureen Church. (Reproduced with permission from FI. Trop. E. Afr. Lobeliaceae: fig. 3.)

lower part 2-4.5 mm long and valves 1-1.5 mm long. Seeds prominently ridged, 0.4-0.6 mm long. Moist places in grassland, bogs, stream-sides. often on open ground; 1300-2730 m. EW TU GD WG SU AR IL; not known elsewhere. Gilbert & Thulin 694; Thulin 1446, 1637. All plants seen from WG and IL are comparatively large and originate from altitudes near the lower part of the range, whereas the highland populations show a re­ markable variation from fairly tall plants with compara­ tively large blue flowers to dwarf plants with minute whitish, often single flowers.

15. L. scebclii Chiov. (1929) type: BA, source of Uebi Scebeli, Basile 107 (TO holo., not seen). Perennial, prostrate or decumbent herb; stems 5-30 cm long, rhizomatous and rooting at least at the lower nodes, glabrous or sparsely pubescent. Leaves with blade very broadly ovate to suborbicular, 6-19 x 6-16 mm. subcordate to truncate or broadly cuneate at the base, distinctly petiolate, dentate-crenate, sparsely pu­ bescent on both surfaces. Flowers scattered in leafaxils; pedicels 9-40 mm long, much longer than leaves. Calyx-lobes 3-5 mm long, entire, glabrous or sparsely ciliate. Corolla 10-12 mm long, blue. Sta­

151. LOBELIACEAE. 1 Lobelia, 2 Monopsis 46

mens with anther-tube 1.5-2 mm long, pubescent on the back; 2 lower anthers each with a tuft of hairs and a hyaline appendage at the apex. Capsule with valves 2-2.5 mm long, about as long as or usually shorter than the lower part of the capsule. Seeds somewhat com­ pressed. 0.7-0.8 mm long, testa almost smooth. Montane grassland and scrub, forest margins; 1600-3100 m. AR BA SD; not known elsewhere. Moo­ ney 7194, 8198; Thulin 1465. 16. L. erlangeriana Engl. (1902) - type; BA, Ladjo, Ellenbeck 1899 (B holo. de­ stroyed). Annual, prostrate to ascending herb; stems 2-10 cm long, often rooting at the lower nodes, glabrous or sparsely pubescent. Leaves w'ith blade more or less broadly ovate to suborbicular. 4 -1 1 x 4 -1 1 mm. broadly cuneate at the base, distinctly petiolate, more or less ob­ scurely dentate-crenate mainly in apical part, glabrous to sparsely ciliate or sparsely pubescent on upper sur­ face only. Flowers in upper leaf-axils; pedicels (3-) 10-30 mm long, usually much longer than leaves. Calyx-lobes 2-3.5 mm long, entire, more or less sparsely ciliate. Corolla 6.5-8.5 mm long, blue. Stamens with anther-tube 1-1.5 mm long, shortly pubescent on the back; 2 lower anthers each with a tuft of hairs and a hyaline appendage at the apex. Capsule with valves 1.5-2.5 mm long, about as long as or shorter than the lower part of the capsule. Seeds somewhat compressed, c 0.6 nun long; testa almost smooth. Afro-alpine grassland, forest margins, often in rocky or moist places; 3200-4100 m. AR BA; not known else­ where. Hedberg 5518; Mooney 7133; Thulin et al. 3702. The type of L. erlangeriana is lost, but the illustra­ tion in Engl.. Pflanzenwelt Afr. 1(1); Fig. 125 (1910) is good enough for an almost certain identification. How­ ever. in the original description the pedicels are said to be sparsely pilose, while in the plants seen for this treat­ ment the pedicels always seem to be glabrous. 17. L. tripartita Thulin (1991) - type; BA, N of Dolo Menna on road to Goba, Gilbert & Ermias D. 8476 (K holo., ETH UPS iso.). Annual or perhaps short-lived perennial, mat-forming herb; stems more or less prostrate, 5-10 cm long, root­ ing at least at the lower nodes, glabrous. Leaves more or less orbicular to triangular in outline. 4-7 x 5-8 mm, di­ vided into usually 3 lobes which are themselves more or less lobed or dentate, more or less sparsely ciliolate, dis­ tinctly petiolate. Flowers in upper leaf-axils; pedicels 10—17(—28) mm long, much longer than the leaves. Ca­ lyx lobes 1.6-2.8 mm long, entire, more or less sparsely ciliate. Corolla 6-7 mm long, blue with white centre. Stamens with anther-tube c 1.2-1.6 mm long, shortly pubescent on the back; 2 lower anthers each with a tuft of iiairs and a hyaline appendage at the apex. Capsule with valves 2-2.5 mm long, about as long as or shorter than the lower part of the capsule. Seeds somewhat com­ pressed, c. 0.5 mm long; testa almost smooth.

Open ground or sides of ditches and earth banks along roads through upland forest; 2100-2300 m. BA; not known elsewhere. Friis et al. 3570. 18. L. holstii Engl. (1894) - types; from Kenya Hildebrandt 2463 (B syn., BM K W isosyn.), and Tanzania, Holst 20 (B syn.) & Holst 8960 (B syn . BM COI G K S W isosyn ). L. scioensis Chiov. (1912) - type; SU. Mt Managasha, Negri 494 (FT holo.). Perennial herb. 10-50 cm tall; stems ribbed, glabrous to more or less sparsely appressed pubescent. Leaves ob­ lanceolate to spathulate, up to 12-45 x 3—9(—12) mm, glabrous or sparsely pubescent, crenate-serrate. Flowers in lax 2 -1 3-flowered pedunculate racemes; pedicels 3—15(—27) mm long. Calyx-lobes 2-4 mm long, entire. Corolla (11-)13-18 mm long, pink, mauve or purple. Stamens with anther-tube 2-2.4 mm long, all anthers densely hairy at the apex, otherwise glabrous or rarely sparsely pilose on the back. Capsule with valves 2-3.5 mm long. Seeds trigonous. 0.8-1 mm long, dark brown. Fig. 151.3. Montane grassland and scrub, forest edges, rocky or disturbed places; 1500-2800 m. SU AR SD, Kenya, Tanzania E Dem. Rep Congo, Rwanda, Burundi Ash 2303; Mooney 7932; Thulin et al. 3372. 2. M ONOPSIS Salisb. (1817) Thulin in Boi. Noliser 132 131-137(1979).

Annual or perennial herbs. Leaves alternate, opposite or verticillate. Flowers usually not resupinate, in racemes or solitary in leaf-axils; bracteoles conspicuous resem­ bling small leaves (in Flora area). Calyx 5-lobed. Co­ rolla split to the base on one side, 2-lipped to subregular. Stamens with filaments free from corolla or attached to it at the base; anthers united into a tube, ail of them with a tuft of hairs at the tip, the two lateral ones somewhat longer than the others and with shorter hairs at the tip. Ovary inferior or almost so, 2-locular, stigma-lobes lin­ ear, becoming revolute and with a ring of hairs present on the style below them. Fruit dehiscing by 2 apical valves or more or less indehiscent; seeds numerous, usu­ ally reticulate. Some 20 species, mainly in South Africa; 1 in the Flora area M. stellarioide.s (Presl) Vrb. (1881) - type; from South Africa, Krebs s.n. (PR holo., B iso). Annual or perennial decumbent hert>; stems 5-60 cm long, with harsh retrorse pubescence. Leaves opposite, linear to elliptic, 10-35 x 2-9 mm, crenate to serrate, usually with a harsh pubescence on both surfaces. Flow­ ers not resupinate. solitary, axillary; pedicels 8-35 mm long, pubescent with retrorse hairs, bracteoles at the base of the pedicel, 6-16 mm long, resembling small leaves. Calyx-lobes 2-6 mm long. Corolla 6.5-11.5 mm long, dirty yellow, dull pink, brownish purple or violet, 2-lipped with the upper lip 3-lobed and the lower lip consisting of 2 spathulate lobes. Filaments free from co-

151. LOBELIACEAE: 2. Monopsis

47

ipetala Thulin is a smallflowered annual restricted to Kenya and Tanzania.

Some 300 species, widely distributed in the northern hemisphere, especially abundant in the Mediterranean region and the Middle East. Many species are of horti­ cultural value. Ragazzi 115 (FT) from Ghinda (EE-EW) is a fragment probably representing C. rapunculus L. This species is widespread in Europe, N Africa and SW Asia and may have been occasionally introduced in Ethiopia.

10. W. hirsuta (Edgew.) Tuyn (1960); Cephalostigma hirsutum Edgew. (1846) - type: from India, Edgeworth 252 (K holo.). Cephalostigma erectum (Roem. & Schultes) Vatke var. coeruleum Chiov., 'mAnn. Bot. Roma 9; 79 (1911) - type: GD, Dembia near Asoso, Chiovenda 2015 (FT lecto.). Annual herb, 4-30 cm tall, with a very short but usually dense pubescence, often mixed with longer hairs. Leaves narrowly oblanceolate to obovate or broadly el­ liptic to ovate, 10-55 x 4-25 mm, margins undulatedentate. Inflorescence lax; pedicels 5-20 mm long. Calyx-lobes 1—3(—5) mm long. Corolla with almost free petals. 1.5-4 mm long, blue to pale blue or whitish. An­ thers 0.5-1 mm long. Ovary 3-locular, semi-inferior; style about as long as or longer than the corolla, glab­ rous or hairy below, subcapitate with 3 short lobes at the apex. Capsule 5-nerved, 3-valved. Seeds trigonous. Fig. 152.4.1-5 Grassland or woodland, rocky places, roadsides, waste or cultivated ground; 1200-1700 m. TU GD GJ SU WG GG SD; widespread in tropical Africa, Mada­ gascar Yemen, India and Nepal. J. De Wilde & Gilbert 293; Gilbert & Thulin 621, 734. 11. W; hookeri (C. B. Cl.) Tuyn (1960); Cephalostigma hookeri C. B. Cl. (1881) - type: from India, Clarke 24796 (K lecto., BM iso.). Annual herb, 4-25 cm tall, hairy at the base. Leaves tending to be verticillate at the middle of the stem, lan­ ceolate to narrowly ovate, 8-35(^18) x 3—10(—13) mm, margins shallowly undulate-dentate. Inflorescence lax; pedicels up to 7-20(-25) mm long, glabrous. Ca­ lyx-lobes 0.5-1.2 mm long. Corolla with almost free petals 1.2-2 mm long, blue to pale violet. Anthers c 0.4 mm long. Ovary 2-locular, subinferior; style about as long as corolla, glabrous below, subcapitate with 2 very short lobes at the apex. Capsule 5-nerved, 2-valved. Seeds trigonous. Fig. 152.2.10 & 11. Grassland or woodland, shady places; 1300-1700 m. WG; Kenya, Tanzania, Cameroun, Dem. Rep. Congo; India. Thailand, Java. Gilbert & Thulin 622, 749.

3. CAMPANULA L. (1753) Annual or perennial herbs, rarely subshrubs. Leaves al­ ternate, simple. Inflorescences panicle-, raceme-, spike-, or head-like, or flowers solitary. Calyx-lobes usually 5, sometimes with reflexed appendages in between them. Corolla campanulate to almost rotate or cylindrical,

1. Plant perennial; calyx-lobes with reflexed append­ ages in between them. 1. C. edulis - Plant annual; calyx-lobes without appendages. 2 2. Stem dichotomously branched; flowers subsessile, with 3-5-lobed bracts; calyx-lobes stellately spreading in fruit. 2. C. erinus -S te m not dichotomously branched; most flowers distinctly pedicelled, with dentate-serrate bracts; calyx-lobes ± erect in fruit. 3. C. dim orphantha

1. C. edulis Forssk. (1775) - type: from Yemen, Forsskal s.n. (C lecto.). C. esculenta A. Rich. (1850); C. rigidipila var. esculenta (A. Rich.) Di Capua, 'mAnn. 1st. Bot. Roma 8: 236 (1904) - type: TU, Oudgerate, Quartin Dillon & Petit s.n. (P lecto.). C. quartiniana A. Rich. (1850); C. schimperi var. quartiniana (A. Rich.) Vatke, in Linnaea 40: 201 (1876), nom. illegit.; C. rigidipila var. quartiniana (A. Rich.) Engl, in Abh. Preuss. Akad. Wiss. 1891; 410 (1892) - type: TU, Memsah, Quartin Dillon & Petit s.n. (P holo.). C. rigidipila Steud & Hochst. ex A. Rich. (1850); C. schimperi Vatke (1874), nom. illegit.; C. schimperi var. rigidipila (Steud. & Hochst. ex A. Rich.) Vatke, in Linnaea 40: 201 (1876), nom. illegit. - type: TU, Oudgerate, Quartin Dillon & Pe­ tit s.n. (P lecto.). C. sarmentosa Hochst. ex A. Rich. (1850); C. schimperi var. sarmentosa (A. Rich.) Vatke, in Lin­ naea 40: 201 (1876), nom. illegit.; C. rigidipila var. sarmentosa (Hochst. ex A. Rich.) Engl., in Abh. Preuss. Akad. Wiss. 1891: 410 (1892) - type: SU, ‘Choa’, Quartin Dillon & Petit s.n. (P lecto.). Perennial; stems 6-70 cm long, decumbent to erect, hairy. Leaves sessile, elliptic to oblanceolate, 10-50 x 3-16 mm, hairy especially on the veins; margin entire to more or less sparsely and obscurely dentate. Inflores­ cence lax, more or less leafy. Calyx-lobes 3.5-12 mm long, hairy with triangular reflexed appendages. Corolla blue to mauve or purple, often with white base, rarely all white, 7-25(-30) mm long, often pilose on the veins, otherwise glabrous. Anthers 1.5-4 mm long. Ovary 3(-5)-locular, style shorter than corolla. Capsule more or less nodding, dehiscing by small basal valves. Fig. 152.5.

152. CAMPANULACEAE: 3. Campanula

Figure 152.5 CAMPANULA EDULIS large-flowered form: 1 - flowering plant x 2/3; 2 - flower, 2 petals re­ moved x 114; 3 - stamen x 6; 4 - flower with corolla and stamens removed x 2 ; 5 - capsule after dehiscence x 3; 6 seed x 24. 1 from Eggeling 2881 & fferlocker 136; 2 -4 from Eggeling 2881; 5 & 6 from Mabberley 610 Drawn by Victoria Goaman. (Repro­ duced with permission from FI. Trop. E Afr Cam panulaceae: fig. 7.)

Upland grassland, forest glades, open ground in mon­ tane evergreen bushland. often on rocky ground, also in pockets on cliffs; (1000-) 1500-3 800 m. EW TU GD GJ WU SU AR GG SD BA HA; Sudan, Djibouti, Somalia E Africa Rwanda; Arabia. Thulin 1367; Gilbert & Thulin 479; Mooney 8101.

hairy, first erect, about as long as the corolla, enlarging and becoming stellately spreading after anthesis. An­ thers c 0.8-1.2 mm long Corolla pale blue to whitish, cylindrical, 3-5 mm long, with some hairs on the veins and at the base. Ovary 3-locular, style shorter than co­ rolla. Capsule nodding, dehiscing by small basal valves.

2. C. erinus L. (1753) - ty pe: from Europe.

Shady places, old walls; 1300-2400 m. EW TU GD; Mediterranean region, SW Asia, Sudan. Friis et al. 6835; Schimper 1231; Ryding & Sileshi N. 1875.

Annual. 5-25 cm tall, dichotomously branched, hairy. Leaves elliptic to ovate or obovate, 7-20 x 4—10(—15) mm. margins crenate-dentate, the upper leaves 3-5lobcd and more or less opposite. Flowers terminal and axillary, subsessile. Calyx-lobes triangular-lanceolate.

Although collected by Schimper in 1853, C. erinus was not included in Cufodontis (loc. cit.). The nearest lo­ calities outside the Flora area are in the Red Sea Hills in Sudan.

152. CAMPANULACEAE: 3. Campanula, 4. Legousia 57

3. C. dim orphantha Schweinf. (1867) type: Sudan. Nubia, Ehrenberg s.n. (B syn. de­ stroyed). Annual, 5-60 cm tall, erect, densely hairy. Leaves ellip­ tic to oblanceolate or the lower ones spathulate, 20-70 x 3-20 mm, margins dentate-serrate. Inflorescence pa­ niculate; pedicels 1-16 mm long; flowers of two types, normal and cleistogainous, sometimes only cleistogamous ones present. Normal flowers: calyx lobes nar­ rowly triangular, hairy, erect, about as long as the coroila: anthers c 2 mm long; corolla pale blue, cylindri­ cal. 4-7 mm long, hairy; ovaiy 3(-4)-locular: style about as long as the corolla or somewhat shorter. Cleistogamous flowers with calyx-lobes, corolla, stamens and style much reduced. Capsule erect or somewhat nodding, dehiscing by small basal valves. Moist places; 1000-2400 m. EW; Sudan, Egypt; S Asia from Afghanistan and Pakistan to China. Terracciano & Pappi 360, 489; Fiori 1712.

Figure 152.6 LEGOUSIA FALCATA. 1 - part of flowering stem x 1; 2 - flower x 2. Both from Ryding & Sileshi N. 1899. Drawn by Yosef Berta.

For the nomenclature of this species, see Thulin in Nordic J. Bot. 7: 419-420 (1987). 4. LEGOUSIA Dur. (1782) Annual herbs. Leaves alternate, simple. Inflorescences panicle-or spike-like, or flowers solitary. Calyx-lobes 5, without appendages. Corolla broadly bell-shaped to disc-sliaped, 5-lobed. Stamens 5, free; Filaments not or scarcely broadened at the base. Ovaiy inferior, more or less cylindrical, (2-)3-locular; style without glands, lobes (2-)3. Capsule dehiscing by lateral valves near the apex. Seeds numerous. About 12 species, mainly in N America, the Mediter­ ranean region and SW Asia; 1 in the Flora area. The genus is a segregate from Campanula of doubt­ ful status and circumscription. McVaugh in Wrightia 1: 28 (1945) placed L. falcata in Triodanis Raf. along with seven American species.

L. falcata (Ten.) Fritsch (1907); Prismatocarpus falcatus Ten. (1811-1815); Specularia falcata (Ten.) A. DC. (1830); Triodanis falcata (Ten.) McVaugh (1945) - type: from Europe. Annual, 10-50 cm tall, somewhat hairy . Leaves ovate to obovate, 15-40 x 7-20 mm, margins crenulate, the lower petiolate. Flowers solitary or in pairs in the leafaxils, sessile, forming a lax spike; flowers at lower fer­ tile nodes often cleistogainous. Calyx-lobes linearlanceolate. about as long as ovary, fruiting calyx enlarged with spreading to recurved lobes. Corolla vio­ let. shorter than calyx-lobes. Capsule 10-20 mm long, often slightly curved. Fig. 152.6. Shady places, dry watercourses; 1350-2200 m. EW; Mediterranean region, SW Asia. Pappi 192, 237; Ryding & Sileshi N. 1899. This is the first record of the genus for tropical Africa.

58

153. GOODENIACEAE by Sebsebe D em issew & Sue Edwards*

Davies, G oodeniaceae in FI. Trop. E. Afr. : 4 pp. (1978); Thulin, G oodeniaceae in FI. Somalia 3: 462-464 (2006). Trees, shm bs or herbs. Leaves alternate, rarely opposite, o r radical, sim ple; blade often obovate to spathulate. w ithout stipules. Inflorescence cym ose or reduced to solitary axillary flow ers. Flow ers bisexual w ith stam ens m aturing first, irregular. C alyx united to the base o f the ovary, upper part free, 5-toothed o r entire. C orolla tube long, split for part or all o f its length, upper part divided into 5 subequal lobes. Stam ens 5, attached to the base of the corolla, free, or anthers occasionally joined together Ovary superior to inferior, 1-2-locular, each containing 1 or several ovules; style em erging laterally from the split corolla tube, bearing a cup-shaped indusium below the sim ple o r bifid stigm a. Fruit a capsule or dm pe. A fam ily o f 14 genera and m ore than 300 species, m ostly in A ustralia; 1 genus, S ca e vo la , w ith 2 species in E and NE A frica, both o f wrhich grow in coastal areas on sand or coral rocks. S. sericea Vahl (1791) is a shrub or sm all tree 2 -3 ( -4 ) m tall characterized by w hitish fm its and leaf-axiJs with dense silky hairs It occurs in coastal areas o f K enya and T anzania (Z anzibar and P em ba Islands), and from South A frica to India and A ustralia. Fig. 153.1.1-4. S. p lu m ieri (L.) V ahl (1791) is a m uch-branched shrub or sm all tree characterized by blue-black fn iits and leaf-axils glabrous or with sparse hairs. It is found in Som alia, Kenya, T anzania (and Z anzibar) as w ell as in the Cape Province o f South A frica, and extends to W est A frica and tropical A m erica. Fig. 153.1.5 & 6. One or both o f these species m ight occur in EE on the coast o f the Flora area.

Figure 153.1 SCAEVO LA SERICEA l l - flowering branch x Vy, 2 - flower x 2; 3 - calyx and pistil x 3; 4 - fruits x 1 S. P LU M IE R I 5 - flower x 2; 6 truit x 1. 1-4, from Faulkner 2681; 5 & 6, from Faulkner 2289. Drawn by Maureen Church. (Modified and re­ produced with permission from F i Trop. E. Afr. Goodeniaceae: fig. 1.)

* T h e N atio n al H e rb a riu m , A d d is A b a b a U n iv e rsity , P.O . B o x 3 4 3 4 , A d d is A b ab a, E th io p ia.

59

154. MENYANTHACEAE Sebsebe Demissew & Sue Edwards* Mackinder. M enyanthaceae in FI. Zamb. 7(4): 51-56 (1990); Thulin, M enyanthaceae in FI. Somalia 3: 462 (2006). Aquatic herbs, w'ith clumped rootstock, or horizontal creeping rhizomes or stolons. Leaves floating or sub­ merged. alternate or in a rosette, simple or compound, entire or crenulate; stipules inconspicuous, rounded, scarious. Flowers in panicles, racemes, fascicles or solitary; regular, generally 5-merous. hermaphrodite or unisexual. Sepals free or almost so. Corolla united, often w'ith a short tube, lobes membranous, often fringed or hairy. Stamens the same number as corolla lobes and alternating with them; anthers opening by longitudi­ nal slits. Ovary superior, 1-locular. Fruit a capsule, dehiscent or not. Seeds often numerous. 5 genera with 40-50 species worldwide; 1 genus w'ith 9 species in tropical Africa, 2 o f which are recorded from the Flora area. This family is sometimes included in the Gentianaceae. but Cronquist, in A n In teg ra ted System o f C lassi­ fication o f F low ering P la n ts : 900-903 (1981), gives a number o f micromorphological and chemical charac­ ters that suggest the M enyanthaceae is a distinctive family, possibly more closely related to the Solanales than the Gentianales. NYMPHOIDES Seguier (1754) A Raynal \x\K4itl. Bot. Staaiss. Munchen 10: 122-134 (1971); in Adansotiia, ser. 2, 14: 227-270. 405^158 (1974); Menyanthaeceae in FI. Afr. Central. 1-16 (1975).

Aquatic herbs, annual or perennial. Leaves in a rosette, or alternate along floating stolons, leaf-blade simple, circular to ovate. Flowers 5(8)-merous, sometimes heterosty lous, in clusters at the nodes, pedicellate, sub­ tended bv a small bract. Capsule developing under wa­ ter, splitting into valves from base or apex. Seeds round to oval, flattened or convex, smooth to tuberculate. A worldwide genus with 9 species in tropical Africa; 2 in the Flora area. Tw o or more species may be found growing together It is possible that there may be other species and that the genus is more widespread than recorded from this ac­ count, as collectors are often reluctant to get into water and sw ampy areas. The flowers are very delicate and easily damaged when making herbarium collections. Collectors should make careful notes of flower colour and hair arrange­ ment, and put the flowers between tissue paper. Pre­ serving flowers in spirit is also helpful. 1. Corolla lobes 7-9 mm long; capsule subglobose. 5-8 mm wide, as long as or longer than calyx. 1. N. brevipedicellata - Corolla lobes 14-18 mm long; capsule ovoid. 4-5 mm wide, shorter than calyx. 2. N. indica 1. N. brevipedicellata (V'atke) A. Raynal (1971); Limnanthemum brevipedicellatum Vatke (1876) ty pe: GD/GJ, Galamotch towards Lake Tana. L. abyssinicum N.E. Br. (1904) - type: Ethiopia, “Abyssinia” without precise locality, Schimper 1358. Nymphoides indica sensu Marais &Verdoom in FI. S. Afr. 26: 243 (1963) p.p. *

T h e N a tio n al H e rb a riu m . A d d is A b a b a U n iv e rs ity , P.O . B ox 3 434, A d d is A b ab a, E th io p ia.

Perennial with numerous floating stolons, 3-5 mm thick, from short thick rootstock. Leaf-blade subcircular. or larger ones somewhat ovate-circular, 6-10 x 8-10 cm; petiole 1.2—1.5(— 4) cm long. Flowers homosty lous; 6-25 clustered at the base o f the petiole; pedi­ cels 2-6 cm long. Calyx 5-6 nun long, lobes appressed to corolla tube. Corolla tube yellow. 7-9 mm long; lobes white, densely and uniformly hairy inside. 7-9 mm long. Capsule 5-8 mm wide, subglobose. as long as the calyx. 15-80- seeded. Seeds 1.6-2 x 1.4-1.7 mm, lentic­ ular to tuberculate. Marshy areas with permanent or semi-permanent pools, rivers with grasses and sedges; c 1800 m. GD GJ; widespread in tropical Africa west to Cameroon and south to S Africa. Gilbert & Getachew A. 3037. 2. N. indica (L.) O. Kuntze (1891) - types: from India and Sri Lanka. subsp. occidentalism. Raynal inAdansonia. Ser. 2, 14(3): 8, t. 18 (1974) - type: from Cameroon. Perennial, with numerous floating stolons from short thick rootstock; stems and petioles 4-8 nun wide. Leaf-blade circular, 5—15(—25) cm long and wide; peti­ ole 1-6 cm long. Flowers heterosty lous; 10-30, clus­ tered at base of petiole, pedicels 3.5—5(—10) cm long. Calyx 5-9 mm long. Corolla tube yellow, 7-9 nun long; lobes white, densely and uniformly hairy inside, 14-18 mm long. Capsule 4-5 mm wide, ovoid, shorter than ca­ lyx, 10- or more seeded. Seeds 1.5—2.2 mm long, lentic­ ular to more or less tuberculate. Fig. 154.1. Marshy areas and permanent to semi-permanent rivers, pools and lakes; c 1750 m. GD GJ; widely distrib­ uted in tropical Africa. Getachew A. 2705. The species as a whole is pantropical. It is very vari­ able in appearance as it can tolerate changes in water lev­ els and even temporary drying up of pools. It reproduces very effectively by vegetative means and can become a weed of waterways.

60

154. M ENYANTHACEAE: Nvmphoides

Figure 154.1 W M P H O ID E S im iC A subsp OCCID ENTALIS. 1 - habit x X i 2 inflorescence, flowers carried above water, capsules below it x 2/ 3; 3 flower x 214; 4 & 5 corolla opened oui to show short- and long-styled flower, respectively, x 5; 6 - capsule wilh calyx x 6; 7 - seed x 6; 1, 2 & 5 from Biegel & Russell 3711, 3 & 4 from Bullock 2162, 6 from Symoens 11196. 7 from Killick & Leistner 3228 Drawn by Eleanor Catherine. (Repro­ duced with permission from FI Zamb 7 tab. 17.)

61

155. POLEMONIACEAE Sebsebe Demissew* & Sue Edwards* Bailey, P olem oniaceae in A M anual o f Cultivated Plants, Revised Ed. : 823-828 (1964); Cronquist, Polem oniaceae in An Integrated System o f Classification o f Higher Plants : 903-906 (1981). Herbs, occasionally shrubs, small trees, or climbers, often with an unpleasant smell. Leaves simple or occa­ sionally compound, entire, or pinnately or palmately divided, alternate, sometimes opposite; without stip­ ules. Inflorescence generally cymose and often appearing corymbose or capitate, terminal, or flowers occasionally solitary and axillary. Flowers usually regular, generally large and colourful, usually with a disc functioning as a nectary between the stamens. Calyx united with 5 lobes, bell- or salver-shaped, persistent in fruit. Corolla united, lobes 5, twisted in bud, bell-, funnel-, or salver-shaped, pleated. Stamens 5, inserted within the corolla tube, sometimes at unequal levels; filaments distinct, sometimes unequal; anthers dehisc­ ing longitudinally, introrse. Pistil with 3 carpels; ovary superior, 3-locular; ovules 1 to many in each locule; style 1, divided into 3 linear stigmas at apex. Fruit a capsule; seeds with seed coat often sticky or m ucilagi­ nous when moistened. A family o f about 13 genera with about 300 species native to Eurasia, Asia, and America. C antua, Co~ baea. Collomia, G ilia and P hlox are cultivated ornamentals found in gardens o f E Africa, and probably also in gardens in the Flora area; 1 genus with I species in the Flora area.

COBAEACov (1791) Climbing plants with leafy tendrils. Leaves alternate, pinnatisect, mostly terminating in branched tendrils. Flowers solitary', peduncles long with flowers hanging down. Calyx large, segments 5, leaf-like. Corolla bell­ shaped, violet to bright green, 5-lobed. Stamens exserted. woolly at the place of insertion. Capsule leath­ ery, 3-valved. Seeds with broad wings. About 10 species, all natives of tropical America; 1 cultivated species in the Flora area. Hutchinson. The Families o f Flowering P lants: 387-389 (1959) put Cobea in its own family, Cobaeaceae.

* T h e N atio n al H e rb a riu m , A d d is A b a b a U n iv e rs ity , P.O . B o x 3 434, A d d is A b ab a, E th io p ia

C. scandens Cav. (1791) Climbing herb with stems growing up to 8 m long, glab­ rous. climbing by means of leaf-tendrils. Leaves pin­ nate; leaflets 2-3 pairs, ovate or oblong, up to 10 x 5 cm, the lowest, close to the stem, with hastate or auriculate base. Flowers solitary, hanging, bell-shaped; pedi­ cels 15-25 cm long. Calyx segments leafy, elliptic or oblong. Corolla c 5 cm long, light violet or greenish to dark purple. Capsule 3-3.5 cm long. Seeds c 8 mm wide, thin. Fig. 155.1. Cultivated in gardens; 2000-2400 m. SU; originally from S America. Amare G. s.n.

62

155. POLEMONIACEAE Cobaea

l i g u r c 155.1 COBAEA SCANDENS: 1 - p art o l (lo w e rin g ste m : 2 disc and p istil; 3 - stamen: 4 - f ru it w ith p e r s is te n t c a ly x ; 5 tra n s v e rs e s e c tio n o f o v a ry ; 6 - s e e d : 7 e m b ry o . ( N o magnifications given and no specimens c ite d in o rig . p u b l.). D ra w n by J. H u tc h in s o n . (R e p r o d u c e d fro m Families o f Flowering Plants. Vol I Dicotyledons . fig. 2 3 8 . 1 9 5 9 .)

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63

156. HYDROPHYLLACEAE Sebsebe Demissew* & Sue Edwards* \ erdcourt, H ydrophyllaceae in FI. Trop. E. Afr.\ 5 pp. (1989).

Annual or perennial herbs, sometimes undershrubs, often scabrid. Leaves usually alternate or radical, with­ out stipules, entire to deeply pinnately or palmately lobed. Flowers regular, in scorpioid or 3-branched cy­ mes, false racemes or clustered, less often solitary. Calyx deeply divided into 5 (rarely 8-12), equal or unequal lobes, or 5 lobes with appendages between. Corolla often blue, variously shaped; lobes usually 5 I rarely 8-12). Stamens as many as the corolla lobes, inserted at the sinuses or low in the tube, filaments filiorm or dilated at the base; anthers 2-thecous, versatile, opening by longitudinal slits, included or exserted. Disk inconspicuous or absent. Ovaiy superior (very unusually inferior), 1-locular or 2(-3)-locular; ovules 2 or more in each locule; style filiform, usually bifid or styles 2; stigmas simple or capitate. Fruit a capsule, isually loculicidal, less often septicidal, 2-valved or splitting irregularly. Seeds oblong, globose or angled, * ith tuberculate, reticulate or rugose testa. About 18 genera with c 300 species, mainly American. H ydrolea with four species (H. sansibcirica Gilg, H. zeylanica (L.) Vahl, H. flo rib u n d a Kotschy & Peyr and H. brevistyla Verde, occur naturally in East Afica. O f these, H. flo rib u n d a Kotschy & Peyr (1867), a herbaceous plant 0.25 to 1 m tall, characterised by linear leaves and glabrous inflorescence (Fig. 156.1), has been found growing in seasonal swamps with %orghastmm stipoides and other species at about 1050 m in Uganda. It is also widespread from the Sudan to I\ ory Coast in W est Africa and might turn up in areas not far from the NE com er o f Uganda, in GG, and ad­ jacent to the Sudan in IL. Species o f P hacelia and W iganda are found as cultivated plants in Nairobi and may also be grown in Addis Ababa, but no material has been seen.

Figure 156.1 HYDROLEA FLO R IBU N D A : 1 - habit; 2 - flower; 3 - part of corolla from inside; 4 - stamen; 5 - calyx and pistil; 6 cross-section of ovary; 7 - fruit. (Magnifications and specimen citations not given in orig. publ.) Drawn by W. E. Trevithick. (Repro­ duced with permission from FI. W. Trop. Afr. 2nd ed. 2: fig. 275, 1963.)

* T h e N a tio n a l H e rb a riu m , A d d is A b a b a U n iv e rs ity , P.O . B o x 3 4 3 4 , A d d is A b ab a, E th io p ia.

64

157. BORAGINACEAE in clu d in g E h r e tia c e a e by H. Riedl* and Sue Edwards** Cufodontis. Emm.. 766-787 (1961 & 1962); Taton, B oraginaceae in FI. du Congo, du Rwanda et du Burundi: 82 pp. (1971); Verdcourt. B oraginaceae in FI. Trop. E. Afr. : 125 pp. (1991); Thulin. B oraginaceae in FI. Somalia 3:31-60 (2006). Annual or perennial herbs, often with woody rootstock, small to large shrubs, or trees, rarely climbers; usu­ ally with rough scabrid hairs, often w ith bulbous bases, especially. on young parts. Leaves alternate, or some­ times subopposite, simple, margin usually entire, sometimes dentate or crenate, without stipules. Inflorescences axillary or terminal, spike or raceme-like, or panicle, the ultimate branches usually scorpioid. Flowers usually regular, parts in 5s, mostly bisexual, sometimes unisexual. ( alyx tubular or bell-shaped, with (2-)4-7 lobes, persistent and often enlarging in fruit. Corolla tubular, bell- or funnel-shaped, or flat; tube short to long; lobes 3-16. usually all the same size. Stamens the same number and alternating with the corolla lobes, usually included in the tube, or sometime cxserted. Nectary disk usually present. Ovary superior, entire or deeply divided into 4 lobes. 2-locular with 2 ovules in each, or 4-Iocular with 1 ovule in each; style usually single, either terminal and persistent on top of the fruit, or separated from the ovary at maturity, or free from the bottom between the ovary lobes, or basal; stigmas various. Fruit a drupe, or, more often, divided into 4 one-seeded nutlets, sometimes separating into 2 two-seeded m encarps, rarely a capsule. Seeds shed in the fruit or nutlet. A large tamily with more than 2000 species in 100 to 145 genera; 19 genera and 62 species in the Flora area including naturalized ornamentals. This family is often treated as two families: Boraginaceae with mainly herbs, and Ehretiaceae with mainly trees and shrubs. The treatment here recognizes the following subfamilies: W ellstedioideae, Cordioideae, Ehretioideae, Heliotropioideae and Boraginoideae. some o f which are also regarded as separate families by various authors. Key to subfamilies 1 Flower parts in 4s; fmit a 1- or 2-seeded capsule. subfamily 1. Wellstedioideae - Flower parts usually in 5s; fmit not a capsule. 2.

Style inserted at the base between the 4 nudets. subfamily 5. Boraginoideae - Style terminal on top of the drupe or nutlets 3 3. Fmit an indehiscent drupe; style forked twice; trees or shrubs. subfamily 2. Cordioideae - Fruit divided either into 2 two-seeded halves or into 4 one-seeded nutlets; style undivided or simply forked; (if annual herb with indehiscent drupe, see Heliotropium supinum). 4 4. Trees, shrubs or subshrubs; style forked or two styles present; fmit usually divided into 2 twoseeded halves. subfamily 3. Ehretioideae - Annual or perennial herbs, sometimes subshrubs; style not forked; fmit a dehiscent dmpe dividing into two to four parts, rarely indehiscent. subfamily 4. Heliotropioideae Key to genera I Flower parts in 4s; fmit a capsule. 1. Wellstedia - Flower parts usually in 5s; fmit a dmpe, or sepa­ rating into 2 two-seeded halves or 4 one-seeded nutlets. ? * Herbarium. Departm ent o f Botany, N aturhistorisches M useum Wien, Burgring 7, A-1014 W ien, Austria ♦* T h e N a tio n al H e rb a riu m , A d d is A b a b a U n iv e rsity , P .O . B o x 3 4 3 4 , A d d is A b ab a, E th io p ia

2. Style terminal on top of the ovary, forming a beak on top of the fmit. 3 - Style inserted basally between the lobes of the ovary, not forming a beak on top of the fmit. 7 3. Style forked twice; small to large trees. 2. C ordia - Style undivided or forked once, sometimes 2 styles present; herbs or shrubs. 4 4. Prostrate heibs; inflorescence of single flowers in leaf axils; styles 2. 5 C oldenia - Erect herbs, trees or subshrubs; inflorescence of 1, 2 or more flowers, axillary and terminal; style 1. 5 5. Inflorescence of spike-like scorpioid cymes; style undivided; stigma forming a ring below the apex of the style, which is sometimes conical 01 cushion-shaped above it; fmit separating into 4 one-seeded nutlets, rarely into 2 halves or undi\ ided with 3 abortive locules. 6. H eliotropium - Inflorescence a corymb or umbel, or few-flowered on dwarf shoots; style forked or bilobed, stigma not forming a ring below its apex; fmit a dmpe. 6 6. Calyx lobes not overlapping, nearly united in bud; style bilobed. not forked. 4 B ourreria -C alyx lobes overlapping in bud; style distinctly foiked. 3 E hretia 7. Stamens with a twisted awn at apex. 9. Trichodesm a - Stamens not awned or awn not twisted. 8

157. BORAGINACEAE: 1. Wellstedia. 2. Cordia

8 Stamens witn a short, straight awn; filaments with basal appendage. 13. Cystostemon - Stamens not awned; filaments without basal ap­ pendages. 9 9. Throat of corolla with scale-like flaps. 10 - Throat of corolla naked, without scale-like flaps. 15 .0. Nutlets globular, depressed at the top, with glochids and/or a wing. 11 - Nutlets ovoid, reniform to cylindrical, not de­ pressed, without a w'ing or glochids. 18. Anchusa 11. Nutlets with a membraneous to firm wing. - Nutlets without a wing, glochidiate or smooth.

12 13

65

1. WELLSTEDIA B a lf f. (1888) Densely haiiy plants with woody base and sometimes also woody branches. Leaves small. Flow ers small, axil­ lary, in a leafy, scorpioid inflorescence, parts in 4s. Sta­ mens with very short filaments. Ovaiy with 2 ovules, becoming a loculicidal capsule with 1 or 2 seeds. Style terminal, forked. Distribution as for the subfamily; only the following species in the Flora area. W. filtuensis Hunt & Lebrun (1975) - ty pe: SD, Mt Filtu, Rippstein 827 (K holo.).

12. Wing of the nutlets with spines or glochids on the outer surface; nutlets with attaclunent scar near their dorsal apex, where they were attached to the short pyramid-shaped base of the style. 11. B randella - Wing of the nutlets without spines or glochids on the outer surface; nutlets fully attached to the columnar base of the style. 12. M icroparacarvum

Small woody herb with loose, prostrate branches. Leaves oblanceolate, up to 30 x 4-5 mm, pale grey, densely covered with stiff liairs on both sides. Flowers in short lateral cymes, that may be branched. Calyx 4lobed, lobes 2 mm long. Corolla tube as long as calyx, 4-lobed; lobes rounded, 1 nun across. Stamens with fila­ ments 0.5 mm long; anthers 0.7 mm long, included in the corolla. Capsule usually 1-seeded.

13. Corolla lobes twisted in bud; nutlets smooth, shiny. 19. Myosotis - Corolla lobes not twisted in bud; nutlets glochidi­ ate. 14

Open dry bushland on thin soil with limestone and calcareous outcrops, often locally abundant; 1300-1700 m. SD BA; Somalia. Friis et al. 11070; Gilbert et al. 7739; Gilbert & Ermias D. 8469.

14. Nutlets with attachment scar near their dorsal apex. where they were attached to the short pyramid­ shaped base of the style. 8. Cynoglossopsis - Nutlets with attachment scar along their whole length or attaclunent scar on a small distinct prominence in the upper part of the nutlet, some­ times additionally with an awn. 10. Cynoglossum 15. Style forked. 16 I - Style simple, not forked. 17 16. Corolla regular; without scales in the tube; inflo­ rescence with small leaf-like bracts. 14. A rnebia - Corolla irregular; with scales in the tube; inflores­ cence without leaf-like bracts. 17. Echium 17. Corolla regular, anthers subsessile, filaments very short. 18 - Corolla usually slightly irregular; anthers with filaments well developed. 7. Echiochilon 18. Nutlets smooth and shining; perennial herbs; co­ rolla with velvety or glandular appendages in the throat. 15. Lithospcrm um - Nutlets tuberculate or granular; annual herbs; co­ rolla with 5 longitudinal lines of hairs in the throat, but no appendages. 16. Buglossoides

Subfamily 2 CORDIOIDEAE Trees or slirubs with more or less leathery leaves. Style terminal, 2-4-branched. Fmit fleshy, indehiscent, 1-4seeded. Most common in tropical and subtropical regions with 3 genera of which Cordia is by far the largest. Only Cordia is found in the Flora area. 2. CORDIA L. (1753) A h m e d M u m in W a r f a , (1 9 8 8 ).

Acta Univ. Ups. 174: 7 8 pp + 5 p a p e rs

Small to large shmbs or woody herbs. Calyx and corolla 4-lobed; stamens 4; style shortly bifid. Fmit a 2-locular capsule.

Trees or shrubs, sometimes climbing or scrambling. Leaves alternate, occasionally subopposite, simple, usu­ ally leathery, often scabrid, margins entire or crenatedentate. Inflorescence a corymb or panicle, with numer­ ous one-sided branches, rarely few-flowered clusters, or flowers solitary. Flowers white, cream, yellow or or­ ange, bisexual or unisexual. Calyx tubular or bell­ shaped, smooth or ribbed, lobed, persisting and enlarg­ ing in fruit. Corolla funnel- or salver-shaped, usually 5Iobed, sometimes 3- or 4-lobed, or 7-10-lobed; lobes erect, spreading or reflexed. Stamens 4-8, well devel­ oped in male flowers, lacking anthers in female ones. Ovary 4-locular, with 1 ovule in each locule, but often reduced to a single seed at maturity; style terminal, di­ vided into 4 branches, base often persistent as a beak on the fmit. Fruits fleshy with a stony endocarp, usually 1seeded.

Subfamily with 1 genus and 3 species in Ethiopia, Somalia, Socotra, W and SW Africa.

About 250 species in the tropical and subtropical re­ gions of all continents; 10 species in the Flora area.

Wellstediaceae is sometimes treated as a family on its own.

The sweet sticky flesh of the fruits of many species is edible.

Subfamily 1. WELLSTEDIOIDEAE M e rx m U ller,

Mitt. Bot. Staat. Miinchen 3: 621 (1 9 6 0 ).

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157 BORAGINACEAE. 2. Cordia

1. Fruiting calyx over 20 mm long, enclosing mature fruit; flowers solitary or a few together in leaf ax­ ils; corolla orange or red; hairs fixed at middle on swollen bases. 1. C. suckertii - Fruiting calyx up to 12 mm long, not enclosing mature fruit; flowers in lax or dense inflores­ cences; corolla white, pale green, cream or pale yellow; hairs fixed at base. 2 2. Woody climber; base of petioles persistent as woody hook-like structures. 7. C. uncinulata - Tree or shrub, not climbing; petiole bases not forming hook-like structures, but sometimes persistent. 3 3. Leaves opposite orsub-opposite. - Leaves alternate.

4 5

4. Petiole 1-1.8 cm long; lower surface of leaves with longer hairs in vein axils and on mid-rib; calyx glabrous to sparsely pubescent; corolla cream or greenish-white. 2. C, sinensis - Petiole 0.7 cm or less: lower surface of leaves with hairs ± evenly scattered; calyx densely pubes­ cent to velvety; corolla pure white. 3. C. quercifolia 5. Calyx with 9 or 10 very prominent longitudinal ribs; corolla (14-)20-25 mm long and wide; an­ thers persisting and turning black after pollen is shed. 4. C. africana - Calyx smooth or with more than 10 fine ribs, not prominent; corolla less than 15 mm long and wide: anthers not turning black after pollen is shed. 6 6. Stigma arms flat and petal-like; fmiting calyx bell­ shaped: stems with raised circular petiole scars. 8. C. myxa - Stigma arms linear, not petal-like, fruiting calyx cup-shaped; stems with sunken petiole scars. 7. Leaves clustered on short shoots, densely hairy and usually very scabrid when older; lateral veins reaching the margins; calyx finely ribbed. 8 - Leaves usually spaced out on the stems, pubescent. glabrous or slightly scabrid when older, lateral veins not reaching the margins; calyx smooth. 9 8. Corolla tube exserted by 1-2 mm from calyx tube; fruiting calyx 14-18 mm wide, irregularly lobed. or lobes indistinct. 5. C. eilenbeckii - Corolla tube not exserted from calyx tube; fruiting calyx up to 10 mm wide, lobes usually distinct and regular. 6. C. monoica 9. Leaf-margins crenate-dentate, particularly in upper half; peduncles and branches of inflorescence velvety, with long hairs. 9. C. crenata - Leaf-margins entire; peduncles and branches of in­ florescence pubescent, only short hairs present. 10. C. sp. —de Wilde & de Wilde-Duyjjes 7335 1. C. suckertii Chiov. (1934) -ty p e: Somalia, Bulo Burti district, between Bulo Burti and El Mocolle, Suckert 61 (FT holo., K iso.). C. suckertii var. exasperata Verde., in Kew Bull 1957: 333-355 (1957).

Shrub or small tree. Bark pale brown to ash-grey or yellowish-brown, young branches and leaves densely scabrid. Leaves alternate or crowded on short branches; blade ovate, elliptic or oblong, 7-30 x 2-6 m m base rounded to truncate, apex rounded or emarginate, mar­ gin entire or slightly undulate, scabrid, hairs on both surfaces with swollen bases fixed at middle, lateral veins 5-7 pairs; petiole 2-6 imn long, tomentose. Flow ­ ers bisexual, or unisexual, solitary or a few together in small clusters, axillary, or terminal on short lateral branches; pedicels up to 5 mm long, softly hairy. Calyx 14-23 mm long, cylindrical; lobes irregular, 3-5.5 x 4-5 mm Corolla deep orange to red; tube 17-35 mm long. 7-10-lobed; lobes broadly ovate to subcircular, 12-15 x 10-14 mm. Stamens inserted at throat of corolla; fila­ ments glabrous. Style 21-38 mm long; stigmas less than 1 mm long. Fruit completely enclosed by the enlarged calyx, woody, 16-25 mm long, ovoid-cylindrical to glo­ bose. shortly acute at apex. Acacia Commiphora bushland; 200-300 m. HA. Somalia. Ellis 349; Gillett et al. 22682. 2. C. sinensis Lam (1792) - type: India, Sonnerat s.n. (P-LAM holo ). C. reticulata Roth (1819), non Vahl (1807). nom. illegit., C. rothii Roem. & Schultes (1819). C. subopposita DC. (1845), based on Cornus sanguinea sensu Forssk. (1775). non L. C. gharaf (Forssk.) Aschers. (1879), based on C ornusgharaf Forssk., nom. invalid., & C. sanguinea sensu Forssk. non L. C. oblongifolia Hochst. nom. nud. Multi-stemmed shrub or tree, 3—10(—12) m tall; bark pale, later on brownish grey, yellowish brown to almost black, longitudinally fissured, rough. Leaves opposite to sub-opposite; blade usually narrowly elliptic-oblong, sometimes oblanceolate-obovate, 1.5-11 x 0.6-4 cm, apex rounded, base rounded to cuneate, margin entire or crenate especially near apex, glabrous to pubescent or slightly scabrid above, long hairs along midrib and in nerve-axils below : petiole 1.3 (—1.8) cm. Flowers in lax panicles: unisexual, peduncles 15-30 mm; pedicels 2 mm Calyx cylindrical to bell-shaped, glabrous or pu­ bescent outside, 4-4.2 mm long together with the short rounded lobes. Corolla creamy white; tube 5 mm long, lobes of male flowers oblong, 4.5 x 1.5 mm, of female flowers 3—4 x 1.3-1.5 mm. Filaments o f functional sta­ mens 3.5 mm long, exserted, 2.2 mm long in female flowers. Style 2.5 mm long, stigmatic branches 2-4 mm. Fruiting calyx funnel- or cup-shaped, 3-5 x 5-8.5 mm, usually with well defined lobes. Fruit yellow to orange or bright red, 0.7-1.3 x 0.7-0.9 cm. flesh sticky Fig 157.1.8 & 9. 1* Along wadis, around temporary pools, in rivenne forest or tangled scnib, sometimes in more open vegeta­ tion with Acacia and Commiphora: 450-1500 m. AF EW TU GD SU GG SD/BA HA; Socotra, Somalia, E Africa, west from Sudan to Chad, Ghana, Togo, Nigeria, south to Angola, Namibia, also Egypt, Saudi-Arabia, Middle East, Pakistan, India, Sri Lanka. Ash 188; Friis et al. 10212; Mesfin T. et al. 4097.

157. B O R AGIN ACE AE: 2. Cordia 67

F ig u re 157.1

C O R D IA U N C IN U L A T A . 1 - m a le flo w e r x 4 . C. M O N O IC A : 2 - m a le f lo w e r x 4 ; 3 - fe m a le f lo w e r x 4 ; 4 fru it x 2. C. E L L E N B E C K II 5 - m a le f lo w e r x 4. C. C R E N A T A . 6 - m a le f lo w e r x 4 ; 7 - fru it x 2. C. S IN E N ­ S IS . 8 - p a rt o f lo w e r le a f s u rfa c e x 10; 9 - in f lo r e s c e n c e x 1. C. Q U E R C IF O L IA . 10 - p art o f lo w e r l e a f s u r f a c e x 10; 11 - in f lo r e s c e n c e

x 1. 1 fro m Songster 167 ; 2 fro m Gillelt 2 0 0 6 6 ; 3 fro m Brenan, Gilleti & Kanuri 1 4 7 3 1 ; 4 fro m Kalende 4 2 6 ; 5 fro m Gillett 1 2 6 7 4 ; 6 fro m Adamson 4 6 ; 7 fro m Synnott 1732; 8 fro m Gillett 1 2 5 7 1 ; 9 fro m Fratkin 2 ; 10 fro m Gillett 1 2 7 0 6 ; 11 fro m Newbould 2 9 5 4 . D ra w n by M a u re e n C h u rc h . (R e p r o d u c e d w ith p e r m is s io n fro m FI. Trop. E. Afr. B o r a g in a c e a e : fig. 4 .)

6 & 7 a re id e n tifie d a s C. crenata s u b s p . meridionalis , b u t th e

N o te ;

s u b s p e c ie s a re d if fe r e n tia te d o n v e g e ­ ta tiv e c h a r a c te r s , an d th e f lo w e r is s im ila r fo r all s u b s p e c ie s a n d is le ft h e re to r e p re s e n t th e ta x o n .

M'

Spreading shrub to small tree, often with a crown wider than its height. 0.3—4.5(—6) m tall, sometimes described as prostrate; bark dark grey to almost black, corky, thick, deeply longitudinally Fissured. Leaves usually op­ posite. rarely alternate on some shoots; leaf-blade ob­ long or oblong-elliptic, 0.9-10 x 0.7-5 cm. apex rounded or slightly indented, with a tiny mucro. base rounded to cuneate. margin entire to indistinctly crenate towards apex, upper surface rough with very short, white hairs, beneath with longer hairs, more or less evenly distributed, or with longer hairs on both surfaces: 3. C. qucrcifolia Klotzsch (1861) type: Mozambique. Tetc, Peters s.n. (B holo. de­ petiole 0.1-0.7 cm long. Flowers white, unisexual, in compact inflorescences, sessile or with a very shod (up stroyed). to 1.5 cm) peduncle; pedicels absent or up to 1 mm long C. nevillii Alston (1931).

C sinensis and C. quercifolia liave both frequently been identified as either C. rothii or C. gharof the latter because this name was used bv Eggeling and Dale in In­ digenous Trees o f Uganda (ed. 2: 48. 1951). and many authors have followed that naming. Turton 85 records numerous uses for this tree with the Mursi. including for making bows, axe handles, fire sticks and pipes. In many places outside the Flora area it is also cultivated and has become naturalized.

157 BORAGINACEAE: 2. Cordia 68

m flower, elongated to 4-5 mm in fmit. Calyx densely pubescent to velvety, becoming glabrescent when older, 3.5-6.5 mm long; lobes very short. Male flowers: co­ rolla lobes up to 6 mm long; stamens exserted for 3-5 mm: style absent or strongly reduced. Female flowers: corolla-tube 3.5-5 mm long; lobes 2-4.5 mm; sterile filaments exserted for 1.5 mm; style 3 mm long; stigmas 2-2.5 mm long, distinctly exserted. Fruiting calyx cup­ shaped. 4.5-5 x 7.5-9 mm; lobes often broken and toothed. Fmit yellow to bright orange, ovoid with per­ sistent style-base, 1-1.7 x 0.7-1 cm. Fig. 157.1 10 & 11. In dry bushland and grassland with scattered tree;., Acacia - Commiphora bushland; 1000-1350 m. AF EW SU GG HA; Somalia, Socotra. E Africa, Egypt, Sudan, west to Mali, Niger, Mauritania, Chad, south to Mozam­ bique, Zimbabwe, also in Arabia, India, and Sri Lanka. Beals 974; Burger 1620; Hemming 1460. This species is sometimes considered an ecological variant of C. sinensis, but Verdcourt (1991) found veiy few intermediaries and treated it as a species The two species are distinct in the Flora area. 4. C. africana Lam. (1792); C. sebestena L. var. fi Poir, in Encycl. Meth. Bot. 7: 45 (1806); C. abyssinica R.Br. (1814): Varronia abyssinica (R.Br.) DC. & A. DC. (1845); Calyptrocordia abyssinica (R.Br.) Friesen (1933) - type: “wanzey”, Ethiopia. Bruce. Travels, 5 (Appendix): 54, t. 17 (1790) (lecto.). Cordia abyssinica R.Br. var. acutifolia A.Rich., in Tent. FI. Abyss. 2: 81 (1850) - type not given. C. harara Beck (1888) - type: HA. Mt Haqim. near Harar town, Hardegger s.n. (W holo.). C. holstii Giirke (1895). C. unyorensis Stapf (1906). Small to medium-sized tree, up to 10 m tall (18 m or more in forests), with a thick spreading crown, rarely a shrub. Bark dark to pale brown, rough, fibrous, peeling off. fissured; slash yellow or white turning green, grey or brown. Young stems brown, velvety-tomentose with interspersed longer liairs. later on glabrescent. Leaves alternate, thick; leaf-blade ovate, elliptic or nearly rounded. 7.5—17.5(—30) x 3.5—9(—22.5) cm, apex rounded to acuminate, base rounded to cordate, rarely cuneate, glabrous to scabrid and dark green above, paler beneath, shortly pubescent with longer hairs on the veins, rarely tomentose. or nearly glabrous apart from a few longer hairs along veins; veins very many and prominent; peti­ ole 1.3-13 cm long. Inflorescence showy, shorter than the subtending leaf, many cymes in a compact cory mb or panicle. Flowers bisexual, sessile, with a sweet scent. Calyx cylindrical to bell-shaped. (5—)7—9 mm long, 4-6 mm wide at throat, with 9-10 conspicuous longitudinal ribs, dark-brown tomentose; teeth 4-5, irregular, or more or less 2-lipped. Corolla white, funnel-shaped, (14-)20-25 mm long and wide; tube 18-22 mm long; lobes spreading, strongly folded looking like cmmpled paper, margin sparsely pubescent outside, crenulated, median veins ending in a short, hairy mucro. Stamens included; filaments 8-10 mm; anthers persistent and be­ coming black after pollen is shed. Ovary glabrous; style 12-20 mm long, divided 7-14 mm from base; stigmas

subulate, 1-2 mm long. Fmiting calyx cup-like, up to 11 mm wide. Fruits yellow, subglobose to ovoid, 10-12 x 6-10 mm. glabrous, with sweet, sticky flesh. Fig. 157.2. Open places in moist montane forest, forest edges, in forest remnants around churches and other traditionally protected areas, as isolated trees in grassland and culti­ vated fields, in villages and public gardens, also in plan­ tations; 700-2550 m. EW TU GD GJ WU SU AR WG IL KF GG SD BA HA; widespread from Sudan in the north to S Africa and Angola in the south, also tropical Arabia, cultivated in India and elsewhere in tropical botanic gar­ dens. Mesfin T. & Kagnew G. Y. 2240; Mooney 9209; Sue Edwards 3820B. The debate over whether the correct name for this species is C. africana or C. abyssinica lias been exam­ ined by Warfa (1988) and been well summarized by Verdcourt (1991). The characters of calyx shape and leaf indumentum used to separate C. harara are also found in specimens of C. africana. Hence it does not seem justified to recog­ nize C. harara as a separate taxon. This is one of the most important timber trees in the Flora area. The wood is easy to work, and is used for making dnims, bee-hi\es. household furniture and con­ tainers. and many otlier items. It polishes well and does not crack easily as long as the wood is seasoned before it is processed. C. africana is also an important forage plant for both domestic animals and bees. 5. C. monoica Roxb. (1796) -ty p e : from India. E. Madras. Roxburgh drawing no. 200 (K lecto ). jC . ovalis DC. & A.DC. (1845) - type: Ethiopia. Gafta, Schimper II 1218 (G lecto., K isolecto ). C. rubra Hiem (1898) - types: from Angola. IVelwitsch 5423, 5431& 4783 (all LISU syn., BM isosyn.) and. Gafta, Schimper 1582, (BM syn., K P isosyn ). C. obovata Bak. (1894), non Balfour f., nom i/le­ git.: C. bakeri Britten (1895) - type: GD. Agow. Dschadscha. Schimper 2180 (K lecto.). C. quarensis Giirke (1895). [c. bequaertii De Wild. (1921), non De Wild. (1920); C. kabarensis De Wild. (1923). C. myxa sensu Richard (1850). non L. Shrub or tree, usually branched from the base. 1.5-8 (-15) m tall. Bark grey, sometimes very light grey or white, smooth or rough, peeling off in long strips; brandies longitudinally striate, with yellow-brown liairs, glabrescent later on, intemodes short and leafscars prominent. Leaves alternate; leaf-blade ovate, obo­ vate or subcircular, sometimes elliptic, (1—)2.5—8(—13) x 1-9 cm, apex rounded, apiculate or emarginate, rarely acute, base cuneate to rounded, margin entire, denticu­ late or rarely dentate, surface usually extremely scabrid. with minute bulbous-based hairs, and a few branched hairs, often densely white-pubescent beneath when young, venation densely reticulate; petioles 0.5-5 cm long, brownish tomentose. Flowers fragrant, unisexual or bisexual, in few-flowered, terminal or axillary inflo­ rescences, 2-6 cm long; pedicels articulate at apex, 0.2-0.8(-3) mm long. Calyx cylindrical to bell-shaped.

157. BORAGINACEAE: 2. Cordia 69

Figure 157.2 CORDIA A F RIC AN A . 1 - flowering branch; 2 - calyx; 3 - corolla opened out to show stamens; 4 - pistil; 5 - longitu­ dinal section through ovary; 6 - cross-section of fruit; 7 - fruit. Drawn by W. E. 1 revithick. (No specimens or magnifications given in orig. publ.) (Reproduced with permission from FI. IV. Trop Afr. (2nd ed.) 2: fig. 276.)

6-8.5 x 5-8 mm. tomentose with simple and branched hairs outside, 3-5-lobed or 2-lipped, or irregularly lobed. Corolla greenish-white to pale yellow; tube cylin­ drical. 4-7 mm long, 3-5-lobed; lobes oblong to nar­ rowly obovate, 3.5-6.5 x 1-2.5 mm, spreading. 1-1.2 cm wide. Male flow'ers: 2 forms, corolla lobes and sta­ mens 5(-6); filaments either 6 mm or 2-3 mm long, pis­ til totally absent. Female and bisexual flowers: corolla lobes 4-5; filaments 2-3 mm long; ovary glabrous, sty le exserted. 6-10 mm long; stigmatic branches 3-5 mm long, recurved. Fruiting calyx shallow, cup-shaped, up to I cm wide. Fruits yellow or orange, sometimes black, ovoid to obovoid, 0.9-2 x 0.6-1.2 cm. Fig. 157.1.2-4. A wide range of habitats: moist evergreen forest, riv­ erine vegetation. Acacia woodland. Acacia - Commi­ phora bushland, thickets in grassland, coastal thickets; 500-1900 in. EW TU GD/GJ WU SU AR IL GG SD BA HA; widespread throughout tropical Africa, also in Ara­ bia and tropical Asia, cultivated in Mauritius. Ensermu K. & Petros E. 1877; Mesfm T & Sebsebe D. 3760; Sue Edwards & Tewolde B.G.E. 3546. A very variable species in which local ecotypes can be recognized. Since the leaves are very rough they are used like sandpaper by local carpenters to smooth wood.

C. ellenbeckii Gtirke (1912) Boran. E llenbeck type. SD, “GalIahochlandv 2057 & 2092 (B syn. destroyed). Shrub, up to 5 m tall. Bark smooth, usually dark grey; sometimes white. Young branches sparsely pubescent or woolly, soon glabrescent. Leaves alternate, mostly crowded on short lateral shoots; leaf-blade almost rounded, obovate or oblong, 0.8-5(-6) x 0.4-3(-5) cm, apex rounded to acute, base tapering towards petiole, margin crenulate-undulate to irregularly dentate, pubescent-tomentose, often velvety on both sides, hairs white, yellow-brown along veins, older leaves become scabrid. Flowers unisexual, fragrant; male flowers in 5-6-flowered inflorescences, in forks of branches, though sometimes looking as if terminal; female flowers solitary. Male flowers: calyx narrow funnel-shaped, 7-11 mni long with irregular lobes, 1-2 mm long, with spreading longer white and shorter rust-coloured hairs outside; corolla cream, glabrous; tube up to 15 mm long, 1-2 mm longer than calyx; lobes 5-6 x 3.5 mm; anthers just exserted from tube, free parts of filaments very short; ovary rudimentary, without style. Female flowers: calyx and corolla similar to male flowers; anthers just exserted, on very short filaments; ovary 2.5 mm long; styles 9 mm long; stigmatic branches about 5 mm long, exserted. Fruiting calyx wide, cup-shaped, 0.6 x 1.8 m m

157 BORAGINACEAE: 2. Cordia 70

mm, finely ribbed. Fruit dark yellow or orange, often nearly globose, up to 12 mm across, obscurely ribbed; caly x cup-shaped. 16 mm wide. 6 mm deep, sessile, en­ closing the fruit. Fig. 157.1.5.

8. C. m yxaL. (1753) > - type: Middle East. Hasselquist s.n. (S-LINN. microfiche 94.5, lecto ). I C obliqua Willd (1794).

Acacia-Commiphora woodland and busliland. on limestone or gravelly soil; 8 5 0 -1400(-1800) m AF WU SD; N Kenya. Friis et al. 2919; Gilbert et al 7701; Mes­ fin T. & Vollesen 4320.

Tree, 6-12 m tall; trunk sometimes twisted: young stems hairy, glabrous later on. Leaves alternate; blade broadly ovate to subcircular, rarely obovate, 3-18 x 3-20 cm. apex rounded to shortly acuminate, base rounded to cuneate. margin usually crenate-dentatc. leathery, not scabrid glabrous above, glabrous to densely pubescent beneath; petiole 0.6-3.5 cm long, glabrous or sparsely hairy. Inflorescence a lax panicle of cymes, often on short lateral branches; pedicels 1-2 inm long, articulate at apex Flowers unisexual, with female flowers generally slightly larger than the male ones. Ca­ lyx tubular or bell-shaped. 4.5-8.5 mm long, usually 3lobcd. glabrous outside, pubescent to tomentose at apex inside Corolla white, tube 3.5-6.5 mm long, 5-lobed: lobes elliptic, c 5 x 2 nun. reflexed or rolled up Male flowers stamens exserted; filaments 1.5-3.5 mm long, liairy at base; ovary rudimentary, without style. Female flowers: filaments 1.542.5 mm long, pubescent; anthers sterile; style exserted. 8-9 mm long, deeply divided into 4 stigmatic branches, 4-5 inm long, flattened and petallike. with irregular or frilled margins. Fruiting calyx cup-shaped. 7-10 x 12—15(—20) mm. Fmit yellow, light orange or blackish, ovoid. 1.2-3.5 cm long, apiculate, 4-locular. but with only 1 seed developing.

Warfa (1988) treated this species as a synonym of C. monoica. Verdcourt (1991) used the original description and recognized this species on the basis of three collec­ tions by Gillett (Gillett 12592, 12674 and 12684). and Gilbert & Thulin 1562. all from the same general area in N Kenya. The three collections cited from the Flora area all agree with Verdcourt’s description, and the species is therefore maintained in this account. Ash 1211 and de Wilde et al. 7335 both have small velvety leaves, and flowers (de Wilde et al. 7335) or fruits (Ash 1211) that agree better with C. ellenbeckii than with C. monoica. Ash 1211 was collected in AF be­ tween Dire Dawa and Mieso. and de Wilde et al. 7335 in WU, near Kombolcha at 1800 in. More material is needed to confirm these identifications. Some collections from Somalia presently named as ( monoica appear closer to C. ellenbeckii. particularly in their fmit characters. 7. C. uncinulata De Wild. (1921) - type; Dem. Rep. Congo. Kivu, Bequaert 3261 (BR lecto., and isolecto ). C. tisserantii Aubrev. (1950). Woody climber, or shrub, up to 10(-15) m tali with long pendulous branches and blunt recurved spurs formed from petiole bases at the nodes. Leaves alter­ nate; leaf-blade ovate to elliptic-ovate. 2.5-17 x 2.5-8 cm, apex attenuate to acuminate, base rounded or cuneatc to subcordatc. margin entire to finely crenate in the upper part, not scabnd. lower surface paler tlian up­ per; petiole 0.5-5 cm long. Inflorescences terminal, much-branched panicles of loose one-sided cymes. Flowers unisexual. Calyx tubular to bell-shaped. 5-7 mm long. 5-7-lobed. finely pubescent, enlarging into a cup up to 10 x 15 mm in fmit. Corolla white, tube 4-5 mm long. 4-5-lobcd; lobes elliptic to narrowly obovate or oblong. 4-6 x 1.3-2 mm. Male flowers with exserted stamens; filaments 5.5-6.5 nun long, hairy at the base; non-functional ovary minute; style absent. Female flow­ ers with sterile stamens; filaments 1-1.5 inm long; ovary well developed. 4 x 3 mm; style 1.5 mm long; stigmatic branches 3.5 mm long. Fmit orange, c 10 x 5-6 mm. usually 2-locular. Fig. 157.1.1. Montane forest with Pouteria (Aningeria), Morus, Antiaris and other tall trees; 1050-1100 m. KF; Central African Republic, Senegal. Nigeria. Dem. Rep. Congo. Friis et al. 3924, 4002. 4036. Not recollected in the Flora area since 1984. C. uncinulata appears to be confined to the moist montane forest dominated by Pouteria altissima. Much of this forest has been cleared for coffee and tea plantations.

Cultivated; 1200 m HA; indigenous to tropical Asia, now cultivated and naturalized in many places. Tadesse E. 720 9. C. crenata Del. (1826) ty pe; Egy pt, cultivated in gardens in Cairo. Delile s.n (MPU holo.) subsp. crenata (' zedambae Martelli (1886) - type: EW. on the slopes of Zedainba (Tsada-Amba). Beccari 115 (FT lecto , K isolecto.) Shrub or small tree, 1.5-9 m. Bark pale buff-brown or grey; young shoots more or less glabrous or velvety, with appressed and usually also long, red-brown, spreading liairs. later glabrescent or liairs persistent in upper parts Leaves alternate, or crowded on short shoots; leaf-blade broadly elliptic, obovate-oblong o ralmost rounded, rarely oblong-lanceolate, 1-11 x 0.8-8 cm rounded, apex emarginate or subacutc. base rounded or cuneate. margin entire or crenate to dentate, puberulous or subglabrous, sometimes slightly scabrid; petiole 0.5—1 cm long. Inflorescences 2.5-3 cm long, sessile or with short peduncle; peduncle and secondary axes velvety with a few long hairs; pedicels 2-3 mm long. Flowers male or bisexual. Male flowers: calyx-tube funnel- or bell-shaped, 3.5—1.5 mm long; lobes broadly triangular. 1.2-1.5 inm long; corolla cream or greenish-yellow, tube 3 5-4 mm long, lobes 4-5, rarely 6. ovate-oblong, 4-4.5(-6) x 1—2.5 nun; filaments 1.5-6 mm long; ovary rudimentary, without style. Bisexual flowers: calyx 4.2-5 5 nun long, lobes 1—1.3 nun long, ovate, glabrous outside, minutely puberulous inside; corolla-tube 4-5 mm long, lobes 4-5(-6), oblong to nar­ rowly ovate. 3-4 x 1.5—2 nun; filaments 1.5-2 mm long;

-

157. BORAGINACEAE: 2. Cordia, 3. Ehretia 71

si vie 2-3.5 mm long, branches up to 1.3 mm. stigmatic branches 7 mm long. Fruiting calyx 6.5-9 mm wide. Fmit red, ovoid-globose. 7-13 x 6-8 mm, apiculate. Fig. 157.1.6 & 7.

bell-shaped. Style divided, persistent on the fmit. Other characters as given for the subfamily.

In villages by wadis and water holes; 900-1100 m. AF EW GG HA; Sudan, Somalia; Yemen, Oman. Iran. India. Carr 318; Gilbert et al. 7832; Tadesse E. 732.

1. Leaves glabrous above, usually smooth, with tufts of hairs in axils of main veins below, or with hairs mainly along the main veins on the lower side. 1. E. cymosa - Leaves hairy throughout at least on lower side, usually rough, without tufts of hairs in axils of main veins below. 2 2. Corolla tube slightly longer than the lobes, or equalling them about 3 or 4 times as long as ca­ lyx; inflorescence glabrous, or with a few, short, red-brown hairs, without glands. 2. E. braunii - Corolla tube markedly longer than the lobes, about twice as long as calyx; inflorescence densely glandular. 3. E. obtusifolia

Subsp. meridionalis Warfa, in Acta Univ. Upsal. 174 •III): 7 (1988) differs by having young shoots densely covered with appressed and long hairs. It is found in Somalia and E Africa in Acacia - Commiphora bushand, and also in river beds and on river banks. Subsp. hinvangensis Verde., in FI. Trop. E. A fr., Boraginiceac: 20 (1991) is only found in Tanzania. It is recoguzed by its narrowly elliptic to oblong-lanceolate leafnlades, and all young growth being covered by velvety ned-brown hairs.

About 75 species in the Old and New Worlds, par­ ticularly Africa and Asia; 3 in the Flora area.

JO. C. sp. = de Wilde & de Wilde-DuyJjes 9779 Small shrub, 5 m tall; young stems slightly pubescent and lined. Leaves alternate; leaf-blade elliptic, 3-6.3 x 1 2-2.8 cm. apex apiculate, base cuneate to oblique, margin entire, both surfaces scabrid, with longer scabrid hairs on the main veins and petioles; petiole up to 1.2 cm long. Inflorescence terminal on short side-shoot. lax, few-flowered: peduncle 2.3 cm long; pedicels articu­ lated. very short. Calyx glabrous with a few scabrid lairs on veins and lobes, funnel-shaped; tube 5 mm long, lobes c 1 mm long, ovate. Corolla cream or white, funnel-shaped; tube extending c 1 mm longer than ca­ lyx; lobes 5, ovate. Fmit not seen. Shrub vegetation, c 60 km south of Kombolcha: c 2000 m. WU. This specimen seems to be nearest to C. obovata. Warfa identified it as C. monoica, but this was queried by Verdcourt. C. monoica typically has densely reticulately veined leaves that are almost bullate; the leaves of this specimen are flat with less distinct veins. The in­ florescence in C. obovata is compact, but in this speci­ men it is lax. Warfa found C. obovata occurring in both Oman and NE Somalia, as well as Socotra. More field work is needed to determine whether or not this speci­ men represents an undescribed taxon or an extension of an established one. Subfamily 3 EHRETIO ID EA E Giirkc in Engler & Prantl (1897); E hretiaceae Lindl. (1830).

Trees or shrubs, rarely subshrubs or herbs, usually with more or less leathery leaves. Mature fruit a dmpe. sepa­ rating into 2-seeded halves (rarely separating into 4 one-seeded nutlets).

1. E. cymosa Thonn. (1827) type: Ghana, Thonning 89 (C holo. & iso., FT P-JU iso.). Slirub or small tree, 3-10 m tall; young branches glab­ rous. pubescent or sparingly hairy, often pale redbrown. Leaves ovate to ovate-oblong, 4-15 x 2-6(-7) cm, apex acute or acuminate, base rounded, smooth, glabrous above, loosely hairy or with tufts of hairs in the axils of the main veins below, margins ciliate; petioles 0.5.3.5 cm long, glabrous or slightly hairy. Inflores­ cence terminal, a dense umbel with many one-sided branches; flowers fragrant, sessile, or pedicels 1-2 mm long. Calyx bell-shaped, 2-2.5 mm long, 4-5-lobed; lobes lanceolate, c 1 mm long, glabrous or pubescent with ciliate margins. Corolla white, 6-9 mm long; tube 1.5-3 mm long, longer than calyx; lobes 5, oblong, acute or obtuse, c 4 x 2 mm. Stamens inserted in throat of corolla; filaments 4-5 mm long. Ovary ovoid; style 4-6 mm long, shortly forked. Fmit fleshy, orange, red or black, 6-7 mm across. Fig. 157.3.1-3. The following 3 varieties have generally been recog­ nized, although the characters separating them are not clear-cut. Riedl in Linzer Biol. Beitrage 17(2): 303-315 (1985) considered E. abyssinica as a species separate from E. cymosa Thonn., but in this account it lias been decided to follow the treatment by Verdcourt in FI. Trop. E. Afr. 1.

Lower side of leaves glabrous except for tufts of hairs in the axils of veins; branches of inflores­ cence glabrous or loosely haiiy. var. cymosa - Lower side of leaves hairy, at least along main veins, tufts of hairs absent or very reduced in the axils of veins; branches of inflorescence with short or spreading hairs. 2

12 or 13 genera confined to tropical and subtropical 2. Branches of inflorescence with short hairs; corolla regions and represented by 3 genera, Ehretia, Bourreria lobes longer than tube. var. divaricata and Coldenia, in the Flora area. - Branches of inflorescence with spreading hairs, corolla lobes shorter than tube. var. sylvatica 3. EHRETIA L. (1759) var. cymosa; E. abyssinica R. Br. ex Fresen. (1838); E. cymosa Trees or shrubs with alternate leaves. Calyx small; lobes Thonn. in Schumach. & Thonn. var. abyssinica (R. Br. overlapping in bud. Calyx and corolla 5-lobed, usually

12

157. BORAGINACEAE: 3. Ehretia

^ J C iM O SA 1 - f lo w e rin g b ra n c h x Vy 2 - flower, side v ie w x 5. 3 - p istil a n d n e c ta ry d isc x 5 E f L S l t u ! [ } 4 ~ f lo w e rin g b ra n c h x 2/ 3.5 - flo w e r, lo n g itu d in a l section x 5. E B R A U N 11. 6 - f lo w e rin g b ra n c h x Vy 7 - f l o w e r s id e v ie w x 5. N o s p e c im e n s c ite d in o n g . publ. D ra w n by W Burger (Mod.fled and reproduced w ith p e r m is s io n fro m W .C B u r g e r ram i lies o f l3—4 mm long, attached to the base of the style; branches of inflorescence not spread­ ing horizontally 4 Nutlets cushion-shaped, more or less evenly cov­ ered with glochids, not forming a distinct rim, glochid-bases not flattened; whole plant densely covered in long silky hairs, leaves smooth. 4. C. densefoliatum - Nutlets flattened, glochids forming a distinct rim, with additional glochids on the upper surface, glochid-bases flattened, whole plant not covered in long silky hairs, leaves often rough. 2. C. coeruleum 1 C. amplifolium Hochst. ex A.DC. in DC. (1846) - type: GD. between Endschetcap and Schoata. , Schimper 111:564 (G BM K UPS W syn ). Perennial, 0.2-2 m tall, with long fleshy taproot; stems branched in the upper part, densely covered with short, irregularly directed hairs, often furrowed. Radical and lower stem leaves ovate-lanceolate, up to 16 x 8 cm, petiole up to 20 cm long, basally widened to an auriculate sheath; upper leaves sessile, base cordate, veins nu­ merous, impressed above, prominent below, liairs sparse, bristly, similar hairs along the veins and short, slightly spreading hairs on the paler lower side. Inflores­ cence branched, without bracts, cymes few-flowered, tightly coiled in bud and flower, elongating as fruits form; pedicels short, up to 6 mm in flower, elongated and bent downwards in fruit. Calyx 2-2.5 mm long in flower, divided to the middle, lobes ovate, persistent and enlarging in fruit, more or less acute or obtuse, up to 3 x 1 mm Corolla usually dark bright blue or even purple, sometime pale blue or even white, tube included in ca­ lyx, lobes 5, nearly orbicular, 5-6 x 5-6 mm; scale-like appendages in throat nearly square. Stamens inside co­ rolla tube. Style very' short, scarcely visible between nutlets. Nutlets free from the style, joined to the gy no­ base by a neck-like structure, flattened, broadly ovale to obovate, up to 6 mm long, with glochidiate liairs all over, but more densely on the margins, or_oply around the margins and along a mid line. 1. Upper surface of nutlets covered all over with glochidiate hairs. var. amplifolium - Upper surface of nutlets without glochidiate hairs. or with only a few along a midline. var. subaipinum

157. BORAGINACEAE: 10. Cynoglossum

91

Figurel57.l3 CYNOGLOSSUM A M P U F O U U M var. AM P LIF O LIU M . 1 - flowering stem x 2/3; 2 - detail from upper leaf surface x 6; 3 - flower x 4; 4 - calyx x 6; 5 - corolla opened out x 6; 6 ovary x 20; 7 - ovary, longitudinal sec­ tion x 20; 8 - fruiting stem x 2/3; 9 fruit x 3; 10 - nutlet x 4. C A M P LI­ FO LIU M var. SU B A L P IN U M : 11 fruit x 3; 12 - nutlet x 4. 1-7 from Battiscombe 505; 8-10 from Verd­ court 1158; 11 & 12 from Verdcourt 3786. Drawn by Maureen Church. (Re­ produced with permission from FI. Trop. E. Afr. Boraginaceae: fig. 30.)

var. amplifolium C. bequaertii De Wild. (1920). C. longepetiolatum De Wild. (1926). Fig. 157.13.1-10.

In open places and partial shade in montane forest, Erica bushland, Helichrysum clumps, also moist grass­ land, 1500-3500 m. TU GJ SU AR IL KF GG; E Africa, Malawi, Dem. Rep. Congo, Mozambique, Zimbabwe. Gilbert & Phillips 9247; Hillier & Lythgoe 505; Mooney 9244. var. subalpinum (T.C.E. Fries) Verde., in FI. Trop. E. Afr. Boraginaceae: 106 (1991);

C. subalpinum T.C.E. Fries (1923) - type: Mt Kenya, R E. & T.C.E. Fries 1331 (S holo. & iso.). Fig. 157.13.11 & 12. By forest paths and in Erica bushland; 2450-3300 m. SU BA; Kenya, Tanzania, Nigeria, Cameroon. Ash 597, 1710; Friis et al. 5552.

2. C. coeruleum Hochst. qxA.DC. in DC. (1846) type: GD, Enschedcap, Schimper 542 (G BM K W iso.). Paracynoglossum afrocoeruleum Mill (1984) 481, nom. illeg.

157. BORAGINACEAE: 10. Cynoglossum 92

Herb 15-120 cm taJi, annual or perennial, with tap root; stems simple or several from the base. Leaves at base, narrowly ovate, elliptic-oblong, linear-oblong, or oblan­ ceolate, 8-21 (-30) x 1-6 cm long, tapering towards base, rough, with sparse to dense short, appressed hairs from white cystolith dots, less dense on lower side; peti­ ole 2-10 cm long; stem leaves similar, linear-lanceolate to ovate, 1.5—17(—30) x 0.2-5 cm, contracted or slightly widened at base, usually sessile. Inflorescence axillaiy or terminal, cymes 1-3 together, finally elongated, 10-25 cm long; pedicels 0.2-0.7(-1.2) cm long. Calyx lobes narrowly ovate to elliptic-obovate, 1.2—3(—3.5) x 0.4-1.2(-2) mm in flower, 3-4 x 1.5-1.8 mm in fruit, acute to rounded, pubescent or ciliate, hairs sometimes with w'hite dots at base. Corolla bright deep blue or pinkish-mauve turning blue, sometimes with yellow or white centre, or white with blue, pink or pale yellow centre, up to 8 mm wide; tube 1.5—2(—2.5) mm long, lobes rounded-oblong, (1.2-) 1.5-4 x 1-4 mm. Style 0.5.3 mm in fruit. Nutlets flattened ovoid-obovoid, (2.5-)3-4 x 3-3.5 miti, arrangement of glochids vari­ able, always some with flattened bases, and usually some forming a distinct rim around the nutlet margin. The subdivision of C. coeruleum as used here owes much to Verdcourt (loc. cit.) and his annotations on the material in Kew. It is difficult to identify infraspecific taxa without mature fruits. 1.

Nutlets covered w'ith many slender glochids, rim of several rows of glochids; stem leaves very rough with conspicuous white cystoliths. often as large as basal leaves. subsp. latifolium - Nutlets with fewer stouter glochids, rim of 1-2 rows of glochids, often with flattened bases; stem leaves with some soft white hairs, w ith or without white cystoliths, smaller than basal leaves. 2 2. Nutlets 3.5-4 x 3-3.5 mm; glochids large, spiny, c 1 mm tall, flattened with a paler margin, 2-5 glochids on upper face; stem leaves with ampJexicaul, almost auriculate base. subsp. coeruleum var. hedbergiorum - Nutlets up to 3 x 3 mm; glochids smaller, up to 0.7 mm tall, base flattened, same colour throughout or with paler head, 6 or more glochids on upper face; stem leaves usually with narrow base, if w ider, not auriculate. 3. Flowers 5-8 mm wide; stem leaves linear-lanceo­ late, widest and parallel-sided at the middle, usually somewhat appressed to the stem; in­ florescence and calyx with at least some woolly hairs; plants of open habitats above 2700 m alti­ tude. subsp. coeruleum - Flowers up to 4 mm wide; stem leaves ellipticlanceolate, widest at or below the middle, usu­ ally spreading; inflorescence glabrescent. calyx ciliate. with or without bulbous-based hairs; plants of shady habitats generally below 2500 m, or up to 3000 m in disturbed areas and juni­ per forest. 4. Plant without woody base, usually with a single stem from base, branching in the inflorescence; terminal inflorescence with 2 diverging branches

up to 20 cm long in fruit; nutlets generally pale, c 4 mm long, with 2 rows of glochids around the rim. and a well-developed midline of glochids, often also with 1-2 glochids in the open areas be­ tween the rim and midline, subsp. geo met ri cum - Plant usually with woody base, with 2 or more stems from base, sometimes single-stemmed, much-branched above; terminal inflorescence with 2 or 3 branches, diverging or not, up to 12 cm long in fruit; nutlets generally darker, c 3 mm long, with a rim of 1-2 rows of glochids and poorly developed midline of glochids or glochids scattered. subsp. johnstonii subsp. coeruleum Afroalpine grassland, rocky areas, field margins and fallow fields, open juniper forest, woodland; 2700-3400 m. GD GJ SU BA, not known elsewhere. Beals B525; de Wilde & Gilbert 121; Mooney 7835. Generally the plants have bright dark blue flowers, 5-8 mm wide, but sometimes white-flowered plants are found, for example Herbert 19, Evans and Hiller 87. Piffard 99 is said to liave ‘greenish-red to rich brick red petals’ This colour is not recorded from the Flora area by any other collector, and may be a mistake for purple’. The type, Schimper 542, comes from the Simen Mountains and this taxon is an afroalpine endemic of the Flora area. In Kenya, the taxon found in a similar ecol­ ogy is subsp. kenyense Verde. (1991). var. hedbergiorum (Riedf) Edwards, stat. nov. C. hedbergiorum Riedl (1985) - type: SU, Ankober. Ash 1016A (UPS holo.. K iso.). Afro-alpine meadow with Thymus, Trifolium, Helichrysum, field banks, grassland, 2800-3350 m. SU ?BA; not known elsewhere. Ash 3747; Boulos & Amare G. 11664; Gilbert & Phillips 9298. Riedl described C. hedbergiorum based on two col­ lections. the type and Boulos & Amare G. 11664. The nutlets on the isotype of Ash 1016 in Kew do not have the characters of the holotype, clearly seen in the cyber­ chrome copy in Kew, and both Verdcourt and Riedl con­ cluded that the isotype was subsp. coeruleum. It would, therefore, appear that Riedl’s description is based on a mixed collection: var. coeruleum is also found in the same geographical area. More material matching the holotype in UPS lias now been found and the recognition of this taxon appears justified. However, there are speci­ mens in var. johnstonii that have similar, but distinctly smaller, c 2.5 mm long, nutlets, and a dark upper surface with only a few paler glochids along the midline, for ex­ ample. Friis et al. 6608 and Stewart E24 from near Jimma in KF. It has, therefore, been decided to recognize Riedl’s taxon as a variety of C. coeruleum. Sue Edwards 25 from 3100 m on Gaysay Mt, BA, has similar nutlets, but is a much larger plant growing to 1 m; the material from SU is not taller than 50 cm. subsp. johnstonii (Bak.) Verde., in FI. Trop. E. Afr., Boraginaceae: 109(1991); C. johnstonii Bak (1894) - type: Tanzania. Kili­ manjaro, 1800 m, Johnston s n (K holo.. BM iso ).

157. BORAGINACEAE: 10. Cynoglossum

Acacia woodland, degraded juniper forest, by paths, roadsides, rocky areas, in fallow land and as a weed; 1800-3000 m. TU GD GJ SU IL KF B A; E Africa, Cam­ eroon, Dem Rep. Congo, Malawi, Angola. Boulos 11353; Friis et al. 6608; Mooney 8488.

3. C. alpinum (Brand) Riedl (1985); C. montanum L. var. alpinum Brand, in Engler Pflanzenr. IV.252 (78): 127 (1921) - type: Ethiopia, Schimper 49 (B holo. destroyed, K W iso ); GD, Mt Guna, Schimper 1194 (B destroyed, K W iso.).

Short or long-lived perennial with a tap root; stems 20-50 cm tall, branched mainly in their upper part, cov­ ered with woolly, soft, more or less spreading hairs. Leaves oblong to lanceolate, 3-15(-20) x 0.4-2 cm, apex usually acute, covered on both sides with soft ap­ pressed hairs, more numerous below, giving a silky ap­ pearance; rosette and lowermost stem leaves with petioles, upper leaves sessile with narrow bases, de­ creasing in size from base to inflorescence. Inflores­ cence much branched, sometimes with one or two bracts; pedicels 1-2 mm long in flower, elongated to 5-8 mm in fruit, bent downwards. Calyx densely woolly, white or grey, 3.5 nun long, divided for 2/3 or more into ovate or linear-lanceolate, acute lobes, about 4 x 2 mm in fruit. Corolla dark bright blue, 3 nun across, cylindrical to bell-shaped; tube about as long as calyx or slightly shorter, lobes cup-shaped; scale-like append­ subsp. latifolium Verde., in FI. Trop. E. Afr., ages crescent-shaped, inserted above stamens. Style Boraginaceae: 112(1991) type: Kenya, Northern Frontier Province, Mt about 1.5 mm, pyramid-shaped, without distinct stigma. Nutlets free from style, 4.5-5 mm long, ovate, with a Kulal, Bally 5549 (K holo., EA iso.). flattened central area surrounded by glochidiate hairs Habitat not given for the Flora area; in Kenya, ever­ with wide, triangular bases, also with similar spines that green forest, thickets derived from formerjuniper forest, are not so much widened towards base. rocky places in grassland; c 1700 m. SD; N Kenya. Gil­ Shade of Juniperus and Erica-, 2000-3250 m. GD SU bert 1333. BA; Somalia. Coady K43; Gillett 5368. Gilbert 1333 is the nearest to subsp. latifolium in the 4. C. densefoliatum Chiov. (1911) Flora area, but the fruits are not mature. The taxon de­ type: GD, at Fasil-Odos in the river Caha, scribed by Verdcourt is confined to a small area on Mt Chiovenda 2411 (FT holo.). Kulal in N Kenya. Further specimens may be found in future collections from SD. Perennial 30-50 cm tall; whole plant densely covered

Subsp. johnstonii is generally much-branched, with inflorescences from most axils of the upper leaves, as veil as terminal; the flowers are generally bright blue and small, 2-2.5 mm wide. There are a number of speci­ mens with small blue flowers and much-branched inflo­ rescences like subsp. johnstonii but having linear lanceolate leaves with soft hairs like subsp. coeruleum-. 4ooney 4751 from dry fallow (SU), E.F. Gilbert 487 from open, sunny areas in juniper forest (SU); Parker E.343 as an occasional weed (GD), Mooney 5151 in grass by a small water channel (AR), Mooney 4837 from •vaste ground (GJ); de Wilde & de Wilde-Duyfjes 7896 in rocky, grassy place (SU). These possibly represent hy­ brids.

subsp. geometricum (Bak. & Wright) Edwards stat. nov. C. geometricum Bak. & Wright (1905); C. lanceolatum Forssk. subsp. geometricum (Bak. & Wright) Brand, in Engler, Pflanzenr. IV.252 (78): 140 (1921); Paracynoglossum geometricum (Bak. & Wright) Mill (1984) - type: Malawi, Mt Chiradzulu, Whyte s.n. (K lecto.). C. lanceolatum Forssk. subsp. geometricum (Bak. & Wright) Brand, var. mannii (Bak. & Wright) Brand, loc. c i t C . coeruleum A. DC. subsp. john­ stonii (Bak.) Verde, var. mannii (Bak. & Wright) Verde., in FI. Trop. E. Afr., Boraginaceae: 110 (1991); C. mannii Bak & Wright (1905). Open woodland, forest margins and paths, with clumps of bushes in grassland; 1750-2250 m. KF SD BA; through E Africa to Cameroon, and south to S Af­ rica. E.F. Gilbert 380; Friis et al. 8232; Mooney 9152. This is the most widespread of the subspecies of C. coeruleum. Verdcourt treated it as var. mannii under subspi johnstonii, which has a similar geographic distri­ bution. The linking character was the elliptic-ovate leaves. Ecologically, these two taxa are distinct in the Flora area with subsp. geometricum only occurring in S Ethiopia where it is confined to more moist and shady habitats.

with soft, white hairs; stems ascending, stout. Leaves very numerous on the stem, densely crowded, especially round the middle, lower ones reduced to a few dry re­ mains at flowering time, sessile with narrow or broad base, lanceolate, acute, 3-10 x 0.5-1.5 cm, widest at or below middle. Inflorescences axillary and terminal, of­ ten several together, flowering before elongating, pe­ duncle elongating up to 12 cm in fruit; pedicels 1-2 mm long in flower, elongated to c 6 mm and curved down­ wards in fruit. Calyx 3 mm long in flower; lobes free to base, oblong, obtuse, enlarged to 5 x 2-2.5 mm in fruit, persistent after shedding of nutlets. Corolla blue, up to 8 mm across, with more or less distinct darker, or lighter, tube, as long as or slightly shorter than calyx; lobes broadly ovate, rounded, 3 x 2-2.5 mm; scales in throat semi-orbicular. Style 1-2 mm long, without distinct stigma. Nutlets equally covered by dense, conical glochidiate hairs all over, ovoid, (3.5-)4 x 3.5 mm, at­ tached to gynobase. Grassland, bushland and pasture, among rocks, on limestone; 1150-3000 m. SU SD BA HA; not known elsewhere. Ash 46; Gilbert 1420; Gillett 5160. 5. C. lanceolatum Forssk. (1775); C. lanceolatum Forssk. subsp. eulanceolatum Brand in Engler, Pflanzenr. IV,252 (78): 139(1921) — type: Yemen, Hadie, Forssk&l s.n. (C holo.).

157 BORAGINACEAE: 10. Cynoglossum, 11 Brandella, 12. M icroparacaryum 94

C. micranthum Poir. (1811). C. abyssinicum Hochst. et Vatke ex Engl. (1892), nom. nud. Herb 20-70 cm high, with or without a woody base, with one stem, usually branched mainly above the mid­ dle, base covered by remains of dried leaves, covered with appressed or slightly ascending soft but rigid liairs. Lowermost leaves stalked, others sessile with usually narrow base, lanceolate, oblong-lanceolate or linearlanceolate, acute, 3-12 x 0 .5 -3 (^ .5 ) cm, covered with appressed hairs on both sides, but more dense on the up­ per side and along the veins of the lower side, hairs sometimes bulbous at base. Inflorescence much branched, terminal cymes nearly always in pairs, strongly divaricate and rigid at fruiting time, without bracts. Flowers numerous; pedicels slender, short, about 1-2 mm long, often recurved in fruit. Calyx-lobes 5, free to base, 1.5(-2) mm long, ovate-lanceolate or ovate-obtuse, hairy Corolla usually white, sometimes blue, 2-2.5 mm across, bell-shaped; tube about as long as calyx; lobes broadly rounded; scale-like appendages inserted in the throat above stamens, much wider than long. Style veiy short, narrowly pyramid-shaped, with­ out distinct stigma. Nutlets attached to style by an awn­ like structure, ovoid, 1.5-2 mm long, densely covered with glochidiate hairs equally distributed all over the surface. Generally disturbed areas in fallow and cultivated fields, evergreen bushland, grassland, by paths; 14502400 m. TU GD GJ SU WG KF GG SD BA HA; wide­ spread in Africa south of the Sahara; tropical Asia, northwards to Pakistan, India. China and Japan. Albers 61069\Ash 856; Friis & Lawesson 5351. 11. BRANDELLA R.R. Mill {1986) Adelocaryum Brand (1915) Mill, Notes Roy. Bot. Garden Edinburgh 43: 477-480 (1986); Riedl, Linzer Biol. Beitr. 24/1: 19-27 (1992).

Annual or perennial hairy herbs. Flowers with 5 scale­ like appendages in the throat. Filaments inserted slightly below scales but tips of anthers surpassing them. Nutlets 4, surrounded by the wing-like, more or less strongly in­ curved margin bearing spines or glochids on its outer surface or by an indistinct wing with marginal glochids. A monotypic genus in NE Africa and tropical Arabia. Very similar in general habit to Cynoglossum, and closely resembling it in fruit and flower characters. B. erythraea (Brand) R.R. M ill (1986); Adelocaryum erythraeum Brand (1921); Cyno­ glossum erythraeum (Brand) Reidl (1971) - type; EE; east of Amba-Tokhan, 398 m. Schweinfurth & Riva 615 (B syn. destroyed, G syn.), upper part of Mogod Valley, 1400 m, Schweinfurth & Riva 1594 (B syn. destroyed, G syn.). Herb; stems hispid, 10-70 cm high. Leaves sessile or shortly stalked, oblong-lanceolate, oblong-elliptic or ovate-lanceolate, 2-8.5 x 0.6-2.2 cm, mostly acute, pat­ ent or more or less appressed hairy, hairs sometimes on tiny calcareous tubercles, rather sparse. Inflorescence

with up to 30 flowers, usually less, fairly lax, elongating after flowering; pedicels 3-4 mm long in flower, 5-10 mm in fruit, slightly deflexed. Calyx divided to the base, 1.2-2 mm long in flower, 3-3.5 mm in fruit stiffly hairy outside, with narrow' lobes. Corolla pale blue to white, 2-4 mm long, 5 mm across, limb as long as tube. Scale­ like appendages in the throat semilunar. Style about 1 mm long. Nutlets 4, broadly ovate in outline, 3-5 x 2.5-4 mm; wing-like margin variable: either strongly in­ curved with very short glochids outside, or narrower, saucer-shaped, with long, flattened glochids on its very edge, scarcely glochidiate outside; disc densely covered with short, slender glochids. Habitat not recorded; 350-1550 m. EE EW GD; Su­ dan; Arabian Peninsula. Fiori 1529; Pappi 3050. 8338. Riedl recognized two forms: f. erythraea with the wing of the nutlets strongly incurved and bullate, or nar­ row and saucer-shapcd, and f. subexalata (Riedl) Riedl (based on A. erythraeum f. subexalata Riedl (1992)) with the wing of the nutlets indistinct, very narrow and nearly flat with m arginal glochids. The type of f. subexalata is Schimper 362 collected from GD (Dschadscha). Material in the Florences Herbarium similar to f. erythraea have also been labelled Cynoglossum hochstetteri Vatke var. calathiforme Chiov. 12. MICROPARACARYUM {Popov ex Riedl) Hilger & Podlech (1985) Paracaryum (DC.) Boiss. sect. Microparacaryum Popov ex Riedl (1967) Annual heifos. Inflorescence lax, without bracts. Calyx divided to base. Corolla funnel-shaped to campanulate, with 5 scale-like appendages in the throat. Stamens in­ cluded in the corolla. Style without a distinct stigma, on a narrowly conical gynobase. Nutlets variable: some with a distinct involute, membraneous or leathery wing; or w ing very narrow, not or slightly curved, central area smooth or with small spines or glochids, whole ventral surface attached to the gynobase. A genus with 2-5 species mainly found in W and S W Asia; 1 in the Flora area IVL intermedium (Fresen.) Hilger & Podlech (1985); Cynoglossum intermedium Fresen. (1834); Omphalodes micrantha DC. (1846); Paracaryum mi­ cranthum (DC.) Boiss. (1849); Paracaryum inter­ medium (Fresen.) Lipsky (1910). Annual, with one or several stems, 5-20(-30) cm high, covered with spreading hairs, stems slender, ascending or upright, branched. Leaves mainly near base, linear-spathulate or oblong, 20-40 x 3-6 mm, decreasing in size towards inflorescence. Inflorescence lax, some­ times with 1-2 bracts near base; pedicels very slender, the lowermost 5-15 mm long, others often recurved Ca­ lyx 1.5-2.5 mm long; lobes oblong-ovate. Corolla campanulate, 1.5-2.5 mm long, blue; limb as long as tube. Scale-like appendages horizontally sickle-shaped or nearly quadrate. Stamens inserted in middle of tube. Style 0.5-1 mm long. Nutlets rounded, 3 mm across, central area smooth or with short warts or small spines.

157. BORAGINACEAE: 12. Microparacaryum, 13. Cystostemon, 14. A mebia

wing membraneous, often bluish-suffused, with almost entire margin. Habitat and altitude not known. EE; Egypt, Somalia; Iraq, Iran. Afghanistan. Pakistan. Caspian area. Central Asia to Tien-Shan, Arabian Peninsula. Sinai. Terracciano & Pappi s.n. This species is included based on the record in Cufo­ dontis. No further material of this species from the Flora .irea lias been seen by either author. 13. CYSTOSTEMON Balf. f (1883) Vaupelia Brand (1914) Miller & Riedl, Notes Rov. Bot. Garden Edinburgh 40 (1): 1-21 (1982).

Herbs or small shrubs. Leaves alternate, entire. Inflores­ cence simple or branched, with bracts. Calyx divided to near base into 5 lobes. Corolla with 5 lobes that are longer than the naked tube and spreading at flowering. Filaments inserted in the corolla tube, sometimes in­ flated, with a basal triangular or oblong appendage sometimes strongly reduced, or with a ciliate, thickened ridge. Anthers longer than filaments, with a terminal, sterile, awn-like appendage. Style elongated; stigma capitate. Nutlets 1 to 4, triquetrous to pyramidate, straight and erect. A genus with 13-15 species in NE to SW Africa, Arabia and Socotra; 2 species in the Flora area. 1. Bristles on stem spreading; annual or perennial herb, rarely shrubby. 1. C. virescens - Bristles below inflorescence appressed; shrubs or subshrubs. 2. C. ethiopicus

95

silky. Ovary glabrous. Style up to 12 mm long. Nutlets not seen. Limestone with Commiphora - Acacia scrub. Com­ miphora woodland; c 900 m. SD; Kenya. Gilbert & Thu­ lin 1492. 2. C. ethiopicus A.G. Miller & Riedl (1982) - type: SD. on the road from Negele to Mega, Thulin, Hunde & Mesftn T. 3572 (UPS holo., ETH K iso.). Shrub or subshrub; whole plant scabrous, yellowish grey. Stems up to 80 cm high, erect or ascending, sparsely branched, covered with antrorse, appressed, tu­ berculate, strigose hairs and shorter, spreading hairs. Leaves narrowly elliptic to elliptic-oblong or nearly spathulate, sessile, 2-6 x 0.3-1.8 cm, margin sometimes revolute, apex acute or obtuse, hairs similar to those on stem, fascicles of smaller leaves often present in axils of the stem leaves. Inflorescence terminal or axillary, 3-9flowered, slightly elongating after flowering. Bracts leaf-like, but smaller. Pedicels (2-)7-9 mm long, erect to spreading. Calyx divided to the base; lobes 6-7 mm long, slightly enlarging in fruit, hairs similar to those on stem, inner surface woolly to silky. Corolla white with pale green centre, 12 mm long; tube 3.5 mm, pubescent between filaments; lobes ovate, narrowing into long ta­ pering linear tips, hairy outside. Stamens 9.5 mm long; anthers 4.5 mm long, tuberculate on back, terminal ap­ pendages about 5 mm. Filaments short, attached 2 mm above base of tube, not much widened; appendages small, woolly. Ovary glabrous. Style up to twice as long as calyx. Nutlets triquetrous, white, with an oblique, acute tip and humped back, minutely warty, up to 3 mm long.

Acacia - Commiphora woodland on limestone, lime­ 1. C. virescens A.G. Miller & Riedl (1982) stone slopes with Pistacia, also Combretum-Terminalia type: Kenya, Northern Frontier Province, Gillett 13267 (K holo.). woodland; 1500-1650 m. SD; not known elsewhere. Thulin et al. 3576. Annual or perennial herb, sometimes shrubby. Whole plant greenish yellow, slightly hispid. Stems 10-40 cm tall, erect, simple or sparsely branched, hispid with spreading, tuberculate hairs and shorter, dense, spread­ 14. ARNEBIA Forssk. (1775) ing hairs. Base sometimes clothed with withered re­ Arnebiola Chiov. (1929) mains of dry leaves. Leaves lanceolate, 2.5-8 x 0.3-1.3 Annual or perennial herbs. Leaves alternate, often with cm, margin revolute and thickened, entire, base longbristly hairs. Inflorescence with small, leaf-like bracts, tapering, upper leaves sessile, lower ones stalked, apex elongated after flowering. Calyx divided either near to acute; hairs similar to those on stem, but comparatively the base or with a short, coherent basal part. Corolla sparse Inflorescence simple or composed of peduncu­ funnel-shaped or with flat upper part, yellow or purple, late cymes, terminal and axillary. Bracts leaf-like, brac­ naked inside, with a ring-like nectary at base of corollateoles sometimes covering the calyx; pedicels erecttube. Stamens either inserted at the same level or at vari­ spreading, shorter than calyx. Calyx 7-10 mm long; ous heights, in some cases heterostyly observed. Style lobes lanceolate to linear-oblong, outside with a few forked once or twice; stigmas capitate. Nutlets 4, scattered antrorse bristles amongst loosely arranged anovoid-pyramidal, sometimes rugose or warty. trorse hairs, inside with dense longer bristles or hairs. Corolla greenish-cream, lobes paler, tube 2.5-3 mm A genus with 25-30 species, distributed mainly in the long, pubescent between filaments; lobes 7-10 mm temperate and subtropical regions of Asia, a few species long, curved upwards, ovate, narrowing sometimes also in N Africa; 2 in the Flora area. abruptly into linear tips 3.5-5.5 mm long, outer surface Arnebia is similar in most characters to Lithosperof tips hairy. Stamens 8-10 mm long, purplish; anthers mum, differing mainly in the pollen grains, the pores of 3.5-5 mm, slightly tuberculate on outer surface, termi­ which are arranged in two series. nal appendage 5-6 mm long; filaments short, attached 1. Flowers yellow. 1. A. hispidissim a 1.5 mm above base of tube, not much widening; basal - Flowers purple. 2. A. purpurascen s appendage inconspicuous, up to 0.5 mm long, woolly to

157 BORAGINACEAE: 14. Amebia, 15. Lithospermum, 16. Buglossoides

1. A. hispidissima (Sieber ex Lehm.) DC. (1846); Anchusa hispidissima Del. (1813) nomen nudum; Lithospermum hispidissimum Sieberex Lehm. (1821) - type: Arabia, Wadi Gamuhs, Sieber s.n. (W holo.). Toxostigma lutea A. Rich. (1851) - type: Abys­ sinia without locality, A. Petit s.n. (P? holo.). Arnebiola migiurtina Chiov. Annual, or sometimes perennial with woody base, 5-50 cm tall, covered with long, rigid, more or less appressed hairs all over, and also with very short hairs. Stem often branched. Leaves 1-5 x 0.2-0.7 cm, lanceolate cr linear-lanceolate, acute. Cymes with bracts, single or in pairs, dense and short when young, elongated later on; pedicels almost lacking. Calyx 5-8 mm long in flower, only slightly increasing later, divided almost to the base; lobes linear. Corolla yellow, 10-15 mm long, hairy out­ side, glabrous or downy inside, with a ring-shaped, haiiy, basal nectary; tube three times as long as upper part, cylindrical, upper part flat or funnel-shaped with broadly rounded lobes. Flowers heterostylous; style simply bifid with 2, often bilobed, stigmas. Nutlets 4, 1.5—2 mm long, ovoid-pyramidate, smooth or with small warts.

varying degree. Nutlets 4, rarely less by abortion, en­ tirely smooth and shiny . A genus with about 60 species, mainly in temperate regions, particularly N America; 1 in the Flora area. L. afrom ontanum Weim. (1940) - type: Zimbabwe, Norlindh & Weimark 5070 (LD holo., K iso ). L. officinale L. var. abyssinicum (Vatke) Engl., in Hochgebirgsfl. Trop. A fr:. 355 (1892), nomen nudum

Dry habitats, rocky outcrops, waste land, 300-2100 m. EE EW TU; widespread in N and NE Africa; Arabia, Iran to Tibet. Bally 6% 16; Friis et al. 6718; Ryding 1449.

Perennial, 75-100 cm tall. Stem upright, hairy, branched in the upper part. Leaves decreasing in size from the base upwards, lower ones 6-8 x 2.5-3 cm, elliptical, only the uppermost stalked, others sessile with broad base, roughly hairy with mostly appressed, s tiff hairs, green on the upper, greyish-green on the lower side. Cy­ mes many-flowered, lax, up to 30 cm long, bracts sub­ sessile; pedicels 1-2 nun long in flower, up to 7 mm in fruit. Calyx divided nearly to the base; lobes tapering to­ wards apex, 4.5-5.5 mm long in flower, up to 10 mm in fruit, roughly hairy. Corolla yellow, tube 7-8 mm long, glabrous outside, papillate inside on trapezoid eleva­ tions; lobes free, 2.5-3 mm long, broadly rounded. Sta­ mens inserted at upper end of tube. Style up to 1.5 mm long. Nutlets 3-3.5 mm long, ovoid, slightly pointed at apex, smooth. Fig. 157.14.

2. A. purpurascens (A. Rich.) Bak. (1906); Toxostigma purpurascens A. Rich. (1850) - type: TU, Adowa, Quartin-Dillon & Petit 87 (K P syn.).

Afromontane Erica scrub, open forest, wooded grassland; 2800-3200 m. SU BA HA, E Africa, Malawi, Zambia, Zimbabwe, S Africa. Hedberg 5512; Mooney 7266; Sue Edwards 4.

Herb with woody base, about 15 cm tall. Stems and other parts covered with rigid, bristly, white hairs. Leaves linear to oblong, spathulate, 3-3.5 x c 0.5 cm, covered with appressed or slightly spreading bristly hairs arising from white tubercles. Cymes numerous, terminal or axillary, many-flowered, dense, bracts lan­ ceolate. Calyx 4-4.5 mm long, divided to the base into 5 narrowly linear lobes. Corolla purple, hairy outside, glabrous inside with basal ring-shaped nectary , 8 mm long; tube cylindrical, about 5 nun long, upper part funnel-shaped, cleft to the middle into 5 broadly rounded ciliate lobes. Flowers heterostylous; style forked; stigmas 2, capitate or club-shaped. Nutlets 4, 1.5 cm long, acute, with ovate, convex outer and angular in­ ner face, whitish, covered densely with tiny warts. Habitat not given; 1900 m. TU; only known from the type collection. A. purpurascens appears to be a colour form of the widespread A. hispidissima. No other material of this taxon has been collected, despite its striking flower col­ our. 15. LITH OSPERM UM L. (1753, 1754) Usually perennials. Leaves alternate, lanceolate to lin­ ear, entire. Inflorescence with bracts, dense in flower, then becoming loose. Calyx usually divided to the base into 5 narrow lobes. Corolla funnel-shaped or campanu­ late, with 5 tiny velvety elevations or 5 velvety folds in the throat. Stamens inserted in tube of corolla. Style fili­ form. bearing two stigmas which are often united to a

The species looks superficially similar to L. offici­ nale L. with which it has been confused. L. afromonta­ num is a woody perennial with very different pollen 16. BUGLOSSOIDES Moench (1974) Johnston tnJourn. A m Arb. 35: 38— 46 (1954).

Annual herbs covered with setulose to strigose hairs; stems usually much branched. Cymes terminal, with bracts. Calyx divided almost to the base, enlarging in fruit. Corolla blue or purple to white; throat with 5 lon­ gitudinal lines of hairs, without scales. Stamens with filaments inserted below the middle o f the corolla tube; anthers not exserted. Nutlets obovate or ovoid with a curved or almost straight beak, tuberculate to granular. A genus with up to 7 species most in Europe and SW Asia; 1 species, a widespread weed, recorded for the Flora area B. arvensis (L .) I.M. Johnston (1954); Lithospermum arvense L. (1753) - type: Europe, Herb. Linn. 181/4 (LINN syn.). Annual. 5-50 cm tall. Stems solitary or several together, with a few branches, covered with more or less ap­ pressed bristly hairs. Leaves oblong, oblong-spathulate to linear, acute with hairs like those on the stem. Cymes solitary or paired; flowers on short pedicels. Calyx di­ vided to the base; lobes linear to linear-lanceolate, acute, often slightly unequal. Corolla 7-8 mm long, campanulate to cylindrical-campanulate, cream-white, rarely pale blue, limb about as long as tube, with broadly

157. BORAGINACEAE: 15. Lithospermum, 16. Buglossoides, 17. Echium

91

Figure 157.14 LITHO SPERM U M AFRO M O NTAN U M 1 - flowering stem x Vy, 2 - detail of upper surface o f leaf x 16; 3 - detail of lower surface o f leaf x 16; 4 - flower x 6; 5 - corolla opened out x 6; 6 - detail of outer sur­ face of corolla x 6; 7 - ovary x 18; 8 longitudinal section through base of pistil x 18; 9 - stigma x 18; 10 - fruit­ ing calyx x 6; 11 - nutlet x 6. 1-3 from Greenway & Kanuri 15048; 4-11 from Verdcourt 3775. Drawn by Maureen Church. (Reproduced with permission from FI. Trop. E. Afr., Boraginaceae: fig. 20.)

rounded lobes; corolla inside with 5 longitudinal lines of hairs, without scales. Stamens inserted below middle of corolla-tube. Style short. Nutlets 4, c 3 mm long, ovoid in outline, with a short beak, brownish, wrinkled, with netlike folds, sometimes also with additional warts. Fig. 157.15. Weed in farmland; c 2100 m. TU GD; widespread in most parts of Europe and SW Asia, and now introduced to most temperate areas of the world. Friis et al. 10698; Schimper s.n. 17. ECHIUM L. (1753, 1754) Annual or perennial hispid herbs or shrubs; bristles in all parts with tubercles at base, sometimes short, stiff,

appressed hairs also present. Inflorescence of scorpioid, lateral cymes, often greatly elongated in fruit. Calyx deeply divided into 5 lobes, enlarging in fruit. Corolla blue, purple, yellow or white, tube broadly to narrowly funnel-shaped, either with 5-10 distinct scales or with a collar-like ring at base; throat oblique, limb oblique with distinctly or slightly unequal lobes, shorter than tube. Stamens 5, often unequal, included or exserted. Style exserted, stigma bifid or capitate. Nutlets 4. more or less triangular at base, usually rugose. A genus with about 45 species, 27 endemic to M a­ deira, Azores, and the Canary Islands, the rest in the Mediterranean region and other parts of Europe; 1 natu­ ralized species in the Flora area.

98

157. BORAGINACEAE: 16. Buglossoides, 17. Echium

Figure 157.15.

BVGLOSSOIDES ARVENSIS subsp ARVENSIS: 1 - flowering stem x 2/ 3; 2 detail of upper leaf surface x 8 : 3 flower x 6; 4 - corolla opened out x 8; 5 - pistil x 30; 6 - ovary, longitudinal scction x 30; 7 - young fruiting calyx x 3 8 - nutlet x 10. All from Abdallah 520. Drawn by Maureen Church. (Re­ produced with permission from FI. Trop. E. Afr. Boraginaceae: fig. 21.)

Many species are very striking plants and several have been cultivated in E Africa (see Verdcourt loc. cit. : 81). It is likely that other species may also be grown in public and private gardens in Addis Ababa and other towns in the Flora area, but no herbarium material of cul­ tivated species is available. E. plantagineum L. (1771) type: Barrelier, Plantae per Galliam exhibitae: 145, t. 1026 (lecto. selected by Gibbs). Erect herb, 20-65 cm tall, with single to many stems. Leaves at base ovate to oblanceolate, 5-28 x 1-3.5 cm, appressed setose, with distinct lateral veins; stem-leaves oblong to lanceolate, the upper cordate or truncate at

base, with appressed, rather soft hairs. Cymes stalked, 25-30 cm long after flowering. Calyx 0.7-1.2 cm long in flower, up to 1.7 cm in fruit. Corolla blue, turning purple, then pink, finally white, 1.8-3.2 cm long, hairy along veins and margins; tube with 10 distinct, hairy lobes in a ring. Lower pair of stamens exserted, the oth­ ers included or only slightly exserted, filaments with long woolly hairs. Styles included, densely hispidulous below, bifid for about 1 mm. Nutlets black or greyishbrown. tuberculate and faintly striate, ovoid-obpyramidal, 2.3-3 x 2-2.5 mm, keels prominent on ventral and dorsal faces; beak short, suberect. Fig. 157.16. Among rocks, beside shallow water, in overgrazed grassland; 2200-2350 m. EW GD SU; native of W

157. BORAGINACEAE: 17. Echium, 18. Anchusa 99

Figure 157.16 ECH IU M P L 4N T A G IN EU M : 1 - flowering stem x Vy, 2 - basal leaf x Vy, 3 - detail o f upper leaf surface x 8; 4 - flower x 2; 5 - part of corolla opened out to show stamens, and style and stigma x 3; 6 - detail of base of corolla x 12; 7 - ovary x 24; 8 - longitudinal section through base of pistil x 24; 9 - part o f style w'ith stigma x 24; 1 0 -fru itin g ealy x x 4 ; 11 - nutlet x 10. 1-3 from Thurston s.n.; 4-11 from K783. Drawn by Maureen Church. (Reproduced with permission from FI. Trop. E. Afr., Boraginaceae: fig. 22.)

Europe, now widespread throughout Europe, S Russia and the Caucasus; naturalized in Kenya, Tanzania, Zim­ babwe, Angola. Friis & Lawesson 5348; Gilbert et al. 8842; Pichi-Sermolli 2129.

rotate limb, sometimes tube sharply bent; lobes unequal in size; throat with scales. Stamens inserted at the same or at different levels. Style filiform; stigma capitate or bilobed. Nutlets 4, upright or nearly horizontal with apex pointing to the centre, with a big basal or ventral attachment-area that is surrounded by a thickened rim.

18. ANCHUSA L. (1753, 1754) Lycopsis L. (1753, 1754)

About 50-55 species in Europe, Asia and northern Africa; 2 in the Flora area.

Perennial or annual herbs. Leaves of various shapes, of­ ten fairly large. Cymes with bracts, usually becoming lax after flowering, bracteate. Calyx with free or basally united lobes, rarely enlarging in fruit. Corolla blue, pink or white, funnel-shaped or with cylindrical tube and flat,

1. Perennial; tube of corolla straight; attachment area of nutlets basal. 1. A. affinis - Annual; tube of corolla bent; attachment area of nutlets ventral. 2. A. ovata

—

157. BORAGINACEAE: 18. Anchusa, 19. Myosotis

100

1. A. affinis R.Br. ex DC. (1846) - type: TU, Gennia in Memsah, east of Adwa. Schimper I: 381 (K iso ). Perennial, 30-60 cm tall; stems stout, herbaceous, usu­ ally simple, covered with slightly retrorse. bristly hairs. Leaves: at base 10-20 x 1.5-2.5(-3) cm, oblonglanceolate or oblong, acute or tapering obtuse; on stem sessile with narrow or wide base, oblong to lanceolate, acute or rarely obtuse 5—10(—12) x 0.7-1.5 cm; all leaves covered with short, more or less appressed. bris­ tly liairs arising from calcareous tubercles on the uppor and along veins of lower side, glabrous or with fewer bristles with or without tubercles on the lower side. Cy mes terminal and lateral in the axils of upper leaves, many-flowered, dense, involute. Calyx nearly sessile, bristly hairy'. 5-6 mm long in flower, 7-8 mm in fruit, much inflated at last, divided to the middle into ovatetriangular, acute lobes. Corolla-tube as long as calyx, straight, limb rotate, blue, 7-10 mm across; lobes obo­ vate or nearly orbicular; scales at the upper edge of throat, densely covered with short hairs, semicircular. Nutlets ovoid, netlike rugose, upright with basal attach­ ment area. In grassy fields, degraded bushland and beside roads, afro-alpine meadows; 2050-3100 m. EW TU WU BA. not known elsewhere. Ash 1650; Friis et al. 10540; Ryding & Mengisteab G. 1504. 2. A. ovata Lehm. (1818) - type: Habitat in Caria (MEL holo ). Lycopsis orientalis L. (1753); A. orientalis (L.) Rchb. (1858). A. milleri auct. non Willd. A. ryssosperma Steud. nom. nud.

1

Annual, 10-50 cm tall, all parts covered by bristles aris­ ing from white tubercles. Stems branched or simple, stout. Leaves up to 1 2 x 2 cm, basal leaves stalked, oth­ ers sessile, lanceolate or oblong-lanceolate, acute or ob­ tuse. margin entire or with a few shallow teeth. Flowers in the axils of upper leaves, on long pedicels. Calyx 5-7 mm long in flower, up to 12 mm in fruit; lobes free to the base, linear-lanceolate, spreading. Corolla 10-17 nun long, pale blue or white; tube short and bent, more or less irregular, limb divided into broadly rounded lobes; throat with scales covered with minute hairs or papillae. Nutlets with an almost ventral attachment area, transversely ovoid, with a dorsal keel, net-like rugose, additionally with fine warts. Newly cultivated fields and irrigated cultivation; 2000-4000 m. TU GD; widespread in E Europe, SW Asia to Himalayas. Schimper 247, II: 740, II: 1142.

No recent collections of this species have been seen. Colville 1 in K, collected from EW (Asmara), is la­ belled A “italica L. This is a large plant nearly 1 m tail with very large flowers. It is probably an introduced or­ namental. 19. M YOSOTIS L. (1753) Herbs, all parts usually covered with white hairs. Leaves lanceolate to oblong, entire. Inflorescences with or with­

out bracts, scorpiod cymes tightly curled up when young, often lengthening into a false raceme with only the upper part curled. Calyx bell-shaped, lobes 5, short or long, persistent. Corolla blue, pink, or white; lube narrow, throat usually w ith scales, often yellow; lobes 5, usually spreading. Stamens 5, with usually very' short Filaments and ellipsoidal anthers, attached at or below middle of tube. Ovary 4-lobed; style usually short, stigma undivided, capitate or depressed capitate. Disc absent. Fruit of 4 nutlets, smooth and shiny, usually slightly compressed dorsiventrally, ovate in outline, with or without a narrow, flattened, wing-like margin in the upper part, acute or obtuse, with very small, basal at­ tachment area, with or without an areole. A worldwide genus of temperate regions with about 50 species, some species also found at higher altitudes in tropical mountain areas; 3 species in the Flora area. 1. Calyx with long spreading hairs, sometimes hooked at their ends; corolla tube longer than calyx; plants usually growing for only one season. 1. M. abyssinica - Calyx with appressed liairs, not spreading or hooked, or caly x nearly glabrous; corolla tube as long as or slightly shorter than calyx; plants usu­ ally growing for more than one season. 2 2. Erect plants, over 5 cm tall, sometimes with sto­ lons; cymes elongating beyond the leaves. 2 . M. vestergrenii - Cushion plants, less than 5 cm tall, cymes hidden in axils of leaves 3. M. keniensis 1. M. abyssinica Boiss. & Reuter (1849) - types: GD, Simien, Demerki, Schimper 11:1146 (G syn., BM P S W isosyn.); M. hispida Schlecht. var bracteata A. Rich., in Tent. FI. Abyss. 2: 88 (1850) TU, Mt Scholoda, Schimper III: 1889 (G syn., BM K P isosyn). M. aequinoctionalis Bak. (1894). Slender heib; stems 5-28(-42) cm tall, with short, spreading and appressed, bristly hairs. Leaves 1-2 cm long, oblong or oblong-lanceolate, lower ones with a short stalk, upper ones sessile. Flowers small, on short pedicels, with bracts in their lower part. Scorpiod cymes compact when in flower, elongating when in fruit. Calyx about 3 mm long, densely covered with spreading hairs, some with hooked ends, divided to the middle into 5 lobes, narrow, acute. Corolla bright or pale blue, or white, bell-shaped, tube a little longer than calyx, lobes opening to 2-2.5 mm across. Fruit with persistent calyx. Nutlets usually 4, elliptical, brown or black, shining, with rounded apex. Fig. 157.17.1-8. In soil pockets on rocks, banks, beside paths, open patches in grassland, in shade; 1800-3800 m. TU GD GJ WU SU AR GG BA; E Africa, Sudan, Cameroon, Bioko (Fernando Po), Rwanda, Dem. Rep. Congo. Gilbert & Phillips 9194; Hedberg 4151; Mooney 8100. 2. M. vestergrenii Stroh (1941) - type: GD, Simien, Demerki, Schimper 1152 (P lecto., BM K S W isolecto.).

157. BORAGINACEAE: 19. Myosotis 101

Figure 157.17 M YO SO TIS A B YSSIN IC A : 1 - flow­ ering plant x Vy, 2 —flower x 8; 3 - co­ rolla opened out x 8; 4 - anther x 18; 5 - stigma x 24; 6 - young fruiting ca­ lyx x 10; 7 - nutlet, side view x 12; 8 nutlet, face view x 12. M. VESTERGRENII: 9 - flowering plant x Vy, 10 - flower x 6; 11 - corolla opened out x 6; 12 - anther x 18; 13 - young fruiting calyx x 10; 14 - longitudinal section through base of pistil x 24; 15 - stigma x 24; 16 - nutlet, side view x 12; 17 - nutlet, face view x 12. M KEN IEN SIS: 18 - flowering plant x Vy, 19 - flower x 8; 20 - corolla opened out x 8; 21 —anther x 18; 22 stigma x 24; 23 - young fruiting calyx x 10; 24 - nutlet, side view x 12; 25 nutlet, face view x 12. 1 from Thulin 6 Tidigs 236; 2-8 from Mabberley 381; 9-12, 14 & 15 from Hedberg 971; 13 from Adamson 472; 16 & 17 from Bogdan 4467; 18-25 from Hedberg 4423. Drawn by Maureen Church. (Reproduced with permission from FI. Trop. E. Afr., Boraginaceae: fig. 24.)

M. sylvatica sensu A. Rich (1850) and Bak. & Wright (1905). Robust, sometimes stoloniferous herb; stems 20-40 (-60) cm tall, with short, scattered, spreading bristly hairs Lower leaves oblong-ovate, shortly stalked; upper leaves sessile, elliptic to oblong-lanceolate, 4-5 cm long. Scorpiod cymes veiy compact and pubescent when in flower, elongating when in fruit, without bracts; low­ ermost pedicels much longer than calyx. Calyx 2-3 mm long, divided for at least two thirds into 5 lanceolate lobes, with short, spreading, bristly hairs along the mar­ gins and on the lower part, persistent. Corolla often starting purple in bud, bright blue on opening, fading to pale blue or white, with paler or yellowish scales in the throat; tube as long as calyx, lobes 5, ovate, 2 x 2.2 mm.

Fruit with persistent calyx and 4 nutlets, with indistinct marginal border, oblong-ovate in outline, tapering to­ wards apex, obtuse. Fig. 157.17.9-17. Mostly on moist ground and in seepage areas (marshy or boggy ground); 2800-3900 m. SU GG AR BA; E Africa, Cameroon, S Africa. Ash 2916; Hedberg 5571; Mooney 6985. 3. M. keniensis T.C.E. Fries (1924) type: Kenya, Mt. Kenya, R.E. & T.C.E. Fries 1399 (UPS holo., BR K S iso.). Cushion herb, with creeping rhizome; stems only about 5 cm long. Leaves crowded and very short, forming a cushion; basal leaves oblong to oblong-lanceolate, ob­

102

157. BORAGINACEAE: 19. Myosotis

tuse, narrowing into a wide petiole; upper leaves smaller, oblong, with very short stalk, or sessile. Cymes few-flowered, almost hidden in axils of longer leaves, nearly sessile or with a very short pedicel. Calyx 2.5 mm long in flower, elongated to 3 mm in fruit, divided to about the middle into 5 lobes, densely hairy along the margins. Corolla tube creamy-white, as long as calyx; lobes rounded, pale blue or purple, c 1 x 0.7 mm. Fruit­ ing calyx 1.6 mm wide, with rounded lobes. Nutlets very broadly rounded at apex, shiny brown or black.

elliptical-oblong, with a distinct, tiny, pointed tip Fig. 157.17.18-25. I] Forming cushion-like clumps in grassland; 42004250 m. BA; Kenya. Hedberg 5575, 5588. The species is confined to the highest altitudes of the mountains and its growth-habit may be explained as an adaptation to the environment. The distribution of hairs on the calyx is distinct as is the shape of the apical region of the nutlets.

158. SOLANACEAE

103

by I. Friis* Dammer, Solanaceae africanae I. - Bot. Jahrb. Syst. 38: 57-60 (1905); Solanaceae africanae II. -B o t. Jahrb. Syst. 48: 224-260 (1912). Cufodontis, Enum:. 855-884 (1963). Hepper in El-Hadidi (ed.), Flora o f Egypt, Family 159, Solanaceae (1998); Thulin, S olanaceae in FI. Somalia 3:197-221 (2006). Herbs, climbers, shrubs or small trees; unarmed or armed with straight or curved prickles; glabrous, or with simple, forked, dendritic (hairs with more or less irregular, tree-like branching, see Fig. 158.21.6) or stellate hairs (for various types o f stellate hairs, see Fig. 158.9.4 & 11, Fig. 158.10.3 & 11 and Fig. 158.12.8). Leaves alternate, sometimes paired, but not opposite or clustered towards the end o f branches, and sometimes heterophyllous (adjacent leaves o f different size), without stipules, petiolate or apparently sessile due to blade being decurrent on the petiole, entire, repand, sinuate, toothed, deeply lobed or pinnatifid, and then the leaflets often o f different size. Flowers single in leaf axils, extra-axillary, or in various forms o f cymose in­ florescences which may sometimes appear to be racemose or umbellate. Flowers bisexual or gynodioecious, 4 - or 5-merous, regular or irregular. Calyx with more or less united lobes. Corolla 4- or 5-merous, rarely with a different number o f parts, united, with very variable corolla-forms, from long tubular to open stellate. Stamens with the same num ber as the corolla lobes, some stamens may be enlarged or reduced to stami­ nodes; filaments free; anthers with 2 thecae, opening by apical or subapical pores or longitudinal slits. Ovary with 2 locules, or 4 or more due to formation o f extra septae; style simple; stigma entire or slightly bilobed; ovules numerous. Fruit a berry, drupe or capsule opening by 2 or 4 valves or by a lid (in H yoscyam us). Seeds winged or unwinged. A family o f about 90 genera and 2600 species, distributed throughout the world, but with most species in tropical America. In the Flora area 4, possibly 5, indigenous genera, and 11 introduced, cultivated or natu­ ralized ones. Several new and partly contrasting classifications of this family have been proposed in recent decades, for example the ones by A.T. Hunziker, South A m erican Solanaceae: a synoptic su rvey in J.G. Hawkes, R.N. Lester & A.D. Skelding (eds.), The biology and taxonomy o f the Solanaceae, Linn. Soc. Sym p. Series 7, pp. 49-85 (1979), and W .G. D ’Arcy, The Solanaceae since 1976, w ith a R eview on its B iogeography, in J.G. Hawkes, R.N. Lester, M. Nee & N. Estrada, Solanaceae III. Taxonom y, C hem istry, E vo lu tio n , pp. 75-132 (1991). Here, the author has used the definition and sequence o f genera in R.W. Purdie, D.E. Symon & L. Haegi, F lora o f A ustralia, Vol. 29, Solanaceae (1982). Some authors now include C yphom andra and L ycopersicum in Solanum . The family includes a number o f very important food plants and stimulants, nearly all o f which are intro­ duced to the Flora area from South America, usually via Europe, at various points in history. These include the smaller, spicy chilli peppers ( C apsicum fru tescen s L.: KARYA when green, GWAMIYA when red-green and BERBEREE when red) and the larger, less hot paprika fruits and capsicums ( C apsicum annuum L.). The former, which must have been introduced at a very early point, are extremely im portant com ponents in the mixed hot spice o f traditional cooking the highlands o f the Flora area, the latter, which has probably been introduced later, are used in European-influenced cooking in towns. Not much is known about the introduc­ tion o f the tomato (L ycopersicon esculentum Mill., TEEMATEEM), but it is now widely used throughout the Flora area. The potato (Solanum tuberosum L., DINISH FERENGI), which was introduced in the 1860s or be­ fore, is now widely grown and eaten throughout the highlands. The leaves and fruits o f annual species of Solanum sect. S o la n u m , e.g. S. am ericanum Miller, are locally eaten as a pot-herb: NECH a w i t , when the fruits are unripe, and KEYE a w i t and TIKUR a w i t for the species with red or black ripe fruits, respectively, but our knowledge about their use in the Flora areas is very incomplete. The egg-plant (S. m elongena L.), is a comparatively recent introduction to the Flora area, which was introduced with the pepino (S. m uricatum Ait.) to Eritrea by the Italians by the end o f the 19th Century. The pepino can now be found in several places o f that country. The Cape gooseberry (Physalis p eruviana L., AUTI), are popular with children. Leaves o f N icotiana spp., especially o f cultivars o f N. tabacum , are widely cultivated for locally produced tobacco; lit­ tle is known about the introduction o f tobacco to the Flora area. * The Botanical Museum and Library, U niversity o f C openhagen, G othersgade 130, D K -1123 Copenhagen K, Denm ark. Jennifer M. Edmonds, U niversity o f Leeds, School o f Biology, Leeds, LS2 9JT, U nited Kingdom , contributed the account o f Solanum, and drafted the account o f Discopodium together with the m ain author.

Solanum sect.

104

158. SOLANACEAE

Most species contain alkaloids; species with a high alkaloid content may be poisonous, or used as drugplants or medicinally. This is particularly true for various species o f Datura 1 and Hyoscyamus. A number o f species o f the genera Cestrum, Datura, Lycium , Nicotiana. Physalis, Solarium and Streptosolen are cultivated as ornam entals in town parks or village gardens; they were probably introduced in the 20 Century. A number o f further genera o f Solanaceae have not yet been recorded from the Flora area, but can be ex­ pected in cultivation in parks or gardens or perhaps as occasional escapes or naturalized:

Brunfelsia L. —B. americana L., a shrub from the W est Indies with lanceolate leaves and white or yellow flowers with a long tube and a radiate limb, is frequently cultivated in the tropics and subtropics. Iochroma Benth. - I. coccineum Scheid., I. lanceolatum (Miers) Miers and I. cyaneum (Lindl.) M L. Greene, tropical American shrubs with tubular purple or scarlet flowers, lanceolate leaves and globose ber­ ries, are now widely grown as ornamentals in hot countries throughout the world. Salpiglossis Ruiz & Pavon —S. sinuata Ruiz & Pavon. viscid annual or shortlived perennial herbs from the Andes in South America with multicoloured, funnel-shaped flowers, is frequently cultivated in most parts o f the world. Schizanthus Ruiz & Pavon - the annual S. pinnatus Ruiz Pavon, “poor m an’s orchid”, a glandular­ haired annual from southern Andes in South America with single or double pinnatisect leaves and pink and violet flowers, is frequently cultivated in many places throughout the w'orld Schwenckia L. - S. americana L., a moderate-sized annual or short-lived perennial herb with subsessile or short-petiolate, lanceolate leaves and long, terminal panicle-likc inflorescences with tubular, pale violet flowers and two-valved capsules, has been used as a fish poison and is now widely naturalized in tropical Af­ rica from Senegal to S Sudan and south to Angola and Zambia Solandra Swartz —S. grandiflora Swartz, a climbing shrub from the W est Indies with evergreen .lanceo­ late leaves and large white, fimnel-shaped flowers, is frequently grown in gardens in warm countries throughout the world. Key to genera

9 . Herbs.

1 Anther dehiscing by terminal pores or short apical slits. 2 - Anthers dehiscing by slits running the entire length of the loculi. 3

~ Shfubs. 2. N icotiana 10. Calyx with segments fused for about half their length or more, capsule with 4 valves (rarely 2). 2. Nicotiana - Calyx with segments free almost to base: capsules with 2 entire or bifid valves. 3. P etu n ia 11. Leaves pinnatisect. anthers with terminal append­ ages. 9. Lycopersicon - Leaves entire or lobed; anthers without terminal appendages 12

2. Connective large, conspicuous at back of anther. 7. C yphom andra - Connective small, inconspicuous at back of an­ ther. 8. Solanum 3. Fertile stamens 4. 4 - Fertile stamens 5. 5 4. Annual herbs. - Woody plants (shrubs or subshrubs).

4. Browallia 5. Streptosolen

5. -

Corolla longer than 20 cm. 16. B rugm ansia Corolla usually not longer than 15 cm. 6

6. Fruit a capsule. 7 - Fruit a berry, sometimes Firm or dryish, but always indehiscent. ' 11 7. Capsule more than 20 mm long, with basal circular rim. irregularly tuberculate or spiny. 15.D atura - Capsule less than 15 mm long, smooth or nearly so. 8 8. Calyx enlarged after flowering, the teeth becoming spiny; capsule opening with a circumscissile lid. 17. Hyoscyamus - Calyx not or scarcely enlarged after flowering; the teeth not becoming spiny; capsule opening with valves, not a circumscissile lid. 9

10

12. Calyx becoming enlarged in fruit, at maturity be­ coming longer than the berry and often com­ pletely enclosing it 13 - Calyx not or scarcely enlarged in fruit, always shorter than the berry. 16 13. Flowers 2 to many together in sessile clusters near the axils of normal leaves; corolla greenish yel­ low. 13 . W ith an ia - Flowers either solitary in leaf-axils or in manyflowered peduncled cymes. 14 14. Calyx lobes at fruiting stage much longer than tube, sagittate at base, corolla blue. 14. N ican d ra - Calyx lobes at maturity much shorter than tube, not sagittate; corolla purple, mauve or yellow­ ish. 15 15. Flowers in terminal or lateral cymes; corolla pur­ ple; calyx firm ly clasping the fruit, never strongly ribbed. 8. Solanum

158. SOLANACEAE: 1. Cestrum, 2. Nicotiana j -

Flowers single in axils of somewhat reduced leaves; corolla yellow, sometimes with dark purple to brown markings; calyx in fruiting stage inflated around the fruit, often very strongly ribbed. 12. Physaiis

16. Corolla stellate, the lobes as long as the tube. 10. Capsicum - Corolla narrowly tubular, funnel-shaped or urnshaped, the lobes much shorter tlian the tube. 17 17. Corolla comparatively short, distinctly urn-shaped. 11. Discopodium I - Corolla longer, tubular or slightly funnel-shaped. 18 18. Spinescent or non-spinescent shrubs with over­ hanging branches; flowers solitary or in small clusters at nodes or at short side-shoots. 6. Lycium - Erect non-spinescent shrubs, flowers in terminal panicles or racemes. 1. C estrum 1. CESTRUM L. (1753) Shrubs or small trees, glabrous or pubescent with sim­ ple, forked or dendritic hairs. Leaves mostly alternate, sometimes paired, simple, entire, petiolate. Inflorescen­ ces usually raceme- or panicle-like, axillary or terminal. Flowers bisexual, sometimes slightly zygoinorphic. of­ ten subtended by small bracts. Calyx tubular, 5-7-lobed. Corolla tubular, variously coloured, limb shortly 5lobed. Stamens 5, equal or subcqual. variously inserted on corolla-tube; anthers bilocular. Stigma capitate. Fruit a juicy berry . Seeds prismatic or ovoid. A genus of c 250 species, indigenous in tropical Cen­ tral and South America, with a concentration of species in Brazil and the Andes. Two species have been recorded in the Flora area as cultivated or as garden escapes. 1. Corolla yellow or pale yellow. 2. C. nocturnum - Corolla red. pink orange or rarely white, but not yellow. 1. C. elegans 1. C. elegans (Brongn. ex Neumann) Schltdl. (1847); Habrothamnus elegans Brongn. ex Neumann (1844) - type: A plant cultivated in Jardin des Plants, Paris, originating from Mexico (?P holo.; specimen not traced, but illustrated in Ann. FI. et Pom. vel Journ. Jard., ser. 2. 2: plate opposite p. 116. 1844). Shrub up to c 4 m high, pubescent on all parts. Leaves alternate, not clustered, usually regularly decreasing in size towards the inflorescence; petiole grooved, 5-30 mm long, densely hairy; blade elliptic to narrowly ovate or lanceolate-elliptic. 7-26 x 2.5-7 cm. base rounded or cuneate, margin entire, apex long acute. Inflorescence a lax. terminal cyme, up to c 12 x 10 cm. Bracts linear, 3-6 mm long. Calyx narrowly campanulate. glabrous or glabrescent. 4-6 x 2.5 mm, with triangular lobes. Co­ rolla deep purple, red or pink, rarely white, tubular, 1.8-2.5 cm long, c 1.5 mm in diameter at base, w idening to c 4 mm upwards, abruptly constricted at mouth: co­ rolla lobes triangular, spreading, mm long, ciliate. Filaments inserted c 6 mm from base of corolla tube; an­ thers and style not exserting from corolla tube. Fruit a berry 1.1-1.6 x 0.7-0.9 cm, red to purple.

105

Cultivated and perhaps sometimes escaped in sec­ ondary evergreen scrub around houses; 1700-2400 m. EW SU SD; indigenous in Mexico; now widely culti­ vated as an ornamental and escaped. Bellini 474; Haugen 1594; Ecbvards & Tewolde B.G.E. 5002. The only Eritrean record of this species is from EW (Sarae), where it was collected 1909 in the surroundings of the tow n of Adi Ugri; the Ethiopian records are recent. This species has frequently, but erroneously been named Cestrum purpureum.

2. C. nocturnum L. (1753) -ty p e : “Habitat in Jamaica, Chili”, unknown col­ lector (BM lecto.), selected by D ’Arcy in Ann. Miss. Bot. Garden 60: 607 (1973). Large shrub or small tree up to c 4 m high. Branchlets sparsely pubescent, most other parts glabrescent. Leaves alternate, not clustered, often greatly reduced in size to­ wards the terminal inflorescence; petiole 5-15 mm long; blade lanceolate-elliptic, 10-15 x 4-7 cm, apex rounded to acute or acuminate, base broadly cuneate, margin en­ tire. pubescent on midrib below. Inflorescence terminal, lax or congested, panicle- or raceme-like, usually with contracted clusters of flowers. Flowers sessile or on pedicels up to 3 mm long, subtended by linear bracts up to 10 mm long. Calyx c 2.5 mm long, lobes triangular, c 0.8 mm long. Corolla greenish-yellow, tubular or slightly w idening upwards, 1.5-2 cm long, lobes c 4 nun long, erect or spreading, often darker than the tube, but with a paler margin. Stamens inserted in upper half of corolla tube, Filaments c 3 mm long, anthers c 0.5 mm long; sty le c 15 mm long; stamens and style not exserted from corolla tube. Fruit a berry, 0.8-1 cm in diameter. Cultivated and perhaps sometimes escaped in sec­ ondary evergreen scrub around houses; 1600-2300 m. EW SU SD HA; indigenous in Central America: now widely cultivated, escaped and naturalized throughout the warm parts of the world. Ryding 1357; Haugen 1218; Demel T. ’s students 1064. 2. NICOTIANA L. (1753) Annual or short-lived perennial herbs or spindly shrubs, glabrous or pubescent with glandular or non-glandular simple hairs. Leaves alternate, radical and/or cauline, simple, entire to sinuate, petiolate or apparently sessile due to blade decurrent on petiole. Inflorescence pani­ cle-like. rarely raceme-like or flowers solitary' in leafaxils. Flowers bisexual, actinomorphic, rarely slightly zygomorphic. Calyx tubular to narrowly campanulate. margins often thin and translucent, 5-lobed. persistent in fruit. Corolla tubular, rarely funnel-shaped, white, green, yellow or pink, limb shallowly or deeply 5-lobed. Stamens 5, equal or unequal in length, often 4 reaching the throat of the corolla-tube, the 5th shorter. Anthers bilocular, dorsifixed. dehiscent by longitudinal slits. Ovary bilocular. Stigma capitate. Fruit an ovoid to glob­ ular capsule with smooth surface, surrounded by the persistent calyx, opening with 2-4 apical valves. Seeds usually reniform. often angled.

106

158. SOLANACEAE: 2. Nicotiana

A genus of 60-70 species, mostly indigenous in South America, but also in North America and West Af­ rica (1 species), Australia and the Pacific region. Many species are of economic importance because of their toxic properties, having a long history of use for smok­ ing. chewing or snuff production, both in areas where they are indigenous, and where they have been intro­ duced. Commercial or home-grown tobacco is manly de­ rived from N. tabacum and N. rustica. both of which are widely grown and escaped in the Flora area. Other spe­ cies are widely naturalized; of these only N. glauca Gra­ ham is known from the Flora area. 1. Perennial, spindly shrub to 6(—10) m high; corolla yellow to red. 3. N. glauca - Annual or short-lived perennial herb; corolla white, greenish-white or pink. 2 2. Corolla c 4 cm long, 0.3-0.4 cm in diameter in lower part of the tube, lobes sharply acute; upper leaves oblong-lanceolate to elliptic, base de­ current on petiole, apex acute; capsule ovoid or ellipsoid, c 2 cm long; surrounded by persisting catyx. 2. N. tabacum - Corolla up to c 2.5 cm long, often shorter, 0.6-0.8 cm in diameter, lobes usually broadly triangular or rounded; leaves broadly ovate to broadly el­ liptic, rounded to broadly acute or truncate at base and apex, petiole up to 4 cm long, not with decurrent blade; capsule globose, about 1.2 cm in diameter. 1. N. rustica 1. N. rustica L. (1753) - type: “In America”, Herb. Linn. (LINN 245.1; specimen seen on microfiche). Annual herb up to 1.5 m tall. Stems viscid pubescent, with simple glandular hairs. Leaves alternate, not crowded; petiole up to 8 cm long, not winged; blade ovate to elliptic, rarely suboibicular, 10-20(-30) x 15-18 (-20) cm, base usually truncate, margin slightly sinuate, apex acute, puberulent on both sides. Inflorescence a lax panicle with 3-5 major branches and numerous flowers. Pedicels glandular, 0.5-1 cm long. Calyx tubularcupuliform, 8-15 mm long; lobes triangular, acute, usu­ ally unequal. Corolla tubular-infundibuliform, greenish white, 12—17(—24) mm long; lobes of limb broadly tri­ angular, apex rounded or obtuse, apiculate, spreading, 3-6 x 2-3 mm. Stamens unequal, inserted in the basal quarter of the corolla-tube. Stamens and style not exserted from corolla tube. Capsule ellipsoid-ovoid to subglobose, 7-16 mm in diameter. Cultivated in or around villages, or apparently as a weed among other crops; (c 200-) 1000-2000 m. (EE, see note below) SU KF SD; indigenous in America (probably originally temperate North America); for­ merly widely cultivated in the tropical and particularly in the warm temperate regions of the world for home con­ sumption of tobacco; now largely replaced by N. taba­ cum, especially in large-scale fanning. Senni 2160; Kuls 631; Straube 11/78. Only the Senni collection has been seen; the two other collections are cited from Cufodontis in Senck. Biol. 38: 412 (1957).

Boselli, in Pirotta (ed ), Flora della Colonia Eritrea'. 248 (1904), states that she has seen material of this spe­ cies cultivated in Eritrea, documented by specimens from seeds from the colonial ministry of agriculture, in­ dustry and trade, and originating from Moncullo (in the plains west of Mitsawa), Samhar, EE, the material lias not been traced. 2. N. tabacum L. (1753) - type: “In America”, Herb. Linn. (LINN 245.3; specimen seen on microfiche). N. lancifolia Willd. ex Lehm. (1818) /V. tabacum L. var. lancifolia (Willd. ex Lehm.) Comes in Atti del R. 1st. Jncoraggiamento Sci. Nat. Napoli, Ser. 5, 1(1); 11. 1899, N. angustifolia Elirh. (1790); N. fruticosa L. var. angustifolia (Ehrh.) Dunal, in DC., Prodr. Syst. Nat. 13(1): 559(1852) iV. latissima Miller (1768). Heib up to 2 m or more tall. Most parts viscid and with numerous glandular hairs. Leaves alternate, not clus­ tered, regularly decreasing in size towards the inflore­ scence; petiole 1-2 cm long, broadly winged, the wings often with a pair o f auricles at the base which clasp the stem, or, due to decurrent blade (often with basal auri­ cles), sometimes apparently absent; blade ovate to ellip­ tic or lanceolate. 15-50 cm (or sometimes more) x 5-25 cm (or sometimes more), not glaucous, membranous, base cuneate or decurrent, margin entire or slightly sinuate, apex acute to long acuminate. Flowers many in much-branched terminal panicles, up to c 25 cm long. Pedicel 0.8-2 cm long. Calyx 1.5—2.5 cm long; teeth un­ equal. acuminate, 0.6-1 cm long. Corolla greenishwhite. often pinkish in upper part, funnel-shaped, 3.5-5.5 cm long, with broadly triangular or rounded, un­ equal lobes, 0.3-0.6 cm long. Stamens unequal, hardly exceeding the corolla tube. Capsules ellipsoid or ovoid, 1.5-2 cm long. Seeds c 0.5 mm in diameter Cultivated in villages in home gardens or in special­ ised tobacco-farms, sometimes escaped; 300-2400 m. EE EW TU GD SU WG KF GG SD HA; indigenous in America (probably originally temperate South Amer­ ica); now widely cultivated, escaped and naturalized throughout the warm parts of the world. Tadesse Ebba 508; Tekle Hagos 63; Friis et al. 2110. 3. N. glauca Graham (1828) - type: plant cultivated 1828 at the Royal Botanic Garden Edinburgh, from seeds sent from Buenos Ai­ res, Argentina (E holo., specimen not seen). Shrub or small tree, usually 1-3 m tall, sometimes up to 6 m, and rarely to 10 m. All parts glabrous and glaucous. Leaves alternate, not crowded, gradually reduced in size towards the inflorescence; petiole 3-12 cm long, not winged; blade elliptic to ovate, 5-25 cm long, base cuneate or rounded, margin entire, apex acute or obtuse, glaucous, often almost leathery. Flowers many in lax panicles up to c 25 cm long. Pedicels 0.5-1 cm long. Ca­ lyx 1—1.5 cm long, tubular, teeth triangular, equal, acute, c 2 mm long. Corolla yellow to red, tubular, 2.5-4,5 cm long, with short blunt erect or spreading co­ rolla lobes usually less than 2 mm long. Stamens sub­

158. SOLANACEAE: 2. Nicotiana, 3. Petunia

107

Figure 158.1. N ICO TIAN A GLAUCA: 1 flowering stem x Vy, 2 - flower, longitudinal section x 2: 3 - cap­ sule with persistent calyx partly removed x 2; 4 - seed x 20. All from Torre 7239. Drawn by Elea­ nor Catherine for FI. Zcimb.

equal, enclosed with the style in the corolla tube. Capsules ellipsoid, 0.7-1.6 cm long. Seeds c 0.5 mm in diameter. Fig. 158.1. Cultivated, escaped or naturalized in waste places and along roadsides; 1300-2300 m. EW TU WU SU HA; indigenous in South America (Argentina); now widely cultivated, escaped and naturalized throughout the warm parts of the world. Burger 3807; W.J.J.O. de Wilde 9825; Ryding 1158.

3. PETUNIA Juss. (1803) Annual or short-lived perennial herbs, pubescent and with numerous simple glandular liairs, viscid. Leaves al­ ternate or the upper ones paired, simple, entire, petiolate or sessile. Flowers solitary in leaf axils, bisexual, usu­ ally single, zygomorphic. Calyx campanulate, deeply 5-lobed. Corolla single (but double in some cultivars), broadly funnel-shaped, white, pink or purple, entire or shallowly lobed, lobes induplicate in bud. Stamens 5, 4

108

158. SOLANACEAE: 3 Petunia

Figure 158.2. PETUNIA IN T E G R IF O L IA , one of the parental species of PETU ­ NIA x H YBRID A. The variation in the garden-petunia covers the mor­ phological variation in the parental species (see further discussion in text): 1 - flowering stem x 1; 2 flower, longitudinal section x 2'/j; 3 - capsule with persistent calyx partly removed x 2'4; 4 - seed x 15. All from Verdcourt 993. Drawn by Eleanor Catherine for FI Zamb.

in two pairs, with the 5th the shortest, inserted in coroiia-tube. Anthers biiocuJar. Stigma capitate. Fruit a smooth capsule, enclosed in the persisting calyx, open­ ing from apex. Seeds globular or angular.

P. x hybrida Vilm. (1863) type: A plant cultivated in Paris (?P holo., speci­ men not traced).

A genus of 30—40 species, closely related to Nico­ tiana, but constantly with n= 14 (in Nicotiana n= 12), in­ digenous in the mountainous parts of South America. Various cultivars are widely cultivated as ornamentals in the Flora area.

Annual or short-lived perennial herb. Stems erect, as­ cending or decumbent, with numerous glandular liairs; the whole plant usually viscid. Leaves alternate or oppo­ site in the flowering parts of stems, somew hat decreas­ ing in size towards the top o f the plant; petiole (0.5-) 1—2(—6) cm long, often winged or with decurrent blade

158. SOLANACEAE: 3. Petunia, 4. Browallia, 5. Streptosolen

on lower leaves; blade ovate to elliptic, 3-6 x 2-4 cm, base cuneate, often decurrent, margin entire, apex acute to rounded. Inflorescence terminal, raceme-like, with flowers in the axils of leaf-like bracts, c 4 x c 1.5 cm, or opposite them. Pedicel 3-7 cm long. Calyx narrowly campanulate to tubular for the lowermost 2-3 nun. with calyx-lobes linear-lanceolate, strap-shaped or leaf-like in some cultivars, 2-3.5 cm long, unequal. Corolla tubu­ lar to funnel-shaped in the 3-5 cm long basal part, limb of spreading corolla lobes. 4—8(—12) cm across, corolla iobes obtuse to rounded, rarely acute, c 3 x 2 cm. subequal, with entire or sinuate margin, plain or multijoloured, white, yellow, pink, red, purple or violet, single or double, usually sweetly scented. Stamens 5, of which 4 in two long pairs, shorter and curved, all in­ serted in lower part of tube; anthers white or yellow, sometime purplish or bluish, partly enclosed. Style partly enclosed with the stamens. Capsule 1-1.5 cm long, elliptic, smooth. Fig. 158.2 shows one of the parental species of the hy­ brids which are the most frequently cultivated forms: see below. Cultivated in public and private gardens; c 2500 in. SU (in gardens and parks in Addis Abeba); widely culti­ vated throughout the world. No herbarium collection seen from the Flora area. The cultivated garden-petunia, usually referred to as P. x hybrida, is a variable plant, which is derived by hybridisation from two wild species, P. integrifolia

?- lrultmg bra'lch x 11 - detail of indumentum x 10 i from Friis et al. 8263. 2 from Carr 875- 1 from Friis et i\ 2047,4-5 from i Cun,per 631; 6 trom Ryding 1536; 7-9 from Gilbert 1387; and 10- M from Espten 103. Drawn by ^ r g a r e t Tebbs

158. SOLANACEAE: 8. Solanum

common; 400-2350 m. TU SU KF GG HA; widespread throughout Africa including Somalia, the Mediterranean area, Europe, Asia, Malesia, Australia, New Zealand, and N, C & S America. Mooney 6232; Carr 875; Burger 378. This species may be native to South America, al­ though other areas suggested include the western Indian Oceanic Islands, the west African coast. Central Amer­ ica and SE North America; it is generally considered to be an adventive in much o f the Old World. It is probably the most widespread and morpholo.gically variable spe­ cies in the section Solanum. and is diploid (2n=2x=24). It is now found throughout the world and a number of ;nfraspecific variants have been formally described. Edmonds {Bot. Journ. Linn. Soc. 75; 143 (1977). recogviised two ecogeographical varieties of this species in South America with var. americanum being pilose and var. patulum (L.) Edmonds being glabrescent. This dis­ tinction is not so apparent in Africa, where both types oc­ cur and are best identified as S. americanum. This species is characterised by umbellate cymes of very small flowers, which are succeeded by small globose berries typically on erect or spreading pedicels. There ire a number of specimens in SW Ethiopia which appear florally intermediate between this taxon and the next species (.S', tarderemotum), which usually has slightly larger flowers and lax cymose many-flowered inflores­ cences in which the fruiting pedicels are reflexed. According to Cufodontis (1957. 1963), who refers to the species as S. nodiflorum Jacq. and to Edmonds & Clnveya (1997), S. americanum is sometimes cultivated in Ethiopia; both the berries and the leaves are eaten in Shashamane (SU), and the Dime people in eastern Omo (GG) use it as a pot herb. 10. S. tarderem otum Bitter (1912) - type: East Africa [Tanzania], Kilimandjaro, Marangu. c 1600 m, Winkler 3865 (WRSL holo ). Erect, procumbent or scrambling herb. 0.4-1.5 m high occasionally with procumbent branches spreading to 3 m. glabrescent to moderately pubescent with eglan­ dular appressed and spreading hairs. Stems smooth or inconspicuously dentate. Leaves alternate, petiole 0.8-3.0(-5.0) cm. winged below from decurrcnt blade; blade usually lanceolate sometimes ovate-lanceolate. 5-18 cm long and 3-7 cm broad, base cuneate. margin usually entire, sometimes sinuate-dentate, apex acute to acuminate. Inflorescence simple, lax extended cymes, rhachis 4-14 mm long fruiting. (6-)7-14-flowercd. Peduncles erect. 10-15 mm flowering. 15—25(—37) nun fruiting. Pedicels 4-5 nun flowering, 6-9 mm fruiting when usually reflexed. Calyx campanulate. 1.3-2.5 nun long, sepals broadly ovate to ligulate. obtuse. 0.5-1.0 nun long and wide, enlarging to 1.3-2.0 mm fruiting, when adherent becoming reflexed. Corolla white occa­ sionally tinged purple, stellate. 3.5-5.3 nun radius; pet­ als 2.0-2.8 mm long, 1.2-1.5 mm wide, spreading. Filaments 0.5-1.0 nun long, pilose to densely villous. Anthers 1.5-2.0 (rarely -2.5) mm long, yellow. Style 3-6 nun. exserted 1-2 mm and often geniculate, lower part pilose; stigma minute, c 0.2 mm diameter. Fmit dark purple or black globose berry. 4-6 mm diameter;

121

seeds 1.3-1.5 mm long; sclerotic granules usually pres­ ent. Fig. 158.6.3. Weed of cultivation, on roadsides, along footpaths, forest- roads or paths, in evergreen cloud or riverine for­ ests or among pebbles; often common; 1100-2600 (7-3000) m. EW GD TU/GD GJ WG SU IL KF SD HA; ?Somalia, E Africa, Burundi, Rwanda. Malawi and Dem. Rep. Congo. Gilbert & Thulin 651; Friis et al. 2047; Westphal & Westphal-Stevels 5499. This frequently collected and fairly well defined taxon lias been recorded from Ethiopia to Tanzania and Malawi, and seems to be conspecific with the protologue of S. tarderemotum although the type specimen of this species lias not yet been located. The vegetative charac­ ters seem to be extremely variable, with the pubescence varying from glabrescent to conspicuously pilose, and the leaf margins from entire to sinuate dentate. It is close to the small-flowered diploid species S. americanum (see above). Work with living material, including the de­ termination of its ploidy level, is necessary to determine whether the variation observed warrants specific recog­ nition. In Dilla (SD) the leaves of this species are eaten after cooking, and in Jimma (KF) the berries are reputedly ed­ ible when ripe. The description of white ripe fmits on Friis et al. 7133 is anomalous and could indicate fruit immaturity; some berries o f these species look whitish and waxy before ripening to their proper mature colour. 11. S. hirtulum Steud. ex A Rich. (1850) type: GD, Semicn. Endschedcap. Schimper (II) 977 (P lecto. selected bv Lester in Bot. Journ. Linn. Soc. 125: 275 (1997));'BM CGE G-DC GOET K MPU OXF P S UPS isolecto.). IS. monactinanthum Dammcr (1912) - type: BA/AR, Jidah in Arussi-Galla. Ellenbeck 1452 (B holo., destroyed). Low or prostrate herb, base usually suffrutescent with branches ascending from woody tap-root, 5-20(-40) cm high, sometimes fomiing a rosette; all parts hirsute with long, white, spreading eglandular hairs. Leaves alter­ nate, sometimes almost sessile; petiole 0.2-1.0 cm long, winged from decurrent blade; blade pubescence denser on lower surfaces, spathulate, obovate or lanceolate, small, 2.5-3.0(-5.0) cm broad and 1.0-1.5 cm long, base cuneate. margin usually entire, occasionally shal­ lowly sinuate, apex acute. Inflorescence leaf-opposed or extra-axillary', simple, umbellate cymes, 2-3(rarely -5 )flowered. Peduncles erect, 12-14 mm long flowering, 10-18 mm long fruiting. Pedicels 4-7 nun long flower­ ing, 7-10 mm long fruiting when spreading or reflexed. Calyx with (1.5—)2—3.5 mm long, densely hirsute, sepals broadly-triangular to -ovate, usually obtuse, c 0.5-1.3 mm long and 0.5-0.8 mm broad, enlarging to 1.5-2.4 mm long and 2.5-3.2 nun broad fruiting, when adherent or reflexed. Corolla lilac blue, purple or mauve, stellate, 7-8 mm radius, petals triangular, 3.5-6.0 mm long and 2.5-5.0 mm wide, usually spreading; buds densely hir­ sute, often appearing white. Anthers yellow to orange, 2.0-2.5(-3.0) nun long. Style 3-4 mm long, pilose ba­ sally. exserted 1-2 mm, often geniculate. Fruit globose. 4.5-7.0 111111 diameter, possibly green at maturity. Seeds

158 SOLANACEAE: 8. Solanum 122

orbicular to renifonn, light brown. 1.5-2.0 mm long: two small sclerotic granules (0.8 mm diameter) present. Fig. 158.6.4&5. Arable land, along river banks, on rough grassland and in forest at high altitudes. 2250-3230 in. TU GD GJ SU HA: not known elsewhere (with the possible excep­ tion of BA/AR, see below), possibly originating in dis­ turbed afroalpine habitats. Mooney 4752; Scott 292; Thulin & liunde 3914. The species S. monactinanthum Dammer. described in Bot. Jahrb. 48: 236 (1912) and collected from the BA-AR highlands [“Galla-Hochland” |, is closely re­ lated to and possibly synonymous with S. hirtulum. The type specimen collected from BA/AR at an altitude of 2600 m was presumably destroyed, but most features in the protologue agree with those charactensing this species including the pubescence on the floral parts and the large flowers. Although the protologue cites oneflowered inflorescences, those in S. hirtulum occasion­ ally appear single-flowered, although rudiments of addi­ tional Howers are often visible. It has provisionally been included in synonymy here pending the location of a du­ plicate Ellenbeck specimen. The mature berry colour and the ploidy level of this taxon has yet to be determined. 12. S. memphiticum Gmel. (1791); S. aegypticum Forssk. var. b) Fructu nigro\ foliis mtegris, vilosissimus, Forssk.. Flora Aegvpt. -Arab.: 46 (1775). nom. inval.\ S. nigrum L. var hirsutum Vahl. Symb. 2: 40 (1791) - type: Cult. Hort. Egypt, Forsskal s.n.; Yemen. Dunal 35 (P syn., MPU isosyn) S. grossidentatum A. Rich. (1850); S. plebeium A. Rich. var. grossidentatum (A. Rich.) Chiov., in N. Giorn. Bot. Ttal. 26: 159 (1919) - type: TU, Tchelikote. Petit s.n. (P lecto. selected by Lester, Bot. Journ. Linn. Soc., 125: 285 (1997); G GOET P Z isolecto.). S. hirsutum Dunal var. abyssinicum Dunal (1852) in DC., Prodr. Syst. Nat. 13(1): 58 (1852); S. memphi­ ticum Gmel. var. abyssinicum (Dunal) Cufod., in Enum.: 872 (1963) - type: TU. Adoa (Adua), Schimper 46 (G-DC syn.. G-Herb Moricand. syn.; CGE G K. MPU P isosyn ). Erect, ascending or spreading small annual or perennial herb or subshrub up to 0.9 in high, branches often aris­ ing from woody base. Stems usually herbaceous, terete, strikingly viscid, all parts covered with pale, long, spreading multicellular villous glandular-headed hairs mixed with eglandular liairs (sand particles adhering to these glandular liairs often makes the indumentum ap­ pear brownish). Leaves moderately to densely villous, hairs as for stems but shorter; petiole 1.2-3.0(^4.0) cm; blade obovate to rhomboidai. 2.5-6.0 (rarely -15) cm long and 1.5—3.6 (rarely -7) cm broad, base long cuneate. margin always sinuate-dentate to incised with many small obtuse lobes, decurrent, apex acute to ob­ tuse. Inflorescence simple, umbellate to lax erect cymes, 3-4(-5) (rarely -6)-flowered. Peduncles extra-axillary, 7-18 mm long flowering, 9—29(—30) mm long fruiting, ercct. densely villous. Pedicels 5-6 mm long and erect flowering. 6-13 mm long and erect or reflexed fruiting, densely villous. Calyx 1.8-2.5(-4) mm long, sepals

spathulate to obovate, 0.8-2(-3) mm long, 0.4-1.2 mm broad, densely villous externally especially on margins, enlarging to 2.5-5 mm long and 2-3 mm broad when stellate and usuallv adherent to ripe berries. Corolla white, occasionally pale purple, often with purple midnb to petals and a yellowish basal star, stellate, 3 5-8 mm radius, petals 1.6-2.5 mm long and 1.3-2.0 mm broad, usually spreading. Anthers 1.5-2.5 (rarely -3) mm long. Style 2.5-4.5 nun long, pilose below, exserted up to 1.5 mm. Fruit purple to black globose b e r n . 5-8 mm in diameter. Seeds 1.5—2 mm long; small apical sclerotic granules sometimes present. Fig. 158.6.7-9. A fairly common weed of cultivation, overgrazed and distuibed land and roadsides, also occurring on bushland. degraded forest, diy stonyr. rocky and gravel areas; 1350-3000 m. EW TU SU AR SD BA HA; Egypt, the Su­ dan, E Africa, sporadic in Dem. Rep. Congo, Malawi and possibly elsewhere in southern Africa; Saudi Arabia, Ye­ men. Gillett 14586; Herbert49;M.G. &S.B. Gilbert 1387. Because of the characteristic stellate fruiting calyx lobes of many Ethiopian specimens, they have often been identified as S. grossidentatum. However, these specimens exhibit a range towards S. hirsutum in many of the diagnostic characteristics, including flower size; those previously identified as S. hirsutum are generally smaller flowered than those identified as S. grossi­ dentatum. Much of this variation apparently could be due to plie notypic responses to some of the more ex­ treme habitats from which they have been collected. Lester JBot Journ. / inn. Soc. 125: 285 (1997)] stated tliat although S. grossidentatum was originally spelt with an “e” by Richard, it should be corrected to S. grossi­ dentatum The earlier suggestion in Edmonds & Chweya (1997) that this taxon might be conspecific with S. retroflexum Dunal [DC., Prodr. 13(1): 50 (1852)) is now considered erroneous This latter purple-berried tetra­ ploid relative originally described from South Africa, is now thought to have a much more restricted distribution, probably not occurring further north than the Flora Zambesiaca region. The ripe berries of S. hirsutum are reportedly eaten in Shashemane (SU/SD). The ploidy level of this taxon is not yet known. The name change to S. memphiticum lias been caused by a recent lectotypification of the taxa besed om Forsskal’s collections from Egypt and Yemen. II subgen. Potatoe (G. Don) D Arcy (1972) Herbs, vines or shrubs, unarmed, hairs (if present) simple or branched, never stellate. Leaves often deeply lobed or sometimes apparently compound. Inflorescence gener­ ally paniculate, pedicels articulate at or shortly above their bases. Corolla rotate to rotate-stellate; anthers stout. Fruit usually a juicy berry, globose or ellipsoid. 4. sect Petota Dumort. (1827) As for the subgenus, bin with underground stolons cariying tubers. Leaves apparently imparipinnate with in­ terstitial leaflets. Hairs various, not “bayonet-hairs” as in next section. Pedicel articulate well above base. Fruit globose.

158. SOLANACEAE: 8. Solanum Only one cultivated species in the Flora area. 13. S. tuberosum L. (1753) - type: presumably cultivated in Europe, Herb. Linn. (LINN 248.12 lecto., designated by Hawkes in Proc. Linn. Soc. London 166: 106, 1956). Perennial herb with creeping, tuberiferous rhizomes, up to e 1 m high. Stems angular or winged, with angles or wings decurrent from leaf-bases. Leaves alternate, de­ creasing in size towards the inflorescence; petiole usu­ ally short and winged, difficult to distinguish from decurrent part of blade; blade usually imparipinnate with winged rachis; primary leaflet-pairs 3-5, decreas­ ing in size from apex to base of blade, usually with in­ terstitial leaflets that are much smaller than the normal leaflets; blade on both sides with scattered simple hairs. Inflorescence a lax cyme with c 10 flowers on bifurcate peduncle. Pedicel c 1-2 cm long, articulate in the middle or upper third. Calyx tubular or broadly campanulate, c 0.5 cm long, with lanceolate lobes. Corolla rotate, 2.5-3.5 cm in diameter, colour varying from white to mauve. Anthers c 5 mm long, subsessile or on short fila­ ments Style exceeding the anthers with 1-2 mm. Fruit up to 2 cm in diameter, globose, pale brown when ripe, but fruits are not always produced. Fig. 158.7. Cultivated for its tuberous rhizomes (“potatoes”); 1800-2600 m. EW SU (and certainly in many other parts of the Ethiopian Highlands); indigenous in the Andes of Peru. Chile and adjacent parts of South America, now cultivated throughout the world in the temperate regions and in the mountains of the tropics. Kiesfin T. 8303; Afeworki G. & M ehretA. 70. Already A. Richard (1851) stated that S. tuberosum was cultivated in Ethiopia; according to Pax (1907) it was first introduced by Schimper, but Chiovenda (1929 & 1937) claimed that it was first introduced by the Ital­ ian missionary Mons. De Jacobis in 1860. Both gentle­ men may have been active potato-introducers and -growers, but the reference made by A. Richard seems to support introduction by Schimper before 1850. 5. sect. B asarthrum (Bitter) Bitter (1913) As for subgenus, but underground parts not with stolons carrying tubers. Leaves various; in the Flora area not ap­ parently pinnate. Hairs usually two-celled, acute, the basal cell the longer, with thicker cell walls, the apical one shorter and tapering (= bayonet hairs). Pedicels usu­ ally articulate at base. Fruit ellipsoid to ovate. 14. S. m uricatum Ait. (1789) - type: cultivated in the botanical gardens of Paris and Kew, introduced by Thouin from Peru or Teneriffe (BM, specimen not traced). S. scabrum Lam. (1796), nom. illeg., non Miller (1768). S. variegatum Ruiz & Pav. (1799). S. muricatum A it var. teleutogenum Bitter in Fedde Repert. 12: 443 (1913). Unarmed or slightly spiny herb or shrub up to c 1 m high. Younger parts puberulous to pubescent with two-celled hairs. Leaves alternate, not clustered, of ap-

F ig u rc 158.7. SO LAN U M s c c t. PETOTA. S. TUBEROSUM 1 - f lo w e rin g b ra n c h x Vr, 2 - f lo w e r x 14; 3 f lo w e r, lo n g itu ­ d in a l s e c tio n x 14; 4 - b e rry x 14; 5 - tu b e r ( p o ta to ) x ' / 3 . A r tis t an d o r ig in a l m a te ria l u n k n o w n . R e a rr a n g e d fro m A . F io ri & G . P a o le tti. Iconographia Florae llalicae ( ) 895-1904).

proximately the same size and shape throughout the plant (in some cultivated forms pinnatisect leaves de­ velop. but this has not been seen in specimens from the Flora area, see note below); petiole 3-7 cm. articulate at the base, slightly ridged or winged, especially in upper part; blade always entire (in the Flora area, see note be­ low), 7—12(—15) x 1.5—2(—5) cm, base cuneate, margin entire or slightly sinuate, apex acute or rounded, with sparse to abundant indumentum of appressed two-celled hairs on both sides, especially on the underside. Inflore­ scence a dense lateral umbellate cluster of 10-12 flow­ ers, all parts finely pubescent. Peduncle 2.5-5 cm long. Pedicels 0.5-0.8 cm. Calyx c 5 mm long, enlarging in fruit to c 0.8 cm. lobes narrowly triangular to linear. Co­ rolla blue, up to 4 cm in diameter. Anthers c 1.5 mm. Style exceeding the anthers by 1—(?)3 mm. Fruit white or pale green, variegated with purple, ovoid. 5-12 cm long, usually seedless. Fig. 158.8.1&2. Cultivated for its fruits, the kpepino \ 'melonchino 'melon berry' or "melon pear'\ grown at c 1800-2600 m. EW SU KF SD HA; widely cultivated throughout the warm regions of the world, e.g. the Mediterranean re­ gion. Australia, New Zealand, and the USA. Scott 58; Tadesse Ebba 555; Burger 2326. The fruits of .S’, muricatum are eaten raw or cooked. The yellow pulp of the fruit is reminiscent of melon. S. muricatum was first taken into cultivation in Ecuador, Peru and Bolivia in pre-Columbian time and became widely cultivated throughout the Andes at altitudes be-

124

158. SOLANACEAE: 8. Solanum

tween c 1000 and 3000 m. The species is not known from the wild. It is presumed that Italians introduced this plant to Eritrea by the end of the 19th Century, and that it was in­ troduced from there to Ethiopia. Hepper reports in FI. o f Egypt, 159 Solanaceae, p. 42 (1998) that it was culti­ vated in Addis Ababa and brought into cultivation in Egypt before 1944 by transporting fruits and young shoots of it to Cairo by aeroplane. Bitter, in Fedde. Repert. 12: 441^144 (1912), hrs proposed various forms of S. muricatum which develcp 3-, 5- or 7-foliolate leaves as var. protogenum Bitter, var. propayanum Bitter and var. praecedens Bitter. Many cultivars have been described on variation in fruit size and abundance. 6. sect. Jasm inosolanum Bitter ex Seithe (1962) Glabrous or glabrescent climbers with simple, lobed or compound leaves, petioles often coiling round objects that may support the plant; inflorescence terminal, cymose, many-flowered, with pedicels articulate almost at the base; fruit globose. One species (cultivated) in the Flora area 15. S. jasm inoides Paxton (1841) - type: cultivated in London [specimen not seen, illustrated in Curtis' Bot. Mag. 8. tab. 5 (1841)). Herbaceous climber with slender stems, growing to a height of c 3 m, sometimes more. All parts glabrescent. Leaves alternate, not clustered, decreasing in size to­ wards inflorescence; petiole 0.5-1.5 cm, often coiling round objects that may support the plant; blade usually entire, up to 2 x 4 cm, base hastate, sometimes develop­ ing prominent basal lobes, truncate, subcordate or broadly cuneate, margin entire, apex acute to slightly acuminate; below with tufts of hairs in the axils of the primary veins. Inflorescence a large, terminal, lax pani­ cle with numerous flowers. Calyx radiating, repandly lobed, c 3 mm in diameter. Corolla stellate, shallowly lobed, 2-2.5 cm in diameter, pale blue. Anthers c 5 mm long, subsessile. Ovary and sty le hidden by anthers. Fruit purple to black, c 0.8 cm in diameter. An ornamental climber, cultivated in parks and gar­ dens in the larger towns; c 2400 m. EW (and probably elsewhere, e.g. SU); indigenous in Brazil, but now w idely cultivated in the warm temperate regions and in the moun­ tains of the tropics. A fw o rki G. & Mehret A. 66. The only herbarium record seen for this account was cultivated in Asmara (EW) at 2400 m. but live plants have been seen cultivated elsew here in town gardens, es­ pecially in and around Addis Ababa. S. jasminoides su­ perficially resembles the indigenous forest climber S. benderianum Dammer, but can easily be distinguished on the often coiling petioles, the usually hastate or trun­ cate leaf-bases, and the tufts of liairs in the axils of the primary veins. III. subgen. B revantherum (Seithe) D 'Arcy (1972) Shrubs or small trees, usually with stellate (or more complexly branched) hairs, but sometimes not. Leaves

enure Inflorescence terminal, Decoming lateral, com­ pound. Pedicels often reflexed at anthesis. Fruits yellow when ripe. Sclerotic granules sometimes present. 7. sect. Pseudocapsicum Bitter (1917) Herbs, shrubs or small trees, glabrescent or pubescent with simple or branched liairs. Leaves entire or sinuate. Inflorescence few-flowered. Corolla mauve or white. Fruits red, orange or yellow; sclerotic granules absent. One cultivated species in the Flora area 16. S. pseudocapsicum L. (1753) [ - type. Madeira. Herb. Linn. (LINN 248.4 lecto.. selected by E. Schombeck-Temesy, FI. Jranica 100: 6 (1972); specimen seen on microfiche). Herb or shrub up to c 2 m high, glabrescent to puberulent. Leaves somewhat irregularly alternate, veiy often two of very different sizes together; petiole 3-7 mm long, difficult to distinguish from the decurrent blade; blade narrowly elliptic or narrow ly lanceolate, smaller leaves of a pair c 1 5-2.5 x 0.7-1.5 cm. larger leaves 6-10 x 1.5-4 cm, up to 5 times as long as wide, base cuneate. margin entire or indistinctly repand. apex acute or acuminate; veins prominent beneath, yellow'. Inflore­ scence lateral, sessile or subsessile cymes mostly con­ densed into a fascicle, with up to 5 flowers. Pedicel 0.5-1 cm long. Calyx c 0.5 cm long, with linear lobes, glabrescent to densely puberulent Corolla w hite, rotatestellate, c 0.8 cm in diameter. Anthers c 2 mm long, subsessile. Style exceeding anthers with c 2 mm. Fruits 1-2 cm in diameter, globose, orange-red. shinv. Fig 158.8.3 & 4. Cultivated; c 2500 m. EW (Asmara) SU (Addis Ababa), and probably elsewhere in the bigger tow ns of the two countries; indigenous in South America, now cultivated as an ornamental shrub in all w arm parts of the world; introduced as an ornamental and naturalized on Madeira and in E Africa. Senni 310; Zemede A. 2005; Afewarki G. is, but they seem to have no other connection with Linne filius). The name is illegitimate because the earlier name D. inermis Juss. ex Jacq. is cited in synonymy . D. tatula L. (1762); D. stramonium L. var. tatula (L.) Toney, yl Flora o f the Northern & Middle section o f the United States'. 232 (1824); Stramonium tatula (L.) Moench (1794) - type: country or origin not indi­ cated; specimen in the Herb. Linn. (LINN 243.2. lectotype, selected by Hadkin. Bye, Brandenburg & Jam s in Bot. J. Linn Soc. 125: 305 (1997); specimen seen on microfiche). D. stramonium L. var. chalybaea Koch. Synopsis Florae Germanicae et Helveticae. ed. 1.; 510 (1838). Datura metel L. var. fastuosa auct.. non (L.) Safford (1921): Wright (1906), based on Schweinfurth & Riva 1235 from Acrour, Eritrea. Shrubby annual herb up to 2 m tall but often much smaller, glabrescent or with few. straight, non-glandular hairs Leaves one or several at the nodes; petiole 1-5.5 cm long; blade broadly ovate to rhomboid or angular. 5-17(-30) x 3-14 cm. asymmetrically cuneate at the base, acuminate at the apex, with inegularly sinuate, lobed or dentate margins. Flowers erect; pedicel c 0.5 cm long. Calyx 2.5-5 cm long, 5-lobed. lobes 6-8 mm long. Corolla white, funnel-shaped, 4.5-10.5 cm long, limb 5-lobed. lobes mucronate at apex, ending in c 10 mm long points. Stamens included; anthers 3-5 mm long, on 3.5-6.0 cm long filaments. Style 4-6 cm long; stigma held below or above the anthers. Capsules erect, globose or ovoid. 3-5 x 2-4 cm, densely spiny or prickly and minutely pubescent between the prickles, spines 100-200, slender, conical, sharp, but capsule sometimes more or less completely devoid of prickles; capsule dehiscent by 4 equal valves; persistent base of calyx up to 10 mm wide. Seeds black, c 4 mm in diame­ ter. Fig. 158.23.8. Weedy and in disturbed places, waste ground, road­ sides, frequent near water holes or in areas of impeded drainage; 600-2800 m. EW GD GJ WU SU WG AR IL KF SD HA; Sudan, Somalia and throughout tropical and South Africa; also widespread in Europe, parts of Asia and America. Straube 674; Chajfey 488; Mooney 8942.

158. SOLANACEAE: 15. Datura, 16. Brugmansia

159

Figure 1 58 .2 3. DATURA inoxia 1 flowering b r a n c h x W3; 2 - detail ol lower leaf surface x 7; 3 - fruit x c Vi. D. M ETEL. 4 - fruit x c Vi ; 5 - fruit, longi­ tudinal section x c Vi; 6 - seed x 4. D. F E RO X: 7 - fruit x c Vi. D. STRA M O N ­ IUM. 8 - fruit x c Vi. 1 - 2 from Aguiar M acc do 5 5 3 2 : 3 from Brum mi it. Banda