Flora of Ethiopia & Eritrea. Vol. 4.1, Apiaceae to Dipsacaceae 9789197128551, 9197128554

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FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 1 APIACEAE TO DIPSACACEAE Editors: Inga Hedberg, Sue Edwards & Sileshi Nemomissa

Addis Ababa, Ethiopia Uppsala, Sweden 2003

Gift from the National Herbarium(ETH) Department of Plant Biology and Biodiversity Management College of Natural Sciences

Addis Ababa University

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 1

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 stmctures. EH

Ertirea East, below and to the cast o f the 1000 m contour

EW - Eritrea west, above and to the west o f 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 o f the 1000 m contour GD - Gonder region GJ - Gojam region WU -W elo region, above and to the west o f the 1000 m contour

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

AR - Arsi region WG

Welcga region

IL -

Ilubabor region

KF - Kefa region GG

Gamo Gofa region

SD

Sidamo region

BA - Bale region HA Harerge region

MAP OF ETHIOPIA AND ERITREA SHOWING MAJOR PHYSIOGRAPHIC FEATURES

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 1 APIACEAE

to

DIPSACACEAE

Editors: Inga Hedberg, Sue Edwards and Sites hi Neniomissa

Addis Ababa, Ethiopia Uppsala, Sweden 2003

Published by The National Herbarium. Biology Department, Science Faculty, Addis Ababa University, Ethiopia and The Department o f Systematic Botany Uppsala University, Sweden Prepared by the Ethiopian Flora Project, funded by Addis A baba University and the Swedish International D evelopm ent Cooperation Agency (SID A ) through the Ethiopian Science and Technology Commission. Project L e a d e r- Dr. Sebsebe Demissew , and European C o o rd in ato r- Dr. Inga Hedberg. Final pages prepared at the Departm ent o f System atic Botany, Uppsala University using a Personal Com puter and a LaserJet 4L printer. Text formatted with Ventura Publisher 8, except some text sections which were prepared in W ord 97. Printed in Tim es New Roman Printed by the Educational M aterials Production and Distribution A gency (EM PDA), Textbook Production Departm ent, Addis Ababa, Ethiopia. © The National Herbarium , Addis Ababa University, 2003 ISBN 91-971285-5-4

Distributed by: In Ethiopia

Outside Ethiopia

The National Herbarium Addis Ababa University P.O. Box 3434, Addis Ababa, ETHIOPIA T e l.+251-1-11 43 23 Fax:+251-1-55 23 50 e-mail: nat.heb@ telecom .net.et

The Swedish Science Press, P.O Box 118 SE-751 04 Uppsala SW EDEN Fax: +46-18-36 52 77 e-mail: info@ ssp nu

Cover illustrations: Front: Psychotria orophila, drawn by Sarah Howard. Back: Four species o f Caralluma drawn by M argaret Tebbs. Spine: M ericaip (outer view) o f Agrocharis incognita drawn by M onica Schaffer-Fehre

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 1 CONTENTS

M ap o f the floristic regions o f Ethiopia and Eritrea M ap o f Ethiopia and Eritrea showing m ajor physiographic features

ii iii

Editorial Team

viii

Editorial Board

ix

C ontributors

X

Forew ord

xi

A cknow ledgem ents

xii

Key to fam ilies

xiii

Flora part 132. A piaceae by I. Hedberg & 0 . Hedberg

1

133. Ericaceae by I. Hedberg & 0 . Hedberg

46

134. Ebenaceae by 1. Friis & F. W hite

49

135. Sapotaceae by I. Friis

54

136. M yrsinaceae by Sebsebe Demissew

64

137. L oganiaceae by A.J.M . Leeuwenberg

70

138. O leaceae by P. Green

79

139. A pocynaceae by A.J.M . Leeuwenberg

87

140. A sclepiadaceae by F. Albers, M. Gilbert, D. Goyder, S. Liede & J. Venter

99

141. R ubiaceae by Ch. P uff

194

142. C aprifoliaceae by Ensermu Kelbessa

283

143. V aleriaceae by J.O. Kokwaro

285

144. D ipsacaceae by I. Hedberg & 0 . Hedberg

286

A G lossary o f Botanical Terms

291

A bbreviations

309

Index to Scientific Names

313

V ernacular N am e for Volum e 4, Part 1

331

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 1 Editorial Team Dr. Inga Hedberg, Department of Systematic Botany, Uppsala University, Sweden Prof. 01 ov Hedberg, Department of Systematic Botany, Uppsala University, Sweden Ms Eva Persson Department of Systematic Botany, Uppsala University, Sweden Prof. Ib Friis, Botanical Museum and Library, University of Copenhagen, Denmark Prof. Sebsebe Demissew, Keeper, The National Herbarium, Biology Department, Faculty of Science, Addis Ababa University, Ethiopia Dr. Ensermu Kelbessa, The National Herbarium, Biology Department, Faculty o f Science, Addis Ababa University, Ethiopia Ms Sue Edwards, The National Herbarium, Biology Department, Faculty of Science, Addis Ababa University, Ethiopia Mr Melaku Wondafrash, The National Herbarium, Biology Department, Faculty of Science, Addis Ababa University, Ethiopia Mr Sileshi Nemomissa, The National Herbarium, Biology Department, Faculty of Science, Addis Ababa University, Ethiopia

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 1 Editorial Board Dr. T ew o ld e B erh an G e b re E g zia b h er, G e n eral m anager, en v iro n m e n ta l P ro tectio n A u th o rity , A d d is A b ab a, E th io p ia E m eritu s P rof. O lo v H edberg, D e p a rtm e n t o f S y ste m atic B o tan y , U p p sa la U n iv ersity , S w eden P rof. S eb seb e D em issew , K e ep er, T h e N ational H e rb ariu m , B io lo g y D e p artm en t, F acu lty o f S cience, A d d is A b a b a U n iv ersity , E th io p ia D r M esfin T ad esse, (a lte rn a te m em b er) T h e N a tio n al H e rb ariu m , B io lo g y D e p artm en t, F a c u lty o f S cience, A d d is A b a b a U n iv ersity , E th io p ia Dr. E n serm u K e lb essa, (a lte rn a te m em b er) C u ra to r, T h e N atio n al H e rb ariu m , B io lo g y D e p artm en t, F ac u lty o f S cience, A d d is A b a b a U n iv ersity , E th io p ia Prof. Ib F riis, B o tan ical M u seu m and L ib raty , U n iv ersity o f C o p en h ag e n , D en m a rk Dr. In g a H ed b erg , D e p a rtm e n t o f S y ste m atic B o tan y , U p p sa la U n iv ersity , S w eden M s S ue E d w ard s, T h e N a tio n al H e rb ariu m , B io lo g y D e p artm en t, F ac u lty o f S cience, A d d is A b a b a U n iv ersity , E th io p ia

FLORA OF ETHIOPIA AND ERITREA VOLUME 4, PART 1 CONTRIBUTORS

Prof. Focke Albers, Institut fur Botanik der Westf. Wilhelms-Universitat, Schlossgarten 3, D-48149 Munster, Germany. Dr Ensemiu Kelbessa, The National Herbarium, Addis Ababa University, P. O. Box 3434, Addis Ababa, Ethiopia. Prof. Ib Friis, Botanical Museum, University of Copenhagen, Gothersgade 130, DK-1123 Copenhagen K, Denmark. Mr Mike Gilbert, Flora of China European Co-ordinator, The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, England. Dr David Goyder, The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, England. Dr Peter Green, The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3DS, England. Prof. Inga Hedberg, Department of Systematic Botany. NorbyvSgen 18D, SE-752 36 Uppsala, Sweden. Emer. Prof. Olov Hedberg, Department of Systematic Botany, NorbyvSgen 18D, SE-752 36 Uppsala, Sweden. Prof. John O. Kokwaro, Botany Department, University of Nairobi, P.O. Box 30197, Nairobi, Kenya. Dr Anthonius J. M. Leeuwenberg, Department of Plant Taxonomy, P.O.Box 8010, 6700 ED Wageningen, The Netherlands. Dr Sigrid Liede, Department of Systematic Botany, University of Bayreuth, 95440 Bayreuth, Germany. Dr Christian Puff, Institute of Botany, Univesity of Vienna, Rennweg 14, A-1030 Vienna, Austria. Prof. Sebsebe Demissew, The National Herbarium, Addis Ababa University, P. O. Box 3434, Addis Ababa, Ethiopia. Dr Johan Venter, Department of Botany and Genetics, The University of the Orange Free State, 339 Bloemfontein 9300, Republic of South Africa. Dr Frank White1, deceased, former Keeper of the Forext Herbarium, Oxford, England.

1. Frank White, former Keeper of the Forest Herbarium, Oxford, died on 12th September 1994. For 40 years he studied and published numerous works on Africa Ebenaceae. His many detailed taxonomic, phytogeographical and ecologi­ cal analyses, as well as carefully considered conclusions, will always associate his name with this family. Like the account of the Ebenaceae for the Flora of Tropical Fast Africa, the account given here is not based on any specific manuscript or work by White, but it has made use of all his published works which deal with the species found in the Flora area.

FOREW ORD T1 t-1 p re s e n t b o o k is th e first part o f V o lu m e 4 o f F lo ra o f E th io p ia a n d E ritrea . T he F lo ra P ro je c t w a s in itia te d in 1980 an d th e v o lu m es so far p u b lish e d are n u m b ers 2 (in 2 p arts), 3, 6 a n d 7. V o l­ u m e 3 p ro v id e s b a c k g ro u n d in fo rm a tio n ab o u t th e w ritin g o f a m o d ern F lo ra , n o tes on th e c o lle c t­ ing an d id e n tific a tio n o f p la n ts, an d in fo rm atio n on th e fo rm a t used for p re se n tin g fa m ilie s, g e n e ra am i sp ec ie s. T h e re a d e r is re fe rre d to th at v o lu m e fo r in fo rm a tio n d ea lin g w ith th ese m atters. A re ­ v ise d v e rsio n o f th e k ey to fa m ilie s is p ro v id ed at the b eg in n in g o f ea ch v o lu m e. V o lu m e 4 p a rt 1 tre a ts 13 fa m ilies, A p ia c e a e to D isp sac ace ae. C o m p a re d to th o se tre a te d in the e a rlie r p u b lis h e d v o lu m e s th is is a co m p a ra tiv e ly sm all n u m b e r o f fa m ilie s, but it in c lu d e s tw o Iai;ze o n es: A s c le p ia d a c e a e w ith 164 tax a and R u b iace ae w ith 132 taxa. A p ia c e a e w ith less th an ha; f th e n u m b e r, 57 tax a, is n e x t in size and w as d iffic u lt to p re p are due to lack o f a re c e n t re v isio n o f i he fam ily . T he fa m ilie s tre a te d co n tain a m o n g o th ers, g en e ra w ith c o m p lic a te d flo w e rs an d a rai ge o f im p o rta n t m e d ic in a l p lan ts (A sc le p ia d a c e a e and M y risin a c e a e ), as w ell as th o se c o n ta in ­ ing d ru g s an d s tim u la n ts (R u b ia c e a e ), th o se o f im p o rtan ce as sp ices an d co n d im e n ts (A p ia c e a e ), tin h e r tre e s (E b e n a c e a e , O le a c e a e and S a p o tace ae) an d o rn a m e n ta ls (A p o c y n a c e a e an d C a p rifo lia c e a e ). T h e se c o n d p art o f th e fo u rth v o lu m e w ill co n tain only one fam ily , th e v ery larg e an d d iffic u lt A s te ra c e a e .

AC K N O W LED G E M EN T S The production o f the F lo ra o f E thiopia a n d E ritrea enjoys the support o f many institutions and individuals, both within and outside Ethiopia. The editors particularly wish to acknow ledge the contributions o f the fol­ lowing who have m ade the publishing o f the present volume possible. The Ethiopian G overnm ent through the Ethiopian Science and Technology Com m ission, in particular the past and present C om m issioners, Mr Asrat Bulbula and Mr M ulugeta Amha, and the Swedish G overn­ m ent through the Swedish International Development Co-operation Agency (Sida), which has provided the financial support for the preparation and publication o f all volum es o f the Flora including this one; the past and present officers o f Addis Ababa University, especially the office o f the President and its current incum bent. Prof. A ndreas Eshete, the current Vice Presidents, Dr Gem etchu M egerssa (A cadem ic V P), Mr M oham m ed Habib (V P for Business and Developm ent), Prof. Endashaw Bekele, A ssociate VP for Re­ search and G raduate Program m e, the past and present Deans o f the Faculty o f Science, especially the pres­ ent Dean, Dr G ezahegn Yirgu, and the past and present Heads o f the Departm ent o f Biology, especially the present Head, Dr Kifle Dagne, all o f whom are gratefully acknowledged for the adm inistrative support they have rendered to the project. T hanks are also due to the Royal Botanic Gardens, Kew, in England, particularly the past and present D irectors, Prof. G illean Prance and Prof. Peter Crane, and the past and present Keepers o f the Kew H erbar­ ium, Prof. Gren Lucas and Prof. Simon Owens, and their scientific and technical staff, for their valuable support and contribution to the present volum e; to the Botanical Museum and Library o f the U niversity o f C openhagen, especially Prof. Ib Friis, both for his contributions and his sound and practical advice on many editorial m atters; to the C arlsberg Foundation for supporting extensive field work which has contributed much to im proving the coverage o f species and their distributions in the present volum e, and to the s ta if ot the Biology Library, U ppsala University, for their indefatigable assistance in tracing relevant literature and titles. Thanks to the courtesy o f various authorities, editors and Editorial Boards, it has been possible tc use illustrations already published in a num ber o f botanical publications. The following authorities and sources are gratefully acknow ledged: Adansonia (Publications Scientifiques du M useum national d ’H istoire naturelle, Paris), B otanische Jahrbucher Syst., Botanical Journal o f the Linnaean Society, Kew Bulletin. N a­ tional Botanic G arden o f Belgium , N ordic Journal o f Botany, The Um bellifers o f the British Isles (The Botanical Society o f the British Isles), The W ageningen Agriculture University Papers, ConspectusyF orae Angolensis, FJore d ’A frique Centrale, Flora o f Somalia, Flora o f Tropical East Africa, Flora o f W est T ropi­ cal Africa, Flora Za'mbesiaca (Flora Zam besiaca M anaging C om m ittee), Flore du Cam eroun, Kenya rees and Shrubs (N ational M useum s o f Kenya), South African Journal o f Botany, Flora Palaestina (The Israel A cadem y o f Sciences and H um anitie), and the M issouri Botanical Garden Press. Last but not least, the editors with their team s acknowledge the support o f their colleagues and friends: Drs Tam rat Bekele, Zem ede Asfaw and Zerihun W oldu, the Herbarium Librarian, M r Solom on K assie, and the technical staff and driver - M r M elkau W ondafrash, M iss Shewangziw Lemma, M iss Sosina Ketema, Mr A ssefa Hailu, M iss Y odit Dagne and M r Ermias Getachew.

KEY TO FAM ILIES1 by Sue Edwards, Ensennu Kelbessa & Sebsebe Demissew2

PRIMARY KEY TO 20 ARTIFICIAL GROUPS (Eich 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 'Dlants 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 lants with leaves less than one metre long, if trees or shrubs then leaves not grass-like nor palm-like. 4 4.

irass-like herbs: leaves alternate or basal, blades narrow and with parallel veins. Group 4 - 1 I mts not grass-like: leaves simple, deeply civided, or compound, usually with pinnate or palmate venation and usually with a petiole. 5 5. 1 owers densely crowded into compact inflores:ences (heads), individual flowers closely rressed against each other or against their ;jbtending bracts and often difficult to separate from each other. Group 5 - F owers not densely crowded into compact nflorescences (heads), individual flowers easy o separate from each other if in a dense inflorescence. 6 6. S imens and pistils naked, without bracts or erianth parts, or on 1 or 2 special bracts ?rming a cone, but not within a cup-like or ubular structure. Group 6 - S imens and pistils covered, surrounded by three r more bracts or perianth parts (sepals and/or ctals), or within a cup-like or tubular structure, but not forming a cone.

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. Flowers 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. 10 10. Each flower with two or more pistils and styles present and arising from separate ovaries, or if only one ovaiy 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 - Ovary on the receptacle, perianth parts attached at base of ovary, or on a disc or raised receptacle, or ovary below the level of perianth and stamen attachment. 12 12. Flower 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 - Flower with a superior ovary, ovaiy 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 Ag ^cultural Experiment Station 2 The National Herbarium, Addis Ababa University, P.O. Box 3434, Addis Ababa, Ethiopia

KEY TO FAMILIES

xiv

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 - Flowers 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 ] 1. Plants of ocean water or of salty water close to the ocean shore - Plants of fresh or of only slightly salty water.

2

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. ]6] 172. Hydrocharitaceae - 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 [6] 178. Zannichelliaceae - 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 flat 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; flowers in a single tubular bract, male reduced to several anthers on a stall 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. (Marsi lea) [1] Marsileaceae - 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. 12,1] 26.Podostemonaceae - Plants floating or submerged. 9 9. Underwater leaves with small (2 to 5 mm) round bladders that catch very small aquatic organisms. (51 163. Lentibulariaceae - Underwater leaves without small bladders. 10 10. Styles 2 or more, coming from separate ovaries or from separate areas on the same ovary, 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 1

i 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 ::,etals 3. pistils 3 or more, separate from each other and from the rcceptacle of the flower. [2,11 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. 14 - Pistils 3, 6, or more; leaves usually near the base of the plant. 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 .eaves in whorls, much divided with thread-like leaflets: pistil 1 with ovary 4-lobed. (Myriophyllum) [2,1] 51. Haloragaceae - _eaves in opposite pairs or alternate, simple, narrow; pistils 4 or more. 16 16. Mant growing erect out of the water, often in dense masses; sepals 4, petals present. (Crassula) [3] 88. Crassulaceae - Mant growing submerged; sepals 3, petals absent. (Zannichellia) [6] 178. Zannichelliaceae 17. lowers in 2-branched spike-like inflorescences. [6] 173. Aponogetonaceae lowers in umbels or racemes, or solitary in leafaxils and in whorls around the stem. 18 18. lowers usually solitary and on a long stalk, with many pistils and stamens in each flower; petals or more; leaves lobed or much divided. [2,1] 5. Ranunculaceae lowers usually in umbels or whorls along the stem; petals 3 or absent; leaves entire. 19 19.; :;imens 3 or 6, or more; fruit an achene. [6] 174. Alismataceae - Siamens usually 9; fruit a follicle. Butomaceae1 20.1

caves alternate and bipinnate; flowers in heads, ;ich flower with 5 or 10 stamens or the asexual owers with 10 petal-like staminodes. (Neptunia) [3] 95. (Leguminosae) Fabaceae

1Rec rded from East Africa but not in the Flora area.

- Leaves 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 thecae 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. Ceratophyliaceae - 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 lgae) - 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.

30 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 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-Iocular 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 33. Leaves opposite or whorled along the stem. - 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 orwith narrow 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. Styles 2, ovary with 2 locules, 1 ovule in each locule; flowers in umbels. [4,1] 132. (Umbelliferae) 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. [2,1] 49. Onagraceae 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. [2,1] 38. Polygonaceae GROUP 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 o f 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 pails of their hosts. 4 - Plants growing from below the ground, attached to the roots of their hosts if parasitic. 5

KEY TO FAMILIES

4 Flowers borne on the stem of the parasite and not directly on the stem of the host; leaves usually present. [3] 115. Viscaceae - Flowers or reproductive structures growing directly from the stem o f the host, the parasite without a stem or with only a short stem and without leaves. [2,1] 14. Rafflesiaceae t Flowers few to very many, if solitary on a stalk 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. Dracaenaceae 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 diy 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 - Ovaiy inferior; fruit with dry perianth or scars where 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. Dracaenaceae - 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] ‘Liliaceae’1 1See 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) [6] 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; inflorescence 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. [6] 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’. [7] 215. (Gramineae) Poaceae - Leaves in 3 vertical ranks or rows (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 structures; 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, leaf curled up in bud from apex to base like a fern leaf; leaflets very stiff, with veins that branch into 2; reproductive structures in cones or on a leaf-like part, young cones not enclosed in a woody boat-shaped structure or in a stiff bract. (1J Cycadaceae1 - 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. [61 214. (Palmae) Arecaceae

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

15. Flowers never present, reproductive structures usually brow n patches or lines on the underside of leaves producing minute powdery spores; petiole usualh with scale-like hairs; young leaves curled up with the tip innermost (ferns) (1) T h e F e r n s - Flowers and/or fruits and seeds usually present. petiole without scale-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 grow ing 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. Araceac - Leaves round in outline; flowers not enclosed in a tubular bract. 1* 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. (Nelumho) [2,1] 9. Nymphaeaceae - 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 (4 mm wide, usually with a hooked tip; branches usually borne in whorls at regular intervals along the main stem. [1 ] Araucariaceae Leaves tamens. 12,1] 31. Aizoaceae - Shrubs or small trees, or herbs without fleshy leaves; petals usually fewer than 8, not grading into non-functional stamens, but staminodes inay or may not be present. 39

39. Stamens 2 or 4; corolla usually 5-lobed and slightly irregular. - Si amens 5; corolla 5-lobed and regular.

40 41

40. C .osule obovate, usually broader above than iclow; seeds 2 to 8, large, on stiff stalks; leaves nd flowers not turning black or dark blue on drying. [5] 167. Acanthaceae - C ipsule usually widest in the middle or near the I ;ise; seeds many, small, not on stiff stalks; saves and flowers often turning black or dark 1 ue on drying. [5] 161. Scrophulariaceae 41. A 1 leaves opposite; inflorescence usually termil al; seeds globular. [5] 146. Gentianaceae - A least some leaves alternate or sub-opposite; i ilorescence usually axillary, but arising to one sine of the leaf axil; seeds flattened. [5] 158. Solanaceae 42. Ch iry with only 1 locule or the locule not visible. 43 - Ch ;ury with 2 or more locules. 46 43. Ov iry and/or style covered with short hairs. - Ov; iry and styles not covered with short hairs.

44 45

44.0 \ pry with 3 styles and many ovules; petals free a* cl borne on the calyx tube. [2,1] 17. Turneraceae - Ov iry with I style and 1 or 2 ovules; petals u ted and forming a tube. [3] 109. Icacinaceae 45. He tfc, climbers, or shrubs, sometimes with white S c o v a ry with 1 or 2 basal ovules or the ovary w hout a locule and non-fimctional. [5] 159. Convolvulaceae

xlvii

- Herbs or cultivated ornamental shrubs, without white sap, ovary with many ovules on 2 or more parietal placentas. ]5] 156. Hydrophyllaceae 46. Inflorescences supported by or enclosed in boat­ shaped bracts with parallel veins; leaf-base clasping the stem, secondary veins more or less parallel with the primary veins; ovary with 3 locules. [6] 207. Commelinaceae - Inflorescence not supported by or enclosed in boat-shaped bracts; leaves with reticulate venation; ovary with 2 or 4 or more locules or 2 separate ovaries each with 1 locule. 47 47. Ovary with 1 to 4 ovules in each locule. - Ovary with 5 to many ovules in each locule.

48 49

48. Herbs, shrubs, or more often climbers, often with white sap; corolla lobes very small and the corolla tube not deeply lobed, or if deeply lobed. usually shrubs with small ( ie short, slender, divergent. Fig. 132.31. iii open woodland and on nverbanks: 1300-2100 m. EE :AV GD GJ AR WG SU SD HA; widespread in tropi­ cal i .J rica. Mali lo Cameroun. cast to Somalia and south to the ransvaal mid SW Africa. Hildebrandt 332: B. Hill s.n. iloonev 5581.

37

Sometimes planted in villages for medicinal uses. Browsed by cattle. 2. S. commiphoroides Thulin (1991) - type from Somalia. Tree or shrub up to 10 m tall. Leaves pinnately trifoliolate or occasionally simple, leaflets ovate or suborbicular 1.55.5(-8) x 1-5.5 cm, serrate-dentate with the teeth ending in bristles up to 1 mm long. Flowering branchlets a few to­ gether at the tips of young twigs, c 3-^.5 cm long. Umbels all male or with the central flower bisexual and the outer male. Rays 4-9, umblets c 10-16-flowered. Calyx teeth small. Petals probably cream. Fruit obovoid, c 11.5-12.5 x 8-9 mm, with 3 prominent dorsal ribs, the outer 2 nar­ rowly winged, pale at maturity. Styles divergent. In woodland and bushland; 300 m. BA; Somalia. Gilbert et al. 8169.

30. FERULA L. (1753) Perennial herbs. Leaves pinnately divided with broad to filiform segments, leaf sheaths widely inflated. Umbels compound, polygamous, bisexual or unisexual. Bracts ab­ sent, bracteoles absent or a few. Calyx teeth absent or minute. Petals yellow. Fruit oblong-elliptic, very strongly dorsally compressed. Each meriearp with 3 dorsal narrow primary ribs and 2 marginal ones developed into wings. Vittae solitary in the grooves and 2 on the commissure to numerous all around the pericarp. Stylopodia shortly con­ ical. disc broad, usually with undulate margin. About 170 species, distributed in the Mediterranean region, Africa, Central and W Asia; 1 in the Flora area. F. communis L. (1753) F. abyssinica Hochst. ex A. Rich. (1847) - types: Oudgerate, Petit s.n. (P syn. not seen); TU without special locality, Ouartin-Dillon s.n. (P syn. not seen); nearTschelatchekanne. Schimper III: 1605 (P syn. not seen, K isosyn). F. ervthraeae Schweinf. ex Penzig (1893) nom. nud. Robust herb. 2-4 m tall. Basal and stem leaves with very conspicuous sheathing bases and long fine linear leaf lobes; upper leaves much reduced, inflorescence leaves mostly reduced to narrowly oblong, amplicaul sheaths, more or less acuminate. Umbels numerous, the lower of­ ten solitary and unisexual, the upper alternate or opposite, mostly 3 on a common branch with a larger central bisex­ ual umbel and 2 lateral male umbels, terminal umbels in groups of 3-5, the central bisexual and the lateral 2—3(-4) male. Peduncle of bisexual umbels obsolete or up to 5.5 cm. of males 2.5-11 cm long. Bracts missing in both. Bi­ sexual umbels with 10-25 rays, 1.5-5 cm long, umblets 10-18 flowered, pedicels 5-12 mm, bracteoles missing. Male umbels with 13-25 rays 1-2.5 cm long, umblets 10-25 flowered, pedicels 3-6 nun, bracteoles missing or up to 3 very narrow, linear. Calyx teeth distinct but fall­ ing off in fruit. Petals bright yellow with an acute

38

132. APIACEAE: 29. Steganotaenia. 30. Ferula. 31 Peucedanum

Figure 132.31. S T E G A N O T A E N IA A R A L IA CEA: 1 - leaves x ‘/ s; 2 - nflorescence x V&; 3 & 4 - mericarp. inner and outer view x 4'/j; 5 - transverse section o f mericarp x 4'/2. 1 from Chr. Smith s.n. and Swytwerion 176; 2 from Biittner, 3 -5 from Godman 203. Drawn by Monika Shaffer-Fehre. (Reproduced with permission from FI. Zamb. U m belliferae. Tab. 164.)

incurved apex. Fruit very strongly dorsally compressed, broadly obovate to broadly elliptic, c 10-17 x 5.5-12 mm, primary ribs prominent, narrow, vittae 3 in each dorsal groove but sometimes reduced to 2 or 1 in some, 4 in the commissure. Stylopodia depressed, crenaternargined. shortly conical; styles long, slender, recurved, c 2 mm. Fig. 132.32. Upland grassland; 1400-3250 m. EE EW TU GD GJ WU SU SD BA HA; S Europe, W Asia, N Africa, Sudan. Uganda, Kenya and Tanzania. S. Edwards et al. 5237; Frits et al. 6682; Mesfin T. 6324.

31. PEUCEDANUM L. (1753) Erect, glabrous, hairy', or tubercled biennial or perennial herbs. Stem almost smooth, striate, or deeply grooved. Basal leaves crowded together at base of stem to form a rosette. Leaves variously pinnately or temately divided into filiform to broad and coarsely toothed segments, leaf sheaths narrow or more or less expanded. Umbels com­ pound, few to many, the terminal ones of stem and branches bisexual, or the inner umblets or their inner flowers male, the opposite or verticillate outer umbels en­ tirely or partially male. Involucre missing or consisting of

132. APIACEAE: 30. Ferula, 31. Peucedanum

39

Figure 132.32. F E R U L A C O M M U N IS 1flowering branch x 2h: 2 - leaf x 2/ 3; 3 - male flower with parts removed x 12; 4 —fruit \ 2: 5 transverse section o f fruit x 6. \ & 3 from Tweedie 2648; 2 from from Eggeling 5617; 4 & 5 from Townsend 2333. Drawn by Christine Grey-W ilson. (Repro­ duced with permission from FI. Trop. E. Afr. U m belliferae, fig. 40.)

I few falling or numerous persistent bracts. Rays 8 or nore. Involucel usually conspicous of numerous bracte•les. Flowers usually polygamous. Calyx teeth absent to •mall but distinct. Petals white, yellow or greenish. Fruit s "ongly dorsally compressed, oblong to elliptic or obcnate, glabrous, truncate to slightiv emarginate at base ■id apex, usually with conspicuous ribs, each mericarp • tli 2 marginal nbs expanded into distinct, broad to i nher narrow thin or thicker wings; 0. 4 or 12 vittae visibie on the dorsal surface, 0, 2 or c 8 visible in the c ::mmissure. Stylopodia conical.

About 120 species in Europe, Asia and tropical and S Africa. 5 in the Flora area. A genus badly in need of revision, especially the spe­ cies P. mattirolii, P. petiticinum and P. winkleri. 1. Leaf blade glabrous, leaf segments less dian 5 mm wide; rest of plant mostly glabrous, but some­ times tubercled near nodes. 2 - Leaf blade hairy; at least along die veins, segments more than 5 mm wide; rest of plant ± haiiy. 3

40

132. APIACEAE: 31. Peucedanum

2. Fruits ± oblong, wings not exceeding stylopodium: stylopodium subtended by a well developed disc 1. P. abyssinicum - Fruits obovate-orbicular, wings exceeding stvlopodiunv. stylopodium not subtended by a well de­ veloped disc 2. P. harmsianum 3. Petals with 3 or more unbranched markings; mencarp always widest above middle. 4. P. petitianum - Petals with 1 central marking or none: mericarp not as above. 4 4. Fruit ovate, mericarp wings tapering towards apex. ending below sty lopodium. 3. P. mattirolii - Fruit elliptic, inencarp wings not tapering, extend­ ing a little beyond base of stylopodium. 5. P. « inkleri 1 P. abyssinicum Vatke (1876) tvpe: GD. Gafat. c 2700 m. Schimper 1863-68: 1147 (BM K iso.). P. silaifolium Hiem (1877) - type: Abyssinia 2500 m, Schimper 1863-68 s.n. (K iso.). Perennial herb, up to 1 in tall, largely glabrous. Leaves vanously pinnate, finely divided: ultimate divisions lin­ ear to linear-spathulate. acuminate. Bracts absent or up to 3. c 3 mm long. Rays (6—)8—13(—15). up to 40 mm long. Pedicels up to c 12. 2-10 mm long. Bracteoles 1-4, c 2 mm long. Petals white with a single central marking, more or less regular. Mericarps ovate-oblong, up to 10 x 8111111. Vittae 4 dorsal. 2 commissural. Stylopodia shorth conical with well developed disc. Sty les very short, deflexed Damp grassland: 2100-2800 m. TU GD SU AR SD: not known elsewhere. Ash 2610; Frtis et al. 3829; Gilbert et al. 8331 2 P. harmsianum MoIff (1927) - type from Kenya. P. muJfivittatum Cuf. (J939). non Maxim. (1887). nom. illegil. - type: SD. Javello. Cufodontis 457 (FT holo.. not seen). Peremual herb, up to 70 cm tall, frequently branched, with thick, cylindrical rootstock. Stem at first tubercled. at and below nodes and at top of peduncles, but becoming glabrous. Leaves finely divided, variously pinnate: ulti­ mate divisions linear to spathulate. acuminate. Bracts up to 6. up to 8 mm long. Rays 8-15. 20-45(-60) nun long Pedicels (8—)13—18. up to 10 nun long. Bracteoles c 3. up to c 3 mm long. Petals white, more or less regular, with a central marking. Mencarps obovate or orbicular, up to 11 x 7 nun Vittae 12 dorsal. 3 between each rib, 6-8 commissural Stylopodia shortly conical. Styles very short, deflexed. Fig. 132.33. Closely grazed grassland: 1800-1900 m. SD; Ken\a and Tanzania. Gilbert & Jefford 442; Gilbert & Sebsebe D. 8808: Mesfin T. & Vollesen 4258. 3. P. m a ttiro lii Chiov. (1929) - type: AR, Bekoji. Galama valley. Basile 61 (TO holo. not seen).

Perennial herb. 1-2 m tall. Stems sometimes with scat­ tered short hairs Branches arising singly or in pairs. Leaves 2-3-pinnate. with rather few divisions, lanceolate to ovate, rather regularly compound-serrate: leaflets haiiy. particularly on veins of lower surface. Bracts 1—3(—5). 3-15 nun long, linear-lanceolate Rays (6-) 8-12(-17). 15-50 mm long. Pedicels 10-15. up to 9 mm long. Bracteoles 1-3, up to 8 mm long, linear, curving Petals yellowish to white, regular, with a central marking Mencarps 7-9 \ 4-6 mm; (ovate or) elliptical, wings ta­ pering at apex so sometimes almost beaked, truncatt to cordate at base. Vittae 4 dorsal. 2 commissural. Stylopo­ dia almost stalked; style short, reflexed. Stream banks, sometimes in Juniper forest; 26003050 m. SU AR SD; not known elsewhere. K. Hylander al. 305; Mesfin 7 . 812; Sebsebe D. 76.

4 P. petitianum A. Rich. (1848) type: GD. Wegera (Ouedgerate). Petit s.n (P holo.). P. ahum Hiem (1877) - type: GD. Gafat Schimper 1214 (BM holo.. K iso ). Peremual lieib, 1-3 in tall, hairy, densely branched to­ wards top of plant: branches whorlcd on stem. Leaves rad­ ical and cauhne. very large ( up to ? 1 in or more long): up to 3-piiuiate or 3-tcmate. with few pairs of leaflets: termi­ nal leaflets shortly stalked or sessile, broadly ovale to lan­ ceolate. (20—)30-80(—140) x (15—)25—35(—45) nun: margin shallowly serrate to crenate. hairy on veins and margins of both surfaces. Bracts and bracteoles 1—1. inconspicuous. Rays (8-)10-18(-24). 15-45 mm long. Pediccls (13 )20 (-25), (5—)6—10(-12) mm long. Petals while, irregular, with 3 or more central markings. Mericarps obovate 8-12 x 5-7 mm. Vittae 2 in the commissure. Stylopodia subtended by undulate disc. Style very short, strongly rcflexed. Thicket margins, sometimes at edges of streams and forests. 1750-3350 m. TU GD GJ SU AR WG KF BA HA. not known elsewhere Gillett 14555, Mesfin T. & Kagnew G. 1678: W. de Wilde 8832.

5 P. winkleril^ ’0/^(1912) - types from Cameroun and Tanzania. Perennial lierb, ( 1-) 1.5-3 m tall, densely branched, with a taproot Branches in whorls on stem and larger branches. Leaves radical and cauline; usually 3-pinnate and then pinnatifid: irregularly divided, so leaflets indefinable, but lobes c 20-65 x 10-20 nun, crenate or bicrenate. hairy on veins and margins of both surfaces. Bracts up to 5. 4(-5-10) mill long. Rays (7-) 11-20, 20-60 mm long. Pedicels (9-)15-20(-22), (5 -)6 -l 1 (—14) mm long. Brac­ teoles up to 4, 3-5 mm long. Petals white with or without a single central marking. Mericarps 6-9(-14) x 4—“(—10) mm, elliptic, often cordate at base; wings extended a little beyond base of stylopodium. Vittae dorsal or in the commissure, not visible externally. Stylopodia cush­ ion-shaped to shortly conical. Styles very short, deflexed

132. APLACEAE: 31. Peucedanum. 32. Lefebvrea

41

Figure 132.33. PEU C ED A N U M H A R M S IA N U M . 1 - flowering plant x 2/ 3; 2 & 3 - meriearp, outer and inner view x 3; 4 - meriearp, transverse section x 6; 5 - narrower form o f fruit x 3. All from Town­ send 2311. Drawn by Christine Grey-WiJson. (Reproduced and modified with permission from FI. Trop. E. Afr. U m belliferae, ftg. 33, as P. harmsianum subsp. harmsianum.)

Wet sites, often among Erica, in Podocarpus or A •mdinaria forest; (1600-)2600-3700 m. GD GJ SU AR KF BA H A . Caineroun. Tanzania. F riis et al. 1606, 3607; Burger 1012.

32. L E F E B V R E A A. Rich. (1840) Biennial or sometimes perennial, glabrous herbs with a I; :)root. Stem hollow, finely striate. Leaf sheaths narrow; le aves usually regularly divided, leaflets linear and entire 0 very sliallowly and remotely serrate, to ovatelai i;eolate and irregularly crenate. or sometimes divided

almost to midrib; veins smooth, rarely minutely verruculose; venation a prominent midrib and a parallel longitudinal vein on each side. U m bels com pound, polyg­ amous or with bisexual flowers only, in a similar way as in Peucedanum. Rays and pedicels long, glabrous. Involu­ cre absent or of up to 8 filiform bracts; involucel of up to 8 filiform bracteoles. Umblets usually polygamous. Calyx teeth obsolete. Petals yellowish or green, more or less flushed with brown or purple. Fruit strongly dorsally com­ pressed, broadly elliptic or obovate, deeply incised at the apex, 3 slightly prominent dorsal ribs, 2 marginal ribs ex­ tended into broad thin wings overtopping the stylopodia. Vittae solitary in the grooves, 2-4 on the commissural

132. APIACEAE: 32. Lefebvera

Figure 132.34. L E IE B V R E A A B Y S S IN IC A 1 - lower pari o f stem with rootstock x Vi; 2 leaf x V2; 3 - flowering branches x Vi, 4 - fruit x 3; 5 - transverse section o f mericarps x 10. 1 from Exell & Mendonca 1727. 2 from Exell & Mendonca 1278; 3-5 from Gossweiler 12308. Drawn by Derek Eras­ mus. (Reproduced with perm is­ sion from Consp. Fl. Angol. IV Tab. XLIII as L. stuhlm annii.)

face, all black and conspicuous. Stylopodia deeply sunk, elongate or shortly conical or cylindrical clavate and ob­ tuse; styles short. 6 species widely distributed in tropical and SW Africa, 1 in the Flora area.

L. stuhlmannii Engl. (1895); Peucedanum stuhl­ mannii (Engl) Drude (1898). P. le/ebvria Drude (1898).

Sparsely branched perennial (sometimes biennial?) heib up to 2.5 m tall. Lower leaves bipinnatisect, or occasion­ ally pinnate, with few pairs of leaflets, median leaves pro­ L. abyssinica A Rich. (1840) gressively less and more narrowly divided, upper leaves type. TU, Yeha. X.1839, Q u a rtin -D illo n s.n. (P reduced to tapering sheaths subtending the flowering syn not seen); Mt Scholoda near Adoua, S c h im p e r s.n. branches. Bracts absent or a few. up to 10 mm long. Rays (P syn. not seen). up to 17, slender, up to 12 cm long. Bracteoles up to 8, c 8 L. a b yssin ica A. Rich. var. a n g u stise cta Engl., in mm long. Umblets up to 25-flowered. pedicels up to 3 cm P flanzenw . Ost. A fr. C: 300 (1895).

£

132. APIACEAE: 32. Lefebvrea, 33. Ervthroselirium

43

Figure 132.35. E R Y T H R O S E L IN U M A T R O P U R P U R E U M : I - part o f flowering plant x 2/ 3; 2 - single leaflet x 2; 3 - lower cauline leaf o f narrower-leaved form x 2/ 3; 4 - single leaflet from same x 2; 5 - petal, ventral surface x 24; 6 & 7 - meriearp, outer and inner view x 4; 8 - meriearp, transverse section x 12. 1, 2 , 5 -8 from D rum m ond & Hemsley 4449; 3 & 4 from Bogdan 3280A. Drawn by Christine Grey-Wilson. (Reproduced with permission from FI. Trop. E. Afr. U m belliferae, Fig. 36.)

loni Petals y ellowish or flushed with greenish, brownish, or p . rple. Fruit obovate, 7—10 x 6-8 mm, primary' ribs obscur. wings extended beyond the stylopodium. Vittae solii iry in the grooves, 2 in the commissure. Stylopodia lon^ narrowly conical, tapering gradually to the erect or divorgent, long and slender styles. Fig. 132.34. Montane grassland and in disturbed places; 17002001 m. TU GJ SU WG KF SD; tropical Africa, south to Zirn labwe and west to Cameroun Friis et al. 190; Gilbert d S .'via Phillips 8854; Mooney 5901.

33.

ERYTHROSELINUM Chiov. (1911)

Townsend in Kew Bull. 42(3); 590-91 (1987). Herbs, probably biennials or short-lived perennials, with more or less thickened rhizome. Stems richly branched, slightly glaucous. Leaves pinnate. Bracts missing or a few'. Umbels numerous, compound, polygamous, with one central bisexual umbel and 1 or 2 lateral male umbels. Rays up to 6. Bracteoles few. Umblets 2-13. Calyx teeth missing. Petals purple, verruculous, with an incurved api­ cal lobule. Fruit elliptic-oblong, with shallow and ill-

44

132. APIACEAE: 33. Erythroselinum, 34. Heracleum

Figure 132.36. H E R A C L E U M A B Y S S IN IC U M 1habit x '/2; 2 & 3 - umbels with devel­ oping fruits x Vi, 4 - leaflet x 1'/3. 5 underside o f leaflet x 3; 6 - umbel with mature fruits x V2; 7 - mericarps, outer and inner view x 3; 8 - meri­ carps in cross section x 8. 1-5 from Robson 389; 6 -8 from Sebald 651. Drawn by Ann Davis. (Reproduced with permission from FI. Zeunb. U m belliferae, Tab. 165.)

defined ribs, dorsaily compressed with a distinct margin, glabrous. Vittae solitary and narrow beneath each dorsal primary rib, solitary and broader ones beneath each groove. Stylopodia cushion-shaped, styles slender.

Description in general as for the genus. Erect herb up to 1 m tall Bracts missing or a few. Bracteoles c 3. Fruit c 5 x 2.5 mm. Styles deflexed, as long as or slightly exceeding the stylopodia. Fig. 132.35.

A monotypic genus occurring in NE and E Tropical Africa.

Open bushland; 1200-1850 m. TU SU SD; also in Uganda and Kenva. M G. Gilbert et al. 8073, 8222. Tewolde B.G.E. 1029.

E. atropurpureum (Hochst. ex A. Rich.) Chiov. (1911); Pastinaca atropurpurea A. Rich.; Malabaila atropurpurea (A. Rich.) Vatke (1876); Peucedanum atropurpureum (A. Rich.) Hiem (1877) - type: TU, near Gapdia. Schimper II: 788 (K iso.). Peucedanum ruspolii Engl. (1897) - type. GG-SD, Dscharibule, Riva 402 (FT not seen).

As pointed out by Townsend (Kew Bull. 42 (3): 587604, 1987) the genus Erythroselinum is very close to both Peucedanum and Lefebvrea. Pending a thorough re­ vision of the whole group the authors prefer to follow Townsend in maintaining Erythroselinum as a separate genus.

132. APLACEAE: 34. Heracleum

34. H E R A C L E U M Z . (1753)

B .nnial or perennial herbs. Leaves simple and lobed to 1 2(-pinnately or temately divided with broad dentate seg­ ments. sheath often large and soft. Umbels numerous, compound, mostly bisexual but sometimes with male flfcwers in the centre. Bracts few or absent. Bracteoles s e . eral. narrow. Calyx teedi small, usually soon falling. Peals white, pink or yellowish-green, often radiating. Fr ut suborbicular-obovate, glabrous to hairy, very suongly dorsally compressed, with 3 filiform but often quae prominent dorsal ribs, and 2 lateral ones expanded in o wings. Vittae solitary in the grooves, conspicuous fn in the exterior. 0-2 on the commissure, sometimes sh rter than the mericarp. stylopodia conical from a br id, crenate-margined. discoid base; styles long and sk nder. About 70 species in north temperate regions and mi untains of tropical Africa; 2 in the Flora area. 1 Petals of outer flowers of outer umblets small (1.5-3 mm); rays and pedicels densely covered with glandular hairs. 1. H. abyssinicum - Petals of outer flowers of outer umblets large (4-9 mm); rays and pedicels pubescent. 2. H. elgonense 1. I. abyssincum (Boiss.) Norman (1936); Malabaila abyssinica Boiss. (1844) - type: TU, Mt Selleuda near Adua, Schimper I: 204 (G holo.. K iso.). M. rn ae Engl. (1897) - type: SD. Biddume-Alghe, y.iva 1229 (FI iso. not seen). Ro :ust biennial or perennial, up to c 1 m tall or more, dei ..;ely pilose. Leaves up to 30 x 7 cm. simply pinnate with 3-7 pairs of ovate-deltate leaflets. Median stem leaves shortly petiolate or sessile on the sheath, uppermost leaves much reduced, often with a veiy small blade. Leaf­ lets dentate. Umbels several. Bracts usually present, sometimes very small, linear. Rays up to 18, sulcate, densely provided with glandular hairs. Bracteoles 4-9.

45

linear. Umblets up to 35-floweied, pedicels up to 10 nun long with dense, glandular hairs. Calyx teeth narrow tri­ angular to linear, sometimes persistent in fruit. Petals white, slightly radiating, the outer ones sometimes emarginate. Fruit obovate to oblong-obovate. c 7-12 x 5-8 nun, pilose. Vittae 4 dorsally, 2 in the commissure, stylopodia conical, not exceeding the apical notch of the fruit, styles long, slender, recurved. Fig. 132.36. Open grassy slopes with Juniperus, Maerua, Rhus; bushiand, Erica-scrub. riverside; 1800-3750 m. EW TU GD GJ SU AR GG BA HA; from Ethiopia through Rep Dein. du Congo, Uganda. Kenya and Tanzania to Malawi. Burger 2399; Friis et al. 1339; W. de Wilde 6393.

2. H. elgonense (WolfJ) Bullock (1932); Malabaila elgonensis H. Wolff (1922) - type from East Africa. Heracleum inexpectatum Norman (1927). Robust, juicy biennial or perennial herb up to 1 m tall. Leaves up to 40 x 8 cm, basal leaves several in a loose ro­ sette, simply pinnate with roundish, or bluntly deltoid leaflets, median stem leaves shortly petiolate or sessile on the sheath, with broadly ovate, dentate leaflets. Umbels several. Bracts absent or a few. Rays 12-20. pilose. Bractcoles up to c 12. at least some exceeding the pedicels in young fruits; pedicels pilose. Calyx teeth triangular, soon falling. Petals white, sometimes externally tinged crim­ son. dramatically radiating, the outer more than 4 times longer than the inner ones, deeply emarginate. Fruit broadly elliptic, c 6-10 x 3.5-6 mm, pilose; vittae 4 dor­ sally. 2 on the commissure, stylopodia conical, exceeding the apical notch of the fruit; styles long, slender, deflexed to recurved. Open areas and in light shade; 3000-4150 m. SU AR BA; Kenya. Uganda. Sudan. Mesfin T. 5815; G. & S. Miehe 700; Mooney 7148.

46

133. ERICACEAE by I. Hedberg* and 0. Hedberg* Pichi-Sermolli & Heimger, Webbia 9: 9-48 (1953); Hedberg, Syrtib. Bot. Ups. 15(1): 140-149 (1957), Nordic J. Bot. 5. 463-467 (1985); Cufodontis, Enum.: 653-654 (1960); Ross, Ericaceae in FI. Zarnb. 7(1): 157-181 (1983); Judd, Joum. Am. Arboretum 65(3): 255-342 (1984); Oliver, S. Afr. J. Bot. 53(6): 455-458 (1987), Bothalia 23: 1-7 (1993), Ke+>Bull. 48(4): 771-780(1993).

Small trees, shrubs or subshmbs. Leaves simple, alternate, opposite or whorled, without stipules, usually ever­ green. Flowers bisexual, actinomorphic or slightly zygomorphic, axillary or in simple or congested, terminal racemes. Calvx and corolla usually 4- or 5-merous. Sepals free or fused, one of them sometimes larger than the others. Corolla usually sympetalous, 3-5-lobed. Stamens equalling the petals in number or twice as many, inserted on a receptacle disc; filaments normally free; anthers with apical pores and often with appendages. Style 1. Car­ pels usually of the same number as the petals, united to form a usually 3—5-locular or occasionally 1-locular ovary, which is mostly superior. Fruit a capsule, drupe or berry. Seeds, in most genera, several per locule. An almost cosmopolitan family o f about 100 genera and c 3400 species, mainly occurring in temperate lati­ tudes and at high altitudes in the tropics and subtropics. 2 genera with 4 species are known from the Flora area. Recent studies by Oliver (Kew Bull. 48) show that the genera Philippia Klotzsch and Blaeria L. can no lon­ ger be kept apart from Erica, and Judd (loc. cit.) reduced the genus Agauria to a synonym of Agarista. Key to genera 1. Leaves in whorls, less than 1 cm long, needleshaped; calyx and corolla 4-merous. 1 Erica - Leaves alternate, more than 3 cm long, ellipticovate; calyx and corolla 5-merous. 2. Agarista 1. ERICA L. (1753) Blaeria L. (1737) Phillipia Klotzsch (1834) Trees, slirubs and subshrubs, usually richly branched, with slender stems. Leaves mostly in whoris of 3-6, needle-shaped, veiy small, with revolute margins meeting in the middle of the abaxial side. Inflorescences of clus­ ters of up to 20 flowers on branchlet tips. Pedicels short and one-flowered, normally each carry ing one bract and two bracteoles. Calyx and corolla 4-merous. actinomorphic and with superior ovary. Sepals free or partly fused. Co­ rolla globular to tubular with the lobes shorter than the tube. Stamens normally 8 but occasionally fewer (4-7) or more numerous (up to 12); filaments free; anthers often with appendages. Style 1; ovary' usually 4-celled; capsule with several seeds per cell. Fruit a cartilaginous, loculicidal capsule which remains inside the persistent calyx and corolla. A large genus with more than 600 species, most of which are confined to the SW part of the Cape region in South Africa, but also extending through highland areas in East Africa and Ethiopia to the Mediterranean region northwards to NW Europe. 3 species in the Flora area. 1. Corolla tubular, calyx pubescent from long. branched hairs; dwarf shrub. 3. E. tenuipilosa - Corolla campanulate, calyx glabrous or with mar­ ginal glands only ; trees or shrubs. * Dept. of Systematic Botany, I'p p sala University, Norbyvagen 18D. SE-752 36 Uppsala. Sweden.

2. Pedicels glabrous with 2 or 3 small bracteoles be­ low the middle; all flowers actinomorphic. 1. E. arborea - Pedicels glandular, with one ± recaulescent bract, making most flowers zygomorphic. 2. E. trimera 1. E. arborea L. (1753) - type in Herb. Bmser XXV.42 (UPS lecto.). E. acrophya Fresen. (1838) - ty pe: Ethiopia sine loc. Rilppell s.n. (FR holo. not seen). Richly branched shrub or small tree up to 8 m tall, with grey-brown stem. Young twigs densely pubescent with both short simple hairs and longer, branched hairs. Leaves in whorls of 3 or 4. linear, 3-5 mm long, glabrous. Flowers campanulate in thyrsoid, terminal or intercalary panicles. Pedicels glabrous, with 2 or 3 minute bracteoles below the middle. Sepals ovate, 1.5 mm long, glabrous, saccate at base. Corolla white, broadly campanulate, 2.5-4 mm long, with erect lobes. Stamens 8. Anthers brown with append­ ages, included in corolla. Ovary glabrous; style stout, con­ siderably overtopping corolla, stigma flattened, white. Fig. 133.1-3. I Steep, rock)' slopes and wooded, often burnt and grazed grassland, sometimes on exposed lava flows, often kept down by burning. Important dominant between 2200-3900 m.’ EW GD GJ WU SU AR KF SD BA HA; Canary Island. Madeira, Mediterranean Region, Somalia, East Africa Gilbert & Tewolde B.G.E. 3274a; Friis et al. 5688; Hedberg & Getachew A. 5424. 2. E. trimera (Engl.) Beentje (1990) Philippia trimera Engl. (1895) - type: Uganda, Ruwenzori, Stuhlmann 2445 (B holo. destroyed). P. abyssinica Pic.Senn. & H eia (1953); P trimera subsp abyssinica (Pic.Serm. & Hein.) Hedb. loc. cit. type: AR, Scire, Milchersich 44 (FI holo. not seen).

233. ERICACEAE: I. Erica, 2. Agarista

S :irub or tree up to 12 m tall, with richly branched stems up to 25 cm in diameter. Branchlets with varying amounts of pubescence of short simple hairs and glandular hairs of different sizes. Leaves in whorls of 3; petiole c 1 mm. pub .'scent; blade linear, 4-6 mm long and up to 1 mm wide, s imetimes pubescent above. Flowers mostly zygomorphic from a fully recaulescent bract, but occasionally mere or less actinomorphic because of an only partly b caulescent bract; the frequency of such flowers varies among different collections. Calyx glabrous or with mar­ ginal glands. Corolla campanulate. c 2 mm long and 2 mm v ide, creamy white, greenish or brownish. Stamens 8, inc uded, anthers brown without appendages. Ovary glab'ous. style stout, overtopping corolla stigma flattened. Fig 133.1.4. On rocky and sometimes boggy' ground from the upper part of the montane forest belt into the lower part of the a iroalpine belt; 3200-4200 m. GJ SU AR GG BA; Rep. C ::m. du Congo, Uganda Kenya, Tanzania. Friis et al. 5 533 ; Gilbert & Tewolde B.G.E. 3274; Hedberg 4250. This species is very variable e.g. in life-form, in freq lency of flowers with only partly recaulescent bract, and in pubescence of twigs. In part this variation may be a inbutable to different stages of regeneration after bumii g It is also worth noting that the largest variation occurs ii Bale mountains, which have a much larger alpine area than any other African mountain.

3. E. tenuipilosa (Engl, ex Aim & Fries) Cheek (1997); Blaeria tenuipilosa Engl, ex Aim & Fries (1924) types; SD, Avara, Ellenbeck 1814 (B lecto. de­ stroyed); Kenya Mt Aberdare. Fries & Fries 2579 (UPS iso.); Cameroun. “Bambutoberg”. Ledermann 1625 (B iso. destroyed. UPS lecto.). B. condensata Hoclist. ex A. Rich. (1850) non Erica condensata Benth. - type; GD, Mt Selki. Schimper 11:667 (P holo., BM S UPS iso.). B. spicata Hochst. ex A. Rich. (1850) non Erica spicata Thunb, - type: GD, Mt Buahit. Schimper 11:749 (P holo.. B FI FR G K RO S UPS W iso.). B.guguensisPic.Serm& Hein. (1953)-type: AR, Mt Gugu, 3000 m.Milchersich 77 (FI holo. not seen).

subsp. spicata Cheek (1997) stat. et comb. nov. B. spicata Hoclist. ex A. Rich. (1850) - type: GD, Mt Buahit, Schimper 11:749 (P holo., B FI FR G K RO S UPS W iso ). Shrub up to 0.4 m tall, usually richly branched from the tl ickened base. Stem covered with short, simple and long, b istly, often branched and sometimes gland-tipped hairs. Leaves 2.2-4.5 x 0.4-0.9 mm, usually dull from a dense C( ver of minute, simple hairs and also provided with long, o len branched and sometimes gland-tipped hairs, partic­ ularly along the margins. Sepals broadly lanceolate/ovate w tth the same type of pubescence as the leaves, 1.3-2.4 x 0 aj-0.7 mm, their length/width ratio variable but usually less than 5. Corolla tubular, 2.8-5.8 x 1.2-2.2 mm, slight­ ly expanded towards the top. Stamens 4; anthers brown

Figure 133.1. E RIC A ARBOREA'. 1 - flowering branch x c Vi; 2 - flower x 7; 3 - stamens dorsal and lateral view x 8. E. TRIM ERA: 4 - flower x 7; E TEN U IP ILO SA subsp. SP IC A T A : 5 - flower x 9. 2 from Hylander et al. 257; 4 from Hedberg 5537; 5 from J.J.F.E.de Wilde 5716 (no specimen given for 1 & 3). 1 drawn by Ingvar Nybom; 2, 4 & 5 drawn by Ylva Stenlund; 3 redrawn from FI. de France 4, PI 207 p.p. (1; reproduced with permission from Bremer et al. Introduction to Phytogeny andSystematics o f Flowering Plants, 6th ed., p. 85.)

with appendices. Ovary pubescent, style slender, overtop­ ping the corolla; stigma flattened. Fig. 133.1.5. In ericaceous scrub and grassland, often on steep and rocky ground; 2300-4000 m. GD GJ SU AR GG BA HA; also in Sudan. Hedberg & Getachew A. 5423; Mooney 5201; P uff et al. 820929-1/3. Subsp. tenuipilosa Cheek occurs in West Africa.

2. AGARISTA D. Don ex G. Don (1834) Agauria (DC.). Hook. f. (1876) Small trees up to 8 m tall, with rough, fissured bark and fragile branches, often regenerating from stumps. Leaves alternate to subopposite, leatheiy. subsessile to petiolate, evergreen, lanceolate to ovate. Inflorescences axillary and terminal racemes. Pedicels short, 1-flowered, subtended by caducous bracts and each carrying a pair of caducous bractlets. Calyx and corolla actinomorpliic, 5-merous and hypogynous. Sepals fused below, persisting in fruit. Co­ rolla obconical, with 5 small, terminal lobes, soon falling. Stamens 10. adherent to the base of the corolla and shed with it. Ovary superior, 5-celled. with few ovules. Fruit a 5-vaIved. loculicidal capsule.

48

133. ERICACEAE: 2. Agarista

Figure. 133.2. A G A R IS T A SA L IC IF O L IA : 1 - flow­ ering branch x Vyy 2 - flower x 4 .3 - ca­ lyx and ovary x 4; 4 - stamen x 8. 5 fruiting branch x 2/ 3; 6 - capsule with calyx and style x 2. All from Robinson 5706. Drawn by Ann Davis. (Repro­ duced with permission from F l Zamb. E ricaceae, Tab. 26 as A gauria salici­ fo lia .)

A genus of 30 species, all in America except for one, restricted to Africa A* salicifolia {Comm, ex Lam.) Don (1834); Andromeda salicifolia Comm, ex Lam. (1783); Agauria salicifolia (Comm, ex Lam ) Hook.f. ex Oliv. (1877) - type: from Reunion. Shrub or small tree up to 8 m tall, branchlets glabrescent. Leaves lanceolate, entire, up to 9 x 3 cm, somewhat leathery, midrib on the abaxial side in younger leaves pu­ bescent; petioles usually c 1 cm but up to 1.7 cm long. In­ florescences 10-20-flowered, up to 11 cm long. Pedicels 1-5 nun long, peduncles and pedicels glabrous or weakly

pubescent. Calyx with 5 triangular segments, 3 nun long, fused for about half of their length Corolla green, yellow­ ish green, ory ellow, c 8 x 4 mm with small c. 0.5 mm long lobes. Style slightly longer than corolla; stigma capitate. Capsule 3 mm in diameter (dry). Fig. 133.2. Steep, rocky slopes in Juniperus - Erica scrub, open grassland with Combretum, Protea, Erica and Helichrysum, bamboo forest, evergreen forest relict in valleys, (1600-)2050-c 2900(-3200) m; SU AR KF; widespread in Africa, also in Madagascar and the Mascarene Islands. Gilbert & Tewolde B.G.E. 3225; Gillett 15058; Mooney 8876. Leaves reported medicinal for scabies.

49

134. EBENACEAE I. Friis* & F. White** C» fodontis, Enum.: 667-669 (1960); White, Ebenaceae, in FI. Zamb. 7(1): 248-300 (1983); Friis, Forests and Forest T i. es of Northeast Tropical Africa, Kew Bull. Add. ser. XV: 215-220 (1992); Verdcourt & White, Ebenaceae, in FI. Tr :ip. E. Afr:. 52 pp. (1996).

Ti ;.*es or shrubs; trunks often with dark brown or black hardwood. Leaves simple, alternate, entire, without stipul VS Inflorescences usually cymose, always determinate, sometimes false racemes, or the inflorescence redi ced to a single flower on a bracteate peduncle. Flowers regular, hypogynous, unisexual, but usually with rudiments o f the other sex, 3-8-merous. Calyx of united sepals, entire and truncate to deeply lobed, always per­ sistent in fruit and usually enlarging. Corolla of united petals, shortly to deeply lobed; tube sometimes fleshy and constricted at the throat; lobes always twisted to the left in bud. Stamens from (2-)3 to over 100, epipetalo is or borne on the receptacle, exserted or included; filaments often very short; anthers basifixed, often o f un­ equal size and often more than one on the same filament. Staminode development very variable, rarely absent. Pi -;tillode very variable in development rarely completely absent. Ovary with 2-8 carpels fused to the base of the styles, each carpel with 2 ovules, sometimes divided by a false septum into 2 uniovulate locules; styles equal in number to the carpels, distinct or united at the base, very rarely completely united; stigmas usually laj ge and conspicuous; ovules apical, pendulous. Fruit usually a berry, rarely a fleshy, tardily dehiscent capsu e. Seeds large, with a distinct circum-peripheral vascular loop; hilum small, inconspicuous. 2 genera and more than 500 species distributed throughout the tropics and subtropics o f both hemispheres, with most species in the Indo-Pacific region, but with a centre of variation in Africa; represented by 5 species fr< m both genera in the Flora area. The position of the embryo in the endosperm is of taxonomic importance and can be used to separate the ge lera. In Diospyros, the embryo is straight or curved on one plane; and the radicle is not in a special ‘pouch’ of thi testa, or only in lower half (Fig. 134.1.4), while in Euclea, the embryo with its cotyledons is twisted at a rit lit angle to the radicle, and the radicle is surrounded by an ingrowth o f the testa for the whole o f its length (F g 134.1.10). K< v to genera 1 Inflorescence cymose or clustered, or flowers solitaiy; fruit usually 2- or more-seeded; seed elon­ gated, very rarely (sometimes in D. abyssinica) subglobose, with 2 lines radiating from apex. 1. Diospyros - Inflorescence a simple or branched pseudo-raceme; fruits nearly always 1-seeded; seed subglobose with 3 radiating lines from apex. 2. Euclea 1. DIOSPYROS L. (1753) W] lie, Bull. Jard. Bot. nat. Belg. 58: 325-448 (1988). Tn trs or shrubs. Leaves nearly always alternate, margin entire. Inflorescence usually cymose, or in bundles, or re­ duced to a solitary flower, axillary or from older wood. Flowers dioecious, the female usually larger than the mate Calyx very variable, 2-8-lobed or cup-shaped and enure, usually persistent and enlarging. Corolla 3-8*

lobed. very variable. Stamens 2-100, included or ex­ serted, solitary or united in pairs or larger groups, joined to corolla-tube or free to base; anthers (in Flora area) de­ hiscent by longitudinal slits. Pistillode variable, absent to well-developed. Disk well-developed to absent, some­ times with fringed margin. Staminodes variable, absent to well-developed. Ovary globose, ovoid or conical, gla­ brous or hairy, consisting of 3-8 carpels, each of which is completely or incompletely divided by false septae. Styles usually partly free, rarely completely united; stigmas usu­ ally fleshy and expanded. Fruit usually a one- to manyseeded berry, rarely a tardily dehiscent fleshy capsule. Seed usually ellipsoid or shaped like a segment of an or­ ange, very rarely subglobose, with 2 lines radiating from the apex. About 500 species widely distributed in the tropics and subtropics; a few species in the warm temperate zone. 3 indigenous species in the Flora area. Diospyros kaki L., the persimmon, is widely cultivated in the warm temper­ ate regions and may also have been tried in Ethiopia.

E-:iianical Museum and Library, University o f Copenhagen, 130 Gothersgade, D K -II2 3 Copenhagen K. Denmark.

** 1tank White died in 1994, but it is assumed that he would have been relatively satisfied with the present account because all the species riive been dealt with also in Friis, Forests and Forest Trees o f Northeast Tropical Africa, 1992, a work o f which Frank White ap­ proved in a kind review.

Key to individuals with male flowers

1. Anthers exserted, visible from outside. - Anthers included, not visible outside the flowers.

2

2. D. mespiliformis

50

134. EBENACEAE: 1. Diospyros

2. Leaves up to 3 x 1.3 cm; lateral veins only faintly visible. 3. D. scabra - Largest leaves larger than above; lateral veins dis­ tinct 1- D. abyssinica Key to individuals with female flowers o r fruits 1. Leaves up to 3 x 1.3 cm; lateral veins only faintly visible. 3. D. scabra - Largest leaves largeT than above; lateral veins dis­ tinct 2 2. Flowering or fruiting calyx glabrous or only mar­ ginally ciliate; fruit ellipsoid or elongate; buds and leaves (on lower surface) glabrescent. 1. D. abyssinica - Flowering calyx hairy to sparsely hairy', fruiting calyx becoming glabrescent fruit almost globular or slightly depressed: buds and leaves (on lower surface) with red-brown appressed or spreading hairs. 2. D. mespiliformis 1. D. abyssinica (Hiern) F. White (1956); Maba abyssinica Hiem (1873) - type: EW. Ainsaba Valley, near Keren, Beccari 55 (K lecto., FT iso.). Small to medium sized tree up to c 35 m tall, but some­ times flowering as a shrub. Trunk usually long, straight, slender, bark dark grey or almost black, rough, reticulate and peeling off on old trees. Leaves somewhat leathery , dry ing grey-green or black; blade 3-12 x 1.2-4 cm, ellip­ tic, oblanceolate-elliptic or lanceolate-elliptic, apex ob­ tuse to shortly and bluntly subacuminate, rarely distinctly acuminate; lower surface glabrescent; lateral veins in 5-12 pairs, venation prominent and closely reticulate on both surfaces. Male flowers axillary' and from older, leaf­ less nodes on the branches, in contracted 10-18-flowered cymes; peduncle c 1 nun long; pedicels c 1 mm long, with rough red-brown hairs; calyx c 2 mm long, shallowly cup-shaped, with 3 or 4 short broadly dentate lobes, gla­ brous outside except for a few minute marginal hairs, gla­ brous inside; corolla 5-6 mm long, somewhat twisted, glabrous, tube c 1.5 mm long, lobes 3 or 4. c 4.5 x 3 mm, broadly elliptic, apex obtuse; stamens 10-15, 2-4 mm long; anthers lanceolate-apiculate; pistillode c 1 mm long or absent, glabrous. Female flowers axillary or from older, leafless nodes on the branches, in (l-2)3-5(-8)-flowered clusters; pedicels c 2 mm long, with red-brown, stiff hairs; calyx c 6 mm long, cup-shaped, glabrous outside, finely stiff-haired towards the base inside, divided almost to the base; lobes 3 or 4, up to 6 x 6 mm, suborbicular, somewhat apiculate, strongly overlapping; corolla slightly shorter than the calyx, otherwise as in male; staminodes 3 or 4, glabrous, c 2 mm long, filiform, exserted, attached to the throat of the corolla and alternating with the lobes; ovary 4 x 2 mm, cone-shaped, glabrous, gradually merging into the short undivided style; locules 6, each with one ovule; stigmatic lobes 3, c l mm long, ascending. Fruit up to 14 x 9 mm, glabrous, ellipsoid or subglobulose, style usually persistent. Seed(s) 1 (very rarely 2), usually c 9 x 6 mm, globose to subellipsoid, black. Fruiting calyx scarcely en­ larging, becoming saucer-shaped. Fig. 134.1.1 & 2.

subsp abyssinica Intermediate and montane humid forest, both broacleaved forest and forest dominated by Podocarpus\ 550-2500 m. EW TU GD GJ SU AR WG IL KF SD BA HA; widespead in tropical Africa from the Flora area ^o Guinea Republic, south to Angola, Zambia, Malawi and Mozambique; rare in the Guineo-Congolian rain forest. Friis et al. 3862, 4138; Gilbert & Sebsebe D. 8803. In Malawi subsp. abyssinica is largely replaced by subsp. chapmaniorum F. White. 2 D. mespiliformis Hochst. ex A DC. (1844) - type: TU/GD, Djeladjekanne, Schimper 655 (G lecto., BM BR K L M OXF P TCD W iso ). Evergreen tree up to c 25 m tall or more, rarely flowering as a shrub; crown dense, rounded. Trunk straight or twisted, sometimes fluted at the base; bark dark brow n or almost black, rough, longitudinally fissured, peeling off in square scales. Young branchlets tomentellous with pink to brown appressed hairs, soon glabrescent. Leaves some­ what leatheiy, old leaves diying dull grey-green above and yellow-green beneath, young leaves diying pale redbrown; blade 6-14 x 2.2^4.5 cm, mostly oblong-elliptic or oblanceolate-elliptic; apex usually acute to subacuminate, rarely obtuse or rounded; base cuneate or rounded, lower surface minuteh puberulous with appressed stiff hairs or spreading flexuose hairs; lateral veins in 15-20 pairs, as­ cending at 45 degrees from midvein, indistinct, together with the tertiary veins forming a reticulum which is slightly prominent on both surfaces. Male flowers subsessile in groups of 3 on 4—6 mm long peduncles arising from axils of reduced leaves at the base of current year's growth or from axils of the first-formed normal leaves; calyx c 3 nun long, lobes 4 or 5, tria n g u la r, 1.5 nun long; corolla c 6 mm long, narrowly um-shaped. silky velvety outside, tube c 5 mm long, glabrous inside and thickened at the throat lobes 4 or 5. triangular, c 1 mm long; stamens c 14, included, c 4 mm long, filaments c 1 mm long, gla­ brous, inserted on the receptacle, anthers narrowly lanceo­ late. apiculate. glabrous except for a few hairs on the connective; pisullode minute, velvety. Female flowers subsessile, solitan or rarely 2 or 3 together, in the axils of reduced leaves at the base of the current year’s growth; ca­ lyx and corolla similar to the male but calyx c 8 mm long and lobes cordate-dentate with undulate-plicate, reflexed margins; corolla 10-12 mm long; staminodes 6-12, filiform, c 4 mm long, glabrous, inserted at the base of the corolla; ovary c 3 mm in diameter, yellow, globose, warty, glabrescent but a few hairs persisting near the base of the persisting style. Seeds 3-6, red-brown. Fruiting calyx sau­ cer-shaped or shallowly cup-shaped, lobes with recurved, strongly undulating margins. Fig. 134.1.3 & 4. Riverine forest, open riparian woodland, or scattered along temporary streams, sometimes at the base of rock}' outcrops; 300—2000 m. EW TU GD GJ SU WG IL KF SD BA; widespread in tropical Africa from the Flora area and Yemen to Senegal, south to Namibia and South Africa (Transvaal) and south Mozambique, but almost com-

134. EBENACEAE: 1. Diospyros, 2. Euclea

Dletely absent from the Guineo-Congolian rain forest. Ash 893 Friis et al. 2533; Gilbert & Sebsebe D. 8693.

3. D. scabra (Chiov.) Cufod. (1969); Maba scabra Chiov. (1951) - type: GG, DandeMarmara, Corradi 8391 (FT lecto.). Evergreen tree up to c 11 m tall; crown dense, rounded. Tnink up to 60 cm in diameter; bark grey, brown or almost black, rough, with deep quadrangular fissures. Young branchlets densely covered with stiff hairs. Leaves some­ what leathery, olive green, drying brown or pale greenbrown; blade up to 3.5 x 1.5 cm broadly elliptic to nar­ rowly lanceolate, apex and base acute to rounded; lower surface densely bristly on midrib and margin, more sparsely so on rest of blade; veins other than midrib virtu­ ally invisible on both surfaces. Male flowers more or less clustered in 2-6-flowered cymules in leaf-axils and on branchlets below the leaves; peduncle c 1 mm long; pedi­ cels slender, 1-2 mm long; calyx 2-3 mm long, cup­ shaped, lobed to about the middie, stiffly hairy on both surfaces, lobes (3-)4-dentate; corolla cone-shaped in bud, c 6 mm long, glabrous, but with scattered papillae; tube 3 < 2.5 mm; lobes 3, ovate-triangular, acute, 3 mm long; sta­ mens c 2, sometimes in pairs, included, up to 2.5 mm long, filaments up to 1.2 mm long, glabrous, inserted at base of corolla-tube; anthers glabrous except for a few minute apical bristles; pistillode up to 3 x 1 mm, glabrous, narrowly and irregularly cone-shaped. Female flowers solitary, in leaf-axils and below the leaves; pedicels 1-2 mm long, calyx and corolla as in male, but corolla with a few minute stiff hairs on the lobes; staminodes not seen; ovary ellipsoidal, c 3 x 1.5 mm, glabrous; locules 6, each with one ovule; styles 3, bilobed, 1 mm long, glabrous, rleshy and stigmatic on inner face. Fruiting calyx slightly enlarging, saucer-shaped, lobes semi-orbicular, apiculate. Fruit c 1~ x 12 mm, ovoid, yellow at first, turning red and rinally black, glabrous. Seed 1, dark brown, c 12 x 8 x 6 mm. Fig. 134.1.5. On banks of seasonal watercourses, at springs or wells, and at the base of rocks; 650-1000 m. GG SD; South Sudan, Uganda, Kenya. Corradi 8398; Friis et al. 9027; Haugen 65. 2. EUCLEA Murr. (1774) Trees or shrubs, usually evergreen. Leaves alternate, subopposite or subverticillate, leathery, margin sometimes wavy, otherwise entire. Inflorescence a simple or branched false raceme, or rarely (outside the Flora area only) flowers solitary. Flowers dioecious, the male usu­ ally larger than the female. Calyx 4- or 5-lobed shallowly ^up-shaped or saucer-shaped, not enlarging in fruit. Co­ rolla campanulate, 4- or 5-lobed, usually divided to below :he middle. Stamens 10-30, basically 2 opposite and 2 al:smating with each corolla lobe, but this arrangement may De modified in many ways; anthers slightly exserted or at least clearly visible from outside, lanceolate or narrowly oblong, usually bristly, often dehiscent at first by large apical pores which later prolong to longitudinal slits; fila­

51

ments usually shorter than the anthers. Pistillode usually very reduced, with or without stylodes, or absent. Ovary globose, hairy or covered with peltate scales, locules usu­ ally 4-6, each with one ovule, occasionally 2 or 3, incom­ pletely septate and then each with two ovules. Styles 2 or 3, free or united in lower half, often as long as the ovary, usually glabrous, ending in a slightly bilobed stigmatic surface. Fruit a small, globose, l(-3)-seeded beriy up to 10 mm in diameter. Seed subglobose, with 3 radiating lines from the apex. Embryo with cotyledons bent at a right angle to the radicle; radicle entirely surrounded by an ingrowth of the testa (Fig. 134.10). About 12 species, confined to Africa, Arabia, Socotra and the Comoro Islands. All species occur in South Af­ rica, and 6 extend into tropical Africa; 2 species in the Flora area. 1. Branchlets, leaves and inflorescence-axes com­ pletely glabrous; corolla mostly urn-shaped, lobed only in upper half; leaves broadest above the middle. 1. E. racemosa - Branchlets, leaves and inflorescence-axes with peltate scales; corolla campanulate, lobed into the lower half; leaves broadest at or below the middle. 2. E. divinonim 1. E. racemosa Murr. (1774) - type: South Africa, Plate 84/1 in Burman, Rar. PL Afr. (1739). Evergreen shrub or small tree up to c 12 m tall. Young shoots glabrous. Leaves dark green and glossy, alternate or subopposite to opposite, sometimes in whorls; petiole 0.1-0.3 cm long; blade very variable in shape and size, up to 11 x 4 cm; lower surface glabrous; secondary veins subprominent on upper surface, less so on lower surface, but often conspicuous because of pink coloration, tertiary veins usually indistinct, especially on lower surface. In­ florescence 15-50 mm long, unbranched, with glabrous axes. Male flowers up to 5 mm long; calyx c 2 mm long, saucer-shaped, glabrous or with minute hairs on the mar­ gin, shallowly lobed or toothed; corolla lobed only in the upper half, glabrous or sparsely bristly along the middle of the lobes. Stamens 10-20, mostly in pairs, filaments up to 1.3 mm long; anthers up to 3 mm long, bristly, espe­ cially near apex. Female flowers similar to male but much smaller; staminodes absent; ovary glabrous or covered with rough hairs. Fruit 6-8 mm in diameter.

subsp. schimperi (A. DC.) White in Bull. Jard. Bot. Nat. Belg. 50: 399 (1980); Kellaua schimperi A. DC. (1842); Euclea kellau Hochst. (1843), nom. illeg.; E. schimperi (A. DC.) Dandy (1952) - type: TU, Adua, Schimper 159 (G holo.). E. latidens Stapf (1906). E. microcarpa Gtirke (1895). E. neghellensis Cufod. (1939) - type: SD, Neghelle, Cufodontis 211 (FT holo.).

52

134. EBENACEAE: 1- Diospyros. 2. Euclea

Fig. 134.1. D IO SP YR O S A B Y S S IN IC A I branch x 1/ 3; 2 - fruit x l ‘/j. D. M E SP IL IF O R M IS 3 - branch x l / 3; 4 - longitudinal section o f seed showing radicle x 1Vs. D. SC AB R .4: 5 branch x 1/3. E U C LE A R A C E M O SA subsp S C H IM P E R I 6 - branch x 1/ 3 , f7 flower x 2*4 & D IV IN O R U M . 8 branch x V 3 ; 9 - part o f inflorescence x 2Vi; 10 - longitudinal section o f Seed showing radicle x l*/2. 1 & 2 from f r i i s et al. 4138; 3 from Friis et al. 4544, 4 after FI. Zamb. 7(1): Tab. 54C; 5 from Friis et al. 9027 ( from photo); 6j& 7 from Friis 1109; 8 & 9 from de Wilde 6364.; 10 after FI. Zamb. 7(1): Tab 54B. Drawn by Brigitte WollcnweberRatzer.

Leaves thinly leathery, obovate or oblanceolate. up to 12 x 4 cm. up to 4 times as long as broad, distinctly broadest in upper half. Ovary covered with stiffhairs. Fig. 134.1.6 & 7. Open montane, usually Juniperus - Podocarpus. for­ est. especially in clearings and along margins, in ever­ green montane bushland as well as in Buxus Acokanthera and Acacia - Commiphora bushland; also in rivenne bushland and in clumps of shrubs in montane grassland: (700-) 1000-2900 m. EW TU GD GJ WU SU KP SD BA HA: widespread in eastern Africa from Egypt. Sudan, through Tropical East Africa, including E Rep. Deni. du Congo. 10 South Africa (east Transvaal, Natal, eastern Cape Province); also in Yemen and Oman. Friis et al. 6 184; Mooney 8002; W.J.J.O. de Wilde 7361. Subsp. racemosa occurs in the remaining parts of Transvaal and in western Cape Province.

2. E. divinorum Hiern (1873) type: Zambia Victcna falls, Kirk s.n. (K le E kenyensis Fries (1925); E. divinorum subsp. kenvensis (Fries) Dale & Greenway in ; rees and Shrubs: 177 (1961). E lanceolata auct., non E Mey. ex A DC. (1844): Cufodontis, Enum.. 667 (1960). Evergreen shnib or small tree up to c 9 m tall (outside the Flora area tip to 15 ra). Bark grey-brown to biack, usually rough and longitudinally fissured Young shoots glabrous except for rust-coloured, peltate scales. Leaves usually op­ posite or subopposite, petiole 0.4-0.6 cm long: blade up to 9 x 4 cm, variable in width, otherwise uniform, some!'hat diamond-shaped to angular-ovate, broadest at or beiow the middle, apex acute, the tip itself rounded, base attenuate or slightly concave, margin often strongly undulate; lower

134. EBENACEAE: 2. Euclea

sicrtace glabrous except for rust-coloured, peltate scalcs; lateral and main veins sometimes slightly raised above, oi iierwise inconspicuous. Inflorescences paired in at least some leaf-axils, unbranched, dense and contracted, up to 1: mm long. Male flowers c 3.5 mm long; calyx sau­ ce -shaped with short broad teeth, glabrous except for nut -coloured peltate scales; corolla deeply lobed, throat wide, each lobe with many bristly hairs along the middle; stamens c 16, bristly; pistillode with 2 simple or bilobed sty lodes. Female flowers like the male, but smaller, with­

53

out staminodes; ovary densely bristly. Fruit c 7 mm in dia­ meter. Fig. 134.1.8-10. Open montane forest especially Juniperus - Podocarpus forest often in clearings and along margins, fre­ quent in montane evergreen bushland. and in semideciduous bushland; 1000-2400 m. SU AR WG KF GG SD; widespread in East Africa from Ethiopia and Sudan to South Africa (Transvaal, Natal). Gilbert & Sebsebe D. 8760; Mesfin T. & Vollesen 4183; M.G. & S.B. Gilbert 1772.

54

135. SAPO TACEAE I. Friis* Cufodontis, Enum:. 663-667 (1960); Hemsley. Sapotaceae. in FI. Trop. E. Afr.: 78 pp. (1968); Kupicha, Sapotaceae, in FI. Zamb. 7(1): 210-247 (1983); Pennington, The Genera o f Sapotaceae (1991); Friis, Forest & Forest Trees of Northeast Tropical Africa, in Kew Bull. Add. ser. XV: 220-228 (1992). Trees or shrubs, generally with milky latex. Leaves alternate, petiolate. simple, entire, sometimes with transluscent dots when viewed against a strong light. Stipules usually falling soon, or absent. Flowers bisexual or unisexual (then usually female) by abortion, solitary or in axillary clusters, som etim es in the axils o f fallen leaves. Calyx with 4—8 free sepals or calyx lobes. Corolla usually cream -coloured or pure white, campanulate tc shortly tubular, with 4 -8 lobes in 1-2 whorls, som etim es each lobe divided into 3 segm ents. Stamens as m am as the corolla-lobes and opposite them , or m ore num erous and in 2 (to several) whorls; anthers 2-thecous, open­ ing lengthwise. Stam inodes som etim es present between the corolla-lobes, variously developed. Ovary superior usually 5- to m any-locular; style simple; ovules solitary in each locule or ascending from the inner angle. Fruit a berry. Seeds with a generally hard, smooth, brown, often shiny, testa; attachm ent area (scar) usually large sometimes covering more than h alf o f the surface area. According to Pennington (loc. cit.), apantropical family o f 53 genera and about 1100 species, mainly distrib­ uted in humid lowland and m ontane forest, but some genera (e.g. Sideroxylon), extend into sem i-arid and arid regions. In the Flora area the fam ily is represented by 5 genera and 9 species, am ongst w hich are im portant trees from the m ost humid forest types o f the area. This family includes im portant tim ber species, many o f which are threatened or endangered both globally and locally. Key based on vegetative and floral characters 1. Calyx normally consisting of 5 sepals, arranged in a single whorl. 2 - Calyx with 6-8 sepals arranged in two distinct whorls, the inner whorl paler in colour and smaller in size. 3 2. Leaves without transluscent dots; style simple or capitate; plants usually spiny. 4. Sideroxylon - Leaves with transluscent dots; style slightly ex­ panded into a 5-papillate stigma; plants not spiny. 5. Pouteria 3. Leaves with very long petioles, usually more than 1/3 the length of blade; short shoots stout (to 1 cm thick) with conspicuous scars. 2 Vitellaria - Leaves and shoots not as above. 4 4. Calyx of 8 sepals arranged in two dissimilar whorls of 4. 1. Mimusops - Calyx of 6 sepals arranged in two dissimilar whorls of 3. 3. M anilkara Key based on vegetative and fruit characters 1. Calyx o f 5 sepals, arranged in a single whorl, per­ sisting or deciduous at fruiting stage. - Calyx with 6-8 persistent sepals arranged in two distinct whorls, the inner series paler in colour and smaller in size. 3 2. Seed-sear lateral, long and narrow, or broad and covering much of the surface; fruits large, usually distinctly fleshy or leathery. 5. Pouteria Botanical Museum and Library, University of Copenhagen, 130 Gothersgade, DK-1123 Copenhagen K, Denmark.

3.

4. -

Seed-scar basal, circular or elliptic; fruit small, fleshy. 4. Sideroxylon Seed-scar large, covering 1/3 to 1/2 of the surface; leaves crowded on swollen, scarred, branch tips. 2. Vitellaria Seed-scar small to narrow, basal to partly lateral; leaves not crowded at branch tips (or, if so, then the fruit less than 3 cm long). 4 Calyx of 8 sepals arranged in two dissimilar whorls of 4. 1. Mimusops Calyx of 6 sepals arranged in two dissimilar whorls of 3. 3. M anilkara

1. MIMUSOPS L. (1753). Unarmed trees and shrubs. Leaves alternate, clustered 10wards the tip of branches or not; venation with many paral­ lel secondary veins. Stipules small, falling soon. In­ florescences axillary, cymose, more or less umbel-shaped. Flowers bisexual. Calyx of 2 whorls of 4 free sepals, the outer whorl more or less valvate. Corolla hairy or glabrous; tube much shorter than the 8 corolla lobes, each of which is nearly always divided to the base into 3 segments, with die median segment clasping the stamen, sometimes pressed in­ wards towards the style, lateral segments widely spreading, entire or more or less deeply divided. Stamens (7-)8 in a single whorl, inserted at top of corolla tube; filaments free or more or less fused with staminodes; anthers opening out­ wards with slits, hairy or glabrous. Staminodes 8, well de­ veloped, alternating with the stamens, usually appressed to ovary and style, often hairy. Ovary (7-)8-locular, hairy, placentation almost basal in each locule, style exserted from the many appressed floral parts or not Fruit 1-6-seeded, irdehiscent, fleshy. Seeds laterally compressed, usually

135. SAPOTACEAE: 1. Mimusops, 2. Vitellaria

55

vith a smooth, rarely wrinkled, shiny, brown or brownblack testa, scar small, often circular or elliptic, basal or nearly so.

An important timber species with hard, heavy wood and edible fruits.

About 40 species distributed in the Old World tropics, mostly in tropical Africa (c 20 species), on Madagascar (c 5 species), on the Mascarene Islands (4 species), and 1 -pecies in Asia and the Pacific; 2 species in the Flora area.

2. M. laurifolia (Forssk.) Friis (1981); Binectaria laurifolia Forssk. (1775) - type: Ye­ men, Forsskdl 559 & 560 (C syn.). M. schimperi A. Rich. (1850) - type: EW/TU, along Tacazze River, Schimper 697 & 873 (both K FHO isosyn.). ?M degan auct., non Chiov. (1916).

1. Flowering and fruiting pedicels distinctly longer than petioles of supporting leaf; leaf apex acu­ minate; buds acute; fruits acuminate. 1. M. kummel - Flowering and fruiting pedicels shorter than peti­ oles of supporting leaf; leaf apex acute; buds with blunt or rounded apex; fruit with rounded apex. 2. M. laurifolia

. M. kummel A. DC. (1844) type: TU, ML Scholoda, Schimper s.n. (K lecto., BM FHO isolecto.). M djurensis Engl. (1904). Imbricaria fragrans Bak. (1877); Mimusops fragrans (Bak.) Engl. (1904). M. stenosepala Chiov. (1940) - type: SD, Borana, Neghelli, Senni 1015 (FT holo.). Small or medium-sized tree, up to 25 m tall. Young branches and petioles with rust-coloured or brown pubes­ cence. becoming glabrous. Leaves: petioles 0.5—1.5(—3) :m long, blade elliptic to ovate-elliptic or oblong-elliptic, 4.5-12 x 2.5-5 cm, apex shortly acuminate, rarely obtuse or einargmate, base narrowly to broadly cuneate; upper surface dark glossy green, lower surface paler with slightly raised, reticulate venation, practically glabrous on lx)th sides or with a few scattered hairs along the midvein. r lowers fragrant, usually in clusters of 2-4 in a leaf axil; LTedicel slender, curved, (15-)20-50 mm long, densely covered with rust-coloured hairs. Outer sepals more or less lanceolate, 9.5-12 mm long, densely covered with lust-coloured hairs externally; inner sepals of similar shape, more or less glabrescent. Corolla creamy white; :ube up to 2 mm long; lobes divided into 3 segments to the use, 2 outer segments often split further into 2 linear seg­ ments; segments 9-12 mm long, Stamens: filaments 2-3.5 mm long. Staminodes linear-lanceolate, 4-6 mm ong, apex extended, densely pubescent on the outside, .'vary densely covered with long, straight, more or less ust-coloured hairs; style slender, tapering, 10-12 mm ong. Fruit an orange or orange-red ellipsoid to ovoid icrrv, up to 2.5 x 1.5 cm, apex acute or even shortly •caked (rarely obtuse). Seed solitary, ellipsoid, up to 1.8 n long; testa brown, hard and glossy; scar obliquely •Asal. Fig. 135.1. 1 & 2. Montane rain forest and other broad-leaved forests, Mrticularly in gullies and veiy frequent in riverine forest; ■iso in riparian woodland and in woody vegetation on lake shores; 550-2500 m. EW TU GD GJ WU SU AR WG IL GG SD BA HA; west to Mali and Guinea, south to Rcp. Dem du Congo, Tanzania and Malawi. Chaffey 308; 1 riis et al 1956; Mooney 8626.

Small or medium sized tree with rounded crown, usually up to c 15 m tall, rarely up to 25 m tall. Young branchlets tend to appear as ‘short-shoots’, c 2-3 cm long, with leaves more or less densely clustered. Leaves: petiole 3.5-5 cm long, usually about half the length of blade; blade somewhat leathery, elliptic to elliptic-oblong, 4-12 x 3-4 cm, apex acute, base bluntly cuneate. Flowers 1—4 in axillary clusters, on stiff erect or spreading, 10-15 mm long pedicels; flower-buds c 5 x 3 mm, blunt. Sepals: outer c 6 x 3 mm, with rust-coloured hairs and a pale stripe along the margin, inner sepals slightly smaller. Corolla white or cream, c 7 mm long, tube c 4 nun long, lobes c 3.5 mm long. Stamens: filaments c 0.5 mm long; anthers c 2 mm long. Staminodes c 2 mm long. Ovary c 1.5 mm in di­ ameter, style c 3 mm long. Fruit a berry, c 3.5x2 cm. Seed c 2 x 1.2 cm; testa pale to dark brown; scar circular, more or less basal, pale. Fig. 135.1.3 & 4. Dry' montane Juniperus forest; also in evergreen and semi-evergreen bushland with Buxus\ 750-1850 m. EW TU WU SU HA; Djibouti, Somalia. Yemen. Friis et al. 6180; Gilbert et al. 7370; M.G. & S.B. Gilbert 1287. Records from the Sudan and Uganda are presumably due to introduction, as must also have been the case with the records from ancient Egypt, where the fruits and leaves were used at funerals and much material has been found in tombs from the Pharaonic times (see discussion by Friis in Kew Bull. 35: 785-792, 1981). The natural range of the species seems to be Yemen, Eritrea, Ethiopia, Djibouti and Somalia and this species, therefore, belongs to the small element restricted to the transition zone be­ tween the Afromontane region and the Somalia-Masai re­ gion.

2. VITELLARIA Gaertn.f. (1805) Butyrospermum Kotschy (1865) Unarmed trees and shrubs with massive twigs. Leaves spi­ rally arranged in dense terminal clusters, blade with a prominent marginal vein, secondary veins parallel; stip­ ules small, falling soon Inflorescences of dense axillary clusters near the shoot apex in the axils of scale-like leaves. Flowers bisexual. Calyx of 2 whorls of (3-)4 free sepals, the outer whorl valvate. Corolla glabrous; tube much shorter than lobes; lobes (6-)8, entire, twisted in bud, spreading at anthesis. Stamens (6-)8, in a single whorl, inserted at the top of the corolla tube; filaments free; anthers opening outwards, glabrous. Staminodes (6-)8, well developed, alternating with the stamens, erect or pressed inwards and forming an envelope round the

135. SAPOTACEAE: 1. Mimusops,2. Vitellaria

from Frits et al. 2513; 6 & 7 from Adam 8844. Drawn by Brigitte Wollenweber-Ratzer

135. SAPOTACEAE: 2. Vitellaria, 3. Manilkara

57

The taxonomy of this species is still in dispute and it lias here been conceived in a broad sense.

pistil, lanceolate, almost glabrous. Ovary (5-)6-locular, h< :.ry, placentation axile; style slightly exserted. Fmit 1- 2-seeded, indehiscent, fleshy. Seeds globose or broadly ellipsoid, not laterally compressed, with rather thin, shiny testa; scar broad, adaxial.

The wood is hard and termite-proof. The fmit is edible and also yields SHEA BUTTER or KARITE, an oil used for cooking and lighting.

\ genus with 1, possibly 2, species; one widespread fr®ra Ethiopia. Sudan and Uganda to Senegal; only this species in the Flora area.

3. MANILKARA A dans. (1763), nom. cons.

The name Butyrospermum lias been widely used for thi> genus and was proposed for conservation in 1962. but the proposal was not accepted and the older, less known name Vitellaria has, therefore, to be used.

V. paradoxa G aertn.f (1807); Butyrospermum paradoxum (Gaertn. f.) Hepper (1962) - type: single seed of unknown origin (P holo.). B. niloticum Kotschy (1865); B. parkii (G. Don) Kotschy var. niloticum (Kotschy) Engl., in Monogr. Afr. Pfi.-Fam., 8, Sapotaceae: 23 (1904); B. parkii (G. Don) Kotschy subsp. niloticum (Kotschy) J.H. Hemsl.. in Kew Bull. 15: 290 (1961); B. paradoxum subsp. niloticum (Kotschy) Hepper. in Taxon 11: 227 (1962). Bassia parkii G. Don (1838); Butyrospermum parkii (G. Don) Kotschy (1865). Tree up to 20 m tall, often much less, with stout trunk and a richly branched, spreading crow-n. Bark usually grey or bLick, often scorched by burning, deeply fissured and splitting mto square or rectangular corky scales. Shortshnots with conspicuous leaf-base scars. Young shoots, pe=ioles and flow'er-buds with rust-coloured hairs of vaiyirn: density. Leaves densely clustered, petiole long, 4-8 cm. 1/3 to 1/2 the length of blade; blade oblong to o\ ite-oblong, 10-25 x 4.5-14 cm. rounded at apex, base acute to broadly cuneate, margin entire, thickened, some­ times more or less undulate; lateral veins 20-30 on each Sid ?, prominent and parallel, slightly curved near the mar­ g in : upper and lower surface of mature leaves puberulous or glabrescent. Flowers fragrant; pedicels up to 30 mm long; puberulous to densely pubescent. Outer sepals free almost to base, lanceolate. 9-14 x 3.5-6 mm, pubescent to cottony on the outside; inner sepals slightly smaller, glabrescent. Corolla cream; tube 2.5-4 mm long, shorter th; n free segments, glabrous or pilose on outside; lobes broadly ovate. 7-11 x 4.5-7 mm wide. Stamens: filaments 7-1 2 mm long; anthers opening with lateral slits, up to 4.5 nun long. Staminodes up to 8 mm long. Ovary 3-5 mm in diameter, pubescent; style 8-15 mm long, exserted from flc wer at anthesis. M ature fruit greenish-yellow , up to 6.5 x ■ 5 cm, more or less glabrous or pubescent in patches, co staining sw'eet pulp surrounding the seed. Seed up to 5 x . 5 cm, ovoid, with large lateral scar. Fig. 135.1.5-9.

In Combretum - Terminalia - Stereospermum - Entada woodland in the w'estem lowlands, penetrating into the highlands along the river valleys: 500-1000 m. IL; from Eti liopia. Sudan and Uganda to Guinea and Sierra Leone. Fr is et al. 2513; Sebsebe D. et al. 5112; Tesfaye A. et al. 19

Unarmed trees, rarely shrubs, nearly always with charac­ teristic sympodial branching. Leaves alternate, clustered at or near the top of shoots; venation with parallel second­ ary veins looping to form an undulating marginal vein; stipules small, falling soon, or absent. Inflorescences axillary clusters, leaves present or fallen. Flowers nearly always bisexual, rarely unisexual and then dioecious. Ca­ lyx of 2 whorls of 3 (occasionally 2 or 4. but mostly so only outside the Flora area), free or slightly united sepals, the outer whorl valvate or slightly overlapping. Corolla nearly always glabrous; tube much shorter than lobes; lobes 6-9, usually spreading at anthesis and usually di­ vided into 3 segments; median segment usually erect, clasping the stamen; lateral segments spreading, usually shorter than median segment, entire or deeply divided. Stamens 6(-l 2). in a single whorl inserted at top of corolla tube; filaments free or partly fused with staminodes; anthers opening outwards, nearly always glabrous. Staminodes (0-)6(-12), alternating with stamens, regu­ larly or irregularly longitudinally divided into 2 segments, sometimes rudimentary, usually erect but not usually closely surrounding the style. Small annular disk some­ times present. Ovary 6-14-locular, hairy or glabrous, placentation axile or nearly basal; style exserted. Fmit a one- to several-seeded berry, indehiscent, fleshy. Seed el­ lipsoid to obovoid. laterally compressed, with hard, shiny, woody testa; scar nearly always narrowly elongate, basiventral. less frequently extending along most of the ad­ axial face, very rarely broad. A genus of c 65 species; c 30 in tropical America, c 20 in Africa and Madagascar, c 15 in tropical Asia and the Pacific, 1 species in the Flora area. M. butugi Chiov. (1940) types: WG, Dulli Forest, Giordano 2465 & Humbi Forest, Giugliarelli 596 (both FT syn.). Manilkara multinervis auct., non (Bak.) Dubard (1915). Atimusops sp. sensu Indigenous Trees o f Uganda ed. 2 (1952). Tall tree with spreading crown, up to 30(-35) m tall, with straight cylindrical or slightly fluted trunk; bark greybrown, rough or finely fissured. Young shoots subglabrous, often with pale raised lenticels. Leaves alter­ nate, usually markedly clustered towards the end of branches; petiole 1—3(— 4.5) cm long, glabrous; blade el­ liptic to oblanceolate, rarely more or less obovate, (5-)8-17(-20) x (1.6—)3—6.3 cm, upper 1/3 of blade taper­ ing to an acuminate apex (especially in crown-leaves), rarely acute or more or less obtuse, base narrowly to broadly cuneate, lateral veins ascending, looped near mar­

58

135. SAPOTACEAE: 3. Manilkara,4. Sideroxylon

gin to form marginal vein; thinly leathery, dull green above, grey-green beneath with sparse indumentum of minute, closely appressed hairs. Flowers in clusters of 1-6, in axils of current leaves; pedicel stiff, erect or hori­ zontal, 8-12 mm long, pubescent. Calyx up to 5 mm long, more or less densely pubescent; lobes fused forming a basal tube up to 2 mm long; outer lobes ovate, c 4 x 3 mm; inner lobes slightly smaller. Corolla pale yellow; tube 1.5-2 mm long; outer lobes narrowly lanceolate, up to 5.5 mm long; median lobe elliptic, up to 5.5 mm long. Sta­ mens: filaments 2.5-3 mm long; anthers up to 2 mm long. Staminodes narrowly oblong to strap-shaped, up to 2 mm long, apex with (1—)2(—3) slender fringed processes up to 2 mm long. Ovaiy depressed globose, pubescent, 10locular, style 6-8 mm long, with base more or less swol­ len and pubescent Fruit subglobose, up to c 3 cm in dia­ meter, glabrous, with milky pulp. Seed shiny brown, more or less obovoid and flattened, 1.3-1.7 x 0.7-0.8 cm; scar lateral, oblique and extending to the base. Fig. 135.2.1-3.

glabrous. Seeds globose, ovoid, oblong or ellipsoid, not laterally compressed, rarely plano-convex when 2 seeds in fruit; testa smooth, shiny, free from pericarp, often sculp­ tured on the adaxial surface; scar nearly always basal or basi-ventral, small, circular, lanceolate or elliptic.

Lowland, transitional and humid montane forest; 1200-2150 m. WG IL KF SD; Sudan (Imatong Mts), Uganda, W Kenya Friis et al. 4078,4506; Mooney 6856.

1. S. oxyacanthum Baill. (1891) Spiniluma oxyacantha (Baill.) Aubrev. (1963) type: GD. Semien, Bayeta, Schimper 1336 (P holo.).

The species has very hard wood which does not decay easily. In Kenya and Uganda it is known as a timber tree used for heavy construction, and for making farm tools and boats. In the Flora area it is used for construction, par­ ticularly doors, windows and flooring. The fruit is edible.

Spiny shrub or small tree, often with stunted growth and rounded crown, up to 10 m tall. Older branches forming 1-3 cm long lateral shoots with retarded growth carrying densely clustered leaves and axillary flowers; a 1-4 cm long spine normally develops at the base of each of these short shoots. Branchlets silky, with a grey indumentum oi medifixed hairs. Leaves alternate, evergreen, on branch­ lets and short shoots, but never on spines; petiole 0.4-0.7 mm long; blade leathery, spathulate to obovate. 1.0-4.5 x 0.8-2.5 cm, apex rounded, more or less truncate, some­ times emarginate, base cuneate, margin entire, veins dis­ tinctly visible, marginal vein formed by arching secondary veins meeting near leaf-margin, glabrous above, silky to glabrous below. Stipules absent. Flowers solitary in leafaxils on short shoots; pedicel 5-7 mm long; flower-buds 4-6 nun long. Calyx with a very short basal tube, 5-lobed. lobes as silky as the branchlets, broadly ovate, c 2.5 x 2 mm. inner ones with hyaline margins. Corolla white, 5-lobed. w ith a 2-3| nun long basal tube, lobes imbricate, ovate, 2-3.5 x c 21mm, usually remaining erect during anthesis. Stamens 5,2-3 mm long, reaching the top of the corolla. Staminodes 5, triangular, fringed, 2-3.5 x c 1 nun. Ovary globular, silky as the branchlets, 1-2 mm in diameter, 5-locular, each locule with one axillary ovule. Style 5-6 mm long, usually exserted. Fruit a black i-seeded berry, c 7 x 5 mm. Seed ellipsoid, ridged, daik brown, c 5 x 3 mm: scar basal, small and circular. Fig. 135.2.4-6.

Two other species could occur in the Flora area: M. multinervis (Bak.) Dubard (1915) in the extreme SW, and M. sulcata (Engl.) Dubard (1915) in the extreme SE.

4 SIDEROXYLON L. (1753) S p in ilu m a (Baill.) Aubrev. (1963)

Sideroxylon sect Spiniluma Baill. (1891) Monotheca A. DC. (1844) Edgeworthia Falc. (1842), nom. illeg., non Meisner (1841) Reptonia A. DC. (1844) Friis, A reconsideration of the genera Monotheca and Spiniluma (Sapotaceae). Kew Bull. 33: 91-98 (1978). Spiny or (rarely in the Flora area) unarmed trees or shrubs Leaves alternate, often clustered on short lateral shoots, venation very variable; stipules absent. Inflorescences of clusters, or single flowers, in the axils of present or fallen leaves; flowers usually bisexual, rarely unisexual and then monoecious, (perhaps sometimes dioecious). Calyx a sin­ gle whorl of 5(-8) free, overlapping, sepals. Corolla cup­ shaped, usually glabrous, tube nearly always shorter than lobes, lobes (4-)5(-8), spreading, entire, rarely divided into 3 segments. Stamens (4-)5(-8), in a single whorl at top of corolla tube, exserted; filaments well developed; anthers opening outwards, usually glabrous, sometimes more or less converted into staminodes and the flower functionally female. Staminodes (4-)5(-8), usually welldeveloped, alternating with the stamens, often lanceolate, pressed against the style. Ovary (l-)5(-8)-locular, hairy or glabrous; placentation basi-ventral or basal; style exserted or included. Fruit 1-2-seeded, fleshy, usually

In the broad concept of the genus proposed by Penning­ ton (loc. cit.) and based on worldwide studies c 50 species occur in tropical America and c 25 species in the Old World; 2 species in the Flora area. 1. Leaves and flowers mostly on older dwarf shoots, never on spines; pedicels 5-7 mm long; ovary 5-locular, seed ellipsoid. 1. S. oxyacanthum - Leaves on spines and shoots with normal growth; flowers in dense clusters on normal shoots, pedi­ cels c 1 mm long; ovary 1-locular, seed globose. 2. S. mascatense

Associated with dry Juniperus and Juniperus - Olea montane forest, persisting in gully forests and in secondary montane evergreen bushland; in hedges or live fences, (1250-) 1850-2800 m. EW TU GD SU AR BA HA; not known elsewhere. Friis et al. 6191; Mooney 5047, 7103. This species and the following belong to a small floristic element restricted to the transition zone between the Afro montane region and the Somalia-Masai region in Eritrea Ethiopia and Somalia. A closely related species

135. SAPOTACEAE: 3. Manilkara, 4. Sideroxylon

59

Fig. 13 1 M A N ILK A R A B V TV G I: 1 - branch x c 1/3; 2 - flower x 3.4; 3 - fruit x 3.4. SIDEROXYLON OXYACANTHUM 4 xIS ? * 3 4 5 M ASC A T E N S E 7 -y o u n g branch x 14; 8 - mature branch x 14- 9 - flower x 3 4- 10- fruit x 3.4. 1 .1. (mm M ayer 8076; 3 from Mooney 6856; 4 & 5 from Gilbert &Jones 196; 6 from Burger 2432 7 from M ooney 5609 8 & 9 from Lt Mon 2356; 10 from Miller & Nyberg 9593. Drawn by Brigitte W ollenweber-Ratzer. ^ ’

60

135. SAPOTACEAE: 4. Sideroxylon, 5. Pouteria

(S. discolor Radcl.-Sm.) occurs in evergreen scrub on the mountains of Socotra. The species is frequently found in the literature as Spiniluma oxyacantha. The plant makes a very effective live fence. The leaves are used to treat eye infections.

2. S. mascatensc (A. DC.) Pennington (1985); Monotheca mascatensis A. DC. (1844); Reptonia mascatensis (A. DC.) O. Schwartz (1939) - type: Oman. Muscat. Aucher-FJoy 4916 (G-DC lecto., K isolecto.). Edgeworthia buxifolia Falc. (1842); Reptonia buxifoiia (Falc.) A. DC. (1844); Monotheca buxifolia (Falc.) A. DC. (1846). Sideroxylon buxifolium Hutch. (1931): Spiniluma buxifolia (Hutch.) Aubrev. (1963). Sideroxylon aubertii A. Chev. (1939). Sideroxylon gillettii Hutch. & Bruce (1941). Sideroxylon corradii Chiov. (1951) - type. SD, Neghelle, Corradi 8252 (FT holo.). Spiny shrub or small tree up to 10 m tall, occasionally re­ corded as a medium-sized tree up to 15 m. Spines in ju ­ venile plants many, thin and leafless, hardly longer than the leaves, in mature plants stronger and longer, and of­ ten carrying small clusters of leaves and flowers. Branchlets grey, silky, with medifixed hairs. Leaves al­ ternate; petiole 0.4-0.8 cm long; blade leathery. 1.0-4.0 x 0.4-2.5 cm, spathulate to elliptic, apex rounded, base cuneate, margin entire, more or less recurved, veins ob­ scure, glabrous above, grey, silky below. Inflorescences dense axillary clusters, sometimes in axils of fallen leaves, usually restricted to older branches, sometimes also on old thorns. Pedicel c 1 mm long. Flowers c 2.5 mm long. Calyx 5-lobed, with a short basal tube, with rust-coloured silky hairs, lobes overlapping, broadly ovate with rounded apex, c 0.8 x 0.6 nun. Corolla 5-lobed. with a short basal tube, yellow, lobes ovate with rounded apex, c 1 x 0.8 nun. Stamens 5, 1-2 mm long, exceeding the corolla, anthers opening outwards. Ovary globose, silky, c 0.5 mm in diameter, unilocular with 5 basal ovules on a short placenta. Fruit a 1-seeded berry, 5-8 mm in diameter Seed almost globose, yellow­ ish-brown, c 5 mm in diameter; scar basal, small and cir­ cular. Fig. 135.2.7—10. Dry Juniperus and Juniperus - Olea montane forest and transition between montane forest and evergreen and semi-evergreen Buxus - Acokanthera Juniperus bush­ land. sometimes also forming isolated clumps in Acacia Commiphora deciduous bushland; 1200-2150 m. SDHA; N Somalia, Djibouti; Oman. Afghanistan, and Pakistan. Burger 3124; Friis et al. 3120; Mooney 5623. The species is often found in the newer literature as Monotheca buxifolia, but the nomenclatural history is complex due to its occurrence in several regions. The ex­ istence of the later heterotypic synonym, Sideroxylon buxifolium Hutch. (1931), prevents a combination under Sideroxylon with the oldest epithet, Edgeworthia buxi­ folia Falc. (1842).

This species and the previous one belong to the small floristic element restricted to the transition zone between the Afromontane region and the Somalia-Masai region in Ethiopia, Djibouti and Somalia, but unlike the previous one it does not occur in Eritrea. However, it is widespread in the mountains of the SE part of the Arabian Peninsula and in SE Asia. 5. POUTERIA Aublet (1775) Aningeria Aubrev. & Pellegr. (1934) Malacantha Pierre. (1891) Hemslev, Notes on African Sapotaceae 3, the genera A ningeria, M a la ca n th a , and Butvrospermum in East Africa Kew B ull 20. 277-283 (1961). Unarmed trees or shrubs. Leaves alternate, not notably clustered towards the end of branches; blades usually with­ out clearly distinct marginal veins, secondary veins parillel. Thinner leaves (in the Flora area) w ith transluscent dots visible when viewed against a strong light. Inflores­ cences as axillary clusters, rarely on short leafless branches. Flowers often unisexual (plants dioecious). Ca­ lyx a single whorl of (4-)5(-6) free, overlapping sepals. Corolla cup-shaped to tubular, tube shorter than, equal, or (in the Flora area) longer than the lobes; lobes (4-)5(-6, rarely -9), usually erect, rarely spreading, simple, some­ times fringed or (in the Flora area) papillose. Stamens (4-)5(-6. rarely -9), Fixed in lower or upper half of corolla lube, rarely free, usually included; filaments generally short; anthers usually opening outwards or laterally, usu­ ally glabrous. Staminodes usually in same number as co­ rolla lobes, sometimes lacking, inserted at top of tube between corolla lobes. Disk present or absent. Ovary l-6(-15)-locular. placentation axile; style included or (in the Flora area) exserted, with faintly 4-5-lobed stigma. Fruit a 1 to seveml-seeded berry'. Seeds broadly ellipsoid, plano-convex, shaped like the segments of an orange or more laterally compressed; testa smooth, wrinkled or pit­ ted; scar adaxial, usually full length, narrow, broad or sometimes covering the entire seed surface. About 325 species occurring in tropical America (c 200 species), tropical Asia, Australia and the Pacific (c 120 species), and Africa (c 5 species). 3 of which are found in the Flora area. Pennington (Joe. cit.) has placed all the species previ­ ously in the genera Malacantha and Aningeria in Pouteria sect. Rivicoa, a section distributed mainly in S America and Africa. His taxonomy, based on a world­ wide review, has been follow ed here, although most Afri­ can Floras have previously accepted Malacantha and Aningeria. 1. Primary veins looped at end and actually forming a marginal vein at the leaf-margin; flowers sessile; staminodes absent; seed-scar narrowly oblong, covering little of the surface area. 1. P. alnifolia - Primary veins looping near leaf-margin so the mar­ ginal vein never reaches the actual leaf-margin; flowers more or less pedicellate; staminodes present; seed-scar oblong-elliptic, covering 1/3 or more of the surface area. 2

135. SAPOTACEAE: 5. Pouteria

2 Mature leaves usually leathery, with opaque blade; pellucid dots can only be seen on shade-leaves. 2. P. adolfi-friederici - Mature leaves papery to slightly leathery, with ex­ tensive system of pellucid dots easily visible when viewed against the light with a hand-lens. 3. P. altissima 1 P. alnifolia (Bak.) Roberty (1954); Chrysophyllum alnifolium Bak. (1877); Malacantha alnifolia (Bak.) Pierre (1891) - type: Nigeria, Barter 1788 (K holo.). Chrysophyllum ferrugineo-tomentosa Engl. (1900); Malacantha ferrugineo-tomentosa (Engl.) Engl. (1904). Small to medium-sized tree, up to 20(-25) m tall, with fluted trunk and slightly buttressed base. Branchlets with dense, rust-coloured, more or less spreading indumentum. Leaves alternate, somewhat crowded towards end of branchlets, apparently not deciduous but change colour to orange and red before falling; petiole 0.8-2 cm long; biade obovate to oblong. 12—22(—36) x 7-12(-20) cm, apex rounded, sometimes apiculate, base cuneate or ob­ tuse, margin entire, lateral veins prominent, 15-26 on each side of midrib, upper surface with scattered hairs, be­ coming almost glabrous, except for dense, short, rough hairs along midrib, lower surface with uniformly scattered eitct hairs, densely arranged on midrib and veins, becom­ ing almost glabrous, except for the veins. Inflorescences a'■Wary or m axils of fallen leaves, bracts and bracteoles large, sepal-like, with dense rust-coloured hairs. Flowers 4-7 mm long. Calyx lobes almost free, elliptic to ovate, up to 6 x 4.5 mm. with dense rust-coloured liairs on outside. Corolla white or cream, up to 7 x 3 mm, with tube longer than lobes; lobes up to 3.5 mm long. Stamens inserted on inside of tube, opposite corolla-lobes, about 1/3 below top of lube: filaments c 4 mm long; anthers opening outwards. Sty le 3—7(—9) nun long. Fruit a red. fleshy, 1-seeded bejnry, c 2.5 cm in diameter, with dense rust-coloured hairs almost to maturity. Seed up to 1.7 cm long, slightly flat­ tened; testa dark brown; scar linear, up to 4 mm wide. Fig. lJfib.1-4. Lowland and riverine forest; 550-1600 m. IL KF GO: widespread in the forested regions from Ethiopia w ok to Senegal, south through Uganda and Kenya to Tanzania and Mozambique. Chaffey 943; Friis et al. 2345, 2446. 2. P. adolfi-friederici (Engl.) Baehni (1965); Sideroxylon adolfi-friederici Engl. (1913); Aningeria adolfi-friederici (Engl.) Robyns & Gilbert (1947) - types: Rwanda, Bugoie Forest, Mildbraed 1447 & 1481; Zaire. Ruwenzori. Butagu valley, Mildbraed 2528 (all B syn. destroyed), j Pouteria ferruginea Chiov. (1940) - types: WG. Huinbi. Giugliarelli 583; Dulli Forest, Giordano 2447; Capuano 2530 (all FT syn.). Pouteria rufiner\’is Chiov. (1940) - ty pes: WG/IL, Mugghi Forest, Giordano 2478; Saio Forest. Giuglia­ relli 564 (both FT svn.).

61

Tall tree of high forest, up to 45(-50) m tall, with long, straight, more or less fluted trunk, and buttresses present at base. Young shoots and leaves with short dense rustcoloured hairs; older branches more or less glabrous. Leaves alternate, more or less scattered along branchlets; petiole 1-2 cm long, more or less twisted, with short rust-coloured hairs; blade elliptic to oblong or obovateelliptic, 4-21 x 2.5-8.5 cm, transluscent dots visible in leaves of saplings or in shade-leaves, mature crown leaves very leathery, apex acute, acuminate or rounded, base rounded or narrowly to broadly cuneate, margin en­ tire, often more or less strongly inrolled, especially prominent in mature crown-leaves; lateral veins 10-25 on each side; upper surface glabrescent, lower surface with varying density of rust-coloured hairs, wearing away so lower surface becomes more or less glabrous with white hairs on veins only. Flowers clustered in leaf-axils or in axils of fallen leaves; pedicels 5-10 mm long, densely hairy. Calyx with 4(-5), almost free calyx lobes (sepals), ovate to more or less oblong, up to 6 x 3 inm. with short rust-coloured hairs on the outside. Co­ rolla cream-coloured; tube much longer than lobes, up to 6.5 mm long, lobes ovate to more or less orbicular, up to 2 mm long. Stamens inserted opposite corolla lobes, near top of tube; filaments c 1.5 mm long, anthers opening outwards, shorter than lobes. Staminodes spike-shaped, sometimes flattened and more or less petaloid. inserted on upper edge of tube between corolla-lobes, up to 1.5 nun long. Style up to 6.5 mm long, slightly exserted. with 4-lobed stigma. Fruit green, maturing yellow to or­ ange. narrowly ellipsoid, up to 4 x 1.5 cm, apex with c 1 cm long beak, pubescent or puberulous with short rust-coloured hairs. Seed ovoid to more or less narrowly ellipsoid, c 3 cm long; testa shiny, brown; scar lateral, pale brown, elliptic. Fig. 135.3.5-7. Emergent or upper canopy species in montane moist forest. 1350-2450 m. SU AR WG IL KF GG SD BA: from Ethiopia through Sudan (Imatong Mts), Uganda and Kenya, west to Rep. Dem. du Congo, south to Zambia Malawi and East Zimbabwe. Chaffey 1273; Friis et al. 3502. 4091. Hemsley (1961) recognized 4 subspecies within Aningeria adolfi-friederici: subsp. keniensis (R.E. Fries) J.H. Hemsl., subsp. usambarensis J.H. Hemsl., subsp. Jloccosa J.H. Hemsl., and subsp. australis J.H. Hemsl., all from East Africa. The trees in the Flora area belong to subsp adolfi-friederici. This is a high value timber tree used for all types of wood work. Traditionally, doors were cut out from the buttresses. Although classified as ‘not threatened’ on a global basis, the species and the high forest associated with it is rapidly being cut down and veiy little is being done to either conserve representative forests or replant the species. Seeds of the species have a veiy short viability period. The seedlings have to be kept well-shaded and planted into established forest if they are to survive.

62

135. SAPOTACEAE: 5. Pouteria

Fig. 135.3. PO U TERIA A L N IF O U A : 1 branch with buds and leaf underside showing a section o f the upper sid ex 1/4,2- b u d sx c 1; 3 - flower x 3 .4 ,4 - fruit x lVi. P. A D O L F II-F R IE D E R IC I 5 flowering branch x V3; 6 - flower x 2.3; 7 - fruit x c 'A. P. A L T ISSIM A 8 - flowering branch x Vi; 9 - flower x 3.4; 10 - young fruit x c Vi. 1, 3 & 4 from Friis 3918; 2 enlarged from Vougham:; 5 & 6 from Chaffex 1142; 7 from Greenway & Hummel 7308: 8 & 9 from Friis 40} 5; 10 from Harris 838. Drawn by Brigitte Wollenweber-Ratzer

135. SAPOTACEAE: 5. Pouteria

3. P altissima (A. Chev.) Baehni (1949); Hormogyne altissima A. Chev. (1917); Sider< xylon altissimum (A. Chev.) Hutch. & Dalz. (1931); / mngeria altissima (A. Chev.) Aubrev. & Pellegr. (1934-35) - types: Guinea Republic, Kaba valley, ( hevalier 13129; tributaiy of Mango River, Chevalier 13141 (both P syn., K isosyn.). Pouteria giordanoi Chiov. (1940) - type: WG, r ::ar Dembidollo, Uaba Forest, Giordano 2455 (FT holo.). Tall :ree of high forest, up to 40(-50) m tall, with clean, straight, cylindrical trunk and pale grey, smooth bark, sligl ilv buttressed at base. Young shoots and petioles finel> pubescent or puberulous, older branches glabrescent : - glabrous. Leaves spirally arranged and not particu­ larly clustered, petiole up to 1.5 cm long, blade thinly leathery, elliptic to elliptic-obovate or oblong-elliptic, 5 -1 ' 1-16) x 3-7 cm, apex obtuse or emarginate, some­ times shortly acuminate, rarely acute, base rounded or abruptly and broadly cuneate, margin entire; lateral veins 14-22 on each side, curving to form wavy marginal vein; almost glabrous on both sides except for midvein and larger veins on lower surface which have an indumentum of lor.g straight hairs. Flowers fragrant, 2-8, clustered in axils of current leaves, rarely in axils of fallen leaves; pedii els 3-6 mm long, pubescent. Sepals free nearly to

63

base, more or less spreading, elliptic to broadly ovate, 3.5-5.5 x 2 .5 ^ mm, pubescent or puberulous on outside with very short hairs. Corolla greenish-cream to pale yel­ low; tube slightly longer than lobes, up to 3.5 mm long; lobes ovate to elliptic-oblong, up to 2 mm long, ciliate. Stamens inserted in upper part of tube, a little below the division into lobes, filaments up to 1.5 mm long; anthers opening outwards. Staminodes spike-shaped, c 2 mm long, inserted between lobes. Ovary densely pilose; style up to 3.5 mm long. Fruit red, obovoid to subglobose, c 2 cm in diameter, fmely pubescent when young, becoming subglabrous at maturity. Seed more or less obovoid, c 1.5 cm long; testa shiny brown; scar pale brown, rough, more or less elliptic. Fig. 135.3.8-10. In the upper canopy of lowland and transitional forest, sometimes also in riverine forest; 1000-1500 m. WG IL KF; from Ethiopia, throughout forested parts of Uganda Kenya and Tanzania to C Africa, Guinea and Sierra Le­ one. Chaffey 1265; Friis et al. 4015, 4145. This is a highly valued timber species, now recognized as ‘vulnerable’ at a global level. Within the Flora area, it is only found in a small area of high forest, which is rapidly being cleared for its timber and for coffee and tea planta­ tions. The species should thus be considered as ‘endan­ gered’ within the Flora area.

64

136. M YRSINACEAE by Sebsebe Demissew* Cufodonlis, Enum.. 655-656 (1960); Kupicha. Myrsinaceae. in FI. Zamb. 7(1) 198-210 (1983); Hallidav, Myrsinaciae. in FI. Trop. E. Afr. 19 pp. (1984); Chen Jie & Pipoly, Myrsinaceae. in FI. China 15; 1-38 ( 1996). Shrubs or trees. Leaves alternate, occasionally opposite or in whorls; stipules absent. Inflorescence cymose. umbellate, racemose or paniculate, axillary or on older wood. Flowers unisexual or bisexual, small, regular, usually marked with dark resinous dots. Calyx 4-6-lobod, usually ciliate Corolla usually with (3 -)4 -6 (-7 ) pet­ als united into a short tube and lobed or occasionally free (Embelia). Stamens equal in num ber to the petals and opposite to them joined to the corolla, free or more or less joined; anthers sagittate, oblong or shortly ovi id. 2-thecous, splitting longitudinally or with apical pores; staminodes in female flow er often alm ost as large as sta­ mens Ovary globose or ovoid, superior or semi-inferior. 1-locular with few to many ovules; style short or long with acute to discoid or capitate, sometimes lobed. stigma. Fruit a drupe or berry, globose to ovoid, l(-few )-seeded. mdehiscent. Seed globose or angular. The family includes about 39 genera, distributed throughout the tropics and subtropics o f both hem ispheres, extendmg to Australia, New Zealand and Japan; 3 genera and 4 species in the Flora area. Many plants in this family are used as anthelmintics.

Key to genera 1. Ovary inferior or senii-infenor; calyx attached to ovary , fmit several-seeded. 1. Maesa - Ovary superior; calyx free from ovary; fmit 1-seeded. 2 2. Inflorescence racemose, rarely flowers in axillary clusters; petals free; leaves usually entire, occa­ sionally sliallowly to distinctly toothed. 3. Embelia - Inflorescence in clusters, cymose or umbellate; petals united into a short tube; leaves usually crenate to serrate. 2. Mvrsine 1. MAES A Forssk. (1775) Trees or shrubs, sometimes with runners or scrambling. Leaves glabrous or sparsely pubescent. Inflorescence usu­ ally axillaiy racemes or panicles; flowers bisexual or funcuonally female, white, pedicellate, with 2 bracteoles at base of calyx. Calyx more or less attached to the ovary, (4-)5-lobed, imbricate. Corolla campanulate, longer than calyx, with spreading lobes, united at base into a short tube. Stamens attached to corolla at throat or below; an­ thers dehiscing by longitudinal slits or aborted. Ovary in­ ferior or semi-inferior, style short or almost obsolete; stigma subdiscoid. often distinctly or indistinctly lobed; ovules few to many . Fmit small, indehiscent, dry or fleshy, globose, obovoid. crowned with persistent style and calyx several-seeded. Seeds small, more or less com­ pressed. globose or turbinate. About 200 species distributed in the tropics of the Old World; 1 in the Flora area.

The National Herbarium, Science Faculty, Addis Abeba University, P.O. Box 3434, Addis Abeba, Ethiopia

M. lanceolata Forssk. (1775) type; Yemen, Oddeim, Forsskal s.n. (C holo , K photo). Ki. picta Hochst. (1843);M lanceolata Forssk. var. serratifolia A. Rich., in Tent. FI. Abyss. II: 20 (1851) type: TU, Mt. Scholoda, near Adwa, 11 June 1837, Schimper I 286 (K isosyn.). Tree or shrub 2-20 m tall. Branches glabrous to sparsely covered with nist-coloured hairs. Leaf-blade narrowly el­ liptic, oblong, lanceolate, ovate, oblanceolate or obovate, 4-21.5 x 1.5—10 cm, obtuse to acuminate at apex, cuneate or rounded at base, subentire or minutely to coarsely and irregularly toothed, glabrous or sparsely pubescent; main veins distinct, midrib often pink; petiole 1.5-4 cm long In­ florescence a richly branched, lax or condensed axillary panicle, many-flowered, 2.5-13 cm long, glabrous to rust-coloured pubescent. Flowers bisexual or functionally female (4-)5-merous, c 3 mm in diameter, pedicels up to 1 mm long, glabrous to pubescent, with 2 bracteoles with cil­ iate margins Calyx campanulate; lobes triangular, ovate to broadly ovate, c 1 inm long, minutely ciliate, occasionally whole surface puberulent. Corolla white or cream, c 2 x 1.5 mm. lobes spreading, rounded. Ovary attached to the ca­ lyx, glabrous; ovules few to many; style exserted. short, stout, cylindrical, up to 1 mm long; stigma obscurely lobed. Fmit a globose berry, 4-5.5 mm in diameter, in clusters, yellowish-brown or red when ripe, with persistent calyx and style, glabrous, with many seeds but few reach­ ing maturity. Seed small, 2-3 x 2-3 mm. with many, irreg­ ularly shaped pits. Fig. 136.1. Gallery forest margin o f evergreen forest along river banks and streams, open woodland and valleys; 13503000 m. TU GD GJ WU SU AR WG IL KF SD BA HA; throughout tropical Africa, extending to south Africa, Madagascar and Arabia. Jackson 711; W.J.J.O. de Wilde 7250; Sebsebe D. 4319.

136. MYRSFNACEAE: 1. Maesa. 2. Myrsine

65

Figure 136.1. M A E SA L A N C E O LATA: 1 - fruiting branch x V2: 2 in­ florescence x 'A; 3 - flower x 6; 4 - co­ rolla spread out x 6; 5 - placenta showing arrangement o f ovules x 14; 6 - young fruit x 6; 7 - mature fruit x 6. 1 & 4 from Banda 325; 2 & 3 from Borges 168; 5 -7 from Forbes 242. Drawn by Margaret Tebbs. (Repro­ duced with permission from FI. Zamb. 7(1) Tab. 38.J

T'ie seeds are used as a vermifuge. Oil extracted from the sti ds is used for greasing the baking plate for iNJERA.a a-cake like bread. An extract of the wood is used in 2. MYRSINE L. (1753) Rapanea Aublev. (1775) HI. SIDA 17(1): 115-162 (1996). or trees Branches slender, glabrous or pubescent, glabrous or pubcscent; blade enure, eremite ot ireiy toothed to serrate Inflorescence axillary, tvered clusters or umbels; peduncles verv short. >vered. Flowers unisexual or bisexual, pedicellate, •merous. Sepais free or united at base, margins usuaiKJ ciliate. Corolla-lobes erect or recurved, margins

often ciliate and speckled. Stamens shorter to longer than the petals anthers sessile or filaments united at base into a collar which is attached to the corolla-tube. Ovary supe­ rior. globose to ovoid; ovuics few io many; style shon or sometimes exceeding petals, stigma large, discoidcapitate. lobed to fringed. Fruit dry or fleshy, globose. 1-seeded with persistent calyx. Seed globose. About 160 species world-wide. 2 in the Flora ;irea. In many previous accounts, the two genera Myrsine and Rapanea were treated as distinct. In Africa, they were separated by the presence (in Myrsme) or absenec (in Rapanea) of a collar uniting the stamens. A recent revi­ sion of the two genera on a world-wide basis has shown that the supposed distinction does not hold. Hence the two genera are united under the older name Myrsine.

136. MYRSINACEAE. 2. Myrsine

Figure 136.2, M Y R S IN E A F R IC A N A. 1 - flower­ ing branch (hermaphrodite) x 1; 2 flowering branch (male) x 1; 3 - her­ maphrodite flower x 12; 4 - pistil x 12; 5 —placenta showing arrange­ ment o f ovules x 18; 6 —male flower x 12; 7 - two stamens x 12; 8 - fruit­ ing branch x 1; 9 - fruit x 6. 1,3 & 5 from Tweedie 1994; 2, 6 & 7 from J.G. Williams in E.A.H. 12333; 8 & 9 from Oteke 66. Drawn by Pat Halliday. (Reproduced with permis­ sion from FI. Trop. E. Afr. M yrsinaceae: Fig. 2.)

longitudinally ribbed on older parts. Leaves alternate or crowded; petiole 0.1-0.3 cm long, glabrous to densely glandular-hairy; blade ovate to lanceolate, rhomboid, obovate or orbicular, 0.6—2 x 0.4—1 cm, acute to acuminate at apex, cuneate at base, crenate to serrate at margin, gla­ brous except for the few translucent hairs at the base of the midrib on upper surface; glands faintly transluscent, or­ ange. Inflorescence consisting of 1-5-flowered axillary clusters on young wood. Flowers functionally unisexual, dioecious, small, 4-merous, pendent, glabrous, gland1. M. africana I. (1753) dotted; pedicels up to 1.5 mm long. Sepals united at base or type: specimen grown in Holland at Clifford’s rarely free# c 1 mm long, margins finely ciliate in male Garden, Hartecamp s.n. (BM-HORT. CLIFT, lecto.). flowers, entire in female flowers. Corolla greenish-white or pink, campanulate with smooth to ciliate lobes, c 2 mm Evergreen undershrub, shrub or tree, 1-5 m tall. Branches long; tube in male flowers as long as the lobes, in female brown or purple, densely minutely hairy on younger parts. 1. Stamen-filaments united into a collar, joined to the corolla tube; corolla lobes erect; stigma discoid, fimbriate; leaves less than 2 cm long with crenate to serrate margin. 1. M. africana - Stamens separate, the anthers subsessile at corolla-throat; corolla-lobes spreading to recurved; stigma conical; leaves 4-18 cm long, with entire margin 2. M. melanophloeos

136. MYRSINACEAE: 2. Myrsine

67

Figure 136.3. M Y R S IN E M E LA NOPHLOEOS: 1 - flowering branch x 2/ 3, 2 - flower x 4; 3 - petal and anther x 8; 4 - pistil and calyx x 4; 5 - pistil x 8; 6 - ovary opened to show ariangement o f ovules x 12; 7 - fruiting branchlets x 2/ 3; 8 - fruit x 4. 1-6 from Thu Iin & Tidigs 247; 7 & 8 from Tweedie 1805. Drawn by Pat Halliday. (Reproduced with permission from FI. Trop. E. Afr. M yrsinaceae: tig. 3, as Rapanea melanophloeos.)

R.

fli vsers twice as long as the lobes. Stamens exserted, su tsessile or with filaments up to 1.5 mm long, united into a : liar which is joined to the corolla-tube; anthers criinso l magenta to blue-violet. Ovary ovoid; ovules few; sr le exserted. 1-3 mm long; stigma cream, discoid, fn i-ied. Fruit a globose berry, pale green to red or purple, tu nmg black when ripe, with a persistent style and calyx, 4- ' nun in diameter, smooth except for raised gland dots, gl. rrous, 1-seeded. Seed globose, c 3 x 3 mm. Fig. 136.2. Forests, hillsides in evergreen tickets, grassland with fo ast remnants, edges of scrub in grassland and by rivers; 190' >-3000(-3800) m. EW TU GD GJ WU SU AR KF SD B/ HA; Somalia and Sudan to South Africa. Angola; also

in Azores, Socotra. Arabia, Afghanistan, India and China; cultivated in Europe. Burger 2260; Mooney 5552; Sebsebe D. & Ensermu K. 1512. Used as a tooth-brush. Fruits edible. 2. M. melanophloeos (/,.) R. Br. (1810); Sideroxylon melanophloeos L. (1767); Rapanea melanophloeos (L.) Mez (1902) - type: South Africa. Cape of Good Hope, van Royen, LINN. 261/3 (LINN holo. K photo). Myrsine simensis Hochst. ex DC. (1844); Rapanea simensis (Hochst. ex DC.) Mez (1902) - type: GD, Mt Silke in Simen, 14 Feb. 1840, Schimper 673 (K iso.).

68

136. MYRSINACEAE: 2. Myrsine, 3. Embelia

Fig. 136.4. E M B E U A S C H IM P E R I 1 - fruiting branch x 3/4; 2 - underside o f leaf showing gland dots x 3/4; 3 - male raceme x 1'/a; 4 - male flower x 4V2\ 5 male flower showing reduced ovary x 6; 6 female flower x 6; 7 - front o f male petal x 714; 8 - back o f male petal show­ ing glands x 7'/2; 9 - fruit x 4'/i, 10 seed showing remains o f membrane x 4 V2 . 1, 2, 9 & 10 from Sebsebe D. 89; 3 -5 , 7 & 8 from Pavlov 97-162; 6 from Lemma G. Selassie 194. Drawn by Sara Howard.

Evergreen tree, occasionally a shrub, 2-20 m tall. Bark rough, fissured, red-brown to grey-white, with prominent lenticels; branchlets purple. Leaves alternate, sometimes in pseudo-whorls at the end of branches; petiole 0.1-1.2 cm long, often w inged, red when young, more or less gla­ brous; blade narrowly elliptic to elliptic, oblong, lanceo­ late to oblanceolate or obovate, 4—15(—18) x 1.5-6(-7.5) cm. obtuse, mucronate to shortly acuminate at apex, taper­ ing towards the petiole at base, leather)', blue-green, glossy above, more or less glabrous, margin entire. Inflo­ rescences axillary', or on short shoots below leaves, in clusters of 3—12; bracts short, broadly rounded, c 2.8 mm long, minutely ciliate at apex; pedicels 3-9 mm long, stout, expanded upwards, glabrous. Flowers bisexual or functionally female, (4-)5(-6)-merous. 7-10 mm in di­ ameter. Calyx cup-shaped, lobes ovate, c 1.5 x 1.5 nun with margins minutely ciliate. rarely w’hole surface puberulent. Corolla white, yellow, green or pink, buds yellow-red or purple. 2-5 mm long, tube c 1 mm long; lobes spreading to recurv ed, ovate to oblong, margins mi­ nutely ciliate. Stamens pale yellow, subsessile to sessile, filaments joined to corolla at mouth of tube, anthers ovate, triangular or sagittate, 1.5-4 mm long, occasionally with

tuft of minute hairs at apex, dehiscing by longitudinal slits. Ovary ovoid, conical or almost globose, glabrous; ovules few; style short, stout, stigma conical, sometimes 2-4lobed. Fmit pink, mauve, blue, whitish-blue or purple, globose, 5-7 mm in diameter. Seed small, globose, w ith cavity at base. Fig 136.3. H a g e n ia - J u n ip e r u s - P o d o c a rp u s forest, open pas­ ture with H yp ericu m , H a g e n ia and E rica , E ric a thickets; 2500-3750 in. TU GD GJ SU AR WG GG BA; south to Southern Africa and Angola, west to Nigeria. Fernando Po and Cameroun de W ilde 9178; G ilb e r t & T e w o ld e B .G .E . 3241; S e b se b e D. & E n serm u K. 4978.

The juice is mixed with other ingredients and drunk to ' improve memory .

3. EM B EU A Burnt, f (1768) Scandent shrubs or small trees. Branches glabrous or pu­ bescent. Leaves petiolate, rarely subsessile, alternate, en­ tire or toothed. Inflorescence a terminal panicle, axillary raceme or cluster. Flowers dioecious (functionally unisex-

136. MYRSINACEAE: 3. Embelia

ial), pedicellate or sessile, 4- or 5-merous. Sepals united lor 1/3 or almost free, usually hairy on inner surface. Sta­ mens longer than petals and joined to them at varying icights, rarely free; filaments long, slender, rarely short or rsent, anthers splitting longitudinally. Ovary of male i twers much reduced, conical, entire or rarely lobed. ruit globose, rarely obovoid, 1-seeded. Seed subglobose, i : vered by membranous remains of the placenta, sornei ines with conspicuous cavity at base.

69

dense axillary raceme. 1-5 cm long, arising from short shoots, or often on bare branches below the leaves, with sparse, rust-coloured hairs, each raceme few to severalflowered. Flowers pedicellate, 4- or 5-merous, 4-6.5 mm in diameter, pedicels 3-8 mm long, with rust-coloured hairs. Calyx cup-shaped, 1.5-3.5 nun in diameter, united to the middle; lobes ovate, spreading, speckled or not. with minutely ciliate margins, glabrous or with sparse rust-coloured hairs. Petals free, greenish-white, yellow, yellowish-cream, speckled, spreading to somewhat reWidely distributed throughout Asia. Australasia, flexed. oblong to ovate, 2.5-3.5 x 1.7-2 mm, minutely pu­ I. lascarene Is., tropical and southern Africa with a total of bescent on inner surface, margin ciliate. Stamens joined to ll2 species; 1 in the Flora area. lower part of petal. 4-4.5 nun long, with slender fila­ ments; anthers creamy-white. Ovary globose or ovoid; E. schimperi Vatke (1876) types: GD/TU, Gerra Abuna Tekla-Haimanot, style cylindrical, 2-2.5 mm long in female flower, shorter in male flower; stigma truncate or 2- or 3-lobed. Fruit orSchimper 1432 (B syn. destroyed; BM E K isosyn.) & Herroe Gottes Georgis near Gaffat, Schimper 1493 (B ange-yellow, reddish-green to red, speckled, globose, syn. destroyed, BM lecto., E K isolecto.). 5-8.5 nun in diameter, with persistent calyx and style, glabrous, 1-seeded. Seed globose, smooth, with cavity at Scandent or climbing shrub or tree, 2-13 m tall. Branches base, 4.5-7 mm in diameter, brown with irregular orange v ith prominent lenticels. Young parts with rust-coloured markings. Fig. 136.4, h lirs, becoming glabrous. Leaves alternate or in clusters Forest edges, secondary forest and scrub, and along a end of branches; blade lanceolate or oblanceolate, rivers; 1700-2800 m. GD GJ SU AR WG IL KF GG SD 3 5-12 x 1.3-6 cm, obtuse, acute or acuminate at the apex, BA; Kenya, Uganda, Tanzania, Burundi, Rep. Dem. du c ineate or more or less rounded at base with entire, Congo, Malawi. Zambia, Zimbabwe, Angola Sudan and s ghtly revolute margins, brittle or leathery, with visible Cameroun. Mooney 8866; Sebsebe D. 89; Thulin & As/aw g and dots: midrib grooved above, with 12-20 or more, H. 3938. c osely parallel, secondary veins on either side; petiole n a. grooved above, 0.4-1.3 cm long. Inflorescence a Seeds widely used as an anthelmintic.

137. LO GANIACEAE by A. J. M. Leeuwenberg* Bruce & Lewis, Loganiaceae in FI. Trop. E. Afr. 47 pp. (1960), Onochie & Leeuwenberg, Loganiaceae, in FI. W Trop. Afr. 34—47 (1963); Cufodontis, E n u m 674-678 (1965). Leeuwenberg, Loganiaceae, in FI. Zamb. 7(1): 327-374 (1983)

Trees, shrubs, climbers or sometimes herbs. Leaves usually opposite, less often in whorls o f 3 or subopposite, simple, pinnately veined. Stipules true, false or absent. Flowers mostly bisexual, 4- or 5-merous (corolla and androecium 8-16-merous \n Anthocleista). actinomorphic (in Flora area). Corolla mostly coloured, petals united, variously shaped; lobes valvate, imbricate or contorted in bud Anthers dehiscent throughout by a longitudinal slit. Pistil simple; ovary superior or sometimes (not in Flora area) slightly inferior, usually 2-celled. Fruit a berry or a capsule. 29 genera and about 570 species mainly in the tropics: 4 genera and 12 species in the Flora area.

Key to genera 1. Candelabrum-shaped tree with often large leaves; inflorescence large, terminal, brittle; corolla tube 25-37 mm long, lobes 8-16. 1. Anthocleista - Trees or shrubs, otherwise branched; inflores­ cence often axillary', corolla tube less than 10 mm long, lobes 4-5. 2 2. Corolla lobes valvate; fruit a berry; leaves mostly 3-5-veined, strictly opposite. 4. Strychnos - Corolla lobes imbricate; fruit a capsule; leaves pin­ nately veined, often in some pairs subopposite, less often in whorls of 3. 3. Stamens included or slightly exserted; fruit a cap­ sule. much longer than the calyx, or fruit a berry; anther cells discrete; inflorescence thyrsoid or loosely paniculate. 2. Buridleja - Stamens well-exserted; fruit a capsule, about as long as. and included in. the tubular calyx; anther cells confluent at the apex; inflorescence more or less paniculate. 3. Nuxia 1. ANTHOCLEISTA Afzel. ex ft Br. (1818) Leeuwenberg, Acta Bot. Neerl. 10: 1-53 (1961). Candelabrum-shaped trees (in the Flora area), unarmed or with paired axillary spines. Stipules intrapetiolar. Leaves often large, each pair often unequal, petiolate or sessile; blade brittle when dried, as is the large terminal inflores­ cence. Sepals 4, persistent, decussate, rounded (in the Flora area), circular, or nearly so. Corolla white or cream (in the Flora area), thick, fleshy; lobes 8-16, elliptic or narrowly so. spreading or recurved. Stamens exserted; an­ thers narrowly elliptic. Ovaiy ovoid or nearly so, 4-celled; style thick; stigma large; ovules numerous. Fruit a bem Seeds numerous, small.

A. schweinfurthii G ilg(l893) - type: Rep. Dein. du Congo, Schweinfurth 3726 (K lecto., S W iso.). A. niamniamensis Gilg (1893). Tree 3-30 m tail, with a terminal inflorescence on each branch. Branchlets without, or occasionally (especially in young plants) with short paired, partially united spines Leaves usually petiolate but often sessile in young plants or on low-level branches: blade dark green above, puler beneath, papery to leathery , narrowly to very narrowly obovate. 7-45 \ 1.5-18 cm. in young plants even narrower and up to 150 \ 40 cm, cuneate at base, mid-vein more or less acute beneath. Sepals green, when dry usually smooth, especially the outer ones, in flower usually appressed to the base of the corolla in fruit often spread­ ing when dried, the outer ones orbicular or slightly broader than long, 8-13 x 9-13 mm. the inner ones usually slightly larger, becoming retuse with the development of the co­ rolla. Corolla in the mature bud 55-61 mm long, white or cream, tube often greenish-white, about 3-4 times as long as the calyx; lobes 10-11, narrowly elliptic, reflexed in open flowers. Berry globose or ellipsoid, rounded or api­ culate at the apex, never shrivelled when dried; wall thick. Fig. 137.1. Transitional rain forest, gallery forest and thickets; 1000-1350 m. IL KF; west through C Africa to Nigeria, south to Tanzania and Angola. Friis et al. 4040; Meyer 8977; Mooney 9213.

2 BUDDLEJA L. (1753) Leeuwenberg, Meded. Landb. Hogesch. Wag. 79(6): 1-163 (1979).

Shrubs or trees, sometimes climbing, monoecious or 14 species in tropical Africa, Comoro Islands and dioecious (in the naturalized B. cordata), mostly w itli a white or pale grey indumentum, usually with stellate hairs. Madagascar, 1 in the Flora area. Leaves opposite or less often subopposite; blade often ser­ rate Inflorescence (in the Flora area) large and manvflowered. Flowers 4-merous. bisexual (or in monoecious species unisexual). Corolla lobes (in the Flora area) imbri­ Wagemngen Branch, Nat. Herb. Netherlands, P.O. Box 8010. 6700 ED cate in bud Sterile reproductive organs present in unisexWageningen. The Netherlands

137. LOGANIACEAE: 1. Anthocleista, 2 Buddleja

71

Figure 137.1. A N T H O C L E IS T A SC H W E IN F U R T H II: 1 - branch: 2 young flower bud; 3 - mature bud; 4 flower; 5 - fruit. All x V2. 1,2 & 4 from Schmitz 3332; 3 from Tisserant 1934; 5 from Louis 3100. Drawn by Yuen Tan. (Modified and reproduced with permis­ sion from Acta Bot. Neerl. 10: fig. 12.)

m J i li 'J

s \

gpR w|v'

i al flowers, but reduced. Anther cells discrete. Ovary 2( r 4-celled. Fruit a capsule or less often (B. madagascarit nsis) a beriy. Seeds obliquely ellipsoid or obliquely iusiform, winged or not. About 90 species in the tropics and subtropics of / merica, Africa and Asia; 4 in the Flora area. Inflorescence thyrsoid or sometimes paniculate (see B. madagascariensis)\ corolla tube cylindrical. 2 Inflorescence paniculate; corolla tube campan­ ulate. i b. cordata Corolla orange or nearly so; stamens inserted near the corolla mouth, barely included. 3 Corolla violet; stamens inserted about hallway down the corolla tube; cultivated. 2. B. davidii Branchlets 4-angled or nearly so; leaf blade usually decurrent into the petiole; corolla tube 3.5-8 mm long, 1.5-2.7 times as long as the calyx; ovary 2-celled; fruit an ellipsoid capsule. 4. B. poiystachya

Branchlets terete; leaf blade not decurrent into the petiole; corolla tube 7.3-10 mm long, 2.5-4.5 times as long as the calyx; ovary 4-celled; fruit a globose berry; cultivated. 3. B. madagascariensis 1. B. cordata H.B.K. (1818) type: Mexico, Humboldt & Bonpland s.n. (P-BO holo.). Deciduous shrub or tree, 2-12 m tall. T ru n k 1 0 -4 5 cm in diameter at the base. Branchlets stellate-tomentose, glabrescent Leaves petiolate; petiole 1-4 cm; blade ovate, narrowly ovate, or narrowly elliptic, 4-23 x 3-14 cm. acute or acuminate at apex, subcondate to cuneate at base, some­ times decurrent into the petiole, entire or serrulate, glabrous or sparingly stellate-hairy and glandular above. Inflore­ scence terminal, paniculate, 6-30 cm long. Flowers grouped into shortly pedunculate cymules, fragrant. Calyx c 2-3 mm long, stellate-tomentose outside; tube 1-1.8 mm long; lobes broadly triangular, 0.5-1.5 x 1.2-1.5 mm. Co­ rolla white, cream or yellow with a flush of orange at the

72

137. LOGANIACEAE: 2 Buddleja

Figure 137.2. BU D D LEJA CORD A T A . 1 - flowering branch x ' ^ 2 female flower x 5; 3 - pistil x 5; 4 opened corolla with staminodes x 5; 5 male flower x 5: 6 - pistillode x 5; 7 opened corolla with stamens x 5; 8 fruit x 5; 9 - seed x 50. 1-4 from J de Wilde 5480; 5 -7 from P. Jansen 7278. 8, 9 from P. Jansen 7279. Drawn by Yuen Tan. (Reproduced with permis­ sion from M eded Landb. Hogesch. Wag. 79 (6): fig. 5.)

throat campanulate, stellate-tomentulose outside on upper two-thirds, pilose inside in upper half of the tube and lower half of lobes, tube 1.5-2.5 mm long, shorter to slightly lon­ ger than the calyx; lobes ovate or nearly so. 1.2-2.2 mm long, rounded, spreading. Stamens slightly exserted, in­ serted just below the corolla mouth; anthers in male flowers 0.6-1 mm, in female flowers about 0.4-0.5 mm. Pistil lon­ ger than the staminodes; pistillode about as long as the sta­ mens. Capsule oblong 3.5-5 x 1.5-2 mm. Fig. 137.2.

shortly petiolate; blade narrowly ovate or narrowly elliptic. 4-20 x 1-7 cm, acuminate at apex, cuneate at base, serrate to subentire, glabrous or nearly so above, whitetomentose (mostly drying brownish) beneath. Inflores­ cence thyrsoid. Corolla violet or lilac, orange-yellow in the throat; tube 6-11.5 mm long, nearly cylindrical, 2-4 times as long as the calyx, 3-5.5 times as long as the lobes Sta­ mens inserted in the middle of the corolla tube. Capsule narrowly ellipsoid, mostly 3-4 times as long as the calyx.

Cultivated and naturalized; 1900 m. HA; native of Mexico. Burger 3111; P. Jansen 5243, 7278.

Cultivated; WG; native of China. Milcherish 126

2. B. davidii Franch. (1888) type: China, E Tibet, Rev. Pere David Aug. 1869 (P holo.). Shrub, 0.5-3 m tall, often sarmenlose. Branchlets more or less 4-angled, stellate-tomenlose, glabrescent. Leaves

3. B. madagascariensis Lam. (1785) -ty p e: Madagascar, Sonnerat s.n. (P-LA, holo.). Sannentose shrub. 2-A m tall, or a climber, 8-10 m long. Branchlets terete, white tomentose with stellate hairs, dry ­ ing rusty. Leaves petiolate, petiole 0.5-2 cm long, stellatetomentose; blade narrowly ovate or elliptic, 2-3 (-4) times

137. LOGANIACEAE: 2. Buddleja

73

Figure 137.3. BUDDLEJA POLYSTA C H YA : 1 & 2 - flowering branches x 16; 3 - flower x 4'/2; 4 - opened corolla x 4'A; 5 - pistil x 414; 6 - fruit x 414; 7 seed x 45; 8-11 - leaves x Vi. 1 ,3 -5 from W. de Wilde 9230; 2 from Schimper 459; 6, 7 from Pichi-Serm olli 1281; 8 from Vatova 1171; 9 from Schweinfurth & Riva 779; 10 from Pappi 1495; 11 from M ooney 5304. Drawn by Yuen Tan. (Re­ produced with permission from Meded. Landb. Hogesch. Wag. 79 (6): fig. 15.)

1

its long as wide, 4-14 x 1.5-7 cm, acuminate at apex, : landed, cuneate, or rarely subcordate at base, entire, with repressed reticulate venation, and glabrous or nearly so i bove (only tomentose when young), white-tomentose ■ tit stellate hairs (drying rusty) beneath. Inflorescence i lvrsoid or paniculate. 5-25 x 2-15 cm. Flowers sessile or hDrtly pedicellate. Calyx campanulate or urceolate, 2-3.5 1.5-2.5 inm. white-tomentose with stellate hairs (drying isty) outside. Corolla dark yellow, orange, or salmon, 5lellate-tomentose outside; tube 7.3- 10 mm long, nearly ; ylindricaL 2.5-4.5 times as long as the calyx, 2.1-3.4 times as long as the lobes. Stamens barely included, in• ..-rtedjust below the corolla mouth: anthers 1-1.4 x 0.3-0.8 i im. Pistil ovary 4-celled; stigma large, clavate, 1.2-1.7 n in long Berry blue-violet or orange, immature white, i lobose or nearly so, 2.5-5 mm in diameter.

Cultivated; 1900 m. HA; native of Madagascar. J.J. Bos 7534. 4. B. polystachya Fresen. (1838); B. saltiana Steud. (1841) - type: Ethiopia, sine loc. Ruppell 14 (FR holo.). B. rufa Fresen. (1838) - type: Ethiopia, sine loc. Ruppell 15 (FR holo.). B. polystachya var. parvifolia Marquand, in Kew Bull. 1930: 193 (1930)-type: EW, Isaba, near Keren, Beccari 69 (K holo., FI WAG iso.). Shrub or tree, 1-12 m tall, with spreading and arching branches; d.b.h. 15-35 cm. Branchlets 4-angled or nearly so, grey- or white-tomentulose with stellate hairs like the leaves beneath, the peduncles, the bracts be­ neath, and the calyx outside; indumentum usually drying

74

137. LOGANIACEAE: 2. Buddleja, 3. Nuxia

rusty. Leaves petiolate; petiole 0.1-1 cm long; blade variable in shape and size, ovate, narrowly ovate, ellip­ tic, or narrowly elliptic, 1-17 x 0.4-6 cm (up to 20 x 8 cm in vigorous shoots), acuminate, acute or, when very small, obtuse at apex, gradually to abruptly narrowed at base, and also mostly decurrent into the petiole, margin entire, senate, or crenate-serrate, surface stellatepubescent with impressed venation above, much paler and stellate-tomentulose beneath. Inflorescence termi­ nal, spicate, mostly interrupted, 3-25 x 1-2.5 cm; lower cymes often pedunculate, upper ones sessile. Flowers sessile, mostly in dense clusters, sweet-scented. Calyx campanulate, 2.2-3.5 mm long. Corolla orange, often with yellow tube, stellate-pubescent outside; tube 3.5-8 mm long, nearly cylindrical. 1.5-2.7 times as long as the calyx, 2.3-4 times as long as the lobes. Stamens barely included, inserted just below the corolla mouth. Capsule ellipsoid, slightly exserted. Fig. 137.3. Montane Juniperus - Podocarpus - Olea forest, par­ ticularly in clearings, degraded woodland and as isolated trees in cultivated fields, in live fences around houses; (700-) 1000-3300 m. ?EE ?AF EW TU GD GJ WU SU AR WG KF SD BA HA, Kenya, Tanzania, NE Sudan: Yemen. Saudi Arabia. Mooney 6864: Ryding & SHeshi N. 1984; Thulin 1562. An important home garden plant grown in living fences. Branches and leaves are used to wash pots, the leaves can provide fodder, the dry twigs can be used to start a fire, and many parts also have medicinal uses.

3. NUXIA Comm, ex Lam. (1791) Leeuwenberg, M eded. Landb. H ogesch. Wag. 75(8): 1-80 (1975).

Shrubs or trees. Trunk often striate. Leaves opposite, in whorls of 3, or in some lateral branches occasionally alter­ nate. petiolate. Inflorescence terminal, thyrsoid. but often dichotomously branched; ultimate branches topped by solitary flowers or by heads of 3 or more flowers. Flowers 4-merous. Calyx campanulate to cylindrical. 4-lobed; lobes much shorter than the tube, triangular. Corolla mostly white or cream, circumscissile; tube usually slightly shorter than the calyx cylindrical, lobes recurved from above the base, concave and rounded to acute at apex. Stamens well exserted, inserted at the mouth of the corolla tube: anthers glabrous, cells 2, confluent at apex, parallel when young, conspicuously divergent after the pollen is shed. Style well exserted. Capsule about as long as. and included in, the persistent calyx, or up to 1/3 lon­ ger. Seed small, fusiform. 15 species in southern Arabia, tropical Africa (inclu­ sive of Madagascar, the Comoro Islands and the Mascarenes). and South Africa: 2 in the Flora area. 1. Leaves opposite, bluntly serrate or less often entire; on river banks. 2. N. oppositifolia - Leaves in whorls of 3, entire, or especially in shade, leaves sharply serrate, in mountains. 1. N. congesta

1. N. congesta R.Br ex Fresen. (1838); Lachnopylis congesta (R. Br. ex Fresen.) C.A. Smith (1930) - type: Ethiopia, Ruppell 13 (FR holo., BM iso ). L. ternifolia Hochst (1843) - type: Ethiopia, Schimper 1159 (FR holo., BM BR BREM CGE FI G GH GOET K L M MO MPU OXF P S UPS US W WAG WU Z iso.). Evergreen or (in the Flora area sometimes) deciduous tree or shrub, 2-25 m tall, often gnarled. Trunk in larger trees striate and irregularly ridged, d.b.h. 5-100(-200) cm; bark pale grey-brown, fissured, peeling off in narrow strips or small thin scales. Leaves in whorls of 3, petiolate; petiole 0.3-2 cm long; blade leathery or somewhat leatheiy, very variable in shape and size, elliptic, narrowly elliptic, obovate, nearly rhomboid, or suborbicular, (1—)2—15 x (0.3—)1.2—7.5 cm, acuminate to emarginate at apex, cuneate or less often rounded at base or decurrent into tiie petiole, margin entire, or sometimes more or less distinctly serrate-dentate or crenate, glabrous to subtomentose with stellate hairs on both sides. Leaves on sucker shoots (mostly in the shade) darker green, herbaceous, serrate, and more hairy. Inflorescence seemingly umbellate to paniculate, congested or rather lax. Flowers solitary or 3 together, fragrant. Calyx green, often viscid, 3-8 x 1.5-2.2 mm. minutely to manifestly pubescent with glandular and mostly also ordinary hairs outside, covered with soft, silklv. appressed hairs inside. Corolla white, lobes oblong, 2-5 x 1-2 nun. Capsule 0.5-1.5 mm longer than the calyx, appressedly pubescent. Montane Juniperus - Podocarpus - Olea forest, partic­ ularly at higher altitudes and with Erica, degraded wood­ land and as isolated trees in cultivated fields, in live fences around houses; (1 100-) 1550-3800 m. EW TU GD GJ WU SU AR WG DL KF SD BA HA, Somalia west to Guinea, south to Angola and S Africa; also Saudi Arabia. Berger 2328; Mesfin T. & Zerihun W. 2828; Friis et al. 527. Grown in living fences round home gardens. An im­ portant bee plant which flowers during the dry months. The wood makes good charcoal.

2. N. oppositifolia (Hochst.) Benth. (1846); Lachnopylis oppositifolia Hochst. (1843) - type: TU, near Djeladjeranne, Schimper 1714 (P holo., BM BR FI G GH GOET K L LE M MO S W WAG iso.). Nuxia dentata R.Br. ex Benth. (1846)-type: Ethio­ pia. sine loc. Salt s.n. (BM holo.). Evergreen shrub or small tree, 1—15(—20) m tall, often gnarled and willow -like, sometimes straggling. Trunk often rather striate, d.b.h. 10-60 cm. Bark reddish-brown, smooth or shallowly and longitudinally fissured. Leaves opposite, less often sub-opposite, or occasionally (on a few of the branchlets) alternate, often sticky when being dried, petiolate: petiole 0.2-1.2 cm long, blade leatheiy when dried, narrowly to very narrowly elliptic, 3-13 x 0.4-3(-5) cm, obtuse or rounded and often mucronulate at apex cuneate at base or decurrent into the petiole, margin bluntly serrate to entire, glabrous or with minute glandular hairs

137. LOGANIACEAE: 3. Nuxia, 4. Strychnos

75

Figure 137.4. N U X IA O PPO SITIF O L IA : 1 - flowering branch x 'A; 2 flower bud x 5; 3 - young flower x 5; 4 opened corolla o f young flower x 8; 5 pistil with corolla base x 7; 6 - fruiting calyx x 7; 7 - seed x 20; 8 - 1 2 -le a v e s x V2 . 1-5 from Mendonqa 3461; 6, 7 from Gossweiler 12636; 8 from Adamson 42; 9 from Leach 11298; 10 from Schlechter 11738; 11 from Torre 3701; 12 from Buchanan in coll. J.M. Wood 7097. Drawn by Louise van der Riet. (M od­ ified and reproduced with permission from Meded. Laridb. Hogesch. Wag. 75 (8): fig. 10.)

A

It; A*

I'

k

I A V j V V

i

on i )th sides. Inflorescence rather congested, ultimate bran.hes and pedicels very short or obsolete. Flowers sligl ly fragrant, 1—3(—8) together. Calyx green. 3.5-5 x 1.2- 1 mm, minutely pubescent with glandular and often alsc ordinary hairs outside, when diy seemingly glabrous and ften shining, silky inside. Corolla white, cream, or son .:times pale lilac, lobes oblong, 1.8-3 x 0.8-1.2 mm. Cap lie aboui as long as the calyx, appressed-pubescent. Fig. 137.4. '-‘graded montane forest and evergreen bushland, alsc in gallery forest and river gorges; 800-2100(-2400) m. f :: EW TL SU AR GG SD HA; west through Sudan to Cor !0, south to W Angola, NW Namibia and S Africa (Nai.il); also Saudi Arabia. Friis et al. 6818; Mooney 974 Schweinfurth & Riva 1690.

4. STRYCHNOS L. (1735) Leeuwenberg, Meded. Landb. Hogesch. Wag. 69(1); 1-316 (1969). Erect or climbing shrubs or trees. Leaves opposite or, on the main axis, sometimes in whorls of 3, entire. Inflores­ cence terminal or axillary. Flowers 4-5-merous. Corolla rotate to hypocrateriform; lobes valvate in bud. Stamens exserted or included; anther cells discrete. Fruit a berry, large or small; wall thin or thick. Seeds large, variably shaped, generally disk-shaped to subglobose. Circuintropical, about 190 species; 4 in the Flora area. Seeds of all species contain the poison stiychnine that is obtained commercially from S. nux-vomica grown in Asia. Several species have good quality timber.

76

137 LOGAN1ACEAE: 4. Strychnos

1. Inflorescence terminal: sepals at least twice as long as wide, long-acuminate or subulate; branches often with some axillary spines; fruits large, glo­ bose, about 5-10 cm in diameter, yellow when ripe. 4. S. spinosn - Inflorescence axillary; sepals about as long as wide: plants unarmed, fruits mostly smaller. 2. Leaves grey-green, rounded at apex, often with pale green reticulate venation on both sides; fmit fairly large, 3-9 cm in diameter, flowers 4-merous; corolla-tube more than 3 mm long. 2. S. innocua - Leaves pale to dark green, mostly acuminate; fruits small, 1-2 cm long, flowers mostly 5-merous; corolla tube up to 2.2 ram long. 3 3. Branches without conspicuous lenticels; leaf vena­ tion reticulate, prominent on both sides. 1. S. henningsii - Branches with conspicuous lenticels; leaf venation not prominent above. 3. S. mitis

Roots, fruit and bark are used in local medicine, the wood is used as firewood, building posts in houses and tool handles. 2 S. innocua Del. (1826) types: Nubia Calliaud s.n. (not seen, apparently not preserved) and TU, Tacazze River valley, near Djeladjeranne, Schimper III: 1817 (P neo.). S. unguacha A. Rich. (1847) - type: Schimper III: 1817 (P holo . BM BR FI G GOET HAL K L L E M MO P S UPS W iso ).

Deciduous shrub or small, often richly branched tree. 2-12 (-18) m tall. d.b.h 7-40 cm. Bark smooth, somewhat pow­ dery. Branches pale grey-brown, sometimes powders , sometimes w ith conspicuous lenticels. Leaves: petiole gla­ brous or pubescent. 0.2-0.7 cm long; blade dull, glaucous and with mostly pale green reticulate veins on both sides, leathery (living and dry' material) or papery (dry), elliptic, narrowly elliptic, obovate, or narrowly obovate, (2-)410(-20) x (1—)2—7(—13.5) cm, rounded at apex, glabrous or 1. S. henningsii Gilg (1893) type. South Africa, E Cape Prov., Pondoland, pubescent on both sides, one or two pairs of distinct sec­ ondary veins from, or from above, the base, curved along near Umnonono, Bachmann 1745 (E lecto.). the margin and often a faint submarginal pair, tertiary ve­ Shrub or small tree, 2-10(-20) in tall, d.b.h. c 40-50 cm; nation reticulate and distinctly prominent on both sides In­ bark rough. Branches pale grey or pale brown, sometimes florescences axillary or from older leafless stems, usually shallowly fissured, without conspicuous lenticels; several together, very short and nearly clustered. 1 x 1-1.5 branchlets conspicuously grooved and often partially x 1.5 cm, few-flowered. Flowers 4-merous. Sepals pale 4-angled when dried, glabrous. Leaves: petiole 0.1-0.3 green, free, ovate, broadly ovate, or suborbicular, 1.7-3.5 cm long, glabrous; blade leathery, very variable in shape x 1.5-2.5 nun, (the inner slightly smaller), rounded, ciliate, and size in a single branchlet, elliptic, oblong, narrowly glabrous or pubescent outside. Corolla in the mature bud elliptic, or ovate, (1.5—)2—6(—10) x (0.6-)l-3(-6) cm, 2.2-4 times as long as the calyx and (6—)6.5—9(—10 5) nun rounded to acuminate at apex, cuneate, rounded, or some­ long, cream or greenish-yellow\ glabrous outside, inside times on main axis subcordate at base, glabrous on both with a brush-1 ike ring of white woolly hairs in the throat sides; one pair of secondary veins from, or from above, and just al the base of the lobes; tube cy lindrical or nearly the base, curved along the margin; tertiary' venation retic­ so^ (3_)3 5_5 5(_6 ) x 1.5-2.5(-3) mm long, 1.6-2.5 times ulate, prominent on both sides, especially in thick leaves. as long as the calyx, 1-1.7 times as long as the lobes; lobes Inflorescences axillary and sometimes also terminal, thick, narrowly triangular, 3-4(-4.5) x 1.3—2(—2.3) mm. much shorter than the leaves, 1 x 1-2 x 2 cm, congested, acute or sub-acute, spreading. Stamens hardly exserted. in­ peduncle often very short. Flowers fragrant, 5-merous, serted at the mouth of the corolla tube; anthers glabrous. sessile or subsessile. Sepals pale green, united at the base, Pistil (4-)5-7.5 mm long. Mature fruit globose, orange or broadly circular or nearly so, 1-1.4 x 1-1.4 nun, rounded yellow, hard, (2.5-)4-7.5(-9.5) cm in diameter with (3-) or obtuse, minutely ciliate, glabrous outside. Corolla 8-50 seeds; wall thick; pulp orange, edible. Seeds pale yel­ 2.8-4 mm long in the mature bud and 2.3-3 times as long low-brown, irregularly curved, flattened or more or less as the calyx, greenish-yellow, cream or white, subrotate plano-convex, obliquely ovate, elliptic, or tetrahedral, and 4-5 mm in diameter when open, glabrous outside, 17-21 x 13-20 x 5-8 mm. with thick, very' short, erect pilose inside or villose at the base of the lobes or some­ hairs, rather rough. Fig. 137.5. times entirely glabrous; tube short, 0.7-1.2 times as long Deciduous woodland; 600-1400(-1600) m. EW TU as the calyx, 0.8-1.5 mm long; lobes thick, triangular to GD GJ SU WG IL GG SD; tropical Africa. Friis et al ovate, 1.7-2.5 times as long as the tube, 2-2.5 x 1-1.6 2507; Gereau 1267; Thulin & Asfaw H. 4031. mm, acute, spreading. Stamens just exserted. inserted at mouth of corolla tube; anthers glabrous. Pistil glabrous, 1.6-2.2(-3) mm long. Fruit ellipsoid, 1 x 0.8-2 x 1.5 cm, yellow, orange, or red, one-seeded; wall thin Seed pale brown, ellipsoid, not flattened, 0.8-1.2 x 0.5-0.7 x 0.5-0.7 cm, glabrous, smooth, with a deep closed groove at one side (like coffee-bean), very minutely pitted. Mixed dry broadleaved woodland; 550-1550 m. IL SD BA; Somalia west to Congo, south to Angola and S Africa. M. G. Gilbert & Friis 8377; Friis et al 1920; P uff et al. 870430-1/16.

The leaves give flavour to local beer and the wood is used for making tools. 3. S. mitis S. Moore (1911) - type: Zimbabwe, Chirinda Forest Swyrmerton 171 (BM holo., K Z iso ). Evergreen tree, 6-35(-40) m tall (rarely shorter), d.b.h. 20-100 cm; bark smooth; wood white, hard. Branches with conspicuous lenticels, with fissured bark; branchlets

137. LOGANTACEAE: 4. Strychnos

77

Figure 137.5. S T R Y C H N O S IN N O C U A : 1 - branch x Vi, 2 - flower x 4; 3 opened corolla x 5; 4 - pistil x 7 ; 5 fruit; 6 - transverse section o f fruit x V2, 7 - seeds x 1; 8 - leaf o f sucker shoot x Vi, 9 - leaf o f branchlet x Vi. 1-4. 7 from Leeuwenberg 7482; 5 & 6 from Leeuwenberg 7586: 8 & 9 from Leeuwenberg 4400. Drawn by Louise van derR iet. (Reproduced and modified with permission from Meded. Landb. Hogesch. Wag. 69 (1): fig. 20.)

gl iorous or sometimes pubescent, grooved when dried. St .pules none. Leaves: petiole glabrous or sometimes pu­ bescent 0.2-0.5 cm long, smooth; blade paler beneath, le;iihery (not thick), narrowly elliptic, oblong, or some­ times ovate or narrowly ovate, 4-11.5 x 1.5-5 cm, ac .iminate, cuneate or rounded at base or decurrent into th. petiole, glabrous or sometimes pubescent on both si>les; one pair of distinct secondary veins from about 1 c r : above the base, and a faint submarginal pair from the base; tertiary venation spreading, not very prominent. Infh nescence axillary and terminal, dense, much shorter th ir the leaves; peduncle usually short. Flowers 4-5m nous, varying even in a single inflorescence, sessile. St pfrls pale green, united at the very base, broadly ovate or ocular, 1.5-1.8 x 1.5—1.8 mm. Corolla in the mature bt :1 2-2.7 times as long as the calyx, 3.5-4 mm long, and ro inded at the apex, cream, yellow, or green, glabrous

outside, densely villose inside at base of lobes; tube campanulate, 0.9-1.3 (-l.5) times as long as the calyx, 0.6-1.2 times as long as the lobes, 1.5-2.2 mm long; lobes triangular or ovate, 1.5-2.5 x 1.1-1.8 mm, acute at apex, often slightly widened above the base, spreading or some­ times recurved. Stamens just exserted, inserted at twothirds from the base of the corolla tube; anthers bearded at base. Pistil glabrous, 2.3-3 mm long; ovary 2-celled. Fruit small, subglobose, 1-2 cm in diameter, yellow or orange, with thin wall. Seeds 1(—2), sub-ellipsoid, usually flat­ tened at one side, 1—1.2 x 0.8-1 x 0.5-0.6 cm, smooth, glabrous, not grooved, very minutely pitted. Afromontane and transitional rain forest and gallery forest; 1250-1900(-2100) m. IL KF GG SD BA; south to Angola and S Africa, also Comoro Islands. Friis et al 1911, 2331; Mesfin T. 4694.

78

137. LOGANIACEAE 4. Strychnos

4. S. spinosa Lam. (1794) type: Madagascar. Herb. Lamarck s.n. (P-LA, holo.). S. lokua A. Rich. (1851) - type: TU, Takazze River valley, Ouartin Dillon et Petit 412 (P holo.). Deciduous shrub or small tree, (0.3-)l-6(-10) tn tall, sometimes flowering on one-year shoots on old fire-cut stumps, d.b.h. 4-15 cm: bark shallowly fissured, not or sometimes thinly corky, often scaly. Branches sometimes deeply ringed at nodes, without conspicuous lenticels. of­ ten with recurv ed or straight spines: branchlets glabrous or pubescent, sometimes terminating in a straight spine. Leaves on main axis sometimes in whorls of 3; petiole 0.2-1 cm long, glabrous or pubescent: blade leathery, young ones thinner and papery when dried, very variable in shape and size, orbicular, elliptic, narrowly elliptic, ovate, or obovate. 1.4—9.5(—13.5) x 1.2-7.5 cm, emarginate, rounded to acute and often apiculate, sometimes acu­ minate at apex, cuneate. less often rounded, or sometimes on main axis subcordate at base, often with hair pockets in the angles of the main veins beneath, glabrous or pu­ bescent on both sides; 1-3 pairs of distinct secondary veins from above the base, curved along the margin In­ florescence terminal, seemingly umbellate, often con­ gested. Flowers 5-merous. Sepals pale green, united at base, narrowly triangular to linear, 1.5-6 x (0.3—)0.5—1.2

mm. elongate at anthesis, acuminate or awl-shaped at apex. Corolla (3.8-)4-5(-6) mm long, 0.8-2.7 times as long as calyx, pale green, greenish, or less often white, cream or yellow, glabrous or sparsely pubescent outside, with a narrow, entire, white-penicellate corona inside al the mouth of the tube; tube (2.2-)2.5-3(—L4) x 2-3(-3.5) mm. 1.2-2.7 times as long as the lobes, urceolate or campanulate, often somewhat contracted at the throat, lobes triangular. 1.2-2 x 1-1.5 mm. acute, ereci or suberect. Stamens included; filaments inserted at 0.4-0.8 mm from the base of the corolla; anthers ciliate with villose or sometimes pilose hairs all around by which jjhey stick together Pistil 1.8—2.4(—3) mm long. Mature fruit globose, hard, yellow, resembling an orange or a grape­ fruit. 7-11 (—15) cm in diameter, with about 10-100 seeds; wall rather thick, pulp yellow, edible. Seed obliquely ovate or elliptic, flattened, more or less planoconvex, usually ir­ regularly curved. 11-23 x6-18 x 2-5 mm, very shortly pu­ bescent. Woodland and gallery forest; 600-650 TU IL; irom Senegal to Ethiopia and south to Cape Province. Friis et al. 2571, 7324; W. de Wilde 10747. The wood is good for making furniture and boxes. Leaves are used as fodder and dry fruit shell as a local mu­ sical instrument All parts are used in local medicine

79

138. O LEACEAE by P.S.Green* Ti mil. Oleaceae in FI. Trop. E. Afr. 31 pp. (1952); Cufodontis. Enum. 1: 669-674 (1960); Kupicha, Oleaceae. in FI Zlrnb. 7(1): 300-327 (1983).

T :es. shrubs or woody climbers, evergreen or deciduous. Leaves simple, 3-foliolate or imparipinnate, oppo­ site or rarely alternate, exstipulate. Inflorescences terminal or axillary-, cymose, paniculate, dichasial. subur bellate or clustered. Flowers bisexual, unisexual, heterostylous in some genera. Calyx small, usually 4- abed, rarely absent. Corolla regular, usually 4-lobed. to 10-lobed in Jasminum, sympetalous, divided to the base in some species of Chionanthus, or absent. Stamens 2 or rarely 4, epipetalous, the connective often extei ded into a small apical appendage; anthers dehiscing longitudinally. Ovary small, superior, 2-celled. each with (l-)2(-several) apical or basal ovules; style terminal, stigma 2-lobed or subcapitate. Fruit a drupe, berry, ca :isule or samara. C osmopolitan. except in arctic areas, with about 28 genera and 900 species, concentrated in E and SE Asia. 4 i jnera and 11 species recorded as indigenous in die Flora area, and a few species of Jasminum introduced as garden ornamentals. K< >. to genera 1 -1 2. -

Leaves simple. 2 Leaves compound. 4 Trees; fruit a drupe. 3 Woody climbers; fruit a paired berry, sometimes single by abortion. 4. Jasminum 3. Corollas with a distinct tube. 1. Olea - Corollas split to the base. 2. Chionanthus 4. Trees; fruit a woody capsule. 3. Schrebera - Shrubs or woody climbers: fmit a paired berry. sometimes single by abortion. 4. Jasminum

1. OLEAL. (1753). Em rgreen trees. Leaves opposite, simple, entire (outside the Flora area sometimes toothed), usually leathery'. Inflo­ rescences terminal or axillary', cvmose-paniculate or dichasial. Flowers usually bisexual. Calyx small, cam­ p a n i l e , with 4 shallow lobes. Corolla united. 4-lobed, ind iplicate-valvate. Stamens 2; Filaments attached near the lop of the corolla tube, exserted at anthesis; anthers rou ided. oblong. Ovary ovoid to conical, 2-locular. with 2 pendulous ovules per locule. or vestigial in male flowers; style capitate; stigma 2-lobed. Fmit a dnipe with hard endoearp. About 30 species, mostly tropical, in the wanner parts of the Old World: from southern Europe and Africa thro.igh tropical Asia to China, New Caledonia and Aus­ tralia: 3 species in the Flora area. 1. Leaves covered below with numerous, overlap­ ping. peltate scales; inflorescences axillary’. 1. O. europaea subsp. cuspidata - ..eaves not so covered; inflorescences terminal. 2

* Hi: Herbarium. Royal Botanic Gardens, Kew. Richmond Surrey TW9 :-AB, England.

2. Leaves 2 to 2.5 limes as long as broad. 6-12 x 2.5-6 cm, apices acute to obtuse, sometimes veiy slightly and shortly acuminate, petioles 0.5-1.5 cm long. 2. O. capensis subsp. m acrocarpa - Leaves about 3 times as long as broad. 8-20 x 2.5-6 cm, apices acuminate, petioles 2-4 cm long. 3. O. welwitschii 1. O. europaea L. subsp. cuspidata (Wall, ex G.Don) Cif. L ’Olivicoltore 19(5): 96 (1942); O. cuspidata Wall, ex G.Don (1837) - type: India. Walhch 2817. O. africana Mill. (1768); O. europaea subsp. africana (Mill.) P.S.Green in Kew Bull. 34:69 (1979). O. chrysophylla Lam. (1791). O. somaliensis Bak. (1902). O. schimperi Gand. (1918) - type: TU. Gennia, Schimper II: 945 (BM K iso.). O. subtrinervata Chiov. (1940) - type: HA. Mt Eliot. Reghini s.n. (?FT holo.). Tree, rarely a shrub. 5-15 m tall, rarely more, bark dark grey, lightly Fissured, fluted with age. Leaves leathery : petiole 0.2-1 cm long; blade nanowly elliptic to elliptic or somewhat lanceolate, (2-)4-7(-9) x (0.8—) 1—1.5(—2) cm base acute, apex acute, rarely slightly obtuse, cuspidate, dark green, somewhat glossy above, pale grey-green below, covered with small overlapping peltate scales, venation somewhat obscure. Inflorescence axillary, cymose-paniculate, 2-4 cm long, many-flowered. Calyx campanulate, 1-1.5 mm long, obscurely 4-toothed. Corolla white to pale-green, globose in bud; tube c 0.5 nun long; lobes 4, broadly triangular-elliptic, c 2 mm long, spreading. Stamens exserted. anthers broadly ellip­ soid. 1-1.5 mm long. Ovary' bottle-shaped, c 1 mm long. Dnipe ovoid-ellipsoid. 5-7 nun long, endoearp dry', dark purple-black when ripe. Fig. 138.1. Evergreen Juniperus - Podocarpus forest: (1250-) 1700-2700(-3000) m. AF EW TU GD WU SU KF GG SD BA HA: Sudan. Somalia, south through East tropical

138. OLEACEAE: l.O lea

80

Figure 138.1. OLEA EU RO PAEA subsp. CUSPID A T A \ 1 - flowering branch x 1 , 2 - leaf underside x 1 ; 3 flower bud x 4; 4 & 5 - flower x 4; 6 corolla and stamens x 4; 7 gynoecium and calyx x 4; 8 — gynoecium x 8; 9 fruit x 1. All from Bally 976. Drawn by D.R. Thomp­ son* (Reproduced with permission from FI. Trop. E. A /r Oleaceae: fig. 2 as O. chrysophylla.)

3XR.T

Africa to South Africa; Arabia, the Himalayas, SW Asia. Zinna Abebe 545, Ash 97; Burger 1643. An important species in evergreen montane scrub, often kept in fields and around churches when the forest is cut. The fruits are edible, and sometimes used to extract oil, though this practice is disappearing with the availabil­ ity of processed edible oils. Leaves, tw igs and wood are used to fumigate pots for milk, TELLA and TEJ, twigs also as toothbrushes. The wood is hard, polishes well and lias many uses, including carving. Subsp. europaea with drupes 2-4 cm long, the source o f OLIVE OIL, is native to the Mediterranean region.

2. O. capensis L. subsp. macrocarpa (C. H. Wright) Verde, in Bothalia 6: 590 (1956); O. macrocarpa C. H. Wright (1909) - type: Grenfell 869 (K lecto.). Olea hochstetteri Bak. (1902); O. capensis subsp. hochstetteri (Bak.) Friis & P.S. Green in Kew Bull. 41: 36 (1986) - type: GD, Mt Aber, Schimper II: 871 (K holo., BM iso ). O. laurifolia sensu Richard (1850), non Lam Evergreen tree 3-25 m tall, bark grey, generally somewhat smooth. Leaves leatheiy ; petioles 0.5-1.5 cm long, blade narrowly ovate to broadly lanceolate, elliptic or oblong, 6-12 x 2.5-6 cm. base obtuse to acute, apex acute to ob-

138. OLEACEAE: 1. Olea, 2. Chionanthus

tu a, sometimes very slightly and shortly acuminate, m cronulate. dark green, somewhat glossy above, pale gr en below, with scattered peltate scales, especially belo' . primary veins generally visible on both surfaces, sli kly raised, usually 6 or 7 on each side of the midrib. Ire Iore see nee terminal, paniculate, many-flowered, 5-8 (- 2) cm long. Calyx cupulate, c 1 nun long, more or less en;re to slightly 4-toothed. Corolla white or creamco >ured, globose in bud; tube c 1 nun long; lobes broadly in; ugular-elliptic, c 2.5 mm long, spreading. Stamens ex: .‘rted, anthers broadly ellipsoid, c 2 mm long. Ovary bo le-shaped. c 1 mm long. Dmpe ovoid to ellipsoid, 10-14 :ni 1 long, pericarp dry. dark, purple to black when ripe. Evergreen Juniperus or Podocarpus forest; 1350— 29 D(—3200) m. GD SU AR IL KF SD BA; W and C Afric , from Guinea Republic to Sudan and south to the No rthem Province of South Africa. C.haffey 1183; Friis et al. 1089; Gilbert 1758. Often an indicator of previously more extensive for­ est An important forest tree providing an attractive timber wluch polishes well. •iubsp. capensis, with petioles 10-20 cm long, is con­ fined to South Africa.

3. O. welwitschii (Knobl.) Gilg & Schellenb. (1913); Mayapea wehvitschii Knobl. (1893); Olea apensis L. subsp. welwitschii (Knobl.) Friis & P.S.

- Leaves pinnate or trifoliolate, occasionally some simple. 6. J. grandiflorum subsp. floribundum 6. Lateral petiolules 1—3(—5) mm long; corolla tube 15-20 mm long. 4. J. abyssinicum - Lateral petiolules (3-)5-8 mm long; corolla tube 20-27 mm long. 5. J. fluminense subsp. gratissimum 1. J. dichotomum Vahl (1804) - tvpe: W Africa, Thonning s.n. (C holo.). J. mathildae Chiov. (1911) - types: TU, “Scire” and GD, “Tzellemti”, Chiovenda 506 (FT) & 3196 (?FT not seen).

Scandent shnib. Leaves dark green, simple, ovate to broadly lanceolate, (4—)5—10(—11) x (2-)3-6(-6.5) cm, base obtuse to rounded, apex acute, shortly acuminate, gla­ brous, 2 basal veins becoming submarginal, without domatia. Inflorescence terminal or axillary, more or less densely corymbose, usually many-flowered, flowers fra­ grant, pedicels 1-3 mm long. Calyx glabrous or sparsely puberulous, tube c 2 mm long; lobes 0.5-2 mm long. Co­ rolla tube 1.8—2.5 mm long, purple on the outside, other­ wise white; lobes 6-9, oblong-elliptic, 0.7-1.6 mm iong. Berries ellipsoid, 8-10 mm long. Forest, forest margins and bus hiand; no altitude given. TU GD; Sudan. E Africa, Rwanda, Burundi, W Africa (from Mali to Cameroun), Rep. D6m du Congo and Zam­ bia. Chiovenda 506. 2. J. schimperi Vatke (1876) - type: GD, R. Repp, Schimper (1863) 1112 (K BM iso.). J. eminii Gilg. Woody climber or scrambler, stems up to 5 m long. Leaves simple, ovate to narrowly lanceolate, (1.5-)2-6(-7) x (1-) 1.5—3(—4) cm, base rounded, apex acute, acuminate, pilose, especially below and on the midrib, rarely glabrous, 2-3 primary veins on each side of the midrib, pilose domatia in their axils below. Inflorescence terminal on side shoots, pilose, cymose 3- to several-flowered more or less compact, pedicels 1-10 mm long. Calyx tube 2-2.5 mm long, lobes filiform, 1—3(—5) mm long, often slightly curled outwards, pilose to rarely glabrous. Corolla tube 15-25 mm long, often tinged purple on the outside, lobes 7-9, narrowly oblong-elliptic, 10-12 mm long, white. Berries globose, c 10 mm in diameter. Gallery forest and secondary scrub, often near streams; 1200-2050 m GD SU KF GG SD HA; Sudan. E Africa, Rwanda, Burundi and Rep. Dem du Congo (often under the name*/, eminii Gilg.) Burger 2868; Gilbert et al 7753; Mooney 9004.

3. J. streptopus E. Mey. (1838) type: South Africa, Natal, Drege s.n. (B holo. destroyed?). ? J bogosense Becc. ex Martelli (1886) - type: EW, Keren,Beccari PI. Bog. 184 (?FI, TFT holo., not seen). ? J. basilei Chiov. 1930 - type: HA, Lago Hida. Duke ofAbruzzi 310 & 316 (?TO syn., not seen). Woody climber, or scrambling shrub. Leaves simple, ellip­ tic, sometimes somewhat narrowly ovate, 2-5 x 1.2-2.7 cm, base acute to obtuse, apex acute, acuminate, pilose, especially below and on the midrib, usually 3 primaiy veins on each side of the midrib, pilose domatia in their axils below. Inflorescence terminal on side shoots, pilose, open, l-3(-5)-flowered, flowers sweetly fragrant; pedi­ cels 3-10 mm long. Calyx tube 1.5-2 mm long; lobes slen­ der. filiform. (3-)4-9 mm long, pilose, rarely glabrous. Corolla tube 10-20 mm long; lobes narrowly oblongelliptic, 8-18 mm long. Berries globose, c 10 mm in dia­ meter.

138. OLEACEAE: 4. Jasminum

85

Figure 138.4. JA S M IN U M A B Y S S IN I­ CUM: 1 - habit x 2/ 3; 2 - detail o f lower leaf surface showing acrodomatium; 3 flower x 2; 4 - calyx x 2; 5 - stamen x 4; 6 - gynoecium x 4; 7 - long-styled flower, semidiagrarnmatic, x 1 ; 8 - short-styled flower, semidiagrarnmatic, x 1; 9 - part o f infructescence x 2/ 3. 1 & 2 from Pawek 5619; 3 -6 from M uller 1593; 7 from Lawton 462; 8 from Torre & Correia 15654; 9 from Fanshawe 7356. Drawn by F. K. Kupicha. (Modified and reproduced with permission from FI. Zamb. 7(1): O leaceae, Tab. 70.)

4nogeissus woodland, thickets in grassland and evergn ::n bushland; 550-1700 m. EW? TU GJ SU IL GG SD; Su lan, Angola, Rwanda, Burundi, Uganda and south to So iLh Africa (Natal). Ash 903; Gilbert & Jefford 4520; Thunn & Hunde 4044. 4. J. abyssinicum Hochst. ex DC. (1844) - type: TU, near Dagabur, Schimper I: 1699 (BM K iso.). J. abyssinicum var. amplifolium Fiori in Nuovo iorn. Bot. Ital. 11,47: 34 (1940) - type: Ethiopia, . unma a Malco, Saccardo s.n. (?FT holo., not seen).

or broadly elliptic, (3-)5-8(-10) x (1 .5 -)2 .8 ^ (-5 ) cm, lateral leaflets somewhat smaller, base acute to obtuse, apex acuminate, with 4-8, usually pilose domatia below in the axils of the midrib and primaiy veins, otherwise glabrous. Inflorescence terminal, also terminating side shoots, finely puberulous, sometimes glabrous, cymosecoiymbose, many-flowered, flowers sweetly fragrant. Calyx broadly campanulate, 2.5 mm long, with 4 teeth, 0.1-1 mm long. Corolla white, exterior flushed red or pink; tube 15-20 mm long, lobes 4-10 x 3-6 mm. Berries globose, c 8 mm in diameter, purple-black. Fig 138.4.

Woody climber. Leaves trifoliolate, petiole 0.5-3 cm lorn; terminal petiolule 0.5-2 cm long, lateral petiolules 0.1 n.3(-0.5) cm long, terminal leaflet ovate to elliptic

Open degraded montane woodland and scrub, forest undergrowth, often near streams; also fences around home gardens and climbing over houses; 1700-2800 m.

86

138. OLEACEAE: 4. Jasminum

TU GD GJ SU AR WG IL KF SD BA HA; R£p. Dem. du Congo, Rwanda, Burundi and Uganda, south to South Af­ rica (Natal, Transvaal). Burger 2464; Friis et al. 1869; de Wilde 5943. The roots are used to dress wounds in animals. 5. J. fluminense Veil, subsp. gratissimum (Defiers) P.S.Green in Kew Bull. 41: 417 (1986); J. gratissimum Defiers (1889); J. abyssinicum var. gratissimum (Defiers) Di Capua in Ann. R. 1st. Bot. Roma 8: 207 (1904) - type: Yemen, Defers 212 (P lecto.). J. fluminense subsp. mauritianum sensu Cufodontis in Bull. Jard. Bot. Etat. 30: 672 (1960), non (Bojer ex DC.) Turrill.

J. tettense sensu Marteili (1886), non Klotzch. Woody climber. Leaves trifoliolate, petiole 0.7-3 cm long, terminal petiolule 1-1.5 cm long, lateral petiolules 0.3-0.5(-0.8) cm long, terminal leaflet ovate to rarely nar­ rowly lanceolate, (4-)6-8 x (1.5-)2.5-4 cm, lateral leaf­ lets somewhat smaller, base obtuse to rounded-truncate, apex acute, long acuminate, domatia below in the axils of primary veins and midrib, glabrous to lightly tomentose, especially below. Inflorescence terminal, cymose-pam­ culate, slightly corymbose, many-flowered, smaller inflo­ rescences in the leaf-axils below the main one, shortly pilose to tomentose, flowers sweetly fragrant. Calyx 2-2.5 mm long, teeth 5, almost imperceptible to almost 0.5 mm long, puberulous to lightly tomentose. Corolla white to cream, tube slender, 20-27 mm long, lobes 5 or 6, oblong, 8—10(—14) x 4-5 mm. Berries broadly ellipsoid, 6-8 mm long. Rocky, seasonally dry river beds, and Acacia bush­ land: 1300-2000 m. EW AF SU KF HA; Sudan and Ye­ men. Burger 2547; Gilbert 5110A; Mooney 8599. Subsp. fluminense, with acute to obtuse leaflets, is widely distributed in tropical East Africa,

6. J. grandiflorum L. subsp. floribundum (R.Br. ex Fresen.) P.S.Green in Kew Bull. 41: 414 (1986); J. floribundum R.Br. ex Fresen (1837) - type: Ethiopia, Ruppel s.n. (FR? holo.). J. steudneri Schweinf. ex Bak. (1902); J. flori­ bundum var. steudneri (Schweinf. ex Bak.) Gilg & Schellenberg in B ot Jahrb. Syst. 51; 83 (1913).

J. floribundum forma decipiens Di Capua in Ann. R. 1st. Bot. Roma 8: 208 (1904); J. floribundum var. decipiens (Di Capua) Fiori in Boschi e Plante Legnose. dell Eritrea: 295 (1912) - types: EW, Pappi s.n. and Terracciano & Pappi s.n. (FT syn.). Woody climber. Leaves imparipinnate to trifoliolate, rarely sometimes simple, petiole 0.5—2 cm, lateral petiolules up to 0.5 cm, terminal leaflet ovate to lanceolate, (1.2—)2.5—3(—6) x (0.6-)l-1.5(-2.5) cm lateral leaflets smaller, base obtuse to rounded, apex acute, long acu­ minate, glabrous or lightly pilose, without domatia In­ florescence terminal, cymose, few- to many-flowered, pedicel of central flower shorter than lateral, often more or less sessile, flowers sweetly fragrant. Calyx tube 1.5-2 mm long, lobes filiform, slightly stiff, 4-8 mm long, gla­ brous or puberulous. Corolla red in bud and on the outside at anthesis, otherwise white; tube 14-20 mm long, lobes 10-15 mm long, 6-8 mm broad. Berries ellipsoid black, c 8 mm long. Dry rocky savanna and scrub; 1600-2800 m. EW TU GD GJ WU SU AR GG SD HA; Sudan, Somalia, Uganda, and Kenya; Saudi Arabia, Oman, Yemen. Getachew A. & Gilbert 648; Chaffey 673; de Wilde 7090. Subsp. grandiflorum, with leaves comprising 7-11 leaflets, occurs in India. 7. J. stans Pax (1907) type: SU, Akaki valley, Rosen s.n. (BRSL? holo.). J. humile sensu Engl. (1906), non L. Shrub to 3 m tall, more or less erect. Leaves alternate, unparipinnate, leaflets (3-)5 or 7(-9), petiole 0.5-1 cm long, lateral leaflets sessile, rachis very slightly winged, termi­ nal leaflet obovoid to elliptic, 1-2 x 0.4-1.2 cm lateral leaflets smaller, base obtuse to acute, apex rounded, some­ times with a very small mucro, glabrous. Infloresence ter­ minal on side shoots, l(-3)-flowered, flowers pendant, without scent, pedicels 5-15 mm long. Calyx tube 2-3 mm long; lobes subulate, 1-2 mm long, glabrous. Corolla yel­ low, tube 15-23 mm long, widening very slightly in the upper part; lobes 8-10 x 4-5 mm. Berries broadly ellip­ soid, 8-10 mm long. Open Juniperus - Erica scrub; 2400-2900 m. SU AR; not known elsewhere. Friis et al. 1177; Gilbert & Tewolde B.G.E. 3242; Mooney 4823. A member of the small, mainly temperate Section Alternifolia.

87

139. A PO C Y N A C EA E by A. J. M. Leeuwenberg* iuber, Apocynaceae in FI. W. Trop. Afr. 51-80 (1963); Cufodontis, Enum:. 683-692 (1965); Leeuwenberg, Kupichaet IL, Apocynaceae, in FI. Zamb. 7(2): 395-503 (1985).

1 rees, shrubs, climbers or sometimes herbs, mostly with white latex. Leaves usually opposite, less often in vshorls of 3 or alternate, simple, pinnately veined. Stipules usually absent or intrapetiolar. Flowers bisexual, -merous, mostly actinomorphic. Corolla mostly coloured, sympetalous, variously shaped; lobes twisted in t ud. Anthers dehiscent throughout by a longitudinal slit, often coherent with the pistil head. Pistil simple; ova­ ries one, entirely or partly 2-celled, or two and 1-celled and then also apically connected by the style; pistil head nasally or laterally stigmatic (receptive) and bearing abilobed, stigmoid apex. Fruit a beriy or a capsule, syn- or apocarpous. Seeds in capsules mostly with l(-2 ) apical tufts of hairs (comas). 165 genera and 1900 species, mainly tropical, a few temperate. 10 genera with 1 species each are indigenous tor naturalized in the Flora area, another 4 species from 4 different genera are cultivated. Key to genera 1. Leaves alternate. 2 - Leaves opposite or whorled. 4 2 Plants not succulent; leaves narrowly oblong, up to 1 cm wide; flowers yellow; cultivated. 12. Thevetia - Plants succulent; leaves variously shaped; flowers usually pink or white. 3 3 Candelabrum-shaped tree; leaves narrowly elliptic, acuminate; stamens deeply included; cultivated. 14. Plumeria - Low, succulent tree or shrub with thick foot; leaves linear to obovate, acute to emarginate; stamens barely included or exserted. 2. Adenium 4 Herbs or low shrubs. 5 - Woody plants, trees, shrubs or lianas 6 5 Creepmg herb; flowers solitary in leaf axis; longpedicellate; stamens with terminal appendage. 13. Vinca - Erect herb; inflorescence terminal though appar­ ently lateral; stamens without appendage. 5. C atharanthus i. Plants armed with straight spines. 4. Carissa - Plants unarmed. 7 '' Liana without tendrils; petiole with two pairs of glands on adaxial side; flowers small; corolla yel­ low-green, tube 2.5-4 mm long; fruits long, slen­ der follicles. 7. Oncinotis - Liana, shrub or tree; petiole eglandular; flowers large; corolla white or cream with tailed lobes, tube at least 5 mm long;fruits variously shaped. 8 Corolla lobes with long, slender tails; follicles with terminal knob. 9. Strophanthus - Corolla lobes not tailed; fruits, if follicular, without a terminal knob. 9 * Leaves whorled. 10 Leaves opposite. 11

J________ ageningen Branch, N a t Herb. Netherlands, P.O. Box 8010,6700 ED W ageningen, The Netherlands

10. Leaves in whorls of at least 4, pale glaucous be­ neath; corolla with cylindrical tube, 5-11 mm long; fruit of 2 very slender follicles. 3. Aistonia - Leaves in whorls of 3, not much paler beneath; co­ rolla with funnel-shaped tube, 12-24 mm long; fmit a thick syncarpous follicle; cultivated. 11. Nerium 11. Leaves not or only partly hairy; stamens included or barely exserted; fruits berry-like. 12 - Leaves pubescent all over on both sides; stamens well exserted; fruits follicular. 10. W rightia 12. Lianas with curled tendrils, or sometimes small shrubs; stamens included; fmit at least 25 mm long, with 2 to many seeds, if few-seeded, then vemicose. 13 - Trees or shrubs without tendrils; stamens barely exserted; fruit less than 25 mm long, with 1-2 seeds, not vemicose. 1. Acokanthera 13. Corolla tube 5.5-12 mm long; fmit vemicose. 6. Landolphia - Corolla tube 16-34 mm long; fruit not vemicose. 8. Saba 1. ACOKANTHERA G. Don (1837) Kupicha, Kew Bull. 37: 41-67 (1982). Trees or shrubs. Leaves leathery. Stipules absent. Flowers in dense, axillary cymes or clusters, sweetly scented. Se­ pals free, ovate to very narrowly elliptic. Corolla white, tinged with red, hypocrateriform; tube cylindrical, pilose inside in the upper half, outside glabrous or pubescent, lobes overlapping to the left, shorter than the tube. Sta­ mens inserted near the corolla mouth; anthers subsessile. Ovary syncarpous with one ovule in each cell. Fruit a globose or ellipsoid berry containing 1 or 2 seeds. Seed plano-convex, ellipsoid, smooth and glabrous. 5 species in tropical Africa; 1 in the Flora area. A. schimperi (A. DC.) Schweinf. (1891); Carissaschimperi A. DC. (1844); A abyssinica K. Schum. (1895) - type: TU, Ado, near Shahagenni, Schimper 254 (P holo., BM BR C GE FI FI-W K iso.).

88

139. APOCYNACEAE: 1. Acokanthera, 2. Adenium, 3 Alstonia

clear or white; tubers rhizomatous or carrot-like. Lea\ es alternate, confined to the apices of branchlets, linear to obovate, cuneate at base; stipules minute or absent, in­ florescence cymose; peduncle very short or absent. Flowers large, showy, slightly zygomorphic. Sepals united at the base, narrowly oblong to narrowly ovate. Corolla ji fundibuliform to hypocrateriform, much widened at the throat, outside more or less pubescent, inside glabrous or pubescent to strigose; lobes obovate or narrowly so, over­ lapping to the right in bud; between the lobes a small scale at the base united to them. Stamens inserted at the apex of the narrow basal part of the corolla tube; anthers narrowly triangular, with a long appendage at the apex. Pistil: car­ pels separate; style cleft at base. Fruit of 2 spreading or re­ curved follicles, united at the base, many-seeded. Seed oblong, with 2 comas. 5 species in Africa, Yemen, Saudi Arabia and Oman; 1 in the Flora area. A. obesum (Forssk.) Roem. & Schult. (1819); Nerium obesum Forssk. (1775) - type: Yemen, Melham. Forsskdl Herb. 235 (C holo.). A. honghel A. DC. (1844). A. speciosum Fenzl (1865). A. somalense Balf. f (1888). A. cogtaneum Stapf (1902). A. dejlersii Schweinf. ex Lewin var. africana A. tricholepis Chiov. (1932). Markgr. in Notizbl. Bot. Gart. Berlin Nr. 77, Bd. 8:464 A. somalense var. caudatipetalum Chiov. in FI. (1923) - type: Ethiopia, Ghinda, near R. Telekit, Somala; 217 (1929). Schweinfurth & Riva 2183 (K lecto.. P isolecto.). A. somalense var. crispum Chiov. in FI. Somala; Shrub or tree, 1-9 m tall. Bark soft, brown; slash cream 217(1929). with fine orange streaks. Young branches glabrous or pu­ Leaves sessile or subsessile, flat or curled; blade 3-12 bescent, conspicuously angled and ribbed, especially (-17) x (0.2-)0.5-5 cm, 1.4-19(-55) times as long as wide, when dried. Leaves: petiole 1—6(—9) mm long; blade ellip­ acute to emarginale and apicuiate to mucronate at the apex, tic. 2.2-10 x 1.5— 4.5 cm, acute, cuspidate or obtuse at often slightly glaucous. Sepals green to white, suffused apex, with a hard mucro, cuneate or rounded at base, with pink, 5-11 x 2-3 mm. Corolla pink to red, tube red­ glabrous, smooth or scabrid. Flowers fragrant. Sepals dish-pink to white, suffused with pink, sometimes red(1—)1.5—2(—2.5) mm long, pubescent or glabrous outside, striped within the throat, 20-45 x 9-17 mm, narrow, basal ciliate. Corolla tube pink or reddish. 8-12.5 mm long, part 9-17 x 4-7 mm; lobes very pale pink to pale red in glabrous or very rarely pubescent inside, lobes white. Sta­ centre, much darker pink to crimson along the margins, mens barely exserted. anthers 1-1.5 mm long. Ovary gla­ 9-28 x 5-25 mm. Stamens barely included or exserted; an­ brous or occasionally pubescent all over or in a ring at the ther appendages densely hairy. Fig. 139.2. apex. Fruit ellipsoid, when dried 10-24 mm long, purple. Semi-desert and dry bushland, on sandy and rocky Seeds up to 13 mm long. Fig. 139.1. soils; 300-1400 m. KF GG SD BA HA; Somalia, Tanza­ Woodland and degraded Podocarpus - Juniperus nia, Senegal, Socotra; Saudi Arabia. Berger 3366; forest, 800-2100 m. EW TU GD GJ WU SU AR SD BA Ensermu.K. & Dessalegn D. 3713; Mesfin T. & TeM'olde HA; Somalia. Djibouti. Kenya, Uganda. Rwanda, E B.G.E. 2813. Congo, N Tanzania; Yemen. Berger 2104; Ryding &

Figure 139.1 A C O K AN TH ER A S C H IM P E R I I - flowering branch x 14; 2 - flower x 2; 3 - fruit x 1. No specimens citcd in orig. publ. Drawn by D. Troupin. (Reproduced with permission from Flore des plantes ligrieuses du Rwanda, fig. 35 p.p.)

Sileshi N. 1974; Mooney 9818. The seed is very hard and has been used for making buttons. The flesh of the fruit is edible, enjoyed by shep­ herd boys. However, the latex is poisonous and has been used to make arrow poison. 2.

ADENIUM R oem . & S ch u lt. (1819)

Plaizier.A/ecfec/. Landb. H ogesch. Wag. 80(12): 1-26(1980).

Succulent shrubs or trees, 0.2-5 m tall; trunk mostly up to 1 m in diameter, bark pale grey or nearly so, smooth; latex

3. ALSTONIA R. Br. (1809) Sidiyasa, Blumea suppl. 11: 1-230. Shrubs or trees. Leaves whorled. Inflorescence compound, cymose, terminal. Flowers small. Sepals suborbicular. Co­ rolla tube cylindrical; lobes spreading. Stamens included; anthers ovate or narrowly so. Pistil: carpels separate. Fruit of 2 slender follicles. 43 species from Central America to tropical Africa, and from the Himalayas and China to New South Wales, Australia, 1 in the Flora area.

139. APOCYNACEAE: 2. Adenium, 3. Alstonia, 4. Carissa

89

Figure 139.2. A D E N IU M OBESU M . 1 branch with leaves x 2/ 3; 2 flower and bud x Vy, 3, 4 - leaves x 2/ 3; 5 - opened up flower x Vy, 6 - anthers and pistil x 2; 7 ,8 -c la v u n c u la x 10; 9 - fruit x 2/ 3; 10-s e e d x 2/ 3. 1, 2 from Greenway 15400; 3 from Paulo 1078; 4 from Khattat 47; 5, 6, 7 from Leeuwenberg 10784; 8 from de Wilde 8726; 9, 10 from D ekker 376. Drawn by Juliet Williamson. (Repro­ duced with permission from Meded. Landb. Hogesch. Wag. 80 (12): fig. 3.)

A. t oonei De Wild. (1914) 11 - type: Congo (Kinshasa), Nala, Uele, Boone 2 BR holo.).

Leeuwenberg & van Dilst, Series of revisions of Apocynaceae XLIX, Carissa. Wag. Agric. Univ. Papers 01.1 (2001).

Big :ree, up to 40 m tall; often as non-flowering shrub in busl . trunk up to 1 m in diameter. Leaves in whorls of 4-8 petiole glabrous, (5-) 10-22 mm long; blade pale grey green beneath, obovate or narrowly obovate, 6-20 (-2 - x 2.5-7.5 cm, apex usually obtuse or rounded; sec­ ond; r> veins in 22-50 pairs. Inflorescence manyflovi ::rsd. Sepals green, ovate, 1.5-2.2(-2.6) mm long, obtu ;e or rounded. Corolla cream or pale yellow, pu­ bescent outside; tube 5-11 mm long; lobes 4-^6 mm long, rour :led. Ovary densely pubescent. Follicles 30-52 cm long >.2-0.3 cm in diameter, densely pubescent. Seeds oblong, 4.5-6 mm long; coma 12-20 mm long.

Shrubs, small trees or climbers, with simple or forked spines. Stipules absent. Flowers in terminal corymbs or 1-5-flowered cymes. Sepals free, imbricate. Corolla hypocrateriform; tube cylindrical, hairy within; lobes overlapping either to the left or to the right, of various shapes, much shorter to longer than the tube. Stamens in­ serted at the middle or towards the top of the corolla tube; anthers subsessile, glabrous. Ovary glabrous, syncarpous, 1-4 (rarely more) ovules in each cell; style slender; stigma below or reaching the androecium. Fruit an ellipsoid or ovoid berry, containing (l-)2-8(-m any) seeds. Seed dis­ coid or compressed-ellipsoid, velutinous.

P )rest; 100-1200 m. EL KF; west to Senegal. Friis et al 3S52, 4137. Tesfaye & Okeach 319.

37 area.

4. CARISSA L. (1767)

species in Africa, Asia and Australia; 1 in the Flora

139. APOCYNACEAE: 4. Carissa

Figure 139.3. CARISSA SPIN ARU M 1 - habit x Vr, 2 - branchlets x Vy, 3 flower x 4; 4 - flowers opened up to show anthers and pistil x 4; 5 - fruits x 1; 6 - seed x 5. 1 from Cadet 2606; 2 from W. D e Wilde 5531, 3 from H agos 6; 4 from Hedberg 4798 and H epper 5968; 5 & 6 from Leeuwenberg 10100. D rawn by E. RiemerGerhardt ( I ) and H. de Vries (2 -6 . (Reproduced and modified with per­ mission from Wag. Agric. Univ. Pa­ pers 01(1), figs 8 -1 0 p.p.)

C. spinarum L. (1767) type: India, sine loc., Koenig s.n. (LINN 295.2 lecto.). Antura (without epithet) Forssk. (1775); Carissa edulis Vahl. (1790) - type: Yemen, Mts Hadie, Forsskdl 234 (C holo., BM S iso.). C. tomentosa A. Rich. (1851); C. edulis var. tomentosa (A. Rich.) Stapf in FI. Trop. Afr. 4,1: 90 (1902) - type: TU Mts Tchelikote, IV-V, Quartin Dillon & Petit s.n. (P lecto.). C. richardiana Jaub. & Spach. (1857) - type: TU, near Adua, Schimper 156 (P lecto., BM BP BR FT G-DC GH K L LG M MEL NY P S UPS W WAG Z isolecto.). C. candolleana Jaub. & Spach. (1857) - type: TU, near Adua, Schimper 209 (P lecto., BM BP BR FT G-DC HBG K M MEL W isolecto.).

C. cornifolia Jaub. & Spach. (1857) - type: GD, Senen, Sanfetch Mt, Schimper 1068 (P lecto., BR G K W WAG isolecto ). Richly branched, sometimes creeping shrub or climber. Young branchlets densely pubescent or puberulent, rarely glabrous. Spines simple, very rarely furcate, 0.4-5.5(-7) cm long. Leaves: blade ovate, elliptic, obovate or suboibicular, 2-7 x 1-4.5 cm, acute or obtuse at apex, with or without mucro, cuneate or rounded at base, thinly leathery, glabrous or pubescent on both sides. Flowers sweetly scented. Sepals 2-4 mm long, very nar­ rowly elliptic, awl-shaped, glabrous, pubescent or woolly. Corolla white inside, pink to crimson outside, tube 9.5-20 mm long, glabrous or sparsely pilose outside, pubescent inside in upper half; lobes very narrowly ellip­

139. APOCYNACEAE: 4. Carissa, 5. Catharanthus

91

Figure 139.4. CA TH ARAN TIIU S ROSEUS: 1 - flowering branch x H; 2 - corolla seen from above x '/2; 3 -5 - leaves x 2/ 3; 6 - calyx with pistil base and disc glands x 6; 7 - follicles x 1.2; 8 - seeds showing adaxial and abaxial sides x 8. 3 & 4 from Leeuwenberg 10887; 5, 7 and 8 from Wild 4423; 6 from Hunziker 11. No specimen or artist given for 1 & 2; 3 -8 drawn by Yuen Tan. [Combined and reproduced with permission from FI. Rwanda 3, Fig. 27 (1 & 2) and Meded. Landb. Hogesch. Wag. 81 (9)fig. 1 (3-8)]

tic, o’ -Happing to iiie right. Stamens inserted near apex ot iiie lube, with their apex 1—1.5 nun below mouth; an­ thers subsessile, 1.5-2 mm long. Ovaiy glabrous; style glabrc us; stigma reaching to base of anthers. Fruit 6-11 (-25) ram long, plum-shaped, green, turning red to blue-black, 2-4-seeded. Fig. 139.3. In upen Acacia woodland, often on termite mounds, and in riverine fringing vegetation; (550-) 1000-2500 m AF EW TU GD GJ WU SU AR WG KF GG SD B A HA; very w idely distributed in Africa from Senegal to Soma­ lia, fix m Sudan to the Transvaal and Namibia, and in Madagascar; also in Asia from Yemen to India and Thai­ land, aivd on the islands of the Indian Ocean. Friis et al. 102, 25 34; Mesfin T. & Tw olde B.G.E. 2705. T hi! species does not appear to be as poisonous as many c iherplants in this family. The ripe fruits are widelyeaten t children. Bushes are grazed vigorously. Spiny

stems are used for fencing fields of growing crops. It also makes a good live fence.

5. CATHARANTHUS G. Don (1836) Van B ergen, W ageningen Agric. Univ. P apers 96(3): 1-46

Herbs or low shrubs. Inflorescence terminal, but appar­ ently lateral, 1-2-flowered. Sepals narrowly to very nar­ rowly oblong, small. Corolla hypocrateriform; tube narrow; limb large and showy. Fmit of two follicles. Seed, without coma, rugose. 8 species, 1 restricted to India and Sri Lanka, all the others endemic to Madagascar. Of the latter group C. roseus is cultivated and naturalized all over the tropics, especially along coasts.

92

139. APOCYNACEAE: 5. Catharanthus, 6. Landolphia, 7. Oncinotis

C. roseus (L.) G. Don (1837); Vinca rosea L. (1759); Lochnera rosea (L.) Rchb. (1828. nomen) ex Endl. (1838) - type: Miller, Fig. Beaut. PI. 2; 1 186 (1757); topotype: Miller 1849 (BM). Low shrub, 30-100 cm tall, usually with white latex. Stems often suffused with dark red. Leaves decussate; petiole 3-10 mm long; blade elliptic, obovate or narrowly obovate, 4-9 x 1.5-3 .5 cm, obtuse or acute at apex, some­ times emarginate, inucronate, veins paler. Sepals 3-5 x 1-1.5 mm. Corolla white with a dark pink or pale yellow centre, or pink with a dark pink centre; tube often slightly greenish Follicles c 5 x 0.5 cm, green. Fig. 139.4. Cultivated and naturalized at low altitude. EW SU SD HA. Haugen 1591 \Mitiku T. 21; Mesfm T. & Sebsebe D. 369. Cultivated for its attractive flowers. The species is also the source of alkaloids used in the treatment of leukaemia. 6. LANDOLPHIA Beauv. (1806)* Persoon et al., Wageningen Agric. Univ. Papers 92(2): 1-232(1992). Erect or creeping shrubs or lianas, with curled tendrils. Stipules absent. Inflorescence terminal or axillary, a more or less compound cyme, few- or many-flowered. Sepals united at the base, imbricate. Corolla tube cylindrical, more or less densely hairy or glabrous within; lobes over­ lapping to the left, much shorter to longer than the tube. Stamens inserted in the corolla tube at various levels; anthers mostly glabrous. Ovary glabrous or hairy, syncarpous, unilocular, many ovules on two protruding pari­ etal placentas; style slender or not; stigma at the same level as the anthers or below. Fruit a variously shaped berry containing one to many seeds. Seed irregularly ovoid or ellipsoid, compressed or not. About 60 species in tropical Africa inclusive of Mada­ gascar, 1 species in the Flora area. L. buchananii (Hall.f) Stapf (1902); Clitandra buchananii Hall.f. (1900) - type: Ma­ lawi, Shire Highlands, Buchanan 220 p.p. (BM lecto ). Creeping shrub or liana. Branchlets more or less densely pilose or glabrous. Leaves herbaceous or leathery; blade ovate, elliptic, or obovate, or narrowly so, 2-14.5 x 0.8-5.4 cm, apex obtuse to acuminate (then acumen ob­ tuse), rounded al the base, glabrous on both sides or sparsely pilose at the base of the midrib and lower second­ ary veins beneath. Flowers fragrant. Sepals 1.2-3.1 nun long, ovate or broadly ovate, glabrous to pilose outside, glabrous inside, rounded. Corolla white, cream or yellow, with an often greenish tube, occasionally tinged pink or reddish; 5.5-12 mm long, glabrous or pilose outside, more or less densely pilose inside below the anthers; lobes nar­ rowly ovate or narrowly elliptic. Stamens inserted below the middle of the tube, 1.3-6 mm included; anthers * by J. G. M. Persoon d o Department o f Plant Taxonomy, P.O. Box 8010, 6700 ED Wageningen, The Netherlands

1.2-2.3 mm long. Ovary glabrous or with some hairs; pistil head reaching anthers. Fruit 2.5-6 cm long, globose or pyriform, brown, orange, green or yellow-green, with grey or light brown, verrucose spots, edible, with 2-20, mi­ nutely pitted seeds. Seed with much endosperm 7-18 x 5-10.5 x 3-8.5 mm. Fig. 139.5. Forests, gallery forests, woodland, often on or near rocks; 1300-1550 m. SU AR WG IL KF SD B A (and prob­ ably GG); widely distributed in Africa from Ethiopia to southern Mozambique, also in eastern Rep. Dem. du Congo, Angola and Cameroun. Friis et al. 223; Mesfin T. & Sebsebe D. 1358; Mooney 6731.

7. ONCINOTIS Benth. (1849) De Kruif, Agric. Univ. Wageningen Papers 85(2): 5—45 (1985). Climbing shrubs or lianas with white latex. Branches un­ armed. Tendrils and stipules absent. Inflorescences termi­ nal or axillary, thyrsoid, rather lax, many-flowered. Flowers small, fragrant. Corolla inside with tufts of stiff hairs alternating with the stamens and 5 corona scales at the mouth alternating with the petals, lobes overlapping to the right. Stamens included, joined into a cone. Pistil: ovary half-inferior, carpels separate, surrounded by disk. Fruits pendulous, of two follicles united at the extreme base, fusiform to narrowly cylindrical, manv-seeded. Seed truncate, with a coma at the apex, directed towards the apex of the fruit. 7 species in tropical Africa including Madagascar. 1 in the Flora area. O. tenuiloba Stapf (l%9&) -type: Congo. Equateur. nearLikasa, Dewevre 883 (BR holo ). O. inandensis Wood & Evans (1899). O. natalensis Stapf (1907). O. chirindica S. Moore (1911). O. oblanceolata Engl. (1910). Motandra erlangeri K. Schum. (1903). Plant climbing, 1.8-30 in high; bark thickly corky, inner bark with rope-like fibres. Brandies with many whitish lenticels; branchlets puberulous or pubescent. Leaves: pet­ iole 5-12 nun long, with 2 pairs of glands along the adaxial side; blade obovate or narrowly so, 4-14 x 1-5 cm, acu­ minate or rarely emarginate at the apex, cuneate at the base, membranous to papery when dried, glabrous or rarely pubescent; domatia consisting of pits with a dense tuft of hispid hairs along the margia Sepals olive-green, elliptic to triangular or nearly so, 1.2-3.2 x 0 .8 -1.5 mm, rusty-brown-pubescent or -puberulous outside. Corolla pale yellowish-green; tube 2.5-4 nun long, barrel-shaped, lobes 4-7 x 0.7-1.5 mm, 1.3-1.9 times as long as the tube. Stamens inserted 0.5—1 mm from the corolla base; anthers 2-2.8 mm long, partially fertile. Style very short. Follicles narrowly cylindrical, 10-30 x 0.4-1.3 x 0.4-1.3 cm, many-seeded. Seed 12-22 mm long with a 22-58 nun long coma. Fig. 139.6.

139. APOCYNACEAE: 6. Landolphia, 7. Oncinotis, 8. Saba

93

Figure 139.5. L A N D O L P H IA B U C H A N A N IL 1 flowering branch x 2/ 3; 2 - leaf, lower side x 1; 3 - opened corolla x 4; 4 - ca­ lyx x 4; 5 - stamen x 10; 6 - pistil x 8; 7 - fruit x 2/ 3; 8 - transverse section o f fruit x 2/ 3; 9 seed x 2 ; 10, 11 —longitudi­ nal and transverse section o f seed x 2. 1, 2 and 4 from C om er 14.9. 1952; 3 & 5 from Robson 588; 6 from de Wilde 6355; 7-11 from Vuyk 426. Drawn by Els Riemer-Gerhardt. (Reproduced with permission from Wageningen Agric. Univ. Papers 92 (2): fig. 2.)

Broad-leaved montane forest, swamp, riverine and secondary forest; up to 1800 in. SU AR IL KF; from Ethiop:i to W Nigeria and South Africa. Ash 1471; Haugen 86S, Mooney 8813.

8. SABA Pichon (1953) Pichcm, Mem 1FAN 35: 302-324 (1953); Leeuwenberg & van Ihlst, Bull. Jard. Bot. Nat. Belg. 59: 189-206 (1989). Liai us with curled tendrils, with much white latex in all pan-; Branches lenticellate. Inflorescence terminal or in the ::orks. Flowers sweet-scented only at night. Sepals free ovate to narrowly oblong, erect. Corolla white or creanay. with a yellow throat and often greenish tube. Sta­ men; included. Ovaiy syncarpous, unilocular. Fruit a larg; edible beriy. Seed irregularly ovoid or ellipsoid. species in tropical Africa inclusive of Madagascar; 1 in the Flora area.

S. comorensis (Boj.) Pichon (1953); Vahea comorensis Boj. (1837); Landolphia com­ orensis (Boj.) K. Schum. (1893), Pacouria com­ orensis (Boj.) Roberty (1953) - type: Comoro Islands, Bojer s.n. (K iso.). Landolphia florida Benth (1849); Saba florida (Benth.) Bullock (1959); Landolphia comorensis var. florida (Benth.) K. Schum. in Engl. Bot. Jahrb. Syst. 15: 404 (1893); Pacouria florida (Benth.) Hiem (1898); Saba comorensis var. florida (Benth.) Pichon in Mem. Inst. Fr. Afr. Noire 35: 309 (1953). Landolphia florida var. leiantha Oliv. in Trans. Linn. Soc. 29: 107 (1875). Branchlets glabrous or sometimes sparsely pubescent. Leaves: petiole 7.5-20 mm; blade ovate, elliptic or ob­ long, 4.5-24 x 3.4 x 12 cm, rounded, obtuse or sometimes apiculate or shortly acuminate at the apex, usually rounded or subcordate at base, glabrous on both sides or with a few hairs on the midrib and secondary veins; ter-

94

139. APOCYNACEAE: 7. Oncinotis, 8. Saba. 9. Strophanthus

Figure 139.6. O N C IN O T IS TENU ILO BA 1 - flower­ ing branch x Vy, 2 - branching node x 2; 3 leaf base with petiole from above x 4; 4 - domatia x 4; 5 - flower x 4; 6 opened flower showing 2 stamens and the pistil; the left stamen detached trom the pistil x 8; 7 - stamen, adaxial side x 8; 8 - follicle x Vy, 9 - seed x Vi. L 3 -7 from Brenan & Greetiway 8103; 2 from G.R. Williams 576; 8 from Drumm ond 10187; 9 from Wild. 2175. Drawn bv W. Wessel-Brand. (Reproduced with per­ mission from Wageningen Agric. Univ. Papers 85(2): fig. 6.)

Liaiy venation, mainly in the basal half, ladderlike. In­ florescence many-flow ered. Sepals pale green, 1.7-4 mm long. Corolla tube 16-34 mm long, pubescent or glabrous outside, lobes oblong or narrowly so, 19-41 x 4-12 mm. Stamens inserted 6-9 mm above the corolla base; anthers 1.9-3 mm long. Ov ary with a setose belt. Follicles orange to dark red. globose, ellipsoid or ovoid, or nearly so. 4-11 cm long. Fig. 139.7. Acacia - Commiphora woodland, riverine forest; 550-1600(-2000) m. GD SU WG IL KF GG BA; wide­ spread in tropical Africa from Senegal to Madagascar and from Ethiopia to Zimbabwe. Ash 1456; Mesfin T. 5959; Mooney 6829; W. de Wilde et al. 8873. In the Gambella area (IL) the fruit is said to be edible.

9 STROPHANTHUS DC. (1 8 0 2 ) Beentje,Meded Landb. Hogesch. Wag. 82(4): 1-191 (1982). Erect or creeping shrubs or lianas. Leaves decussateor less often in whorls of 3. Corolla with corona. Stamens in­ cluded or paniv exserted, inserted where corolla tube widens. Fmit of two follicles united at the extreme base. Seed with a decidous basal and a beaked apical coma. 38 species in tropical Africa and Asia; 1 in the Flora area

S. mirabilis Gilg (1902) - type? Kenya, Gove L ibia near Wonte, Ellenbeck 2205 (K lecto ). Shrub. 1-3 m tall, occasionally with climbing branches. Flowers; appearing before or with the leaves. Leaves

139. APOCYNACEAE: 8. Saba, 9. Strophanthus, 10. Wrightia

95

\ \ \ \ \

Figure 139.7. SA B A C O M O R E N SIS: 1 - flower­ ing branch x 1; 2 - flower x 3; 3 opened corol la tube x 6; 4 - calyx and pistil x 6; 5 - gynoecium x 6. 1 from Le Testu 9203; 2 -5 from P. Wit 2346. Drawn by Yuen Tan. (Reproduced with permission from Bull. Jard. Bot. Nat. Belg. 59: 189-206, fig. 1.)

subsi 5sile or shortly petiolate; blade narrowly elliptic, 8-36 x 3—10(—14) mm, rounded or obtuse at apex, cuneate at base or decurrent into the petiole, nearly glabrous or hispi l iious on the midrib and margins. Inflorescence l(-3; -flowered. Sepals erect or spreading, 6-15 x 1-3 mm. mucronate. Corolla tube white and suffused w'ith pink, turning yellow on both sides, 6.5—13 mm long and wide iing at 1/2 —4/5 of its length into a cup-shaped upper part, -11 mm wide at the mouth, hirtellous or puberulus on both sides; corona lobes spreading, yellow and red- or brow i-streaked, awl-shaped, 2.5-6.5 x 0.5 nun; corolla lobes ivhite, turning pale yellow on both sides, with or­ ange tails turning red outside, paler inside; lobes 4-8 x 3-5.5 nun, narrowing into the 1-2.5 nun wide tails, lobes including the tails 42—85 mm long. Follicles divergent, 16.5-12 x 1-1.8 cm, long-tapering towards the apex and ending in a small or minute knob. Fig. 139.8.

Acacia - Commiphora woodland; up to 600 m. Kenya and Somalia close to the Ethiopian border. So far not col­ lected in the Flora area but expected to be found in SD BA and HA.

10. W R I G H T I A R. B r. (1809) Ngan, Ann. M issouri Bot. Card. 52: 133 (1965); Leeuwenberg, Agric. Univ. W ageningen P apers 87(5): 33— 43 (1988).

Shrubs or trees, occasionally climbers. Leaves petiolate. Inflorescence terminal, aggregate, few- to manyflowered. Sepals almost free, imbricate, with 1-2 rows of alternate glandular hairs or small scales. Corolla tube cy­ lindrical to campanulate; lobes overlapping to the left; co­ rona variously shaped. Stamens exserted (in Flora area); anthers narrowly triangular, partly fertile. Ovary of two

%

139. APOCYNACEAE: 9. Strophanthus, 10. Wnghtia

Figure 139.8. ST R O P H A N T H U S M I R A B IU S 1 flowering branch x Vy, 2 - section o f flower x 4; 3 - fruit x 2/ 3; 4 - seed x 2/ 3. 1 from G illett 13322; 2 & 4 from Green­ way 84); Pagen, A gric. Univ. W ageningen P apers 87(2):

1-^13 (1988). Shrubs or small trees, 1-6 m tall. Branches flexible, bear­ ing a terminal inflorescence subtended by 3 equal branch­ less Branchlets slightly triangular at apex. Leaves in whorls of 3, leathery; blade very narrowly elliptic. Flowers showy. Fruit a bicarpellate follicle. Seed tomentel ous with upcurved hairs grading into a coma.

N. oleander L. (1753) type: cult. Netherlands, near Haarlem. Herb Cliff. 76 (BM lecto.). N. indicum Mill. (1768). Leaves: petiole 3-10 mm long; blade 5-21 x 1-3.5 cm. 4-8 times as long as wide, acuminate or acute at apex. Sepals narrow, 3-10 x 1-3 mm. Corolla hypocraterifonn, usually pink (sometimes white, pale pink, dark pink, wine-red. pale yellow or salmon), often with darker, longitudinal lines in the throat, single or double; tube 12-22 nun long; lobes obovate or nearly so, 13-30 x 8-25 mm; corona in the co­ rolla mouth of 5 epipetalous, variously lobed, parts. Sta­

98

139. APOCYNACEAE: 11. Nerium, 12. Thevetia, 13. Vinca, 14. Plumeria

mens barely included; anthers with long apical lursute appendages. Follicles 7.5-17.5 x 1-1.3 x 1-1.2 cm. Widely cultivated as an ornamental in public gardens, and along streets, sea level (Mitsiwa) to 2450 m (Addis Ababa, Science Faculty campus); cultivated throughout the world. Demel T. & students 1035; O. Rvding 1186; Sebsebe D. 1420. An extremely poisonous plant; one leaf is sufficient for a lethal dose. 12. THEVETIA L. (1737)

Low herb. Steins creeping and/or erect, glabrous. Leaves opposite, petiolate; petiole 4-15 mm long, glabrous; blade ovate, 2-71 x 1.5-4.5 cm, obtuse or acute at apex, rounded or subcordate and at the same time cuneate at base, ciliate, with or without short hairs on the midrib and secondary veins above. Flowers solitary, axillary, longpedicellate. Sepals linear, 10-15 x 1 mm, acuminate, gla­ brous or with a few long hairs outside, long-ciliate. Co­ rolla blue-violet; tube obconical, 15-17 mm long; lobes spreading, obovate, 15-25 x 10-20 mm. Stamens in­ cluded. Follicles erect, pod-like, 30 x 5 mm, acuminate, adaxially dehiscent.

Shrubs or small trees. Leaves alternate, blade of various shape. 1-veined or pinnately veined. Flowers large, yel­ low. in terminal cymes. Sepals 5, inside glandular at base. Corolla infundibuliform with cylindric tube abruptly expanded into the campanulate throat. Fruit a compressed drupe, broader than long. Seeds large with acute edges.

Indigenous in southern Europe. Cultivated all over the world in temperate and subtropical regions, in the tropics especially at higher altitude. EW SU. Getahun s.n., O. Rvding 1687.

Indigenous in tropical America. Cultivated all over the world in the tropics and even sometimes in the subtropics. IL. Chaffey 383; ACB Thomerson 608.

Plumeria rubra L. (1753) - ty pe from America. P. acuminata W.T.Ait. (1811); P. acutifolia Poir. (1812).

13. VINCA L. (1735)

Small tree, 2-10 m tall, repeatedly branched as a candela­ brum, with white latex in all parts. Branchlets succulent, 2-3 cm in diameter. Leaves alternate, petiolate, petiole 2-8 cm long, blade glaucous, narrowly elliptic, 12-40 x 3-12 cm, acuminate at the apex, with many straight sec­ ondary veins forming an angle of 70-90° w ith the mid­ vein and a submarginal vein. Inflorescence terminal, later surrounded by branchlets. dense when young, later lax, long-pedunculate. Sepals minute, c 1 x 2 nun. Corolla white with pink, pink, wine-red or canary yellow, usually withayellowthroat. tube narrowly cylindrical, 15-20 mm long; lobes elliptic, 25-40 x 15-25 mm. Stamens deeply included, inserted at the base of the tube. Follicles forming a very wide angle, c 12 x 1.5 cm, adaxially dehiscent. Seeds winged Frequently' cultivated in towns and villages all over the tropics. EW HA. Demel T. 1025; O. Ryding 1208.

14. PLUMERIA Tourn. ex. L. (1735) 9 species in S America, one of which, T. peruviana. is Trees or tall shrubs, usually with stout brandies. Leaves al­ widely cultivated also in the Flora area. ternate; petioles usually long, resinous at base without ex­ ternal glands, secondary veins numerous, straight, T. peruviana (Pers.) K.Schum. (1895) connected by a more or less conspicuous marginal vein, - type from South America. axillary stipules missing. Flowers white, pink or yellow­ ish, in cymes in terminal, often umbelliform, cory mbs, Shrub or small tree 1-4 m tall, with white latex in all parts, supported by often large, caducous bracts. Calyx small glabrous all over Leaves alternate, shortly petiolate; w ithout glands on the inside. Sepals 5, almost free, usually blade very narrowly elliptic to linear. 6-12 x 0.5-0.9 cm, broad and obtuse sometimes partly suppressed. Corolla acute to acuminate at the apex, decurrent into the peUole; infundibuliform, tube cylindric, slightly widened at the secondary veins obscure. Inflorescence axillary, fewbase; lobes 5, broad, oblong, straight or more or less flowered. Sepals narrowly triangular, 5-10 x 1.5-2 mm, twisted. Stamens in the widened base of the corolla-tube. acuminate, spreading or recurved. Corolla bright yellow, Style very short, columnar. Mericarps follicular, follicles infundibuliform. 50-65 nun long; basal part of tube cylin­ widely divergent, elliptic to linear, leathery . Seeds oblong drical, 10-15 mm long, upper part 10-17 nun long; lobes or fusiform, flattened, winged at the apex or all around suberect obovate, 20-40 mm long. Stamens inserted at the apex of tlie narrow part of the corolla tube, hidden by About 17 species indigenous to tropical America some lanate scales. Drupe obtriangular or nearly so, 25 x 30-35 widely cultivated, 1 in the Flora area. x 20 cm, bluntly apiculate; mesocarp fleshy.

Low dwarf shrubs or herbaceous perennials, usually with trailing vegetative shoots and ascending flowering stems. Leaves opposite. Flowers solitary in leaf axils, longpedicillate. Corolla blue, violet or w hite, hvpocrateriform with 5 broad asymmetric lobes and a gradually widened tube with a zone of hairs above the insertion of the sta­ mens. Lobes oblique, as long as the tube, overlapping to the left in bud. Stamens inserted half-way up the corolla tube; anthers with the connective expanded above into broadly triangular, liaiiy flaps. 7 species in Europe. North America and Central Asia; 1 species cultivated and naturalized in the Flora area. V. m ajor I. (1753) - type from Europe.

99

140. ASCLEPIA DAC EAE by D. J. Goyder et al. N s. Brown, Asclepiadeae in FI. Trop.Afr. 4(1): 132-503 (1902-3); Bullock, Periplocaceae&Asclepiadaceae in FI W. 71, p. Afr. ed 2, 2: 80-103 (1963); Bruyns & Forster, Taxon 40: 381-391 (1991); Liede & Kunze, PI. Syst. Evol. 185: 2 ‘1:-284 (1993); Liede & Albers, Taxon 43: 201-231 (1994); Swarupanandan et al., Bot. J. Linn. Soc. 120: 327-369 ( 96); Liede, Taxon 46: 233-247 (1997); Venter & Verhoeven, Taxon 46: 705-720 (1997), Ann. Missouri Bot. Gard. 88: 5^i*-568 (2001); Endress & Bruyns, Bot. Rev. 66: 1-56 (2000).

F rbs, shrubs or climbers with white, yellow or clear latex, mostly perennial, but frequently with shoots ren . ved annually, often with tuberous roots, rarely whole plant annual. Stems strongly succulent to herbaceous o voody. Leaves simple, usually opposite but occasionally in whorls o f 4; stipules absent or reduced to an ann ciIar fringe or ridge. Inflorescences terminal, axillary or extra-axillary, cymose, frequently reduced to umbellif< rm or racemiform clusters, or much branched with many fascicles. Flowers pentamerous, mostly a tinomorphic, bisexual. Sepals free, usually with colleters at their inner base. Corolla sympetalous, lobes valvate or imbricate in bud. Corona present or absent, arising from the corolla, the gynostegium, or both. Stan ens usually inserted at the base of the corolla tube; filaments free or fused into a tube surrounding the ovaries; ar.thers 2- or 4-locular, fused to the stigmatic head, with or without a membranous apical appendage; pollen in te ads or pollinia, dispersed on translators or in pollinaria. Ovary superior or semi-inferior; carpels 2, free, ex­ cept at base and apex; placentation axile, ovules mostly numerous; styles fused towards the apex and dilated into a stigma head. Fruit o f 2 unilocular follicles, frequently only one developing. Seeds usually numerous, rr : stly compressed, crowned by a coma of long silky hairs. A diverse family with approximately 240 genera and 3400 species, widely distributed throughout the world b t centred in the tropics and subtropics. The family is most diverse in eastern and southern Africa. 170 species ir 43 genera are native to the Flora area. Although readily recognisable by the complex structures of the flowers, it is now generally regarded that the A: ;lepiadaceae as treated here represents the more derived end o f a broad Apocynaceae. The 3 subfamilies recc L;msed here, w'hich are defined largely on the basis o f floral structures and arrangements leading to progres­ sively more precise control over the transfer of pollen, are perhaps best regarded as subfamilies within an e: panded Apocynaceae. Note on floral structures within the Asclepiadaceae Fig. 140.1. T e 5 stamens are fused apically to the expanded stigma h( id and together form a compound structure called the g} ostegium. The ovary is therefore almost entirely conce. iled within a ring of stamens (the staminal column) wi ose filaments are usually fused into a tube (free in snbfam. PERIPLOCODDEAE). Only the sterile apex of th stigma head remains visible, level with or extending be ond the anthers; the receptive portions of the stigma he id are on its underside behind the frequently sclerified m ;gins of the anthers which form a chamber for the de­ pt ition of pollen. The pollen transfer apparatus is formed b\ secretions from the stigma head and. in Periplocoid gt era, generally consists of a spathulate translator onto w ich pollen tetrads, or more rarely pollen masses (p ilinia), from adjacent anthers are shed, in the two re­ in ;ning subfamilies, the secretions form a central cc rpusculum linked by caudicles or translator arms to 2 or 4 ollinia of adjacent anthers - the unit is transferred in its er irety from one flower to another and is called a pc inarium

The Herbarium, Royal Botanic Gardens, Kew. Richmond, Surrey TW9 3AB, UK

The corona presents many diagnostic characters for generic and specific recognition. Following the system devised by Liede & Kunze (1993), it may be coralline (de­ rived from the corolla) or gynostegial (from the staminal column). Coralline coronas can be divided into those oc­ curring in the corolla lobe sinuses, and those forming an annulus in the corolla tube. Gynostegial coronas again have two basic elements, a staminal corona attached dor­ sally to the stamens, and an interstaminal corona, and these elements can be combined in a number of ways: e.g. staminal corona lobes only ; a fused ring of staminal and interstaminal lobes; fused staminal and interstaminal co­ rona with additional staminal lobes.

Key to genera 1. Staminal filaments free; pollen in tetrads or occa­ sionally aggregated into pollen masses (pollinia). deposited onto a spathulate pollen carrier (trans­ lator) lying on the stigma head between adjacent anthers. (Subfain. PERDPLOCOIDEAE) - Staminal filaments, when present, united into a tube or annulus; pollen of each anther cell aggre­ gated into 1 or 2 pollinia the pollinia from adja­ cent anther locules attached directly or indirectly to a single translator situated between the anthers.

140. ASCLEPIADACEAE

100

Figure 140.1. Parts o f an Asclepiadaceae flower (Asclepias syriaca). Note that this model applies strictly to most genera o f the tribe Asclepiadeae - please re­ fer to individual figures for species in other parts o f the family 1 - com­ plete flower, showing reflexed corolla and gynostegium complete with 5 staminal corona lobes; 2 gynostegium viewed from above showing, from the outside 5 corona lobes each attached to the base of in anther, ring o f 5 stamens enclosing the stigma head; 3 - gynostegium viewed from the side, with the co­ rona lobes removed to expose the staminal column; 4 - single anther viewed from within, showing posi­ tion o f pollen masses (pollinia) and translators; 5 - pollinarium consist­ ing o f 2 pollinia joined by a pair of translator arms to the central corpusculum, 6 - paired ovaries fused above into the shared stigma head. (Modified from Rosatti (1989). J A rnold Arb. 70: 307-401, 443-514: fig. 8.)

2. Four minute pollinia attached to each pale, soft

translator, shrubs or lianas. (Subfam. SECAMONOIDEAE) -

7. Secamone

Pollinia attached in pairs to each translator, transla­ tors hard, horny, mostly dark brown or black; herbs, shrubs or lianas. (Subfam. ASCLEPIADOLDEAE) 8

3. Corolla showy, pink, to c 6 cm long; corona of five lobes in mouth of corolla tube. 6. Cryptostegia - Not as above. 4 4. Corona lobes fused basally into an annulus; pollen in pollinia 5. Schlechterella - Corona lobes free from one another, pollen in tetrads. 5 5. Corolla tube distinct and campanulate. 4. Raphionacme - Corolla rotate with tube indistinct and shallowly saucer-shaped. 6. Corolla lobes glabrous, without any swelling or dark midvein area. 3. Tacazzea - Corolla lobes with a dark, adaxial swelling at their bases or adaxially hairy. 7. Stamens glabrous. - Stamens hairy. 8

L eav es a b se n t o r re d u ced to scales.

-

Leaves well developed.

I. Buckollia 2. Periploca 21

9. Corolla tube well developed, much longer than wide, with a distinct basal swelling around the co­ rona; corolla lobes mostly shorter than corolla tube, remaining attached at tips to form a ‘cage like structure. 34. Ceropegia - Corolla not as above; if corolla tube longer than lobes, then stems with prominent angles or rows of tubercles. 10 10. Non-succulent shrub or small tree; corona coralline. 32. Leptadenia - Plants clearly succulent; scramblers or low herbs; corona gynostegial. 11 11 Stems smooth or striate, mostly scrambling or twining; latex white. 12 - Stems with prominent angles or rows of tubercles, prostrate to erect but never scrambling or twining; latex clear or at most slightlycloudy. 13 12. Flowers with a short outer corona ring and hood­ shaped staminal corona lobes; stems smooth, uni­ formly green. 26. Sarcosteinma - Flowers with a corona of highly fused staminal and interstaminal parts, without hood-shaped stami­ nal lobes; stems finely striate, somewhat glau­ c o u s /" ! 25. Cynanchuro 13. Stems with 6-20 angles or rows of tubercles; epi­ dermis often very minutely papillose. 14

140. ASCLEPIADACEAE

Stems with 4 or 5 angles or rows of tubercles, rarely unusually vigorous stems with 5 or 6 an­ gles: epidermis smooth, tessellate or bullate. 15 |4. Stems repent, ascending or erect, (6-)8-20-angled. angles divided into distinct tubercles or ± flat and tessellate; corolla reflexed to urceolate, lobes spreading to erect. 38. Echidnopsis Stems completely prostrate, 6-angled, angles very well-defined and only indistinctly divided into tubercles; corolla globose, lobes inflexed and hidden within corolla lube. 39. Pseudopectinaria 15 Stem leaves apparently absent, tubercles conical with sharp brown thorn-like tips, stems creeping; corolla more than 5 cm diam.; corolla lobes short and reflexed. 41. Edithcolea Stem ieaves well-defined or, if apparently absent and tubercles callosed, tip blunt and stems erect or ascending; corolla up to 5 cm diam.; corolla lobes conspicuous. 16 1f Stems with 4 angles, tubercles poorly defined or if prominent then not tapered, never marked by dark blotches or streaks though sometimes with more regular paler markings; pollinaria with corpusculum wings very inconspicuous to ab­ sent. 17 Stems with 4 or 5 ranks of prominent conical tuber­ cles tapering to fme point, often marked with ir­ regular darker blotches or streaks, rarely prostrate with 4 rounded angles and relatively small tuber­ cles; leaves poorly differentiated from rest of tu­ bercle; pollinaria with corpusculum wings prominent. 19 Stem solitary', subglobose; epidermis conspicu­ ously tesselate. 40. Pseudolithos Stems numerous, elongated; epidermis smooth, minutely papillose or bullate. 18 Stems ascending or erect, with prominent tubercles and/or conspicuous angles; surface smooth. 36. Stems rigidly spreading, sometimes horizontal, with low, obscure tubercles and rounded an­ gles; surface conspicuously bullate. 37. Rhytidocaulon 1 i Inflorescences subapical, sessile; stems with 4 ranks of tubercles. 42. Pachycymbium Inflorescences from near base of stem, often elon­ gating with age; stems usually w ith 5 ranks of tu­ bercles, occasionally with 4 ranks and then ± prostrate. 20 20 Corolla tube shorter than corolla lobes; corona stipitate. 43. Duvalia - Corolla tube well developed, about as long as or lon­ ger than corolla lobes; corona sessile. 44. Huernia 2 11. Slender, erect, ephemeral herb with linear leaves; plant of desert environment. 33. Conomitra Perennial herbs, shrubs or vines. 22

101

22. Corolla tube well developed, generally much lon­ ger than wide, usually with a distinct basal swell­ ing around the corona; corolla lobes mostly shorter than mbe and remaining attached at tips to form a 'cageMike structure, rarely separate at tips and rigidly spreading; plants with clear latex. 34. Ceropegia - Corolla tube, when present generally shorter than wide, if longer then plant woody; corolla lobes never united at tips; plants with clear or white latex. 23 23. Shrubs up to 5 m tall; glaucous, mostly obovate leaves to 26 x 15 cm. 10. Calotropis - Erect perennials or twiners, if shrubs then no more than 2 m tall; leaves much smaller than above. 24 24. Herbs, frequently with erect annual stems arising from a perennial rootstock, or tufted slirubby herbs woody at the base only, with ascending or erect branches and linear or narrowly lanceolate leaves; stems never twining. 25 - Large woody climbers, small shrubs or slender vines; stems mostly scrambling or twining distally, if not. then leaves broad. 33 25. Corona arising from the corolla mbe; dwarf peren­ nial tomentose herbs branching from the base. 8. Glossonema - Corona arising on the gynostegium. 26 26. Corolla united into a tube basally; corona in two series; pollinia ± exposed on top of stigma head; latex clear. 35. Brachysteima - Corolla lobed ± to the base; corona of one series; pollinia pendulous in vertical anther thecae and obscured by the margins to the anther; latex w'hite. 27 27. Plants shrubby, stems persistent sometimes woody at base; roots fibrous or woody, never tuberous. 28 - Plants herbaceous, stems arising annually from pe­ rennial tuber or woody rootstock. 29 28. Inflorescence an indeterminate condensed spiral; corolla with shaggy indumentum apically; plants rheophytic, always found in temporary or perma­ nent watercourses. 11. Kanahia Caralluma - Inflorescences mnbelliform; corolla lobes gla­ brous; plants not rheophytic. 12. Gomphocarpus 29. Corona lobes solid, without cavity, lobes, keels or teeth. 16. XvsmaJobium - Corona lobes pouched or laterally compressed with a cavity or sinus, or dorsally flattened with lobes, keels or teeth apically or on the inner face. 30 30. Flowers erect; margins of anther wings convex; translator arms with an obovate membranous proximal portion and a filiform distal portion: fruiting pedicel straight erect. 15. Stathmostelma - Flowers in secund or globose umbels: margins of anther wings not curved; translator arms not as above; fruiting pedicel contorted. 31 31. Leaves narrowly linear, less than 2.5 mm wide; sessile or subsessile. 9. Aspido^lossum - Leaves lanceolate to ovate, at least 5 mm wide, mostly much broader; petiole at least 2 mm long. 32

102

140. ASCLEPIADACEAE: l.Buckollia

32. Leaves softly pubescent; corona lobes subglobose. upper margins fimbriate or dentate. 14. Trachycalymma - Leaves with stiff, scabrid hairs on both surfaces; lateral margins of corona lobes inflexed to form wings or keels, apex entire or 3-Iobed. 13. Pachycarpus 33. Large woody climbers. - Slender vines or small shrubs.

34 37

34. Coralline corona present in corolla lobe sinuses; gynostegial corona minute or absent. 35 - Coralline corona absent; gynostegial corona of fleshy lobes on back of anthers. 36 35. Corolla rotate, lobes pubescent to lanate; stigma head enclosed by anthers; pollinia with translu­ cent marginal germination zone. 32. Leptadenia - Corolla campanulate to urceolate, lobes glabrous; apex of stigma head exserted from staminal tube, pollinia without translucent germination zone. 30. Gymnema 36. Leaves strongly cordate at the base, follicles slen­ der, mostly single, smooth, if paired then held at an acute angle; corolla glabrous within. 29. Gongronema - Leaves truncate, rounded or obtuse at the base, never strongly cordate, follicles robust, mostly paired at c 180°, conspicuously winged or with longitudinal wrinkles; corolla pubescent on tube or lobes within. 28. Dregea 37. Plants richly branched woody shrublets, young shoots frequently twining. 38 - Plants scrambling, climbing or decumbent, some­ times basally woody, but not forming richly branched shrublets. 40 38. Plants with white latex; stigma head long-rostrate. well exserted from anthers. 31. Stigmatorfaynchus - Plants with clear latex; stigma head short, not or barely exserted from anthers. 39 39. Corona in two series: a spreading outer corona ring appressed to the corolla and conspicuous rectan­ gular lobes radiating from the backs of the an­ thers. 23 Diplostigma - Corona of slightly fleshy, ovate scales adnate to the back of the anthers; outer ring absent. 22. Blyttia 40. Flowers at least 1 cm diam., broadly campanulate; corolla lobes fused for at least 1/2 of their length, white with dark red pattern. 27. Oxystelma - Flowers smaller, not broadly campanulate; corolla lobes fused for less than 1/2 of their length, brown, cream yellow or purple, but not white with dark red pattern. 41 41. Latex sparse, clear. - Latex white, usually abundant.

42 45

42. Corona united into a tube; leaves cordate or lobate; stems succulent; gynostegium long-stipitate. 19. Schizostephanus - Corona of staminal lobes only; leaves rounded, truncate or acute basally, rarely cordate; stems not succulent. 43

43. Corolla lobes twisted; corona of inconspicuous scales only; stylar head extended into a long ap­ pendage. 20. Pleurostelisia - Corolla lobes not twisted, staminal corona lobes conspicuous; stylar head not extended into a long appendage. 44 44. Guide rails conspicuously extended along the co­ rona lobes. 2 1. Pentatropis - Guide rails not extended along the corona lobes. 18. Tylophora 45. Corolla lobes ciliate, follicles generally paired. thin-walled and covered with soft spines. 17. Pergulaha - Corolla lobes smooth; follicles usually single thin-walled and smooth or thick-walled and tuberculate or with a few soft spines. 46 46. Staminal corona lobes fused at the base only, or highly fused but with clearly visible sutures: folli­ cles thick-walled, smooth or with soft spines. 24. Pentarrhinum - Staminal corona lobes fused for at least 1/2 of co­ rona length but without visible sutures; follicles thin-walled, smooth or keeled. 25. Cynanchum

PERJPLOCOIDEAE Perennial climbers, shrubs or herbaceous geophytes; roots often tuberous; latex white. Leaves with interpetiolar lines, ridges with dentate colleters or frill-like stipules In­ florescences terminal or axillary. Corolline corona of 5 free or fused, antisepalous lobes; gynostegial corona ab­ sent. Interstaminal antipetalous discs often present Sta­ mens glabrous or hairy; filaments free, sometimes fused to corona lobe bases and/or interstaminal discs; pollen in rhomboidal. decussate or linear tetrads or in pollinia shed onto spathulate pollen translators alternating with anthers. 31 genera and 181 species in Africa Madagascar, Asia and SE Asia of which 18 genera and approximate 1> 90 species occur in Africa. Some authorities treat this group as a separate family, the Periplocaceae. 6 genera and 13 species are present in the Flora area. The sequence of genera follows the tribal classifica­ tion proposed by Venter & Verhoeven (1997), with the following tribes represented in the Flora area: Periploceae (genera 1-3); Gymnantherae (genera 4 and 5). The third tribe. Cryptolepideae, is represented by the naturalized genus Cryptostegia (6). 1. BUCKOLL1A Venter & R.L. Verh. (1994) by H. J. T. Venter* Venter & Verhoeven, S. Afr. J. Bot. 60. 93-98 (]994). Woody climber, latex wliite. Stems twining, bark verrucose. Leaves opposite or in clusters, petiolate, blade nar­ rowly to broadly ovate, hairy, with secondary veins divaricate and parallel. Inflorescences monochasial, or di• Dept, of Botany and Genetics, University of the Orange Free State, 339 Bloemfontein 9300, South Africa.

140. ASCLEPIADACEAE: 1. Buckollia. 2. Periploca

ch; iiaJ with two monochasial branches, each bearing up lo 1 flowers. Flowers with gynostegium exserted from cor :>ila tube Sepals free, with paired colleters at inner ba: os. Corolla rotate; tube shallow, saucer-shaped; lobes wii i an adaxial glandular swelling at their bases. Corona lot os arising at corolla mouth, simple, filiform, distinct, lnt rstaminai discs distinct. Stamens arising at and fused lo i ler base of corona lobes, glabrous; filaments filiform; anthers ovate, apex acute, pollen in tetrads. Stigmatic he; d pentangular-ovoid. Follicles paired, divergent to ho ontal. long cylindrical-ovoid, tapering to a blunt ap; x. puberulent and verrucose. 2

species in E and NE Africa, from Ethiopia to Tanza­

103

berulous above, densely white-tomentose with midrib rusty-tomentose below. Inflorescences densely tomentose to pubescent; peduncles 7-10 mm long; pedicels 4-5 mm long. Sepals triangular to ovate. 1.5-2 x 1 mm. Corolla tube 0.5 nun long; lobes 5-6 x 3 mm. ovate to oblongovate, apex obtuse, greenish to cream coloured, tomen­ tose on the outside, sparsely puberulous within, glandular swelling violet-brown. Corona lobes 3-4 mm long, filiform, puberulous. Follicles 130-175 x 9 mm, brown, sparsely verrucose. Fig. 140.2.1 & 2. An uncommon species of semi-arid Combretum Acacia - Commiphora scrub; 1400-1800 m. GG; north­ eastern Uganda. Gillett 15070; Haugen 158.

nia 1

-

1.

Most leaves clustered on short lateral shoots, green, pubescent; corolla lobes glabrous; follicles densely verrucose. 1. B. volubilis All leaves opposite, discolorous, dark green and puberulous above, densely white and rusty tomentose below; corolla lobes puberulous above, white tomentose below; follicles sparsely verrucose. 2. B. tomentosa I. volubiiis {Schltr.) Venter & R.L. Verh. (1994); Raphionacme volubilis Schltr. (1895); Tacazzea iolubilis (Schltr.) N.E. Br. (1902); Curroria volubilis iSchltr.) Bullock (1955) - type; Central Africa, Scott ’ lliot s.n (BM holo., K iso.).

Ro ms tuberous. Stems up to 4 m x 10 mm. scrambling, braiuhes divergent to horizontal, lateral shoots mostly stu led Leaves in clusters on stunted lateral shoots or oppo: ne on normal shoots; petiole 1-3 mm long; blade narrov. L to broadly obovate, 35-50(-65) x 7-15(-24) mm, ap’ hillsides, perhaps restricted to limestone. EW TU; northern Somalia. Saudi Arabia, Yemen and Oman. Bally 6683; Gilbert & Getachew A. 2666. 3. G. phillipsiae (N.E. Br.) Goyder (19%); Asclepias phillipsiae N.E. Br. (1895) - type: So­ malia, LortPhillips s.n. (K holo.). Gomphocarpus jruticosus var. angustissimus Engl. (1892) - types: TU, Scholoda 2200 m, Schimper 1862: 938 (B destroyed); EW, Keren, Steudel 747 (B destroyed, K); Masai highlands, Hohnel 35 (B de­ stroyed); Kilimanjaro, Kersten (B destroyed). Stems (0.2-)0.5-1.5 mtall, erect, much branched from the base, slender, densely spreading-pubescent. Leaves nar­ rowly linear, 2.5—5(—10) x 0 .1-0.2(-0.5) cm. apex acute, mucronate. Inflorescences with 4—7 flowers; peduncles 0.5-2(-3) cm long; pedicels c 1.5 cm long. Sepals 3-5 x

1-2 mm, triangular, acute. Corolla reflexed, white or cream, frequent!} tinged pink outside; lobes 5—7 x 3-5 mm, ovate, subacute. Corona lobes cream or green, teeth and margins frequently pink. 2—3 x 2—2.5 mm, more or less as tall as the column, the upper margins rounded and with a distinct notch at base of proximal teeth, proximal margins produced into a pair of broad, often denticulate teeth 1-1.5 mm long, pointing back along the upper mar­ gins of the lobe and curved down into the cavity, cavity without teeth or projections. Follicle 4-7 x 1.5-2.5 cm. ovoid, tapering gradually into an attenuate beak, inflated, pubescent and with pubescent filiform processes. In open rocky ground and disturbed areas; 1400-2200 (-2800) m. EW TU ?GD SU AR KF GG SD BA HA; So­ malia, Kenya, Uganda, Tanzania, Rwanda and eastern Rep. Dem. du Congo. Gilbert et aJ. 419; M Gilbert 3399; Thulin et al. 3803. 4. G. purpurascens A Rich. (1851), non Asclepias purpurascens L.; G. Jruticosus var. purpureus Schweinf. (1867); Asclepias pubiseta N.E. Br. (1902) - type: TU, Tchelikote, Petit s.n (P holo., K iso.). Asclepias albida N.E. Br. (1895) - type: TU, Schimper 27 (K holo.). Stems 0.6-2 m tall, erect, much branched from base, stuut, densely spreading-pubescent or tomentose. Leaves nar­ rowly linear, 5-15 x 0.1-0.5 cm, apex acute, mucronate. Inflorescences with 5-11 flowers; peduncles 1-3 cm long; pedicels J 5-3 cm long. Sepals 4-5 x 1 mm, triangu­ lar. acute. Corolla reflexed, white, cream or green, gener­ ally tinged purple outside; lobes 8—10(—12) x 4-6 nostegium c 1.7 \ 1.8 mm on a bulge of 0.4-1.3 mm; anther wings c 0.6 mm long. Sty lar head c 1 x 1 mm. upper part equal ling the lower part in height, depressed-conical, lollicles single, occasionally paired, pendulous, c 6 cm long. 8-10 nun in diameter, obclavate. keeled. Frequent and widespread in forest margins, thickets, roadside slirubbery; 1000-2600 m. EW GD SU SD BA HA; E Africa Gilbert & Jefford 4316, de Wilde & de Wilde-DuyJjes 7020. 10386.

Used as a relish (Perdue & Kibuwa 11341); leaves used to make tea as a tome (Matthews 6354). The degree of hairiness varies greatly from densely tomentose to almost glabrous. As there is neither a sharp limit between glabrous and haiiy forms, nor a correlation with other characters, the separation of var. tomentosum Oliver does not seem advisable.

4. C. altiscandens K. Schum. (1895) -type: Tanzania. Holst 9078 (B holo. destroyed. K lecto. designated by Liede. 1996). C. mensense Schweinf. ex K. Schum. (1895) type. EW. Gheleb. Schweinfurth 1505 (B holo de­ stroyed. M lecto designated by Liede. 1996).

5. C. clavidens N E. Br. (1895) - type: Somalia, James & Thrupp s.n. (K holo.). Erect, small shrub up to 2.5 m tall, with long twining shoots. Leaves with petiole 6-25 mm long, blade liasrate, 10-75 x 4-35 mm. basally truncate to cordate or lobate. Inflorescences sciadioidal. sessile: pedicels 3-10 mm long. Corolla lobes 3-5.5 x 0.8-1.2 mm, horizontal to decurved, linear to lanceolate, green or creamish green. Corona white, abaxiallv papillose towards tlie apex, tubu­ lar to urceolate for 2/3 to 3/4 of its length, free staminal parts erect, ovate or ovate-oblong, producing a pro­ nounced convex fold; free interstaminal parts erect to re­ flexed. ovate to oblong, with laterally involute margins. Gvnostegium sessile; anther wings 0.5-0.8 nun long.

140. ASCLEPIADACEAE: 25. Cynanchum, 26. Sarcostemma

>tylar head white. 1-2.5 mm high, upper part uinbonate or .:longated-corucal. Follicles nonrially 1 per flower, penlulous, 6-10 cm long, 10-15 mm in diameter, obclavate. wrongly beaked apically. 1. Stylar head well exceeding the corona; flower buds 6-6 5 mm long. subsp. clavidens Stylar head not or liardly exceeding the corona: flower buds 3-4 mm long. subsp. hastifolium .ubsp. clavidens |

)ry savanna. Acacia - Commiphora busliland; 250-1400 HA; local but widespread in Somalia, Kenya and Tanama. De Wilde 5983. Fruits edible.

- ubsp. hastifolium (N.E.Br.) Liede, in Ann. Missouri Bot. ,'iard. 83: 306(1996); C. hastifolium N.E.Br. (1895) - type: TU. near Djeladjeranne, Schimper 1690 (K holo., P iso.). C. hastifolium K. Schum. (1895)-ty p e not listed. Acacia seyal - Balanites bushland; 200-1600 m. EE 'J SU GG SD; Upper Volta, Mali, Niger. Somalia. . cnya and Tanzania. Friis et al. 1018; Gilbert & Gelahun 123; Gilbert et al. 7399. The only known material of Perianthostelma Baill., a ; i>ecimen in the Paris herbarium with the unpublished i ime P. abyssinicuni, represents Cynanchum clavidens i ubsp. hastifolium. 1 C. falcatum Hutch. & E.A. Bruce (1941) -type: Somalia,boundary,Gillett4114(Kholo ). ( Umber, 1-1.5 m high. Leaves with petiole 2-10 mm lo tig; blade triangular to falciform, 20-40 x 2-16 mm. In1 irescences bostrychoid to sciadioidal, 5-12-flowered, 4 10 flowers open at the same time; rachis missing or up l 2 mm long; peduncles missing or up to 5 mm long, put .scent; pedicels 2-5 mm long, densely pubescent; flower t :ds 2.2-3 x 1.5 mm, ovoid. Corolla lobes 2-3 x 1.3 nun, p iient, oblong to lanceolate, apically obtuse to acute, c eamish green to yellow. Corona wliite, cyathiform, 1 sed for 3/4 of corona length, c 2 mm high, equalling the £ k’nostegium in height; staminal lobes broadly triangular. I apically reflexed; interstaminal lobes ovate to bifid, r Hexed, with straight, entire margins. Gynostegium c 2 x 1 5 mm; anther wings c 0.7 nun long. Stylar head 1 x 1.2 r m; upper part higher tlian the lower part, clavate. Follic 2S 1 per flower, pendulous, 5-6.5 cm long, obclavate. a ically strongly beaked, keeled. Open Acacia - Commiphora busliland; 250-1500 m. 5 SD; northern Kenya, Somalia. Friis et al. 3169; ■ ’istphal & Westphal-Stevels 1466; J. de Wilde 6331. Closely allied to C. heteromorphum, but leaves triang Jar. corona more highly fused and stylar head clavate. 1 be type is an atypical specimen with very slender, falcif nn leaves - probably caused by disease. Most speci­

147

mens exhibit leaves more like those widespread in the ge­ nus. while the floral structure is the same. 7. C. heteromorphum Vatke (1876) - ty pe: TU. Hamedo plain. Schimper 940 (B holo. destroyed. K lecto. designated by Liede, 1996). C. fraternum N.E. Br. (1895) - type: TU. near Djeladjeranne. Schimper 1802 (K holo.). C. pleianthum K. Schum. (1898). Climber. Leaves with petiole 6-8 nun long; blade ovate. 18-32 x 8-14 inm. apically acute to acuminate, basally rounded to obtuse. Inflorescences bostrychoid, 8-10flowered. 3-5 flowers open at the same time; peduncles 7-9 mm long, densely pubescent; pedicels 4-6 mm long; flower buds ovoid, c 2 x 1.5 nun. Corolla lobes patent, ob­ long, apically obtuse. Corona cyathiform, 3 mm high, ex­ ceeding the gynostegium but not obscuring it, tube reaching a little less than 1/2 of total corona length, staminal lobes elongate-triangular, apically reflexed; interstaminal lobes shorter than staminal ones, oblong, flat, reflexed. Gynostegium sessile, c 1.8 x 1.8 mm; anther wings c 1.0 mm long. Stylar head c 1 x 0.5 mm high; equalling the lower part in height, umbonate. Follicles 1 per flower, pendulous. 5-6 cm long, obclavate. apically strongly beaked, keeled. The species was rarely collected in the 19th century' and not at all more recently. Probably endangered. EW TU; Somalia. Pappi 7300; Schimper 940 & 1802. The flowers of this species are quite similar to those of C. falcatum; however, the clavate stylar head and the pronouncedly triangular leaves of the latter are unmis­ takable.

26. SARCOSTEMMA R. Br. (1810) by S. Liede Liede & Meve, Bot. J. Linn. Soc. 112: 1-15 (1993); Meve & Liede, Bot. J. Linn. Soc. 120: 21-38 (1996). Plants scrambling or twining, often dichotomously branched; shoots uniformally green, smooth, apically of­ ten with a more or less dense indumentum or glabrous, ba­ sally often corky, then up to 5 cm in diameter; latex white. Leaves reduced to sessile, triangular, persistent scales. In­ florescences one per node, extra-axillary, multi-flowered, sciadioidal. Corolla rotate, lobes basally fused. Corolline corona absent. Gynostegial corona double, consisting of inner free staminal parts and an outer ring of connate staminal and interstaminal parts. Gynostegium sessile, anthers with apical appendages and clearly differentiated lateral anther wings consisting of a distal ridge only. Pollinia pendulous, characteristically golf-club shaped. Stylar head forming a protrusion at the upper end of the corpusculum; upper part umbonate to conical. Follicles obclavate or elongate. Seeds glabrous or with unicellular hairs. 14 species in the Old World (S. viminale with numer­ ous infraspecific units); 2 in the Flora area.

148

140. ASCLEPIADACEAE 26. Sarcostemma

1. Plants not taller tlian 50 cm. sometimes with some much longer trailing shoots, with subterranean rhizomes; corolla brow n or purple. 1. S. vanlessenii - Plants taller than 50 cm. erect or climbing, with or without trailing shoots, without subterranean rhi­ zomes; corolla predominantly yellow. 2 S. vim in ale 1 S. vanlessenii Lavranos (1975) type Yemen. Lavranos & van Lessen 1832 (K holo., PRE iso ). S. subterranea Adams & Holland (1978). Plant erect to decumbent, occasionally overhanging or trailing on bushes and trees, non-twimng. sometimes pro­ ducing a single main stem; stems succulent, 0.1-1 in long. 2-4 mm in diameter, frequently rooting at the very slightly constricted nodes, light to dark green glabrous (or nearly so); rhizomes 4-10 cm long, up to 10 111111 in di­ ameter. pale brown. Leaf scales c 1-1.5 x 1-1.5 mm. deltoid, acute, glabrous (or nearly so), quickly withering Inflorescences 2-8-flowered. terminal on short or sub­ sessile lateral shoots. 1-12 nun long, c 2 mm in diameter, sparsely pubescent: pedicels 4-9 mm long, pubescent Flowers scentless or nearly so. opening successively. Co­ rolla lobes 3-4 mm long, ovate-deltate. ascendingspreading. bright pink to dark purple. Corona white. rarel\ with a pinkish tinge, outer ring 3^t x 2-3 111111; staminal lobes 1-2 mm long, higher than the gynostegium, apicalh strongly incurved, claw-shaped. Sty lar head conical. Fol­ licles slender fusiform, up to 5 cm long, densely pubes­ cent when young. Stony ridges, only at higher altitudes, usually between 1500-2300 111. SD. Kenya. Tanzania; Saudi Arabia. Ye­ men. Thulm et al. 3452. 2. S. viminale (L.) R. Br. (1811); Euphorbia viminalis L. (1753) - type: Alpinus. Plantis aegypti 190, t. 190 (1735) (holo.); Iittora Africae’, Bassi s.n. [LINN 308.1 epi.. selected by Liede & Meve (1993)]. Plant erect, scrambling or twinmg up to 5 m or more over trees and shrubs, with or without corky main stem: shoots succulent, glabrous or pubescent when young. Leaf scales triangular to deltoid. Inflorescences sessile or on short lat­ eral shoots, one or two per node. 2-30-flowered; pedicels up to 30 mm long, glabrous or pubescent; flowers at least faintly scented Corolla lobes 2-7 x 1.3-2.5 mm, cream to yellow. Corona white; outer corona annular, 1.5-3 mm high, 1/3—1/2 of gynostegium length: staminal corona lobes 0.6-1.8 111111 long, shorter or longer than the gy no­ stegium, as broad as the anthers, claw-shaped. Stylar head conical or depressed-conical. Follicles single or paired. 5-18 cm long, fusiform, glabrous. The taxa of the S. viminale complex are still in need of further research in NE Africa. As many diagnostic fea­ tures readily apparent in the field are difficult or impossi­ ble to observe in herbarium material, two keys are presented to the subspecies as currently understood.

Key to living material 1 Plants forming a conspicuous main stem; branches twining like a corkscrew. 2 - Plants erect, or scrambling, but not twining like a corkscrew. 3 2 Inflorescences 011 short lateral shoots; young shoots glabrous or subglabrous. pedicels 15-30 mm long, less than 1 111111 in diameter; usually both side-branches developed, resulting 111 a ladder-like appearance. subsp. odontolepb - Inflorescences sessile at the nodes, terminal or extra-axillary; at least young shoots with indu­ mentum; pedicels 4-10 nun long. 1 111111 or more in diameter, usually only one side-branch devel­ oped subsp. suberosum 3. Plants forming dense tangles: lateral shoots at least 4 cm long; inflorescences 10-20-flowered. glo­ bose: flowers intensely sweet scented. subsp viminale - Plants forming dense mats with most shoots erect. some shoots arching and rooting but never twin­ ing; lateral shoots not more than 2 cm long; inflo­ rescences 2-8-flowered: flowers only faintly scented. subsp. stipitaceum Key to herbarium material 1 Pedicels 15-30 mm long. - Pedicels not more than 10 nun.

subsp odontolepi.s 2

2 Inflorescences sessile at the nodes, terminal or lat­ eral. subsp. suberosum - Inflorescences on conspicuous lateral branches 3 3. Lateral shoots at least 4 cm long; inflorescences 10-20-flowered. globose subsp viminale - Lateral shoots not more than 2 cm long, inflores­ cences 2-8-flowered. subsp. .stipitaceum subsp viminale Sea level-1000 111 EE; Kenya. Madagascar. Seychelles. Aldabra Hemming 1264. Meve & Liede (1996) suggest tliat the unusually long follicles (up to 18 cm) of this subspecies may be diagnostic subsp stipitaceum (Forssk.) Meve & Liecle (1996); Asclepias stipiiacea Forssk. (1775); Sarcostemma [ stipitaceum (Forssk.) Schult. (1820) - type: Yemen. Forsskal 277 (C holo.). Widespread and regionally abundant in dn or very dr\ rocky places; 200-2000 m. EE SD HA: Sudan, Somalia. Kenya, Tanzania, South Africa; Saudi Arabia. Yemen. Oman. Burger 2693 Schweinfurth 1432. subsp suberosum Meve & Liede (1996) - type: South Africa. Kirk 97 (K holo ). Fig. 140.30. Open country, tree savanna, thombush or disturbed grass­ lands ; 1000-2000 m. EW TU; eastern Africa from Enirea

140. ASCLEPIADACEAE: 26. Sarcostemma, 27. Oxystelma

149

Figure 140.30. SA R C O ST E M M A V IM IN A L E subsp. SU BE R O SU M : 1 - corky base o f main stem x 1/7 and sketch o f growing habit; 2 - apex of young shoot x 2>4; 3 - part o f flower­ ing stem x 2 V2 , 4 & 5 - flowers x 314; 6 - gynostegium and corona x 25; 7 pollinarium x 32. 2 & 6 from Erwee et al. 540; 3 from Albers et al. 562; 4 from M eve & Liede 545; 5 & 7 from Liede & Newton 3215. Drawn by U. Meve. (Reproduced with permission from Bot. J. Linn. Soc. 120: 36,1996.)

nd Somalia to South Africa; possibly more widespread. "niovenda 353; Corradi 8185; Terraciano & Pappi 2182.

campanulate; lobes fused for c 1/2 of the total length, white with dark red markings. Outer corona ring corolline, strongly hairy. Gynostegial corona of staminal subsp. odontolepis (Balf. J'.) Meve & Liede (1996); lobes with subulate apices. Anther connectives with api­ S. odontolepis Balf. f. (1877) - type: Rodriguez,cal appendages. Translator arms veiy short, pollinia I Balfour s.n. (E holo ). clavate; stylar head flat. Follicles single or paired, inflated or not; seeds very small, ovate, comose. C jastal and lowland forest; sea level-800 m. Tlie only uterial seen from Ethiopia is collected at c 760 m in the 2 species in Africa and Asia, both found in the Flora area. leaden. HA; Somalia?, Kenya, Tanzania, Rodriguez. 1. Corolla c 1.5 cm in diameter, staminal corona lobes . pt, Sudan; widely distributed in tropical and subtropi­ cal Asia and Australia. Ash 1149; Schimper 2305. According to Goyder (1992), Oxystelma secamone (L.) Karsten, a combination retrieved by Rahman (1990), is invalid because the basionym Periploca secamone L.. refers to Secamone alpini Schult.

140. ASCLEPIADACEAE: 28:.Dregea

28. DREGEA £. Mey. (1838), nom. cons. Pterygocarpus Hochst. (1843) Wattakaka Hassk. (1857) by F. Albers & S. Gtlldenberg* Bullock, Kew Bull. 9:349-373 (1954) & 11: 503-522(1957). Pubescent, somewhat woody lianas; latex white. Leaves opposite, petiolate, ovate to lanceolate. Inflorescences ex:ia-axillaiy, sessile or pedunculate clustered umbels. Se­ pals ovate, acute or obtuse. Flowers sweetly scented. Corolla more or less succulent, white, yellow or green, obed usually to more than half-way; tube campanulate; obes linear, contorted, basally adnate to gy nostegium. ;rorolline corona absent. Gynostegial corona of 5 ovoid or lattened fleshy staminal lobes, humped, attached to anhers basally, shorter or longer than anthers, apex free. Anthers often long and leaf-like, anther membranes idnate to the style and reaching tlie stylar head; stylar lead as long as or longer than anthers, conical or acute, ivnostegium conical to cylindrical. Pollinia erect, clubshaped or rounded, basally attached to translator arms; corpusculum narrowly ovate. Follicles paired and spreadng at an angle approaching 180°, thick, conspicously vinged or weakly ribbed; seeds flattened, convexconcave, with a broad marginal wing, glabrous, brown. About 15 species in tropical and subtropical parts of Africa, Madagascar, Arabia and Asia; 4 in the Flora area. Dregeafalls into synonymy w ithM arsdeniaR. Br., if the latter is treated in a broad taxonomic sense. 1 Whole plant pinkish tomentose; stylar head at most as long as the anthers. 2 - Plant not tomentose; stylar head longer than the an­ thers. 3 2. Corolla lobes adaxially villous only towards mar­ gins: staminal corona lobes at most as long as the gynostegium; follicles with 8 or more wrinkled wings. I D. abyssinica - Corolla lobes adaxially wholly tomentose; stami­ nal corona lobes overtopping the gynostegium; follicles with 4 unwrinkled wings. 2. D. rubicunda :. Flowers in clusters, inflorescence ± sessile; flower buds rounded; follicles apically hairy, 4-winged. 4. Flowers not clustered, inflorescence clearly pe­ dunculate; flower bud apices elongated; follicles completely pubescent, unwinged. 3. D. schimperi 1 D. abyssinica (Hochst.) K. Schum. (1895); Pterygocarpus abyssinicus Hochst. (1843); Hoya africana Decne. (1844); Dregea africana (Decne.) Martelli (1886); Marsdenia abyssinica (Hochst.) Schltr. (1913) - types: GD, near Mt Sabra, Schimper II: 1294 & 1366 (K isosyn). Dregea rubicunda Hiem (1898), non K. Schum. Marsdenia spissa Moore (1901).

_________________________________ * nstitut filr Botanik, Universitat Munster, Schlossgarten 3, D 48149 MQnster, Germany.

151

Plant scrambling up to 16 m; young shoots pale green; leaves and inflorescence covered with pinkish-red tomen­ tose hairs. Leaves with petiole 1-8 cm long; blade 6-11 x 3.5-8 cm, ovate or elliptic-ovate, apex acute, broadly cuneate to rounded at base. Inflorescences up to 4 cm in diameter, 10-30-flowered; peduncles 1—2 6 cm long, woody; pedicels 1.5-2 cm long. Sepals c 4 x 4 mm, broadly ovate, sub-acute. Corolla rounded in bud, more than 1.2 cm in diameter, tube c 1.3 mm long; lobes 4.5-6 x 2.6-3.9 mm, linear to lanceolate, truncate, spreading or reflexed, villous at the margins. Corona lobes c 2.6 mm long, shorter than staminal column, fleshy, cylindrical, apically rounded, white. Stigma head conical, about as long as the anthers and largely obscured by the anther ap­ pendages. Follicles 4-9 cm long, 2.5^1 cm in diameter, ovate, with at least 8 wrinkled, longitudinal wavy ridges, papery when dry. On light coloured soil in riverine habitats to 1650 m; EW TU GD SU IL KF GG SD HA; widespread in drier parts of tropical Africa also in Yemen. Gilbert et al. 7336; Siegenthaler 20; Smeds 1407B. 2. D. rubicunda A". Schum. (1893); Marsdenia rubicunda (K. Schum.) N.E. Br. (1903) - types: Sudan, Schweinfurth III: 33 (B syn. de­ stroyed) & 1255 (B syn. destroyed, K iso.); Kenya Hildebrandt 1944 (B syn. destroyed) & 2024 (B syn. destroyed, K iso.). Plant scrambling up to 10 m; young shoots covered with pink-maroon tomentose hairs. Leaves with petiole 1.52.5 cm long; blade ovate, oblong, elliptic or elliptic-ovate, 3.5-8.5 x 1.8-6.5 cm, apex sub-acute, truncate or appear­ ing mutilated, base broadly cuneate, truncate, or cordate. Inflorescences 2.5-3 cm in diameter, 10-40-flowered; pe­ duncles 1-1.5 cm long; pedicels 0.8-1 cm long. Sepals c 4.3 x 2 mm, ovate or elliptic, obtuse. Corolla rounded in bud; tube c 2.5 mm long, campanulate; lobes c 7 x 3.3 mm, oblong, apex asymmetrical, sub-acute, outside smooth, inside completely tomentose. Corona lobes 3.9-4.5 mm long, overtopping gynostegium, basal part with a wide free margin, apical part ending in a free subulate or linear apex, incised or bilobed. Stigma head more or less conical, about as long as the anthers. Follicles D. stelostigma bright pale yellow when ripe, c 11 x 4 cm, with 4 notched, or gently toothed, but not wrinkled, longitudinal wings c 1 cm wide, stiffly leathery, glabrous. Fig. 140.32. In wooded bushland in dry country, on sandy to loamy soil; to 1350 m; AF GG SD HA; drier regions of Central African Republic, Sudan, Somalia Kenya and Tanzania Brown 105; M. Gilbert 2391; Schweinfurth 1255. The flower scent is described as sweet but also as un­ pleasant (herbarium label). 3. D. schimperi (Decne.) Bullock (1957); Cynanchum schimperi Hochst. in sched. (1840), nom. nud.\ Marsdenia schimperi Decne. (1844) type: TU, Gennia Memsach region, Adua Schimper I: 260 (P holo., K iso.).

152

140. ASCLEPIADACEAE: 28. Dregea

Figure 140.32. D REG EA RL'BJC VN D A. 1 -habit x '/i; 2 - fruit x 3 - flowers x 3; 4 - flower from below x 3; 5 - gynostegium with corona x 5; 6 - ovaries and stigma head x 7; I p pollinarium x 20. 1 from Jeffrey 388; 2 from Thomas 3091; 3 -7 from Faulk­ ner 735. Drawn by D Erasmus.

Gymnema macrocarpum A. Rich. (1851) - types: TU, Beless, Prov. Schire, Quartin-Dillon (P syn.); SU, Shoa, Petit s.n. (P syn.). Traunia albiflora K. Schum. (1895).

liead with a long apical beak overtopping the anthers by c 4 mm. Follicles c 14x2 cm, ovate, tapering gradually to a blunt apex, unwinged, surface somewhat wrinkled, sparsely to densely pubescent, shiny green.

Plant scrambling up to 10 m. Leaves with petiole 0.7-2.6 cm long; blade broadly-ovate or ovate-elliptic, 5-12 x 2.5-7 cm apex shortly acute, base rounded or truncate. Inflorescences up to 6 cm in diameter, up to 50-flowered; peduncles 1.3-4 cm long; pedicels 0.8-1.5 cm long. Se­ pals 3.3-5.2 mm long, ovate or ovate-lanceolate, sub­ acute. Corolla elongated in bud, c 1.5 cm in diameter, tube 3.3-^1 mm long, globose-campanulate, inside slightly hairy ; lobes frequently somewhat tw isted, 4-5 nun long, narrowly oblong, glabrous. Corona lobes 2.6 mm long, erect, lanceolate, acute, base with a thick free margin, shorter than the adjacent adnate anther membrane. Stigma

Mostly in forest edges, open forest or in disturbed ground; 1600-2400 m; EW GD SU WG AR KF SD BA HA; Nigeria and Cameroun in the west to the Arabian peninsula, Tanzania. Gilbert & Jefford 4320; Moonev 5537; W. de Wilde & B. de Wilde-Duyjjes 6661. 4. D. stelostigma (K. Schum.) Bullock (1957); Marsdenia stelostigmo K. Schum. (1903); Stigmatorhynchus stelostigma (K. Schum.) Schltr. (1913) - types: SD. Tarro Gumbi, Boran, Ellenbeck 2080 & 2086 (B syn. destroyed). M. stefaninii Chiov. (1916).

140. ASCLEPIADACEAE: 28. Dregea, 29. Gongronema. 30. Gymnema

Plant scrambling up to 3-4 m; flowering branches at least :• cin long. Leaves with petioles 0.5-1.5 cm long; blade 1 - 5 x 0 7-3.2 cm, broadly ovate, acute or obtuse, base -ounded to weakly cordate. Inflorescences 1 cm in diame­ er, 5-10-flowered, sessile or subsessile; pedicels rudi­ mentary. c 1 nmi long. Sepals c 4 mm long, oblonganceolate, sub-acute. Corolla rounded in bud, c 5 mm in diameter, tube c 2 nun long, campanulate; lobes c 4.5 mm ong, oblong or more or less lanceolate, obtuse, inside white-villous in the middle. Corona lobes c 0.8 x 0.1 mm, oblong-lanceolate, sub-acute; corona basally rectangular, apically triangular. Stigma head with a thread-like apex, c 2 mm long, exserted from the staminal column, with an ornamented surface, often furrowed or branched. Foili,:ies 4-wingecL veiy unevenly developed, often longitudi­ nally asymmetrical, woody, with short, rigid liairs. On sandy soil; 800-1200 m. SD HA; Somalia, Kenya. Burger 2957; Gilbert et al. 7668; Haugen 2146. 29. GONGRONEMA Decne. (1844) Gymnema subgen. Gongronema Endl. (1838) |Marsdenia auctt. p.p., non R. Br. (1809)]. by F. Albers & S. Gtildenberg Bullock, Kew Bull. 15: 193-206 (1961). Pubescent lianas becoming woody with age; latex white, leaves opposite, heart-shaped, leathery, smooth. Flowers in many-flowered, extra-axillary umbels, sessile or pe­ dunculate, one per node. Sepals ovate. Corolla campanu­ late, lobed to middle; lobes recurved or reflexed, inangular, throat and tube glabrous. Corolline corona ab•;ent. Gynostegial corona of 5 rectangular, f -shy staminal lobes basally adnate to the cylindrical gynostegium, end­ ing in a long subulate apex. Stigma head enclosed by coiona, apex conical, 5-angled; Pollinia erect, ovate or linear, basally attached to translator arms, which are long and kinky; corpusculum linear. Follicles tapering, smooth; seeds with a tuft of liairs. About 12 species in Africa and Asia; 1 in the Flora area. The African species of Gongronema must be included uiMarsdenia R. Br.. if this is understood in the broad taxpnomic sense currently applied for the New World and ;,apuasia. G. angolense (N.E. Br.) Bullock (1961); Marsdenia angolensis N.E. Br. (1895) - type: An­ gola, Welwitsch 4245 (K lecto. designated bv Bullock. 1954). M. gondarensis Chiov. (1911) - type: GD, Dembia. Chiovenda 1741 (?FT holo. not seen). Plant scrambling up to 8 m, roots thick and fleshy. Leaves ith petioles up to c 6 cm long; blade broadly ovate, 4-12 v 2.5-10 cm, apex shortly pointed, base cordate, upper surface pubescent, lower surface more or less tomentose. inflorescences c 4 cm in diameter, 5-14-flowered, with up p 4 branches and 8 cymes; pedimcles c 4 cm long; pedi:els c 0.8-1.3 cm long. Sepals c 1.3 x 1.1 cm. ovate:-blongto elliptic-oblong, obtuse. Corolla white to yellow

153

to green, c 6 mm in diameter; tube c 3.9-5.2 nun long, campanulate; lobes spreading, c 4.4 x 1.3 mm, ellip­ tic-oblong to linear, obtuse, slighty fleshy, outside pubes­ cent, inside glabrous. Staminal corona lobes c 3.9 nun long, equal to or longer than the anthers, linear-oblong, narrowed in the middle, basal half with two keels, broadly winged. Gynostegium including stigma head c 2.3 nun long. Follicles mostly single, c 10-15 cm long, 4-8 nun in diameter, linear, apex more or less densely pilose. Fig. 140.33.1 In forest edge or open forest; 1300-1700 m. GJ KF; widespread in tropical Africa. Mooney 9238; J. de Wilde & Gilbert 286;

30. GYMNEMA R. Br. (1810) Bidaria (Endl.) Decne. (1844). by F. Albers & S. Guldenberg Pubescent shrubby twiners; latex clear or whitish. Leaves opposite, petiolate, elliptic or ovate, base rounded or cuneate. Inflorescences extra-axillarv. ses­ sile or shortly pedunculate, many-flowered umbels, of­ ten 2 opposite umbels at the same node. Sepals small, obtuse, ovate or elliptic, erect. Corolla pale yellow' or whitish, 5-lobed to half-way; tube campanulate or urceolate; lobes fleshy, reflexed. oblong or ob­ long-ovate, minutely notched at apex, glabrous on both sides, base adnate to gynostegium. Corolline corona of 5 fleshy, linear lobes, which are channelled on face and adnate up to mouth of corolla-tube. Gynostegial corona absent. Gynostegium cylindrical; stigma head often lon­ ger than staminal column, conical, papillose. Anthers rectangular. Pollinia erect, small, ovate to linear-ovate; corpusculum more or less as long as the pollinia. Folli­ cles single, slender, acute, smooth; seeds depressed with a clear marked margin, glabrous. Because of the very shortened peduncles, which are often shorter than the pedicels, the two-branched umbel­ late cymes at a node appear like two separate umbels. Therefore, in cases of opposite umbels at a node, they ap­ pear like 4 umbels. More than 25 species in SE Asia and Australia; 1 in .Africa.

G. sylvestre (Retz.) R. Br. ex Schult. (1820); Periplocasylvestris Retz. (1781) - type: Komg s.n. (?BM iso.). Cynanchum subvolabile Schum. & Thoim. (1827); Gymnema subvolubile (Schum. & Thonn.) Decne. (1838). Asclepiasgeminata Roxb. (1832) Gymnema rufescens Decne. (1838). Cynanchum senegalense Sieb. ex Decne. (1844), nom. nud. Gymnema humile Decne. (1844); G. fruticulosum Hochst. (1844), nom. nud. - type: TU. near Gapdia. Schimper 807 (P holo., K iso.). G. m ’kenii Harv. (1868), nom. nud.

140. ASCLEPIADACEAE: ^29. Gongronema, 30. Gymnema, 31.Stigmatorhynchus

154

Figure 140.33. G ONGRO NEM A A N G O L E N SE . 1 - flowering branch x V4. G YM N EM A S Y L V E S T R E 2 - flowering branch x Vr, 3 flower x 10; 4 - fruit x ‘/2. No specimens given in orig. publ. Drawn by N. Labeye (1)& D. 'I roupin (2-4). (Reproduced with permission from FI. du Rwanda IE, A sclepiadaceae, fig. 32 p.p.)

Plant up to 3 m tall, stem 4-15 mm in diameter. Leaves with petioles 0.4-2.6 cm long; blade elliptic, ovateoblong to ovate-lanceolate or rhombic, 2-8.8 x 1-5.5 cm upper surface green, lower surface pale green. Inflores­ cences up to 10-flowered, umbels 1.5 cm in diameter, peduncles 0.2-1.3 cm long; pedicels 0.2-0.8 cm long. Se­ pals 0.7-2.6 mm long. Flowers sweetly scented; corolla up to 5.5 mm in diameter, tube 2-2.6 mm long; lobes c 2.5 mm long. Corona lobes c 1.3 mm long, on inside with a small channel in centre, with a rolled obtuse apex, densely ciliate on both sides. Gynostegium including stigma head c 2.1 nun long. Follicles 5.2-10 cm x 0.6-0.8 cm erect, more or less fusiform, beaked, green-brownish Fig. 140.33.2-4. On sandy to loamy soil, often also at the edges of river beds, sea level to 1600 m. EW TU GD IL GG; widely dis­ tributed in tropical Africa, Madagascar and the dry areas of India. Bally B6621; Gereau 1364; Gilbert & Getachew A. 2898. 31.

STIGMATORHYNCHUS Schltr. (1913) by F. Albers & S. Gvlldenberg

Bullock, Kew Bull. 9: 349-373 (1954). Branched shrubs, sometimes with twining shoots, short shoots veiy leafy; latex white. Leaves shortly petiolate,

ovate, ovate-lanceolate, elliptic, orspathulate, softly pu­ bescent Inflorescences extra-axillary, sessile or shortly pedunculate umbels or clusters with up to 12 flowers; pedicels very short. Sepals ovate or elliptic, softly pu­ bescent. Corolla lobed to middle, campanulate, outside smooth, inside densely hairy, base adnate to gyno­ stegium Corolline corona absent. Gynostegial corona lobes staminal, basally adnate to the back of the anthers, dorso-ventrally flattened. Anther membranes ellip­ tic-lanceolate, more or less pointed. Stigma head elon­ gated and tapering. Pollinia erect, ovoid, attached basally to horizontally oriented translator arms; corpus­ culum elongated. Carpel with two seed chambers for each follicle. Follicles single or paired at 120°, smooth or with an irregular knobbly ridge near the base, beaked; seeds usually one per follicle. 2 or 3 disjunct species in tropical and subtropical Af­ rica, 1 in the Flora area.

S. sp. =J. de Wilde 7216 Shrub 0.5-3 m tall, c 1 m in diameter, branches 30-40 cm long, twining above, becoming woody with age, whole plant with short hairs. Leaves with petioles 1-8 mm long; blade 0.5-1.5 x 0 2—1 cm. Inflorescences 1-12- flowered, sessile, pedicels c 1 mm long. Sepals c 2 mm long, ovate-oblong, adaxially hairy/. Corolla white to cream;

140. ASCLEPIADACEAE: 31. Stigmatorhynchus, 32. Leptadenia

tube 1-2 mm long, campanulate, with 5 densely hairy ar­ eas llong a circumference half-way up tlie tube; lobes eret r. and with a reflexed apex, 1-2 x 0.8 mm. oblong, obtust. basal half or more covered with long recurved hairs, out ide glabrous. Corona lobes shorter or longer than the apk es of anther membranes, basal half wide triangular, api( al half with triangular apex lying on the back of antlieiG ynostegium including stigma head more than 1 mm long; stigma head forming a thread-like apex 0.5-3 mm long, apex sometimes divided in two. [illy dry steppes; 1500 m. HA; central and northern Son alia J. de Wilde 7216. i ollections of this taxon have considerably smaller lea\ ss than the florally more or less identical S. umbelliferu Schltr., a species apparently endemic to Tanzania. The- also have sessile inflorescences as opposed to the shoitlv pedunculate umbels of the latter species. This combination of characters matches the Namibian endemic S. h< reroensis Schltr., which, in addition to tlie features noted above, differs from S. umbelliferus in tlie form of the : :)!licle. In S. hereroensis this is smooth, while in S. umbe.Uferus there is a conspicuous irregular knobbly ridg< :owards the base. Unfortunately Ethiopian and So­ mali material lacks even immature fruit, so its affinities remain obscure. 32 LEPTADENIA R. Br. (1810) by D.J. Goyder

155

1. L. pyrotechnica (Forssk.) Decne. (1838); Cynanchum pyrotechnicum Forssk. (1775); Sarco­ stemma pyrotechnicum (Forssk.) Schult. (1820); Microlomapyrotechnicum (Forssk.) Spreng. (1825) type: Yemen, Forsskal (not traced). Leptadenia spartium Wight. & Am. (1834). L. jacquemontiana Decne. (1838). L. gracilis Decne. (1838). Much branched leafless shrub or small tree 2-4(-6) m tall, rarely with linear leaves on young shoots; branches slen­ der, erect, terete, minutely pubescent at tip, otherwise gla­ brous. Inflorescences minutely pubescent with up to 10 flowers at one time, flowers developing successively and forming a floral axis 1-4 mm long; peduncles 1-2 mm long; pedicels 1-2 mm long. Sepals c 1 mm long, ovate, subacute, minutely pubescent. Corolla cream oryellowish green; tube c0.5 mm long, glabrous within; lobes 1.5-2 x 1 mm, ovate, acute, densely pubescent on both surfaces apart from a glabrous, non-fleshy triangle at the base of tlie lobes witliin; corolla lobe sinuses with squat, fleshy, corolline corona lobes c 0.3 x 0.5 mm. Gynostegium c 0.5 mm long. Follicles 9-12 x 0.5 cm, narrow'ly fusiform, gla­ brous. Sandy desert and dry river beds at low altitudes; grow­ ing around sea level in Eritrea, recorded from c 700 m in Harerge. EE AF HA; widely distributed around the mar­ gins of the Sahara and eastwards through Arabia to Baluchistan and Sind. Ash 714; Burger 3817; Mooney 8568.

Bullock, Kew Bull. 10: 287-292 (1955). Leaftess shrubs or woody twiners with well developed leaves latex clear and watery. Inflorescences with many flow rs in subsessile or pedunculate, extra-axillary umbi Is Corolla rotate with a short campanulate tube and flesh ii. pubescent or lanate lobes with slightly revolute margins. Corolline corona lobes present in tlie corolla lobe inuses. Gynostegium largely enclosed by the corolla tube and corolline corona lobes, with or without an annu­ lar corona at the base. Anthers leaning over the stigma he ld, lacking apical appendages. Pollinaria subhorizontal or su'orect; corpusculum minute, reddish brown; transla­ tor aims translucent, flattened and obtriangular; pollinia somewhat flattened and with a translucent apical germinatio ' zone. Follicles developing singly, fusiform with a long- irtenuate beak, slender or stout; seeds flattened, smoo h. with a silky coma.

2. L. arborea (Forssk.) Schweinf. (1912); Cynanchum arboreum Forssk. (1775) - type: Ye­ men, Forsskal 271, 273 (C syn.); Forsskdl s.n. (LD syn.). C. heterophyllum Del. (1826); Leptadenia delilii Decne. (1844). L.forskalii G. Don. (1837). L. abyssinica Decne. (1844) - type: TU/GD, Tacazze stream. Schimper 632 (P holo., K iso.). L. clavipes S. Moore (1912)-type: BA/HA, Schebelli, Donaldson Smith (BM holo.).

Tw ining shrub; branches minutely pubescent at tip, other­ wise glabrous. Leaves with slender petiole (0.5—)1—3 cm long; blade extremely variable in shape even on tlie same shoot, elliptic to broadly ovate or rarely hastate, 2—6(—9) x (0.5—)1—5 cm, apex subacute to attenuate, base cuneate to 4 species bordering the Sahara and extending east­ truncate or cordate. Inflorescences with 5-20 flowers in ward Ho Arabia, the Indian subcontinent and Madagascar, an umbel, minutely pubescent; peduncles 5-12 mm long; 3 spe :ies in the Flora area. pedicels 5-10 mm long. Sepals 0.5-1 mm long, ovate, subacute, densely pubescent. Corolla cream or yellowish 1. E nect shrubs, leafless or with few linear leaves on green; tube 0.5-1 mm long, glabrous within; lobes 2-2.5 x oung shoots only. 1. L. pyrotechnica 1 mm, oblong, acute, densely pubescent on both surfaces - 1 * ining shrubs with broad, w'ell developed petioapart from a glabrous, non-fleshy triangle at the base of late leaves. 2 tlie lobes within; corolla lobe sinuses with minute, fleshy 2. C irolla lobes 2-2.5 mm long; sepals shorter than erect corolline corona lobes c 0.4 mm long with an apical he corolla tube. 2. L. arborea tuft of erect liairs. Gynostegium c 1 mm long. Follicles - C irolla lobes 4—5 mm long; sepals longer than the 6-8 x 1 cm, lanceolate in outline, stout, smooth or some­ oroUatube. 3. L. hastata what vemicose. glabrous.

156

140. ASCLEPIADACEAE: 32 Leptadenia. 33. Conomitra

Figure 140.34. L E P T A D E N IA H A S T A T A 1 £ 2 habit showing polymorphic leaves x 14; 3 - fruit x Vi\ 4 - flower x 4; 5 -p o llin a r­ ium x 20; 6 - seeds x 2. 1 ,2 ,4 & 5 from Twee die 2114; 3 from Mwangangi & Gwynne I 148; 6 from Newbould 6935. Drawn by E. Papadopoulos.

Scrambling over bushes in hot dry regions, sometimes in dry river beds. 250-1400 m. EE AF EW TU G D G J SU SD BA HA: widely distributed from Niger to the Arabian peninsula. A sh 1930; E d w a r d s & R a o 3534; G ilb ert et al 7619. 3. L. hastata fPers.) D ecne. (1844); C ynanch u m h a sta tu m Pers. (1805); C. la n ceo la turn Poir. (1811) - type: ‘Africa’ (L holo.). C. lancifo liu m Schum. & Thonn. (1827); L e p ta ­ d enia la n cifo lia (Schum. & Thonn.) Decne. (1838). C ynanch u m sca b ru m Schum. & Thonn. (1827). L. la n cifo lia var. sc a b ra Decne. in DC., P ro d r 8 628 (1844).

densely pubescent. Corolla white to cream or orange, be­ coming darker with age; tube c 1 nun long, glabrous within; lobes 4-5 x 1 inm. oblong, acute, upper surface lanate with white hairs, lower surface pubescent; corolla lobe sinuses with fleshy corolline corona lobes c 1.5 nun long, obiong, sublanate, erect or leaning over head of gy nostegium Gy nostegium c 1.5 nun long. Follicles 8-11 x 1-1.5 cm. lanceolate in outline, stout, glabrous. Fig. 140.34. Scrambling over bushes in hot dry regions, sometimes in di> river beds; 300-1600 m. TU GD GJ SU IL KF GG SD B A ; widely distributed across the semi-and belt south of the Sahara from Mauretania to northern Kenya and Ethiopia F rits et al. 2515; G ilb e rt et al. 7320; J. de W ilde & G ilb ert 418.

Twining shrub; branches minutely pubescent. Leaves with petiole 1-2 cm long; blade extremely variable in shape, elliptic to broadly ovate or rarely hastate, 2.5-8 (-12) x 1.5-4.5 cm. apex subacule to attenuate, base cuneate to truncate, minutely pubescent. Inflorescences minutely pubescent, w'ith c 20 flowers in a crowded umbel; peduncles 5-12 mm long; pedicels 2-5 mm long. Sepals 1.5-2.5 mm long, oblong-lanceolate, subacute.

33. C O N O M IT R A F e n zl (1839)

by D. J Goyder Field, Kew Bull 37 344-347 (1982). Ephemeral heib. latex not recorded but probably clear. Flowers 1-2. extra-axillary. Corolia with a short tube;

140. ASCLEPIADACEAE: 33. Conomitra

157

Figure 140.35. C O N O M ITRA L IN E A R IS: 1 & 2 habit x V2, 3 - flower x 10; 4 —flower bud x 10; 5 - pollinarium x 20. 1 & 4 from Gilbert & Thulin 1260A; 2 ,3 & 5 from Stantuird t' adpressed hairs. Leaves: petiole 1.3-3 cm; blade lance as long as mbe overall, c 2.2 mm w ide, upper part of tu t.! c 1 nun wide, interior pale with dark band at top of basal swelling, throat with pairs of whitish stripes oppo­ site lc bes; mouth oblique; corolla lobes strongly recurved, lineai revolute for most of its length, c 4.5-8 nun long, black :h-green. margins strongly revolute, very minutely velve ■papillate, otherwise glabrous. Fig. 140.36.11-13. Sliaded sites in Oxytenanthera thickets and degraded decid H-us woodland; 1300-2200 m. WG GG; not known elsew lere. Gilbert & Thulin 500, 627. 753. 7. C. nicrogaster M.G. Gilbert (2003) - type: WG, 23 km from Ghiinbi on road to Asosa (9§2D’N;35C*38’E). Gilbert & Thulin 775 (K holo., ETH UPS iso.). Twin igor trailing herb. Rootstock a flattened globose tu­ ber uj !lo 5 cm wide, sometimes forming chains of ovoid tuber; Stems with two lines of short adpressed hairs. Leave-r petiole 1.8-3.5 cm; blade ovate, 4.7-7 x 2.6-4 cm. b i:-e cordate, apex acuminate, thinly pubescent. Infloresci ncc a in,my-flowered. subsessile pseudumbel; pedunclc up to 2 mm; pedicels 6-11 mm. Sepals up to 3 mm long, acuminate. Corolla distinctly zygomorphic, 9-11 nun la mg, glabrous outside, pale grey; corolla mbe 7-8 mm, J -shaped, ventricose, basal swelling more than half as lon;! as tube overall, c 3.5 mm w ide, with pale interior, upper j iirt of tubeshaiply delineated, c 1.5 mm wide, with black) h interior, throat with pairs of whitish stripes oppo­ site lobes; mouth oblique; corolla lobes spreading to slightly reflexed, triangular, c 2 x 2 mm, blackish, margins slight! revolute, purple-pubescent around sinuses, other­

Erect to twining herb; all parts except corolla glabrous. Rootstock a more or less globose tuber. Leaf blade nar­ rowly lanceolate. 5-7.5 x 3.5-0.5 cm. base attenuate into poorly defined petiole, apex acute. Inflorescence a pe­ dunculate pseudumbel; peduncle up to 15 nun; pedicels 3-6 nun. Sepals 2.5-3 mm. Corolla ascending, 17-28 nun; corolla tube 13-23 mm, pale green, mostly mottled with dark red outside, dark red inside, basal swelling subglobose. 4-4.5 mm wide, tube c 2.2 nun wide, widen­ ing to 4.5-6 5 mm at mouth; corolla lobes more or less lin­ ear. 5-8 mm. purple veined at base, upper part uniformly green with darker margin and brown ciliate hairs. Corona wliite, outer lobes shallowly emarginate with acute to slightly acuminate comers, slightly higher than bases of inner lobes; inner lobes connivent-recurved Fruit not seen. Fig. 140.36.1-3. Variously in degraded Terminalia brownii w'oodland. deciduous busliland with Dodonaea. and on massive gra­ nitic outcrops with Euclea; 1650-2100 m. EW TU SU HA; Somalia, ?Kenya. Arabia. M. & S. Gilbert 2200; Gilbert Getachew 2839; Gilbert & Thulin 19. 9. C. inomata Masinde (1998) - type: Kenva. Masinde et al. 838 (EA holo.. K MSUN iso.). Twining herb; all parts glabrous. Rootstock a subglobose tuber, or sometimes a chain of tubers, each formed at the node of a rhizome. Stems slender. Leaves ovate, fleshy. Inflorescence a 1- or 2-flowered pedunculate cyme. Co­ rolla: basal inflation conspicuous, subglobose. pale green, rest of corolla pale yeliow. sometimes flushed reddish; co­ rolla tube more or less cylindrical, c 1/3 as wide as basal inflation and widening only slightly towards mouth in­ side glabrous; corolla lobes narrowly oblong-spathulate. remaining attached at tip to form narrowly obovoid ‘cage’, narrower than the basal inflation. Outer corona lobes each with 2 triangular teeth, c 1/5 as long as inner lobes, ciliate; inner lobes connivent, erect, dark at base. Fig. 140.36.4 & 5.

162

140. ASCLEPIADACEAE: 34. Ceropegia

Deciduous bushland; 1000-1400 m. SU GG BA; Kenya. M. Gilbert 3936, 3980; Gilbert & Thulin 253. The very fleshy glabrous leaves and the tendency to produce chains of tubers distinguishes vegetative material of tliis species from all other Ethiopian members of the ge­ nus. M. Gilbert 3936 differs from the other material by the wider corolla lobes which form a vcage' almost as wide as the basal inflation. 10. C. sp. = M & S. Gilbert s.n. - type: GJ, Tississat (Blue Nile) Falls, grown in Addis Abeba.M & S. Gilbert s.n (photos). Hysteranthus herb, possibly pyrrhophytic. Rootstock a subglobose tuber. Vegetative stem trailing. Leaves linear lanceolate (not preserved). Flowering stem erect, more or less leafless. Corolla erect, outside glabrous, grey, faintly streaked and blotched with dull red. smelling strongly of lemons at anthesis; basal swelling ovoid, c l/5tli length of tube as a whole, mouth of tube about as wide as basal swelling; corolla lobes triangular-lanceolate, c 1/2 as long as tube, rotate to slightly reflexed, revolute, lobes and mouth of corolla tube yellow with large black blotches which decrease in size towards tips of lobes, lobes densely pilose. Growing amongst grasses in broad-leafed deciduous woodland. GD/GJ boundary; known only from a single collection. This species is related to a group of Southern and West African species including Ceropegia dinteri Schltr. and C. campanulata G. Don. It is the only Ethiopian species with obviously (pleasantly!) scented flowers. Unfortu­ nately the only material was a single flower, preserved in spirit, that lias been lost, which means that it is not possi­ ble to formally name this extremely distinctive species. 11.C. subaphylla/^. Schum. (1903) - type: Somalia, near Dadab. Ellenbeck 190 (B holo. destroyed). C. nuda Hutch. & Bruce (1941) - type: Somalia. Afard (10 10’N 44 08’E), Gillett 4484 (K holo.) Twining herb. Rootstock a more or less cylindrical tuber which narrows only gradually into the stem. Stems slen­ der but fleshy. Leaves lanceolate, up to 1 cm long, very rapidly lost. Inflorescence a sessile 1- to few-flowered cyme. Corolla up to 6 cm long overall, outside puberu­ lous, glabrescent; tube 15-35 mm long, basal swelling cy­ lindrical. l/4th length of tube, narrowing more or less abruptly into tube, which then widens gradually to mouth

about as wide as basal swelling, colour not recorded, inte­ rior ciliate; corolla lobes 10-25 mm, narrowly triangularlanceolate, narrowly auriculate at base, ciliate inside to­ wards base, remaining attached at tips to form a corucal-ovoid, slightly acuminate cage Fig. 140.36.19& 20. OpenAcacta - Commiphora busliland overlying lime­ stone; c 500 m. HA; Somalia. Gillett 4190. The combination of the clear to more or less cloudy la­ tex. fleshy stems w ith reduced leaves and the rather elon­ gated basal tuber, which is not sharply differentiated from the stem enables the identification of sterile material within this complex, which is probably more widespread in the Ogadeu region than the current lack of records sug­ gests. Ceropegia subaphylla was described simulta­ neously with ('. botrys K. Schum. from what appears to have been a mixed collection. The primary difference be­ tween them is the presence of hairs on the outside of the corolla in C. subaphylla, in contrast to the glabrous out­ side in the better known C. botrys, for which there is ex­ tant type material and which is relatively well know n in Arabia (Bruvns, Notes R. Bot. Gard. Edinburgh 45(2): 287-326, 1988). 12. C. variegata Decne. var. com igera Huber in Mem. Soc. Brot. 12: 141 (1957); Ceropegia devechii Chiov. (1932) - type Soma­ lia. Senni 459 (FT holo ). Climbing herb, glabrous except for the corolla. Base of stem swollen in seedlings but otherwise no basal tuber or fusiform roots Stems very fleshy, up to 1 cm thick, smooth, usually irregularly blotched with pinkish-brown, tw ining only towards tip. Leaves reduced to fleshy ovate scales, soon falling. Inflorescence a pedunculate, fewflowered, raceme-like cyme; peduncle longer than pedi­ cels. Sepals more or less linear. Corolla up to 6 cm long, glabrous outside; corolla tube up to 45 mm, basal swelling double, the lowermost ovoid, uniformly grey, demarkated from upper swelling by a well-defined constriction mark­ ing position of internal annulus, upper swelling spotted with red, the 2 together c 1/2 as long as tube, which w idens gradually to mouth, interior glabrous except for band of cilia at top of upper chamber, corolla lobes up to c 16 mm but with the sinuses between them produced into very prominent oblong lobes about as long as corolla tube, out­ side grey spotted with red. inside brilliant white w ith dark red tips to lobes, uniformly pilose. Outer corona of 10 slender, erect lobes; inner corona lobes similar to outer lobes. Follicles fusiform-cylindrical with blunt, more or less swollen tips. Fig. 140.36.21.

Figure 140.36. CERO PEG IA species, mostly with tubers (sp 12 without tuber, sp. 27 with swollen roots). Habit and base o f plant xVs; leaves x 4/s; corollas x l'/j, coronas x 11. C. V1GNALDIANA: 1 - base o f plant with subglobose tuber; 2 - leaf; 3 - corolla. C. IN O RN A 7 /1 :4 - leaf; 5 - corolla. C RIN G EN S: 6 - range o f leaf forms, 7 - corolla, C. M IC R O G A STE R : 8 - leaf; 9 - corolla; 10 corona. C. R EC U R V A TA . 11 - leaf; 12 - corolla; 13 - corona. C A B Y SSIN IC A . 14 habit showing characteristic erect stem. 15 leaf; 16 corolla. C. LO R A N T H IF L O R A : 17 - leaf; 18 - corolla. C SU BA P H YLLA : 19 - base o f plant with tapered tuber; 20 - co­ rolla. C. V AR IE G A TA var. C O RN IG ERA: 21 - corolla. C. B A R B IG E R 4 2 2 -p o rtio n o f stem show ingstriationsand scale leaves x 3; 2 3 -corolla. 1-3 fromM. & S. Gilbert 2200; 4 & 5 from Gilbert & Thulin 2 5 3;6& 7 from Gilbert & Getachew 2850; 8-1 0 from Gilbert & Thulin 775; 11-13 from Gilbert & Thulin 627; 14 from Ash 2142; 15 & 16 from A/. Gilbert 4024; 17 & 18 from Friis & Lawesson 5333; 19 from Gillett 4190; 20 from Ellenbeck 190; 21 fronvphotographs o f cultivated plant grown from material collected between Harar and Jigjiga, 22 & 23 from clonotype material grown under RBG Kew Acc. 443.77.04023. Drawn by M argaret Tebbs.

140. ASCLEPIADACEAE: 34. Ceropegia

164

140. ASCLEPIADACEAE: 34. Ceropegia

Acacia - Commiphora busliland; c 1100 m. HA; So­ malia. Kenya; Arabia. Burger 1749.

linear with short triangular base, apical part ciliate. app,irently cohenng to form a slender beak.

A species with a corolla that is remarkable even by Ceropegia standards. The Ethiopian material is very uni­ form and a good match for the Somalian type. However, matenal from .Arabia and Kenya shows a lot of variation in the length of both corolla lobes proper and the sinus lobes, connecting it to Ceropegia variegata Decne. var. variegata, which differs only by the much shorter sinus-lobes and is recorded from Arabia and Kenya.

Acacia - Commiphora - Kirkia busliland overlying limestone: 1350-1400 m. SD BA; not known elsewhere. Friis et al. 2751, 3679.

13. C. somalensis Chiov. (1916) - type; Somalia. Paoli 889 (FT holo.). Ceropegia somalense forma erostrata Huber, Mem. Soc. Brot. 12: 90 (1957) - type: HA/Somalia boundary. 8°37’N; 45°08’E, Gillett 4192 (K holo.). Twining herb; most parts minutely puberulent. Tuber or fusiform roots absent. Stems succulent rapidly glabrescent. Leaves: petiole up to 6 mm; blade ovate, more or less fleshy, up to 11x11 mm on flowering shoots, base cordate to rounded, apex rounded-cuspidate to acuminate. Inflorescence a pedunculate pseudumbel; peduncle up to 11 mm; pedicel up to 10 mm. Sepals c 3.5 x 1 mm. Corolla ascending, usually strongly curved near base, up to 40 mm densely puberulent outside, inside ciliate at base and upper part of tube, glabrous between; corolla tube funnel-sliaped. up to c 20 mm. basal swelling slight up to 3.5 mm wide, tube c 2.5 mm wide for half length, then widen­ ing to c 15 mm at mouth, pale green with purplish blotches outside, basal half deep purple inside; lobes as long as tube, with triangular base and linear-spathulate tip. lower part strongly revolute, finely purplish-mottled, upper part uniformly greenish, sometimes remaining attached at apex only and forming more or less cylindrical ‘cage', other times basal part forming conical ‘cage’, and spalhulate tins cohering to form a clavate beak, sometimes apical part absent (forma erostrata). Outer corona lobes short, erect. 2-toothed; iiuier corona lobes connivent into col­ umn. recurved at Ups. Fig. 140.37.14 & 15. Volcanic ash with sparse cover of slirubs; c 10001100 m. SU HA; Somalia, Kenya. Gilbert & Thulin 200; Gilten 4192, 4212. The corolla often lias the lobes cohering below the tip to form a characteristic club-shaped extension to a more or less conical cage’ but sometimes the lobes are com­ pletely free and the cage is then cylindrical with a blunt apex. 14. C. sp. = Friis et al. 2751 SD. 60 km SE Negele on track to Wolensu. Closely related to C. somalensis. Leaves: petiole up to 11 mm; blade oblong-ovate, up to 6 x 3 cm, base rounded, apex rounded-cuspidate. Peduncle up to 35 mm; pedicel up to 12 mm. Sepals c 3 x 1 mm. Corolla c 20 mm, de­ scribed as “w hite”, glabrous outside, ciliate inside, appar­ ently straight; corolla tube cy lindrical, only slightly wider at base and apex, unmarked; corolla lobes as long as tube.

Known only from one collection with a single bud. and another with very poorly preserved flowers. It is also rather similar to the West African species Ceropegia rhynchantha Schltr. 15. C. aristolochioides Decne. (1838) - type: Senegal, Heudelot M l (P holo.). subsp. aristolochioides Ceropegia beccariana Martelli (1886) - type: EW, Abita, Keren, Beccari 306 (FT holo., K photo, draw­ ings & fragment) Twining herb, often also w ith arched stolons. Tuber or fusiform roots absent. Stems succulent rapidly glabrescent Leaves petiole up to 11 mm, puberulous, blade ovate, up to 35 x 25 mm. base rounded to shallowly cordate, apex acute to slightly acuminate, more or less fleshy, glabrous except for margins. Inflorescence a pedunculate pseudumbel. most parts glabrous; peduncle 20-40 mm. descending vertically , pedicels 5-10 mm. Co­ rolla ascending to erect. (16—)27—32(—39) mm, yellow or pale green flushed dark red on upper part of tube and on lobes, glabrous outside, ciliate inside, tube (12-)20-27 mm. basal swelling ovoid, up to c 1 x 6 mm. tube 2.5-3 mm wide, w idening to 7-10 mm; corolla lobes ovate, 4-8(-14) mni, strongly revolute, remaining attached at apex to form rounded cage'. Corona as in C. somalensis. Outer corona lobes erect notched fore 1/3 length, c 2/3 as long as inner lobes, purple, ciliate, inner lobes white Fol­ licle up to 12 cm long Fig. 140.37.8 & 9. Acacia woodland, evergreen bushland with Euclea, overlying granites and basalts, 1200-1700 m. EW AR GG; Uganda, west to Senegal. Gilbert & Getachew 2744; Gilbert good match with the type from Soma­ lia, and indicates that Bruyns was probably wrong to dismiss this very distinctive plant as just a variant of E. planiflora.

140. ASCLEPIADACEAE: 38. Echidnopsis

181

^ E C H ID N O P SIS M O N T A N A . A - stem tip x 6; 2 & 3 - buds x 3 Vi; 4 - corolla side view x 3 '/ 3; 5 - corolla top view x 3 '/ 3; I*®. —corona an/3; 7 - two corolla lobes seen from side, showing opening between them x 4; 8 -c ilia te hairs from inside of corolla x 20; 9 - corona side view x 7; 10 - corona top view x 7; 11 & 12 - corona longitudinal section x 7; 13 pollinarium x40. 1-10, 12-13 drawn from material cultivated at Karoo Botanic Garden, most probably clonotype material o f Lavranos 6721; 11 from W. de Wilde & M. Gilbert 443. Adapted from Bruyns. Bradleya 6: 44. fig. 29 (1988).

Flowering material which shows that this is a distinct species most closely related to P. cubiformis (Bally) Bally has recently been collected by Maurizio Dioli. He intends to describe this species as Pseudolithos gigas.

saccate: inner lobes erect with swollen rips, spinytuberculate on their backs. A single, variable, taxonomically isolated species en­ demic to the Somali-Masai region including tropical Ara­ bia.

41. EDITHCOLEA N.E. Br. (1895) by M. G. Gilbert Stems creeping, much branched and mat-forming, branches divaricate, not articulated; tubercles spirally arranged in 5 ranks, conical with thom-like, brown, callused tips. Latex clear. Leaves not apparent. Inflorescen­ ces extra-axillary from near tip of stem, sessile, 1-(or 2)-flowered; pedicels moderately long. Corolla very large, rotate to slightly reflexed with a relatively very small tube, divided about half-way to the centre with a broad limb and short, usually strongly recurved, acu­ minate lobes, small recurved sinus lobes present, outside smooth, inside strongly concentrically rugose. Corona enclosed within corolla tube; outer lobes more or less

E. grandis N.E. Br. (1895) types: Somalia, Henweina Valley, Cole s.n (K syn.) & Lort-Phillips s.n (K syn.). Stems usually flushed dark red with very sharp yellowbrown tips to the tubercles. Corolla up to 10(—12.5) cm wide, usually predominantly purplish brown with intri­ cately arranged yellowish spots and streaks, less often predominancy yellowish with relatively few purplish markings; tube c 6 mm wide with prominently raised mar­ gin; limb with radiating lines of clavate, purpie hairs op­ posite the sinuses; lobes with centres patterned as the limb but with the strongly recurved tips smooth, uniformly pur-

140. ASCLEPIADACEAE: 41. Edithcolea ,42. Pachycymbium

pie-brown or yellow and sharply delineated by a line of hairs. Follicles broadly fusiform, narrowly divergent. Deciduous busliland. mostly on volcanic substrates; 1 100-1300 m. SU GG SD HA; Somalia, Kenya, Tanza­ nia; Yemen. Gilbert & Phillips 8886. Sometimes called the “Persian Carpet Flower’ bese of the very large, ricliiy patterned corolla.

+

42. PACHYCYMBIUM Leach (1978) Angolluma R. Munster (1990). by M. G. Gilbert CVilbert, Cacl. Succ. J. Gr. Brit. 40: 39-50 (1978) & Bradleva £ 20-28 (1990); Plow es, Excelsa 16: 103-120(1993).

I .'warf succulent herbs with clear latex. Stems erect or as­ cending, usually forming dense clumps, less often sobo­ liferous with solitary stems or prostrate and mat-forming; si sms with 4 (rarely 5 or 6) ranks of tubercles, often narked with dark streaks and blotches; tubercles conical ttapered. Leaves subulate, forming soft point to tubercle, often very poorly differentiated, rarely w ith minute stipels at base. Inflorescences extra-axillaiy from near tip of ssm , rarely from lower part, usually sessile, few- to seve nil-flowered, flowers usually opening in succession; f ;.‘dicels short to well developed. Corolla more or less rot ite to deeply tubular, glabrous to setose, rarely with ciivate hairs along margin; corolla tube very short to lon­ ger than lobes, sometimes with narrow annulus; corolla lobes ovate to linear revolute. Corona double; outer cor >na more or less annular or w'lth saccate outer lobes, r i.rely with outer lobes bifid; inner lobes staminal. leaning cm. anthers, occasionally w ith basal horn alternating with cuter lobes, rarely with dorsal projections. Pollinaria c implex: corpusculum prominently winged; translators attached to wings; pollen masses D-shaped, large. A genus of c 35(-41) species with the greatest diver­ sity of species in Ethiopia, Kenya, Somalia, and tropical • .rabia, but with outlying species in extreme western Afr ca (Senegal and Morocco) and south tropical Africa to i doth Afnca; 9 species in the Flora area. As with Caralluma, Plowes takes a more divisive ^ new than the author of this account. He places all the I Ihiopian taxa in the genus Angolluma. The differences used to separate the two genera are all essentially quanti­ tative and none are unique: Pachycymbium in the very i ;irrow sense preferred by Plowes just represents a particiilarlv distinctive combination of characters. Bruyns (Aloe 37: 72-76, 2000) lias suggested a change in the opposite ( rection and included Pachycymbium within a consideraWy enlarged Orbea Haw.. a genus otherwise entirely 5outhem African except for one species in East Africa and anl anomalous species in Somalia, recently placed in it's ( wn monotypic genus. 1 Corolla yellow to brown or dark red, glabrous, mi­ nutely asperulous or with prominent hairs on mar­ gin; corona pink, brown or yellow; inner corona lobes entire or w ith short bluntly triangular teeth. 2

-

185

Corolla lime green, lobes with long colourless hairs terminating prominent fleshy papillae; co­ rona white; inner corona lobes produced into 2 or more slender teeth. 7. P. circes

2. Corolla rotate to very shallowly campanulate with the corona exserted from the corolla tube. 3 - Corolla deeply campanulate to tubular with the co­ rona included within the corolla tube. 8 3. Corolla lobes lanceolate to ovate, only slightly revolute near tip; inner corona lobes without dor­ sal projections; stems in dense clumps. 4 - Corolla lobes linear-revolute; inner corona lobes with one or more dorsal projections at base; stems soboliferous, solitary or in small groups. 6. P. baldratii 4. Outer corona ± annular or with flat lobes exten­ sively fused laterally to each other. 5 - Outer corona lobes deeply saccate. 7 5. Corolla with very small tube completely filled by the corona and delimited from corolla limb by a narrow' but distinct annulus; corolla epidermis rugose to velvety papillose, papillae sometimes tipped by minute hairs; margin of outer corona fleshy, smooth and rounded. 6 - Corolla rotate to very shallowly bowl-shaped, without any trace of an annulus; corolla epidermis smooth, glabrous; margin of outer corona thin, ir­ regularly crenulate. 3. P. gemugofanum 6. Corona more than 7 mm wide; inner corona lobes concealing stigma cap; pollen masses greenish. 2. P. laticoronum - Corona less than 6 mm wide; inner corona lobes barely longer than anthers and leaving white stigma cap exposed; pollen masses yellow. 1. P. sprengeri 7. Pedicel shorter than calyx; buds patent. -

4. P. gilbertii Pedicel more than twice as long as calyx; buds deflexed 5. P. abayense

8. Corolla tubular, erect, glabrous or uniformly puberulent; stems in dense clumps. 8. P. saccu latum - Corolla campanulate, horizontal or deflexed, mar­ gins with prominent daik club-shaped hairs; stems markedly soboliferous, mostly solitary' w'ith few branches. 9. P. vibratilis 1. P. sprengeri (N.E. Br.) M.G. Gilbert (1990); Caralluma sprengeri N.E. Br. (1895); Angolluma sprengeri (N.E. Br.) Plowes (1993); Orbea sprengeri (N.E.Br.) Bniyns - type: TU. near Adowa (Adwa), Petit (K holo.). Steiijs erect to procumbent, up to 15 cm tall, sometimes soboliferous. Buds horizontal to deflexed, acute to rounded. Corolla rotate to more or less reflexed, deeply divided; tube very' shallow' and hidden by corona, delim­ ited by veiy narrow, often poorly defined annulus hidden by corona; lobes lanceolate to ovate, sometimes revolute or recurved to slightly reflexed, yellowish brown to dark

140 ASCLEPIADACEAE: 42. Pachycymbtum

186

chestnut brown, variously smooth, minutely velvety papillose or rugulose, sometimes minutely hairy. Outer corona annular, yellow or dark brown; inner corona lobes procumbent on, and barely longer than anthers, leaving white stigma cap exposed. 3 subspecies are recognized. 1. Corolla glabrous, lobes 9.6-12(-13.4) mm. - Corolla minutely hairy (epidermal cells tipped with simple hairs), lobes (13.5-)16-20 mm long. subsp. sprengeri 2. Corolla lobes reflexed, never revolute, smooth, minutely velvety-papillate. subsp. foetidum - Corolla lobes revolute, never reflexed, rugose but lacking distinct papillae. subsp. ogadense subsp. sprengeri Stems erect or ascending up to 15 cm never soboliferous. Buds acute. Corolla lobes lanceolate, 13.5-20 x 5.7-7 mm, slightly revolute, uniformly light greyish-brown to tan, minutely papillate, each papilla terminated by minute colourless hair. Outer corona 4.3-5.2 mm wide, dull yel­ low. Pollen masses yellow. Figs 140.47.1; 140.48.1-3. Dry evergreen bushland, often dominated by Euclea schimperi; 1600-2000 m. EW TU; not known elsewhere. M. Gilbert (photo). subsp. foetidum (M.G. Gilbert) M.G. Gilbert (1990); Caralluma sprengeri subsp. foetida M.G. Gilbert in Cact. Succ. J. Gt. Brit. 40: 42 (1978); Angolluma foetida (M.G. Gilbert) Plowes (1993); Orbea sprengeri subsp. foetida (M.G. Gilbert) Bruyns in Aloe 37: 76 (2000) - type: HA, 8 km S of Babile along track to Fiche, cult. Addis Abeba, M. Gilbert 2297 (K holo., ETH iso.). Stems procumbent up to 7 cm tall, sometimes shortly soboliferous. Buds truncate to broadly acute, up to 5 in a cluster. Corolla lobes ovate, 11-13.4 x 6.2-7.1 m m flat or slightly recurved, light brown to chocolate brow a some­ times with a paler ring around corona, apparently smooth and glabrous. Outer corona 4.7-6 mm wide, very dark brown. Pollen masses dark yellow, almost orange. Acacia bushland; 1100-1600 m. SU HA; not known elsewhere. subsp. ogadense (M.G. Gilbert) M.G. Gilbert (1990); Caralluma sprengeri subsp. ogadensis M.G. Gilbert, loc. cit.: 43 (1978); Angolluma ogadensis (M.G. Gilbert) Plowes (1993); Orbea sprengeri subsp. ogadensis (M.G. Gilbert) Bruyns in Aloe 37:76 (2000) - type: HA, 6 km SE of Jijiga on road to Degeh Bur, J. de Wilde & Gilbert 6481 (WAG holo., ETH iso.). Stems procumbent up to 7 cm tall, usually soboliferous, underground stems up to 20 cm. Buds suberect, subacute. Corolla lobes lanceolate, 9.6-11 x 5-5.5 mm, spreading to slightly erect, revolute, dark reddish brown with distinct narrow pale ring adjacent to corona, markedly rugose.

glabrous. Outer corona 4.5-4.9 mm wide, yellowish or pale pinkish brown; inner corona lobes yellowish with pinkish tips. Pollen masses yellow. Open bushland overlying limestone, with shrubby Lamiaceae and Acanthaceae; 1500-1800 m. HA; Soma­ lia. M. Gilbert 2041a. 2. P. laticoronum (M.G. Gilbert) M.G. Gilbert (1990); Caralluma sprengeri subsp. laticorona M.G. Gilbert, loc. cit.: 43 (1978); Angolluma laticorona (M.G. Gilbert) Plowes (1993); Orbea laticorona (M.G. Gilbert) Bruyns (2000) - type: SU, c 5 km S of Debra Zeit (Bishoftu), Arenguardi Hayk (‘Green Lake’), cult Addis Abeba, M. & S. Gilbert 1383 (K holo., ETH iso.). Stems procumbent up to 10 cm tall, never soboliferous. Buds horizontal to deflexed, blunt, rarely apiculate. Co­ rolla rotate to more or less reflexed, 2.5-3 cm wide, me­ dium to light brown; tube Filled by corona; corolla lobes 8.4-9.3 mm, often with narrow annulus Corona rotate, 7-8 mm wide, very dark brown, margin thickened, glis­ tening with nectar at anthesis; inner corona lobes inter­ locking over and concealing stigma cap. Pollen masses greenish or yellow Figs 140.47.2; 140.48.4 & 5. Acacia woodland on basalts and volcanic ash; 14002000 m. SU AR; N Kenya, Sudan. Edwards & Tewolde B.G.E. s.n.: Gilbert & Tewolde B.G.E. 2472; M. & jC. Gilbert 3968. 3. P. gemugofanum (M.G. Gilbert)M.G. Gilbert(1990); Caralluma gemugofana M.G. Gilbert (1978); Angolluma gemugofana (M.G. Gilbert) Plowes (1993); Orbea gemugofana? (M.G. Gilbert) Bruyns (2000) - type GG, E of Arba Minch, base of escarp­ ment along track to pumping station, cult. Addis Abeba,M. & S , Gilbert 1731 (K holo., EA ETH iso.). Stems more or less erect less often procumbent, excep­ tionally up to 22 cm, usually less; tubercles long attenuate, up to 22 mm. Pedicel 3.5-5 mm. Sepals 3.2-5 mm. Co­ rolla rotate to very shallowly bowl-shaped, 2-2.5 cm wide, yellow to light tan, smooth and glabrous; corolla tube 7.8-8.3 mm wide, (1—)2.5—3.5 mm deep; corolla lobes 4-5 mm wide, slightly revolute. Corona exserted from corolla tube, c 5.5 mm wide, dull pink to dark red; outer corona discoid, margin thin, obscurely sinuatedenticulate; inner corona lobes leaning on and barely ex­ ceeding anthers. Pollen masses yellow. Follicles 11-16 x 1-1.2 cm; seeds 7.5-8 x 3.5-4 mm, light brown. Figs 140.47.3 & 4; 140.48.10 & 11. Acacia - Commiphora bushland. often on dark soils; 1200-1500 m. GG SD; Kenya, Uganda. M. Gilbert 2290, 3306; Edwards & Gilbert 200. 4. P. gilbertii (Plowes) M.G. Gilbert (2003); Angollumagilbertii Plowes (1993) - type. TU, be­ tween Mekele and Kwiha, M. Gilbert s.n. (K holo.). Stems procumbent, forming loose mats, occasionally shortly soboliferous, up to 5 cm tall in the wild, much

140. ASCLEPIADACEAE: 42. Pachycymbium

larger in cultivation; tubercles up to 7 mm high. Pedicels 3 ! mm long. Sepals 3.5-6 mm. Buds patent to more or less erect. Corolla rotate to slightly reflexed, 2.2-3.5 cm v. de, brown; corolla tube filled by corona, sometimes de­ limited by narrow, raised annulus; corolla lobes 7-9 mm wide. Corona 6.2-8 mm wide, 2.5-^ mm high; outer co­ rona lobes deeply saccate, emarginate; inner corona lobes lanceolate, about as long as anthers. Fig. 140.48.8 & 9. Degraded bushland with Euclea schimperi and Aloe camperi s. lat. overlying limestone; 1800-1900 m. TU; nc I known elsewhere. This taxon was initially identified (Gilbert 1978) as Cc '-alluma commutata Berger, a species based on culti­ vated material of unknown origin. No material exactly m tching Berger’s figure has been seen but there are colle< :ions from Sudan and Saudi Arabia that could equally we 11 match, the best fit perhaps being “Angolluma” sun’anica Plowes. Bruyns does not include this species in his account of Or ttea, apparently because he believes it not to be separabh from P. abayense. This view is definitely not accepted by -.he present author.

187

emarginate margin lower than stigma cap; inner corona lobes leaning on and only slightly longer than anthers, with prominent ascending basal horn alternating with outer lobes, dorsal horn usually div ided into three or more teeth and often with other teeth along back of lobe, tips of lobes irregularly 3- or 4-toothed. 1. Corolla lobes linear revolute. subsp. baldratii - Corolla lobes lanceolate to ovate, only slightly revolute towards tips. subsp. subterraneum subsp. baldratii Corolla lobes linear-revolute, two-coloured. Bushland to deciduous woodland on basement com­ plex. EW; Sudan. subsp. subterraneum (Bruce & Bally) M.G. Gilbert in Bradleya 8: 24 (1990). Caralluma subterranea Bruce & Bally (1941); Orbea subterranea (Bruce & Bally) Bruyns (2000) type: Kenya, Boy Joanna in Bally S(4) (K holo.).

Corolla lobes lanceolate to ovate, only slightly revolute towards tip. Figs 140.47.5; 140.48.17 & 18. 5. P. abayense (M.G. Gilbert) M.G. Gilbert (1990); Caralluma abayensis M.G. Gilbert (1978); Angol­ luma abayensis (M.G. Gilbert) Plowes (1993); Orbea abayensis (M.G. Gilbert) Bruyns (2000) - type: SU, Abay (Blue Nile) Gorge, near km 205 marker on Addis Abeba-Debra Marcos road, cult. Addis Abeba, V/. Gilbert 3891 (K holo., ETH iso.). Closely related to P. gilbertii. Tubercles much more prc minent, up to 20 nun high. Pedicels 8.5-9 mm. Buds de:lexed. Corolla reflexed at least towards tips of lobes, c 3 cm wide; tube delineated by narrow annulus. Corona c 2.5mm high, dull brown, fleshier; outer lobes shallower, almost annular. Fig. 140.48.6 & 7. Evergreen or semi-evergreen bushland on steep sloncs, overlying basalt and gypsum-rich limestone; 14(K»-2400 m. SU; not known elsewhere. 6. P. baldratii (White & Sloane) M.G. Gilbert (1990); Caralluma baldratii White & Sloane (1937); A ngolluma baldratii (White & Sloane) Plowes (1993); Orbea baldratii (White & Sloane) Bruyns (2000) type: figures 200 & 201 of protologue, based on mate­ rial from Eritrea, Addi Keyili (‘Addi Che’) (lecto., designated by Gilbert, 1990). Stems strongly soboliferous with slender white under­ ground portion, solitary or forming small clumps; tubercle triangular conical to acuminate. Leaves relatively stout, sometimes with a pair of minute stipels at base. Pedicel ven variable in length. Corolla rotate with very shallow tube;, lobes linear revolute to ovate and only slightly revolute at the tip, uniformly minutely setose, usually pur­ plish or brownish, less often yellow, sometimes 2coloured. Corona exserted from corolla tube, almost blai a ; outer corona lobes saccate with denticulate to

Habitat not recorded (in Kenya amongst short grass on rocky outcrops). SD; Kenya, N Tanzania. G. Ball photo! This subspecies is known in Ethiopia from a popula­ tion found in Sidamo, near Kibre Menghist, with lanceo­ late corolla lobes somewhat intermediate between the two subspecies. The exact identity of these plants needs more detailed investigation based on more extensive material. The Kenyan populations are notoriously variable and disjunct from the southern Ethiopian record. Bruyns has described a further subspecies from Somalia, as Orbea baldratii subsp. somalensis. 7. P. circes (M.G. Gilbert) M.G. Gilbert (1990); Caralluma circes M.G. Gilbert (1978); Angolluma circes (M.G. Gilbert) Plowes (1993); Orbea dummeri subsp. circes (M.G. Gilbert) Bruyns in Aloe 37: 75 (2000) - type: SD, 6 km E of Wachile, cult, in Addis Abeba, M. Gilbert 2288 (K holo., ETH iso.). Stems erect, forming dense clumps, up to 20 cm tall in cultivation, not soboliferous; tubercles up to 20 mm high. Inflorescence rarely more than 2-flowered; pedi­ cels c 8 mm. Sepals c 6 mm. Buds more or less erect. Co­ rolla campanulate with half-erect lobes, c 3 cm wide, lime-green; corolla tube 12 mm wide, 5 mm deep, virtu­ ally smooth inside; corolla lobes c 6 mm wide, promi­ nently papillate, each papilla ending in prominent one-celled hair up to 1.3 mm. Corona c 4.5 x 4.5 mm, stipitate; outer corona lobes small, deeply saccate, en­ tire; inner corona lobes with apex divided into 2 or 3 subulate, erect teeth and with a transverse row of 3 or 4 short conical teeth and a pair of pits immediately above them. Pollen masses pale yellow, overlapping when pollinaria removed from gynostegium. Figs 140.47.6; 140.48.14-16.

188

140. ASCLEPIADACEAE: 42. Pachycymbium

Figure 140.47. PAC H YCYM BIU M : corollas (1 ,2 ,4 -8 ) x 2 '/s, flowering shoot (3) x c 1 1 - P. S P R E N G E R I subsp. S P R E N G E R I: 2 P L4TIC O RO N U M : 3 & 4 - P. G EM U G O F A N A ; 5 - P. B A L D R A T II subsp S U B T E R R A N E U M ; 6 - C C IR C E S '; 7 - /> SACCULA TUM\ 8 - C VIBRA TILIS. 1 from photo o f cultivated material collected from near Mekele; 2 from photo o f material prop­ agated from the type collection A/. e£ 5. Gilbert 1383; 3 & 4 from photos o f material propagated from the type collection M. & S. Gilbert 1731; 5 trom photo o f cultivated material collccted near Kibre Mcnghist; 6 from material propagated from the type collection M. Gilbert 2288; 7 from cultivated material originally collected near ErcrG ota: 8 from photo o f plant growing in N Kenya. Drawn by M ar­ garet Tebbs.

140. ASCLEPIADACEAE: 42. Pachvcymbium

189

f e d e ^ ie w p x 2 V F - SP K E N G E R Isu bsp. S P R E N G E R I 1 - corona top view; 2 - cotorona side view P C I I R F R T n TI C0R0NlJl M 4 - c° mna alia. are the only stapdiad genera with substantial representations in both sout hem Africa and tropical Africa to Arabia. 1. Stems usually erect or ascending, 5-angled, form­ ing dense clumps; tubercles prominent; buds uni­ formly pale; inner corona lobes with smooth tips. 2 - Stems ± prostrate, 4(-7)-sided; tubercles almost obsolete; buds red marked with pale lines; inner corona lobes with papillose tips. 3. H. recondita 2. Corolla campanulate to tubular, similar in colour and texture to corolla lobes. 3 - Corolla limb reflexed and expanded to form smooth reddish brown annulus contrasting strongly with pale, prominently papillate corolla lobes. 4. H. somalica 3. Corolla campanulate; papillae on interior of corolla uniformly coloured or with bands of colour; inner corona lobes blunt, straight to, at the most slightly recurved. 1. H. macro carp a - 'orolla tubular, papillae on interior of corolla white with minute dark spots; inner corona lobes long acute, meeting in centre and then clearly re­ curved. 2. H. holeana

1. H m acrocarpa (A. Rich.) Sprenger (1892); Stapelia macrocarpa A. Rich. (1851) - type: TU, ( lure, Ouartin-Dillon s.n. (P holo.); epitype: EW, Gheleb, Penzig s.n., cultivated in La Mortola, Italy (K epi ! designated by M. Gilbert in Brodie. Kew Mag. 15 9, 1998). Stems erect or ascending, densely tufted, to 7.5 cm tall, 1.5 cm thick; tubercles in 5 ranks, conical acuminate, up to If mm high. Leaves poorly differentiated from tubercle prop-■r fmely tapered, persistent. Pedicel 5-10 mm. Se­ pals I —10(—12) mm. Corolla campanulate, 15-20 mm wide outside whitish, smooth or minutely asperouspapL late, inside uniformly brown to blackish purple or very pale cream variously marked with dots or concentric lines sparsely papillate, papillae small, conical, blunt; co­ rolla obes acuminate-triangular, 5-7 mm long. Corona blacl sh purple except usually for bright yellow spots on dors; 1gibbosities of each inner corona lobe; outer corona conic :il to almost flat, lobes broadly oblong with ob­ scure iy emarginate tip; inner corona lobes with prominent transverse dorsal gibbosities, upper part straight or sligh Iv reflexed, more or less meeting in centre, tip blunt, very i iiinutely tuberculate. Fruiting pedicel up to 6.5 cm; follic es up to 14 cm long.

191

1. Outside of corolla minutely papillate; inside uni­ formly dark brown, less often reddish brown or wliitish with conspicuous concentric dark lines. subsp. m acrocarpa - Outside of corolla smooth; corolla always whitish, usually with concentric lines of spots, less often these confluent into continuous lines. subsp. harerghensis

subsp. m acrocarpa H. penzigii N.E. Br. (1892); H. macrocarpa var. penzigii (N.E. Br.) White & Sloane in The Stapelieae 3: 855 (1937) - type: EW, Gheleb, cultivated in La Mortola, Italy (K holo.). H. penzigii var. schweinfurthii Berger in Stap. und Klein:. 142 (1910) - type: EW, Acrur, Schweinfurth 1340 (P iso.). If. penzigii var. schimperi Berger loc. cit. 143 — type: GD, ‘Choum-Delaque’, Schimper 945 (P lecto., designated by Brodie, 1998). Corolla outside veiy minutely asperous-papillate, inside uniformly dark brown, sometimes almost black, or, less often bicoloured: whitish with prominent dark brown con­ centric bands; corolla lobes acuminate. Fig. 140.49.1-3. Open evergreen bushland, often with Euclea schimperi, usually in shelter of shrubs and Aloe, overlying basalts and basement complex granites and schists; 1600-2400 m. EW TU WU; Sudan; Saudi Arabia? Ryding 1389. There has been considerable recent confusion in the naming of this taxon, mainly originating from a lack of knowledge of Huernia in the northern part of the Flora area. The type specimen is in fruit and clearly represents a member of this genus. Any doubt as to the identity of the taxon has been removed by epitypification (Brodie, loc. cit.). Leach (Excelsa 4: 107-114, 1988) did not examine the specimen and his rejection of the species as insuffi­ ciently known, even though the name had been used by White and Sloane (The Stapelieae 3: 854, 1937), can not be supported. Having seen numerous populations in Tigray and Eritrea, the author is convinced that there is only one species in the region, which has two colour forms, one a uniform dark brown, the other two-coloured, white with prominent concentric dark bands. There is no evidence of morphological or ecological differences be­ tween these, and thus they are not formally named, but H. macrocarpa var. schimperi was explicitly based on the bi­ coloured plant and is available if a formal distinction is thought essential.

subsp. harerghensis M.G. Gilbert in Nordic J. Bot. 22(2): 205-210(2003) type: HA, 4 km W of Grawa, M. Gilbert 4044 (K holo., ETH iso.). H. macrocarpa subsp. concinna sensu Gilbert loc. 2 jbspecies in the Flora area, one extending into Su­ cit. 12, fig. 9 (1975), ncn sens. str. dan arid possibly Arabia. Huernia penzigii'War”, Leach, loc. cit.: 114, fig. 68.

192

140. ASCLEPIADACEAE: 44. Huemia

Figure 140.49. H U E R N L i M A C RO C AR P A subsp. M AC RO C ARPA 1 - flowering shoot x 9/io; 2 - Corona side view x 3; 3 - Corona top view x 3. //. M AC RO C ARPA subsp. H A R E R G H E N SIS. 4 Corona side view x 3; 5 - Corona top view x 3 //. B O LE A N A 6 flowering shoot x 9/io; 7 - Corona side view x 3; 8 Corona top view x 3. //. REC O ND ITA. 9 - flowering shoot x 10 - Corona sid e view x 3; 11 - Corona top view x 3. R. SO M A L IC A . 12 - flowering shoot x *1% 13 -C o ro n a side view x 3; 14 - Corona top view x 3. I drawn from photograph o f cultivated plant originally collected near Weldiya; 2 & 3 from cultivated material originally collected near Axum; 4 & 5 from cultivated material originally collected from near Harar. 6 from photograph o f clonotvpe material; 7 & 8 from clonotvpe material; 9 from photograph o f holotype; 10 & 11 from clonotype material; 12 from photograph o f Lavranos des” in the descriptions) represent a special case of erminal inflorescence: the terminal, inflorescenceuroducing shoot is pushed to tlie side and overtopped by a solitary (strongly developed) lateral axis (sympodialnonochasiai growth). The latter may thus be mistaken for a terminal shoot, and the terminal inflorescence for an ixillary one ( Fig. 141.36.1). Careful investigation is re­ quired! Also in the case of solitary (or paired) flowers, :are should be taken in the interpretation of their position: in certain groups (e.g. members of the Gardenieae), seem­ ingly axillary, solitary flowers turn out on closer investi­ gation to be terminal on rather contracted shoots. OVARY, FRUIT and SEED. The number of locules, the number of ovules (one. several, many per locule), ovule structure and position on the placenta, the placentation it­ self (predominantly axillary', but also parietal: various ]ardenieae\), and tlie attachment of the placentas to the septum (base, middle or top) are often important charac­ ters distinguishing genera or entire tribes (compare, for jxample. Figs 141.4.6. 141.15.4. 141.38.6 and 141.54.8). i: is often essential to investigate this character by section­ ing ovaries. A wide array of different fruit types occur in the fam­ ily. ranging from fleshy or diy indehiscent fruits to dry, dehiscent fruits, and the development of mericarps. Fleshy indehiscent fruits need to be opened to see whether rJaey contain pyrenes (“stones”, drupes) or seeds (berryNce fruits). In tlie case of dehiscent fruits (typically cap­ sules), it is necessary to carefully observe the exact mode af dehiscence. In some taxa the number of developed seeds is important. In certain taxa. the seeds need to be sectioned to see whether the endosperm is entire or rumi­ nate (e.g. Fig. 141.38.8-9), or to observe the position of

195

the embryo within the seed and the size ratio of radicle to cotyledons in the embryos. LEAVES and STIPULES. Bacterial nodules, either scat­ tered over entire leaf blades or restricted to certain areas of tlie blades, occur in several (woody) genera, although not necessarily in all species of a genus; their presence or ab­ sence may be of systematic value. Moreover, there may also be (hairy) domatia which are typically found in the vein axils of leaf-blades (e.g. Fig. 141.37.8). Stipules ex­ hibit a wide range of forms and sizes; they may be of diag­ nostic value in certain alliances (compare, for example, Fig. 141.25.2, 141.26.2 and 141.38.2). In some woody groups, stipules may only be observable on the youngest parts of the branches as they fall off soon. In herbaceous genera, there is mostly a stipular sheath bearing one to many teeth, setae or fimbriae (cf. Fig. 141.15.2). Genera occurring in the Flora area belong to 4 subfamilies and 18 tribes [classification based on Robbrecht in Opera Bot. Belg. 1: 1-271 (1988)]. Key to genera 1. Plants armed with thorns. 2 - Plants unarmed. 5 2. Thoms reflexed (climbing hooks), inflorescences globose. 26. Uncaria africana - Thoms straight, inflorescencesnot globose. 3 3. Thoms paired, decussate. 51. Meynatetraphylla - Thoms not paired. 4 4. Thoms alternate above paired, leafy or flowering, short shoots. 33.Catunaregam nilotica - Thoms in whorls of 3, only on long shoots. 46. Canthium oligocarpum 5. Inflorescences spike-like, up to 7—15(—20) cm long, subtended by a pair of bracts; capsules with winged seeds; small tree. 27. Hymenodictyon floribundum - Inflorescences different. 6 6. Flowers in globose heads, 1-5 cm in diameter. 7 - Flowers not in distinctly globose heads. 10 7. Plants herbaceous. 10. Agathisanthemumglobosum - Plants woody, small to large trees. 8 8. Flowers forming a head fused by their ovaries; fused fruits forming a fleshy infructescence. 23. Sarcocephalus latifolius - Flowers forming a head but ovaries entirely free from each other. 9 9. Leaves decussate, broadly elliptic to suborbicular; flowering heads arranged in long-pedunculate umbellate to paniculate inflorescences. 25. Hallea rubrostipulata - Leaves in whorls of 4, lanceolate; flowering heads solitary on long stalks. 22. Breonadia salicina 10. Leaves and leaf-like stipules in (pseudo-)whorls of 4 to many; corolla always small, rotate or less commonly subcampanulate; pfants herbaceous. (Tribe Rubieae) 11 - Leaves decussate or, if in true whorls of (3)4 plants woody and corolla not ± rotate. 13

196

141. RUBIACEAE

11. Leaves and leaf-like stipules petiolate. in whorls of 4; corolla mostly 5-merous. 19. Rubia cordifolia - Leaves and leaf-like stipules (sub)sessile. in whorls of 4 to many , corolla usually (3)4-merous. 12. Leaves and leaf-like stipules in whorls of 4. 1-veined; small annuals with short, 3-flowered axillary cymes 21. Valantia hispida - Leaves and leaf-!ike stipules in whoris of 6 to many (if in whorls of 4, leaves 3-veined); inflorescen­ ces terminal and more extensive than above and plants perennial. 20 Galium 13. Leaf-blades with spot-like, dark bacterial nodules scattered throughout the surface. 14 - Leaf-blades without bacterial nodules (but sometimes w ith do matia) 15 14. Flowers heterosty lous. corolla lobes valvate, style with bifid stigma included or somewhat exserted: fruits with 2 thick-walled semi-globose pvrenes. 1. Psychotria kirkii - Flowers not heterosty lous, corolla lobes contorted. anthers and sty le with clavate stigma long ex­ serted; fruits with 1 (rarely 2) papery pyrene(s). 42. Pavetta (in pan)

21 . Corolla tube cylindrical or narrowly to broadly

funnel-shaped, at least 20 mm long, often consid­ erably longer -Corolla tube less than 20 mm long. 31

22. Woody plants 23.

r 25 26.

’21

15. Climbers. 16 - Erect to prostrate herbs, shrubs or trees. 17 16. Evil smelling plants, fruits flattened, with a thin. brittle exocarp containing 2 winged, compressed pvrenes; flowers white or greenish with reddish purple centre 17. Paederia pospischilii 28 - Plants not evil smelling; fruits fleshy (± hard, indehiscent in dry state), with many small seeds: flowers bright yellow with an orange-red star of hairs around the throat. 29. Mussaenda arcuata 17. Calyx lobes of some flowers of an inflorescence much enlarged, white and showy, similar in shape 29 and size to foliage leaves. 30. Pseudomussaenda flava - Calyx different, if lobes unequal or foliaceous not with the above characteristics. 18 18. Calyx eccentric, entire or shallowly lobed, coloured 6. Carphalea glaucescens - Calyx not as above, usually with distinct lobes or 30 teeth. 19 19. Flowers small, wind-pollinated, bisexual or uni­ sexual: bisexual and female flowers without style but with long, much exserted hairy stigmas: bi­ sexual and male flowers with large anthers exserted. dangling on long, slender filaments. 31 (Tribe Anthospermeae) 20 - Flowers animal-pollinated, small or larger, mostly bisexual; distinct style with bilobed. clavate or capitate stigmas; anthers exserted or not. never 32 dangling on long, slender filaments. 20. Fruits dry, splitting into two indehiscent mericarps; leaves small, lanceolate to ± linear 18. Anthospermum - Fruits fleshy, drupes; leaves large, leathery, ever­ green. Coprosma lucida (cultivated!

Herbaceous plants. '$» Small shrub with very brittle wood: the bulk of leaves clustered on short shoots; fruits capsular. 9. Shrubs or small trees: no leafy short shoots; leaves wider, fruits indehiscent ± fleshy. 24 Few- to many-flowered, seemingly axillary inflo­ rescences solitary at nodes 36. Oxyanthus Flowers solitary or paired or terminal in several to many-flowered inflorescences. 25 Inflorescences several to many-flowered, terminal.26 Flowers mostly solitary or paired. 27 Flowers 4-inerous: one ovule per locule, fruit 2-seeded. 42. Pavetta (in part) Flowers 5-6-me ro us; ovules numerous, fruit many-seeded 32. M acrosphvra longistyia Corolla salver- or funnel-shaped; flowers 6-9merous. terminal on short lateral branches. 34. Gardenia (in part ) Corolla broadly funnel-shaped or cylindrical be­ low and ± campanulate above: flowers 5-merous, often seemingly axillary but actually terminal on short abbreviated branches. 35. Rothmannia urcelliformis Fruits indehiscent hard, one ovule/seed per locule: calyx lobes unequal; flowers bright blue to pale blue or whitish, heterosty lous. 54. Pentanisia (in part) Fruits capsular, ovules/seeds numerous; calyx lobes unequal or not; flowers never bright blue, heterostylous or not. 29 Corolla 4-inerous: anthers included and style with stigma exserted (plants not heterostylous); calyx lobes ± equal: flowers axillary. 8. Conostomium Corolla 5-merous; anthers exserted and style with stigma included or vice versa (flowers hetero­ stylous); calyx lobes unequal; inflorescences ter­ minal. 30 Corolla with a conspicuous crimson or purplisbred ‘eve'; inflorescences elongated, spike-like in fruit up to 30(-40) cm long. 5. Otom eria Corolla without an ‘eye’; inflorescences some­ times ± elongated in fruit but if so. only a few cm long. 4. Pentas (in part)

Corolla lobes valvate in bud; herbaceous or woody plants. 32 Corolla lobes contorted or imbricate in bud, always woody plants 58 One ovule per locule. 33 Two to mam ovules per locule. 50 33 Ovule pendulous from near the apex of the locule. 34 Ovule attached to near the base of the locule or to the lower half or middle of the septum. 44

Pent

141. RUBIACEAE

197

3 Perennial herbs with terminal inflorescences and flowers with unequal calyx lobes. 54. Pentanisia - Shrubs, small trees or occasionally subshrubs with axillary inflorescences, calyx lobes not unequal. 35

46. Pyrenes without obvious dehiscence; seeds red­ dish. 1. Psychotria - Pyrenes opening in the upper half by 2 slits (along the margins of the ventral face); seeds pale. 2. Chazaliella

35 Stigma lobes filiform, flowers heterostylous; fruits

47. Fruit circumscissile about its middle, upper half coming off like a lid; seeds with an X-like groove on ventral side. 16. M itracarpus hirtus - Fruit splitting open differently or separating into two indehiscent mericarps; seeds without X-like groove. 48 48. Capsule opening from the base upwards, the valves, held together by the persistent calyx, com­ ing off as a unit, leaving behind the septum. 15. Spermacoce sphaerostigma - Capsule opening from the top downwards or fmit splitting into two mericarps. 49 49. Capsule splitting open to release the seeds. 15. Spermacoce - Capsule splitting into two mericarps, each contain­ ing one seed. 14. Diodia sarmentosa 50. Fruits indehiscent, berry-like, many-seeded; co­ rolla bright yellow with an orange-red star of hairs around the throat. 29. Mussaenda arcuata - Fniits capsular or splitting into twomericarps:co­ rolla not yellow. 51

indehiscent. mostly 1-seeded by abortion of sec­ ond ovule. 45. Craterispermum schweinfurthii Stigma a conspicuous cylindrical, subglobose or capitate structure, well separated from the style below; flowers not heterostylous; fruits indehiscent. with 2-5 hard pyrenes. (Tribe Vanguerieae) 36 36 Subshrubs; leaves in whorls of 3(4), markedly discolourous. 50. Fadogia cienkowskii Shrubs or small trees, sometimes scandent; leaves decussate or, if in whorls of 3, not markedly discolourous. 37 37 Ovary 4-5-locular. 38 Ovary 2-locular. 39 38 Flowers few to several, clustered on short shoots; calyx lobes c 0.5 mm long, entire; corolla 3-5 mm long; mature drupes 10-15 mm in diameter: plants with paired thorns. 51. Mevna tetraphylla Flowers in axillary inflorescences borne on long shoots: calyx and corolla larger; mature drupes to 40 mm in diameter: plants unarmed. 53. Vangueria 3"i Stipules villous inside; corolla lobes ± apiculate; flowers in simple, unbranched, axillary inflores­ cences. 52. Rytigynia neglecta Stipules not villous inside; corolla lobes not ± api­ culate. 40 4) Leaves leathery, evergreen, glossy' above, gla­ brous: flowers many in ± sessile or pedunculate cymes or umbel-like inflorescences. 48. Psydrax Leaves thin to slightly leathery', mostly not ever­ green; flowers in very few- to many-flowered in­ florescences. 41 41 Plants dioecious, female flowers solitary, male flowers in groups of 1-6. 42 Plants not dioecious, bisexual flowers many in pedunculate, much-branched cymes or corym­ bose inflorescences. 43 4 1 Flowers 4-merous, subtended by paired bracts; buds entirely enclosed by these bracts; plants gla­ brous. 49. Pyrostria phyllanthoidea Flowers 5-merous, not subtended by paired bracts; plants hairy 46. Canthium pseudosetiflorum 4 : Style with ± long and ± cylindrical stigmatic knob about tw ice as long as corolla tube. 47. Keetia Style with shorter, rounder and ± capitate stigmatic knob much less than twice as long as corolla tube. 46. Canthium 4 » Ovule erect from the base. (Tribe Psychotrieae) 45 Ovule attached to lower half or to near the middle of the septum. (Tribe Spermacoceae) 47 4 1 Forest floor herbs; stems rooting at nodes, with paired, reniform leaves. 3. Geophila repens - Shrubs or trees. 46

51. Seeds winged; cultivated/ornamental trees. 24. Cinchona pubesccns (cultivated) - Seeds not winged; herbs or small shrubs. (Tribe Heyotideae) 52 52. Corolla mostly 5-merous. 53 - Corolla mostly 4-merous. 55 53. Small annual; inflorescences axillary, 1-3flowered. 12. Thecorchus wauensis - Perennial herbs or subshrubs; inflorescences termi­ nal or (pseudo-)axillary, but then very lax, elon­ gated, and with more flowers. 54 54. Erect herbs or subshrubs (occasionally decumbent. forest floor herbs); inflorescences terminal. 4. Pentas - Decumbent herb of wet places; flowers in very lax, elongated (pseudo-)axillary inflorescences. 11. Pentodon pentandrus 55 Anthers and/or style with stigmas exserted (flow­ ers heterostylous); corolla cylindrical to funnelshaped. 13. Oldenlandia (in part) - Anthers and style always included. 56 56. Calyx lobes enlarged, foliaceous in fruit, up to 5 mm long; inflorescences axillary. 1-3-flowered. 12. Thecorchus wauensis - Calyx lobes not enlarged in fruit, smaller, inflores­ cences axillary or terminal. 57 57. Anthers always inserted above the level of the stigma (but included in the corolla tube); corolla tube narrowly cylindrical. 7. Kohautia - Anthers overtopped by stigma; corolla tube cylin­ drical to funnel-shaped. 13. Oldenlandia (in part) 58. Fruits capsular. - Fruits indehiscent.

59 60

198

141 RUBIACEAE: 1 Psychotria

59. Seeds winged. 28. Crossopteryx febrifuga - Seeds not winged (or very inconspicuously so). 31. W endlandia arabica 60. Deciduous shrubs, leaves appearing with or after the flowers. 61 - Evergreen plants. 63 61. Flowers with style and stigma included, corolla tube 15-18 mm long; fruit 2-seeded, seeds with ventral groove (like coffee-bean). 44. Psilanthus leroyi - Flowers with style and stigma exserted. corolla tube less than 10 mm long; ovary with 2 or more ovules per locule. 62 62. Flowers 6-merous, calyx cup-like; fruits in axillary clusters; plants not always deciduous. 37. Tricalysia niamniamensis - Flowers 5-merous, calyx lobes lanceolate; fruits distinctly pedicellate, paired or solitary at lower­ most nodes of newly produced lateral branchlets. 39. Feretia apodanthera 63. Calyx reduced to a short indistinct rim, occasion­ ally shallowly lobed; fruit 2-seeded, seeds with ventral groove. 43. Coffea arabica - Calyx with distinct lobes or at least with a distinct. truncate tubular part; fruits 2- to many-seeded, seeds without ventral groove. 64 64. Inflorescences axillary' (or sometimes supraaxillary), paired at nodes. 65 - Inflorescences terminal on main and lateral branches (occasionally appearing axillary due to much-contracted lateral branches). 67 65. Flowers pedicellate, in paired axillary pedunculate cymes; corolla subrotate, tube up to 1 mm long. J 38. Galiniera saxifraga - Flowers in (sub)sessile axillary clusters; corolla not subrotate, tube longer. 66 66. Leaves with pilose domatia; flowers 6-merous. an­ thers exserted; seeds with entire endosperm. 37. Tricalysia niamniamensis - Leaves without domatia; flowers 4-merous, an­ thers included; seeds with ruminate endosperm. 40. Polysphaeria 67. Flowers 5-12-merous. mostly solitary or paired, terminal on (very) short lateral branches; ovary 1-celled, with 2-9 parietal placentas, ovules numerous; fruits hard (often Fibrous-woody), usually very many-seeded, mostly large (up to 10 cm in diameter). 34. Gardenia - Flowers 4- or 5-merous, in pedunculate to sessile corymbose to capitate inflorescences; ovary 2celled, with 1—15 ovules; fruits not as above, mostly less than 10 mm in diameter. 68

RUBIOIDEAE Herbaceous (perennials or annuals) or less often wood) plants with raphides. Stipular sheaths often fimbriate o; bifid. Flowers often heterostylous, corolla lobes valvate in bud; numerous, few or only 1 ovule on each placenta, solitary ovules erect. Fruits fleshy or dry dehiscent or not. Seeds mostly with thin seed-coa (without very conspicuous thickenings) and with oil> or homy endosperm. PSYCHOTRIEAE Dumort. Mostly woody plants, rarely herbaceous. Inflorescences mostly terminal. Ovary 2-5-locular; 1 ovule per locule, erect from base. Fruits mostly drupes, with 2 to several pyrenes. 1. Shrubs or trees. - Forest floor herbs.

3. Geophila

2. Py renes opening in the upper half by 2 slits (along the margins of the ventral face); seeds pale with entire endosperm. 2. Chazaliella - Pyrenes without obvious dehiscence, seeds red­ dish. endosperm entire or ruminate. 1 Psychotria 1. PSYCHOTRIA L. (1759), noni. cons. Cephaelis Sw. (1788) Shrubs or small trees, less often (and not in the Flora area) subshrubs or lianas. Leaves decussate, usually petiolate, either with domatia (mostly in vein axils) or with bacterial nodules (scattered throughout the leaf blade or variously restricted), occasionally both absent. Stipules usually de­ ciduous, entire or divided above. Flowers 4-5(-6)merous, small, heterostylous, in terminal paniculate, often many-flowered inflorescences, or in capitate inflorescen­ ces subtended by an involucre of bracts. Calyx lobes mostly small to minute. Corolla tube mostly short cylin­ drical. haiiy at throat lobes valvate in bud. Anthers exserted. on long filaments, in short-styled flowers; shorter filaments in long-styled flowers. Sty le w ith mostly bilobed stigmas included or exserted. Ovary mostly 2-locular. one erect ovule per locule on placenta attached to near base of septum. Fruits drupes, with 2 (or by abor­ tion 1) py renes. Seeds often plane on ventral side, convex dorsally; testa reddish, endosperm entire or ruminate. A huge pantropical genus (probably more than 1000 species); over 200 in Africa, 5 in the Flora area. 1. Leaf blades with bacterial nodules scattered over their entire surface. 5. P. kirkii -

Leaf blades without bacterial

nodules.

2

2. Inflorescences capitate, subtended by a conspicu­ ous involucre of bracts. 4. P. peduncularis 68. Flowers 4-merous. tube often longer than lobes; fruits 2-seeded, pyrenes with a central circular t - Inflorescences paniculate, several- to manyhollow on ventral side. 42. Pavetta flowered. 3 - Flowers (mostly) 5-merous. tube more or less as 3. Stipules 10-20 mm long; pyrenes 5-groovcd on long as lobes, fruits several- to 1-seeded, pyrenes dorsal side. 3. P. orophila not as above. 4 41. Tarenna graveolens subsp. graveolens* - Stipules less than 10 mm long.

141. RUBIACEAE: l.Psvchotria

- Much-branched panicles to 150 mm in diameter, ca­ lyx cup-like, truncatc above or very indistinctly lobed: pyrenes not ribbed or grooved on dorsal side. 1 P. capensis Inflorescence ± umbel-like, to 75 mm in diameter, calyx distinctly 5-lobed; disk prominent on imma­ ture fruits. 2. P. sp. = Mooney 9252 1 P. capensis (Eckl.) Vatke (1875); Logania capensis Eckl. (1830); Grumilea capensis (Eckl.) Sond. (1865) - type from South Africa. subsp riparia (K. Schum. & K. Krause) Verde, in Fi Zarnb. 5: 13 (1989); Grumilea riparia K. Schum. & K. Krause (1907); Psy­ cho tri a riparia (K. Schum. & K. Krause) Petit (1964). Grumilea bussei K. Schum. & K. Krause (1907). var. puberula (Petit) Verde, in FI. Zamb. 5: 14 (1989); Psy>chotria riparia (K. Schum. & K. Krause) Petit var. puberula Petit in Bull. Jard. Bot. Brux. 34: 46 I (1964) - lype from Kenya. S lirub or small tree up to 3 m tall, stems with rust-coloured pubescence. Leaves elliptic to obovate-elliptic, 3-20 x 1-6.5 cm, acute to more or less acuminate at apex, cuneate aijbase, rusty-pubescent; bacterial nodules absent: petioles 0,5-4 cm long. Stipules triangular or ovate-triangular. C 3-0.8 cm long, thin, soon falling. Flowers 5-merous, in r luch-branched, many-flowered panicles up to 150 mm in c iameter: peduncles 50-100 nun long, pedicels up to 1 mm ( '- 5 mm long in fruit); bracts minute. Calyx glabrous or I ubescent. cup-like, up to I mm long, lobes indistinct, mini i:e or absent. Corolla yellow, greenish-vellow or whitish, tabe 3—4.5 nun long, lobes 2-3 nun long. Drupe red. usu­ ally somewhat pubescent (when young), subglobose. 6-7 mm in diameter, not grooved in dry state; pyrenes 2. 1;:mi-globose, not grooved or ribbed on dorsal side. Seed with ruminate endosperm. Riverine forest; 1150-1300 m. IL BA; tropical E Africa I :>Mozambique. Friis et al. 1974,3278; PuJJ& Ensermu K. #61104-2/3. The species (as a whole) occurs from Ethiopia south to outh Afnca. Subsp. capensis, with longer fniiting pedi| sis, is restricted to South Central and southern Africa, •ubsp. riparia var. riparia. with mostly glabrous parts, ocurs in tropical E and South Central Africa. I P. sp. = Mooney 9252 ; hmb (or small tree?), 2-3 in tall, stems glabrous. Leaves elliptic to obovate-elliptic. (4-)9-23 x 1.5-9.5 cm, acute to more or less acuminate at apex, cuneate at base, glabrous, eathery; bacterial nodules absent; petioles 0.7-4 cm long. Stipules triangular, 0.6-0.7 cm long. Flowers unknown; oung fruits many in more or less umbel-iike paniculate inlorescences c 75 mm in diameter; bracts minute, up to 1 nm long. Calyx glabrous, 5-lobed. c 1 nun long. Young ruit glabrous, with prominent disk and crow tied by lobed ::alyx. Lowland rain forest; 1350 m. KF. Mooney 9252.

199

The only available collection of this plant is too poor for a proper identification. It may be near P. capensis. 3. P. orophila Petit (1964) - type: Kenya, Elliott 27 (K lecto.). Grumilea elliottii K. Schum. & K. Krause (1907). Psychotria sidamensis Cufod. (1965) - type: SD, forest area of Maleko, north of Adola, Haberland 2057 (FR holo.). Shrub or small tree 1.5-^4 in tall, with glabrous stems. Leaves elliptic or elliptic-oblong, 7-20(-25) x 3-9 cm. acute to more or less acuminatc at apex, cuneate at base, veins beneath (at least mid-vein) often (reddish) hairy, blades otherwise glabrous; bacterial nodules absent; pet­ ioles 1-4 cm long. Stipules ovate(-elliptic), 1-2 cm long, usually shortly bifid above, thin, falling soon. Flowers 5-merous. in much-branched, many-flowered panicles, 30-80(-100) mm in diameter; peduncles 40-100 mm long, pedicels absent or very short (c 1 mm long in fruit); bracts small to minute. Calyx cup-like, up to 1 mm long, lobes more or less absent. Corolla white or creamy white, tube 2.5-5 nun long, lobes 1.5-4 mm long. Dnipe red. ellipsoid, (5—)6—7 x 4-5 mm, more or less grooved in dryr state; pyrenes 2, semi-ellipsoid, 5-grooved on dor­ sal side. Seed w ith ruminate endosperm. Fig. 141.1 Montane evergreen forest; 1600-2600(-2700) m. SU AR WG IL KF GG SD BA HA; tropical E Afnca, E Rep. Dem. du Congo. Puff et al. 820924-1/1; P uff & Ensermu K. 820815-1/1. 861 111-1/2. 4. P. peduncularis (Salisb.) Steverm. (1972); Cephaelis peduncularis Salisb. (1808) - type from Sierra Leone. var. ciliato-stipulata Verde, in Kew Bull. 30: 689 (1976); Uragoga ciliato-stipulata De Wild. (1936), nom. invalid. - ty pe from Rep. Dem. du Congo. Cephaelis sp. in Cufod. (1965). Shrub up to 1.5 m tall, unbranched or sparsely branched. Leaves narrowly to broadly elliptic, 5—15(—17) x 3-7 (-8) cm, acute to acuminate at apex, mostly cuneate at base, mostly glabrous; bacterial nodules absent; petioles up to 3 cm long. Stipules oblong to lanceolate, 1-2 cm long, bilobed at apex, rusty-hairy on margins. Flowers 5-merous. several to many in involucrate, capitate inflo­ rescences; pedicels absent or very short (up to 10 mm in fruit); inflorescences usually solitary, on peduncles up to 15 cm long; bracts fonning involucre more or less free or joined, white to greenish. 6-10 mm long. Calyx cup­ like, c 1 mm long, lobes more or less absent. Corolla white, tube funnel-shaped, 3.5-6.5 mm long, lobes up to 3 mm long. Drupe mostly shiny biue(ish), ellipsoid. 5-7 mm long, more or less grooved in dry state; pyrenes 2, semi-ellipsoid, grooved on dorsal side. Seed with entire endosperm. Montane evergreen forest; on forest floor in deep shade; 1500-1700 m. IL KF; S Sudan, W Kenya, SW

200

141 RUBIACEAE I Psychotna

Figure 141.1.

PSYCHOTRIA OROPIIILA

I -

branch with inflorescence in bud and in fruit x Vs; 2 - bud x 3: 3 - open flower, short-styled x 3; 4 - calyx, ovary, style with stigma lobes x 3; 5 - fruit x 3. 1 & 5 from P u ff & Enserm u K 861110-2/12; 2-4 trom Friis et al. 2157. Drawn by Sarah Howard.

5. P. kirkii Hiern (1877) - type from Mozambique. P. petroxenos K. Schum (1907).

triangular. up to 1 cm long, bifid at apex. Flowers 5merous, in several- to many-flowered panicles, in more or less umbel-like or in compound, paniculatcly arranged more or less umbel-like inflorescences; peduncles 10-30 (-50) mm long, pedicels 1—3(—6 in fmit) mm long; bracts minute Calyx minute, less than I mm long or more or less absent. Corolla white, creamy wliite or greenish, tube 2.5-4 mm long, lobes 2-2.5 mm long. Drupe red or orange-red. more or less subglobose, 5-6 x 6-8 mm, often 2-lobed. with 2 semi-globose pyrenes; sometimes only 1 pyrene developed. Seed with entire endosperm.

Shrub up to 3 m tall or, (not in the Flora area) subshrub with glabrous to densely velvety pubescent stems Leaves elliptic-lanceolate, elliptic to more or less ovate. 2-10 (-14) x (0.5-)0.8—4.5 cm, acute to rounded at apex, cuneate at base, glabrous to densely velvety pubescent, bacterial nodules spot-like, numerous, scattered over en­ tire blade, petioles up to 1(—2) cm long. Stipules ovate-

An extremely variable E and SE African species which was devided into 8 ill-defined varieties by Verd­ court in Kew Bull. 30: 262 (1975) and FI. Trop. E. Afr.. Rubiaceae 1: 93 (1976). The species is veiy closely allied to the coastal E African P. punctata Vatke (1875) and it is doubtful whether the two can really be considered sepa­ rate species. However, a considerable number of name

Uganda, E Rep Deni, du Congo. Friis et al. 2004, 2238, 3879. Numerous varieties, distinguished by differences in hairiness, pedicel and peduncle length and. in part, of un­ certain relationships, occur in tropical Africa: of these, only var. ciliato-stipulata extends into the Flora area.

141. RUBIACEAE: 1. Psychotria, 2. Chazaliella, 3. Geophila

cnanges would become necessary if they were to be com­ bined into one species. Verdcourt’s treatment is therefore for the present followed. The following two varieties are, in the Flora area it­ self, easily distinguished and would qualify for a higher category' than variety, but they are too variable elsewhere to justify a change in rank. . Leaves and younger parts of stems mostly ± gla­ brous; inflorescences few-flowered, often ± umbel-like. var. nairobiensis Leaves and y ounger parts of stems densely (vel­ vety) pubescent; inflorescences many-flowered, paniculate. var. taram bassica

var nairobiensis (Bremek.) Verde, in Kew Bull. 30: 262 1(1975); P. nairobiensis B remek. (193 3) - type from Kenya. P. marginata Bremek. (1933). P. punctata Vatke (1875) var. hirtella Chiov. in Miss. Biol. Borana, Racc. Bot.: 231. Fig. 72 (1939); P. ciliatocostata Cufod. (1948) - types: SD, Javello, Cufodontis 387 & M ega Cufodontis 626 (both FT syn.). Wooded grassland, woodland; often in rocky places; 92IJ-1920 m. SU WG KF GG SD; tropical E Africa to Malawi. Mesfin T. & Kagnew 2307; P uff & Ensermu K. 811217-3/4; P uffet al. 870428-7/3. Variable characters are leaf margin (“crinkly'’ or not), leaf size and shape, and extent of the inflorescence. var. tarambassica {Bremek.) Verde, in Kew Bull. 30: 263 (1975); P. tarambassica Bremek. (1961), p.p. - type from Kenya. Woodland, dense thorn scrub or open busliland; often at the base of large boulders; 1200-1800 m. IL SD BA; N Uganda, N Kenya. Friis et al. 2423; P uff & Ensermu K. 821225-2/17; P uff el al. 870509-2/1.

2. CHAZALIELLA Petit & Verde. (1975) Shmbs or small trees; stems usually 2-ribbed and covered with pale brown soft cork. Leaves decussate or in whorls of 3 or 4, petiolate or subsessile, usually deciduous, with or without small, white-pubescent domatia, never with bacterial nodules. Stipules small, entne, bifid or some­ times with a few teeth. Flowers (4)5(6)-merous, small, hdKrostylous, in tenninal sessile or pedunculate head- or pa.nicle-like inflorescences. Calyx lobes small to minute, oi calyx truncate. Corolla tube short, cylindrical, hairy at throat, lobes valvate in bud. Anthers exserted, on long ftlaien ts, in short-styled flowers; shorter filaments in long-styled flowers (anthers included or only tips ex­ serted). Style with 2 thick stigma lobes, either included or exserted. Ovary 2-locular, one erect ovule per locule on

201

placenta attached to near base of septum. Fmit a drupe, with 2 pyrenes; pyrenes more or less flat on ventral side and mostly more or less 3-ribbed or 3-lobed on dorsal side, opening in the upper half by 2 slits (along the mar­ gins of the ventral face). Seeds with pale-coloured testa, endosperm always entire. A tropical African genus of 24 species, previously in­ cluded in Psychotria but also Chassalia (the latter not in the Flora area); 1 species in the Flora area. C. abrupta (Hiern) Petit & Verde. (1975) var. abrupta; Psychotria abrupta Hiern (1877) - type from Mo­ zambique. Psychotria coaetanea K. Schum. (1895). Shrub or small tree up to 4.5 in tall, stems with greyish to whitish thin cork-covering (except for youngest parts) and with 2 longitudinal ridges in the intemodal regions. Leaves elliptic to ovate-lanceolate. 1—17.5(—20) x 1-7.5 (-10) cm, acute to acuminate at apex, cuneate at base, usu­ ally fully expanding only during fruiting stage, glabrous or finely pubescent; petioles 1.4(-3) cm long. Stipules ovate or triangular. 0.2 cm long, shortly bifid or with sev­ eral teeth above. Flowers (4)5(6)-merous, in small 6-20flowered heads; peduncles 7-35 mm long, pedicels up to 1 mm long; bracts minute. Calyx glabrous to pubescent, truncate or lobes indistinctly to distinctly triangular. 1 mm long. Corolla yellow, tube c 3 mm long, lobes 1.5 mm long. Dmpe ellipsoid, glabrous or slightly pubescent, 6-9.5 x 4-6.5 mm, slightly ribbed in dry state, pyrenes semi-ellipsoid. Seeds pale brown Lowland (evergreen) forest; 600 m. IL (Gog forest); tropical East Africa, south to Zimbabwe and Mozam­ bique. Gilbert & Friis 8375; Friis et al. 7285. A second variety, var. parvifolia Verde., is restricted to coastal areas of Kenya.

3. GEOPHILA D. Don (1825), nom. cons. Carinta W.F. Wight (1905) Perennial creeping forest-floor herbs. Leaves decussate, ovate-cordate to rounded-reniform, (long) petiolate. Stip­ ules ovate, entire or bilobed. Flowers 4-7-merous, some­ times heterostylous, in tenninal clusters (sometimes subtended by involucral bracts) or solitary on long peduncles or pedicels. Calyx lobes lanceolate to subulate. Corolla cylindrical to funnel-shaped, lobes valvate in bud. Anthers included or exserted. Style with bilobed or subcapitate stigma included or exserted. Ovary 2-locular, one erect ovule per locule on placenta attached to near base of septum. Fmit a dmpe, with 2 one-seeded pyrenes. plano-convex, dorsally compressed. A pantropical genus of 10-15 species; 1 in the Flora area. G. repens (L.) I. M. Johnston (1949); Rondeletia repens L. (1759) - type from Jamaica. Psychotria herbacea Jacq. (1760).

202

141. RUBIACEAE: 3. Geophila, 4. Pentas

Figure 141.2. G EO P H ILA R E P E N S 1 - habit x */2; 2 - flower, longitudinal section x 2' j. 3 - flower, longitudinal section showing limited heterostvlv x 2' i. 4 —pvrene. two views x 3' 2 1-3 from Biegel 2822; 4 from Chapman 6092. (Reproduced with permission from FI. Zamb. R ubiaceae lab 4 p.p.)

P. herbaceo L. (1762). non Jacq. (1760). nom. illegit. Geophila renifomiis D. Don (1825). G. uniflora Hiem (1877). Creeping herb, stems (rhizomes) rooting at nodes, with erect flowering shoots up to 40 mm long at regular inter­ vals, each with one pair of leaves. Leaf blades rounded reniform. 1.5-3.5(-4.5) x 1.5-3.0(-4) cm. petioles 1.5-4 cm long. Stipules rounded. 0.1-0.2 cm long. Flowers 5-merous. not truly heterostylous. solitary or (rarely) in pairs on peduncles 15-30(—40) nun long. Calyx lobes linear-lanceolate up to 3 nun long. Corolla white, tube cylindiical. 5-10 mm long, lobes lanceolate, c 5 mm long. Anth,. s and style with stigma included. Ovary up to 2 mm long. Drupe light orange or orange-red. 6-10 mm in dia­ meter. crowned by persistent calyx. Pyrenes 3-4 mm in diameter, dorsal side sliallowly convex, rugose, ventral side flat. Fig. 141.2. Forest floor in low - or upland rain forest: 1150-1600 m. IL KF BA; west to W Africa, south to Z im b ab w e, Madagascar, also tropical Asia and America. Friis et al. 3519, 4127; P uff & Ensermu K. 861108-1/1

HEDYOTIDEAE Cham. & Schlechtd.

Mostly herbaceous (perennial or annual) plants, less commonly also woody (shrubs or small trees). Inflores­ cences frequently terminal but also axillary and paired at nodes. Ovary mostly 2-locular with numerous (rarely few) ovules. Fruits usually dry, capsular, occasionally inde­ hiscent. 1. Calyx eccentric, entire or shallowly lobed. col­ oured. 6. Carphalea - Calvx different, usually w ith distinct lobes or teeth. 2

2. Corolla tube cylindrical or narrowly funnelshaped, at least 20 mm long, often considerably longer 3 - Corolla tube less than 20 nun long. 6 3. Small shrub with very brittle wood; the bulk of leaves clustered on short shoots, leaves ± lanceo­ late. 1—3(—8) mm wide. 9. Pentanopsis - Herbaceous plants. 4 4 Corolla 4-merous; anthers included and style with stigma exserted (plants not heterostylous); calyx lobes ± equal, flowers axillary 8. Conostomium - Corolla 5-merous. anthers exserted and style with stigma included or viceversa (flowers hetero­ stylous); calyx lobes unequal; inflorescences ter­ minal. ? 5. Corolla with a conspicuous crimson or purplishred "eye” : inflorescences elongated, spike-like in fruit, up to 30(-40) cm long. 5. Otomeria - Corolla without an “eye”; inflorescences some­ times ± elongated in fruit, but if so, only a few cm long. 4 Pentas 6. Corollamostly 5-merous 7 - Corollamostly 4-merous 9 7 Small annual herbs with linear leaves 10-30 x 1-3 mm: inflorescences axillary, 1-3-flowered. 12. Thecorchus - Perennial bert>s or subshrubs with larger leaves; in­ florescences terminal or (pseudo-)axillary, but then very lax. elongated, and with more flowers 8 8. Erect herbs or subshmbs (occasionally decumbent forest floor herbs), inflorescences terminal. 4 Pentas - Decumbent herb of wet places; flowers in very- lax, elongated (pseudo-)axillary inflorescences. 11. Pentodon 9. Capsule opening both septicidally and loculicidally. 1° Agathisanthemum - Capsule opening only loculicidally. 10 10. Anthers and/or style with stigmas exserted (flow­ ers heterostylous). 13 Oldenlandia - Anthers and style always included 11 11. Calyx lobes enlarged, foliaceous in fruit, up to 5 mm long; inflorescences axillary. 1-3-flowered 12 Thecorchus - Calyx lobes not enlarged in fmit, smaller, inflores­ cences axillary or terminal, many- to fewflowered. 12 12 Anthers always inserted above the level of the stigma (but included in the corolla tube); corolla lube narrowly cylindrical. 7. Kohautia - Anthers overtopped by stigma; corolla tube cylin­ drical to funnel-shaped. 13. Oldenlandia

4. PENTAS Benth. (1844) Perennial herbs, dwarf shrubs or shrubs. Leaves decussate or in whorls of 3-5. Stipular sheath with several setae. Rowers mostly 5-merous. usually heterostylous (ifaostly di-. seldom tristylous), in terminal, usually branched.

141. RUBIACEAE: 4. Pentas

rrany- to lew-flowered cymose inflorescences. Calyx lc »=s subequal to unequal with 1-3 lobes enlarged, either coloured or green, foliaceous. Corolla shortly cylindrical tc long, tubular, hairy in the throat, lobes valvate in bud. ov ate to oblong. Anthers (long) exserted or included in widened uppermost part of tube (long-styled flowers). S^yle with bilobed stigma included (short-styled flowers) O’ exserted. Ovary 2-locular. ovules numerous on pla­ centa attached to septum. Fruits crowned by persistent calyw capsular, beaked, opening into 4 valves at beak, or sometimes splitting septicidally. Seeds minute, often ir­ regularly shaped, brownish to blackish. An Afro-Madagascan genus of close to 40 species; 1 species extends to SW Arabia; 7 species in the Flora area. Species 1-3 belong to subgen. Vignaldiopsis Verde. (1953) having flowers with (sub)equal. subulate to linear calyx lobes and rusty reddish-brown indumentum. Spe­ cies 4 belongs to subgen. Longiflora Verde. (1953) with corolla tubes long, calyx lobes subequal and leaves often inwhorls of 3-5. The remaining species belong to subgen. Pvntas Verde. (1953) in which the flowers have 1-3 en­ larged. elliptic to more or less deltoid foliaceous, green cafiyx lobes. 1. Leaves in whorls of 3(4). 4 P. decora var. triangularis Leaves decussate. 2 Leaves 0.2-1 (-2) cm wide; corolla scarlct-red. 6. Leaves usually over 1 cm wide: corolla never bright red. 3 Corolla tube (40-)45-65 mm long, calyx lobes veiy unequal: flowers not heterostylous. 5. P. caffensis Corolla tube much shorter, up to 35 nun at the most; flowers heterostylous. 4 Calyx lobes veiy unequal, the largest lanceolate. 5-13 mm long, the smallest 1-3 mm long. 7 Calyx lobes (sub)equal, often ± linear. 1.5-10 nun long 5 Shmb or woody herb 0.5-3 m tall: stipules large, with 6-10 bristles 5-15 mm iong: inflorescence many-flowered. ±corymbose. 40—100(—150) mm wide, all parts velvety-rusty (rarely whitishyellowish) pubescent; calyx lobes (3—)5—10 nun long 1. P. schimpcriana Perennial herb up to 0.5(-0.75) m tall: stipules small, with 3-5 bristles 1-6 mm long; inflore­ scence not as above: calyx lobes 1.5—5 mm long. 6 Leaves 2-6 cm wide, with whitish hairs. 2. P. tenuis Leaves 0.5-2.5(-3) cm wide, with rusty-brown hairs on veins below and whitish hairs on the sur­ faces. 3. P. concinna 1. P. schimperiana (,4. Rich.) Vatke (1876) subsp. schimperiana; Vignaldia schimperiana A. Rich. (1848) - type: TU. Cliire [Sliirej, Quartin Dillon 126 (P lecto.. K isolecto.).

203

Pentas schimperi Engl. (1895), nom. superfl. P. thomsonii Scott Elliot (1896); Neurocarpaea thomsonii (Scott Elliot) S. Moore (1905). Mussaenda schimperi Hochst., nom. nud. Shrub or woody herb 0.5-3 m tall, soft-w'ooded, younger parts densely rusty-brown pubescent. Leaves decussate, ovate to ovate-lanceolate, (5—)6—15(—22) x (1.5-)2-6 (-10) cm, mostly acuminate at apex, cuneate at base, rusty-brown hairy at least on veins below, very rarely glabrescent or pubescence whitish-yellowish; petioles ab­ sent or up to 1(—3.5) cm long. Stipular sheath large, conspicuous, with 6-10 bristles. 5-15 mm long. Inflore­ scence many-flowered, branched, more or less coiymbose. 40—100(—150) nun wide, all parts velvety-rusty (rarely whitish-yellowish) pubescent. Flowers hetero­ stylous. Calyx lobes (sub)equal. linear, (3—)5—10 mm long. Corolla white tinged pink on the outside or pinkish, tube 5-12 mm long, funnel-shaped (long-styled flowers) or cylindrical (short-styled flowers), lobes more or less lanceolate, 3-6 x 1—2(—3) mm. Capsule 4-6 mm long and wide. Fig. 141.3. Montane scrub, bushland, edge of montane rainforest, in open places in more or less disturbed forest; (1800-) 2000-3000(-3200) m. TU GD GJ SU AR CL KF GG SD BA HA; south to Malawi and Zambia. Mesfm T. et al. 3597; Puff et al. 811001-1/1; P uff & Ensermu K. 821219-1/1. Plants prefer open, more or less sunny habitats, but are P. parvifolia sometimes found inside dense evergreen forest; these “shade forms" seem to flower rarely and may look some­ what atypical having abnormally large and glabrous leaves up to 22 x 10 cm and petioles up to 3.5 cm long; e.g. Friis et al. 2104. A second subspecies, subsp. occidentalis (Hook, f.) Verde., with shorter calyx lobes, occurs mostly in W Afnca. P. Verde. lanceolata 2. P. tenuis (1966) - type: BA. Hclgol, Mooney 8454 (K holo.. ETH iso.). Perennial herb up to 30-50(-75) cm tall, single- or fewstemmed, often more or less erect; stems thin, rustybrown hairy Leaves decussate, ovate to elliptic. 3-10 (-13) x 2-6 cm. acute at apex, gradually narrowed to base, thin, sparsely covered with whitish hairs; petioles 0.5-2.5 cm long. Stipular sheath small, with 3-5 bristles to 6 mm long. Inflorescence manv-flowered. made up of 3-4 slightly elongated cymose branches; inflorescence branches elongated to 50 mm in fruit. Flowers hetero­ stylous. Calyx lobes (sub)equal. more or less linear. (2.5—)3—5 nun long. Corolla white, tube 5-7(-8) mm long, funnel-shaped (long-styled flowers) or cylindrical (short-slvled flowers), lobes oblong-lanceolate. 2.5-4 x 1-1.5 mm. Capsule 3-5 mm long and wide. Evergreen, often riverine forest, frequently in wet to moist shady places; 1300-2000 in. SU WG KF SD BA; endemic. Mesfm T. 574; P uff & Ensermu K. 820811-1/1, 820819-3/4. ’

204

141. RUBIACEAE: 4. Pentas

Figure 141.3. P E N T A S SC H IM P E R IA N A subsp S C H IM P E R I A N A : 1 flowering branch x W, 2 - short-styled flower x P/2; 3 - long-styled flower A opened corolla with subsessile an­ thers x 1Vi; B - calyx, ovary and style with stigma lobes x l'/i; 4 capsule with persistent calyx x IV2; 5 — capsule with calyx removed to show beak x \ V2. 1 from living spec­ imen and Honeybee Flora o f Ethio­ pia p 189; 2 & 3 from Puff & Ensermu K 861109-1 /3; 4 & 5 from Gilbert. Thulin & Getachew A. 491. Drawn by Sarah Howard.

4 P. decora S. Moore (1910) var. triangularis (De Wild.) 3. P. concinna K. Schum. (1903) (non sensu Verde. I erdc., in Bull. Jard. Bot. Brux. 23: 291 (1953): (1953)1 P. trianguJaris De Wild. (1914) - type from Rep types: HA. Gara Mulata. Ellenbeck 563 & Ml Dem du Congo. Hakim, Ellenbeck 881 (both B, syn. destroyed). P. globifera Hutch. (1921). Perennial up to 40 cm tall, several-stemmed, woody at P. verticillata Scott Elliot var. pubescens S. Moore base; stems rusty-brown hairy. Leaves decussate, ovate to in Journ. Bot. 48: 220 (1910), nom. inval. elliptic, (2-)3-6(-7.5) x 0.5-2.5(-3) cm. acute at apex, Perennial up to 1.5 m tall; stems erect, usually unbranched thin, upper suface sparsely, lower surface more or less and single from a w oody base, pubescent above. Leaves in densely covered with whitish hairs, hairs on veins below whorls of 3(4), ovate-lanceolate or lanceolate, 4—9(—1i) x rusty-brown: petioles 0.2-1.1 cm long. Stipular sheath 1.5-4(— 4.5) cm. 1 acute at apex, pubescent, subsessile. with a few bristles 1-2 mm long. Inflorescence several- to Stipular sheath \Vitli several bristles c 7 mm long. Inflore­ many-flowered, made up of c 3, slightly elongated, cy­ scence many-flowered, made up of 3-5 congested mose branches, 30-40 mm long and wide. Flowers branches, more or less capitate, 30-50 mm in diameter heterostylous. Calyx lobes (sub)equal. linear. 1.5-4 nun (excl. corolla); inflorescence branches slightly elongated long. Corolla pink and white, tube 6-8(-10) mm long, in fruit Flowers not heterostylous, style with bilobed lobes 3-4(-5) mm long. Capsule 2.5-5 mm long. stigma always exserted, anthers included. Calyx lobes Limestone cliffs; 1400-1800 m. IL HA; endemic. (sub)equal, narrowly triangular to more or less linear, Burger 3153. 4-8(-10) mm long Corolla wliite, velvety pubescent out­ side. tube 60-100 mm long, lobes elliptic-lanceolate, Poorly known and obviously very' close to the previ­ 8-12(-15) x 2-4(-5) mm. Capsule obovoid. 7-10 x 5-8 ous species, but both ecologically and geographically well mm. with a distinct beak, pubescent, ribbed. separated.

141. RUBIACEAE: 4. Pentas

Grassland, wooded grassland, bushland: 1400-2000 m SU WG IL KF: west to Nigeria, south to Tanzania (? and Malawi). Gilbert & Thulin 611: Puff & Ensermu K. 82 >813-3/1. 820818-6/1. Two other varieties, differing primarily in having mostly glabrous leaves, are recognized: var. decora (rather widely distributed in tropical Africa) and var. la.s.ocarpa Verde. (SW Tanzania).

5. P. caffensis Chiov. (1940) - type: KF. nr. Gogeb [Gojcb] River, Paiella 275 (FT holo ). Perennial up to 1(—1.5) m tall; stems erect, usually unbr&nched and single from a w oody base, pubescent above. Leaves decussate, ovate-elliptic or lanceolate. 3—10(—11) x (if-) 1.5-5 cm. acute at apex, cuneate at base, pubesccnt. sut sessile. Stipular sheath with several bristles up to 7(—10) mm long. Inflorescence many-flow ered, made up of several congested branches, more or less capitate. 20- M)(-60) mm in diameter (excl. corolla): inflorescence brarehes slightly elongated in fruit. Flowers 5(6)-merous. not heterostylous. style with bilobed stigma always exserted, anthers included. Calyx lobes 5-7. very un­ equal. 1-3 lobes foliaccous. up to 10 x 2.5 rnin. the others much smaller. Corolla white, hairy outside, tube (40-) 45^'»5 111111 long, lobes lanceolate. 5-10 x 2-3 mm. Cap­ sule more or less subglobose. c 5 mm in diameter. Grassland, wooded grassland, woodland, woodlandbamboo mosaic; 1400-2000 m. SU WG IL KF SD BA, endemic. Hildebrandt 161; Afoonev 9266; Puff 2—3.5 mm (bisexual, male) or less than 1 mm (female) long; lobes 2-2.5 mm (bisexual, male) or less than 1 mm (female) long. Anthers exserted (male, bi­ sexual). Stigmas exserted, up to 10 mm long (bisexual, fe­ male). Fruits yellowish-brown to reddish-brown, with a more or less broad longitudinal groove between the meri-

141. RUBIACEAE: 18. Anthospermum

233

Figure 141.19. A N T H O SP E R M U M H E R B A C E U M 1 - habit x 2/ 3; 2 stipule x 2; 3 - male flower; position o f rudimentary stigmas and detail o f co­ rolla lobe x 4; 4 & 5 - female flowers with long exserted stigmas x 4; 6 - fruit with carpophore in side view x 8; 7 carpophore with mciiearp (ventral side) x 8; 8 - seed x 8. 1 ,2 , 6 -8 from Milne-Redhead & Taylor 10452; 3 from Polhill & Paulo 1414; 4 from M ilneRedhead & Taylor 9399; 5 from Rounce 422. Drawn by Ann Davis. (Reproduced with permission from FI. Trop. E. Afr. R ubiaceae: fig. 46.)

cmps, crowned by obscure minute calyx lobes or calyx lobes absent, surface “warty” or more or less smooth, n::ver hairy; each meriearp 1.7-2.5 x 1-1.5 mm, oblong to n ore or less obovate. Fig. 141.19. Grassy slopes, grassland with bush clumps, farmland, s :rub, edge of montane evergreen forest; often in more or 1»SS disturbed places; 2000-3000 m. EW TU GD WU SU AJR KF GG SD BA HA; south to South Africa; SW Arab a. Mooney 9991; P uff & Ensermu K. 821219-2/8; Puff e al. 810916-3/1. Variable in growth form, leaf size and shape. Someti ties approaching the following species in these characU rs but easily distinguished by its calyces, ovaries and fruits.

2. A. pachyrrhizum Hiern (1877) type: TU, from Maigoigoi to Debra Sina, Quartin Dillon & Petit s.a (P lecto.). A. hirsutum sensu A. Rich. (1848), non DC. (1830). A. unisetum Hochst., nom. nud.

1

Rounded or more or less cylindrical dwarf shrub or subshrub up to 0.4 m tall. Leaves decussate (rarely in whorls of 3), often seemingly in much larger numbers at nodes due to much-contracted leafy short shoots, narrowly lan­ ceolate to linear-lanceolate, (0.5-)0.9-I.8(-2.5) x (0.05-) 0.1—0.2(—0.3) cm, acute at apex, narrowed to base, mostly glabrous except for margins, upper surface often with large, conspicuous epidermis cells. Stipular sheath with a

141. RUBIACEAE: 18. Anlhospcrmum. 19 Rubia

234

small seta. Flowers 4-merous, male, female (seldom also bisexual; plants often dioecious), subsessile to very shortly pedicellate (female), in several- to few-flowered axillary clusters. Calyx lobes often distinctly unequal. Co­ rolla greenish-vellow to pale yellow, sometimes purplish outside, tube funnel-shaped, c 1 mm (male) or up to 0.5 mm (female) long; lobes 2-2.5 mm (male) or less than 1 nun (female) long. Anthers exserted (male). Stigmas exserted, up to 10 mm long (female): ovary densely hairy (well discernible rudimentary ovary in males). Fruits red­ dish-brown densely covered with white, spreading hairs, crowned by unequal calyx lobes, the larger ones up to (1.3—)1.7 mm long; each meriearp 2 x 0.8 mm, more or less oblong. Cracks of rocks in cliffs (mostly basalt), occasionally in rocky' grassland or in disturbed sites (steep, rocky noadbanks. etc.); 1700-3100 m. EW TU GD GJ SU; Su­ dan; Yemen Arab Republic. Selassie 672; Puff et al. 811005-4/1; Tewolde B.G.E. 1099. 3. A. villosicarpum (Verde.) Puff( 1986); A. herbaeeum L.f. var. villosicarpum Verde, in Kew Bull. 30: 299 (1975) - type from Kenya. Dwarf shrub or subshrub with woody base. Stems up to 0.6 m long, erect or ascending, younger parts hairy. Leaves in whorls of 3. but seemingly in much larger num­ bers at nodes due to much-contracted leafy short shoots, narrowly elliptic-lanceolate, 1-1.3 x 0.1—0.3 cm. acu­ minate to mucronate at apex, narrowed to base, with bris­ tly hairs above and below, particularly on midrib, margins revolute. Stipular sheath with a small seta. Flowers 4-merous. bisexual, female (plants bisexual or bisexual + female), subsessile in several- to few-flowered axillary clusters. Calyx lobes subobsolete. Corolla greenish to greenish-vellow; tube more or less narrowly funnelshaped. c 1 111111 (bisexual) or less than 1 mm (female) long: lobes c 2 inm (bisexual) or c 0.5 nun (female) long. Anthers exserted (bisexual). Stigmas exserted. up to 2.5 mm long. Fruits brown to reddish-brown, densely shortly hairy. not crowned by distinct calyx lobes; each meriearp 2.5 x 1.2-1.3 mm. obovate. Rocky places in Olea Juniperus scrub; 1800-2000 m. SD. N Keny a. Gillett 14448. RUBIEAE

Mostly herbaceous perennial or annual plants with true leaves and foliaccous stipules forming pseudo-whorls In­ florescences mostly terminal Ovary 2-locular; one ovule per locule. attached to septum. Fmits fleshy or dry. split­ ting into 2 indehiscent mericarps. 1.

2.

Leaves and leaf-like stipules distinctly petiolate; flowers mostly 5-merous. 19. Rubia Leaves and leaf-like stipules (sub)scssilc; flowers usually (3)4-merous. 2 Leaves and leaf-like stipules in whorls of 4. 1veined. small annuals with short. 3-flowered axillary cymes. 21. Valantia

-

Leaves and leaf-like stipules in whorls of 4 to many (if in whorls of 4, leaves 3-veined, inflorescences terminal and more extensive than above, and plants perennial). 20. Galium 19. RUBIA L. (1753)

Scrambling, creeping or climbing perennial herbs, son ewhat woody near base, roots quite woody, reddish (y ield­ ing a red dye). Stems branched. 4-angled, more or less brittle, mostly beset with recurved prickles. Leaves and leaf-like stipules in whorls of 4-8(-12), distinctly peti­ olate or, (not in Flora area) sessile, blades cordate or broadly ovate, ovate-lanceolate to linear, with l-5(-7) prominent veins. Flowers 5(rarely 4 or6)-inerous, bisex­ ual. maxillary and/or terminal cymes. Calyx lobes obso­ lete. Corolla rotate to (sub)campanulate. greenish yellowish-green lo yellow, lobes more or less triangular, (long-)acuminate Anthers exserted. Style branches 2, joined below; stigmas capitate. Ovary crowned by minute disk. 2-locular. one ovule per locule. Fruits fleshy, gla­ brous. consisting of 2 round mericarps, each with a single seed, round on dorsal side, plane to convex on ventral side, one inencarp often aborted. An Old World genus of c 60 species; 3 species occur in Africa south of the Sahara, only the following in the Flora area R. cordifolia L. (1786) subsp. conotricha (Gand.) Verde, in Kew Bull JO 323 (1975); R. conotricha Gand (1918) - ty pe: South Africa Schlechter 6550 (LY holo., K iso). var discolor (Turzc.) K. Schum. in P.O.A.C.: 395 (1895); R. discolor Turzc. (1845)-type: TU, Mt Scholada. Schimper 24 (CW ?holo., BM K UPS iso ). R. cordifolia sensu A. Rich. (1848). Perennial herb, stems up to several metres long, with quite massive recurved prickles on the angles, often also pu­ bescent Leaves in whorls of 4, blades lanceolate to broadly ovate. (1—)1.5—6(—8) x 0.5-3 cm, acuminate at apex, cordate to (less often) rounded at base, 3-5(-7)veined. scabrid pubescent above, densely velvety v illose orv\ holly hairy below, margins often with recurved prick­ les; petioles 0 5-7 cm long, with recurv ed prickles, often also pubescent Inflorescences numerous along the stems, lax to more or less dense, usually many-flowered; pedi­ cels up to 5 mm long Corolla yellow, creamy yellow, yel­ low ish-green or greenish, often reddish or purplish tinged in bud. mostly distinctly campanulate. tube 1-1.7 mm long (seldom less), lobes less than 1—1.5 nun long, apiculate or acuminate Each meriearp 2-3 mm in diameter Fig 141.20 Scrub, bush clumps, remnants of montane forest, in more or less disturbed areas; (1000-) 1650-2850 m. EW TU GD GJ WU SU AR KF SD BA HA; N Somalia, scat­ tered in tropical E Africa. P uff et al. 810913-1/1. 811007-2/3. 820921-1/1.

141. RUBIACEAE: 19. Rubia, 20. Galium

235

Figure 141.20.

RUBIA CORDIFOLIA subsp CONOTRICHA var DISCOLOR 1 - habit x Vi, 2 - magnification o f leaf tip showing pubescent under­ side and scabrid upper x 15; 3 - bud x 6; 4 —flower x 3; 5 —ovarv with styles and stigma x 6; 6 - fruit x 3. 1 from Petelin 7 1, 2 5 from P u ff et al. 820918-4/1; 6 from Ensermu K. B583. Drawn by Sarah Howard.

Variable in leaf shape and size, and in petiole lengths b ii not or only to a limited extent in leaf indumentum and corolla shape. Subsp. conotricha, referring to African material of the widely distributed Afro-Asiatic/?, cordifolia , occurs from NE to S Africa. Var. discolor seems to be centred in Ethiop a var. conotricha, with more or less rotate corollas, is c< mmon elsewhere. 20. GALIUM L. (1753) P :renrual herbs or annuals. Stems erect, creeping or c i ibing, often distinctly 4-angled. Leaves and leaf-like stipules in whorls of 4-10. linear to (ob)ovate, mostly with a prominent midvein (1-veined), (sub)sessile. F o wers 4-merous, mostly bisexual, in axillary and/or terir uial man\- to few-flowered cymes. Calyx lobes obso­ lete Corolla mostly rotate, greenish, whitish, white or y llow, lobes ovate to triangular, acute to acuminate. An­ ti ::~s exserted. Style branches 2. joined below; stigmas c;;:itate. Ovary crowned by an often 2-lobed disk, 2loc liar, one ovule per locule. Fruits dry. rarely fleshy, gla­

brous or variously hairy, dehiscing into two mostly (sub)globose mericarps, each with a single seed, round on dorsal side, more or less hollow on ventral side; one meri­ carp occasionally aborted. A cosmopolitan genus of several hundred species,

mostly in extratropicai regions; 8 species in the Flora area. 1. Leaves (and leaf-like stipules) 3-veined. always in whorls of 4; margins never prickly. 1. G. thunbergianum - Leaves (and leaf-like stipules) 1-veined. in whorls of 4-8; margins sometimes prickly. 2 2. Leaf margins with ± recurved prickles or small teeth. 3 - Leaf margins without recurv ed prickles. 9 3. Mericarps very densely covered with long, yellow­ ish to brownish hairs. 2. G. chloroionanthum - Mericarps ± sparsely covered with short, some­ times hooked hairs, or glabrous. 4 4. Mericarps w ith short, sometimes hooked hairs. - Mericarps glabrous.

5 7

236

141. RUBIACEAE: 20. Galium

5. Flowers mostly solitary on peduncles/pedicels, up to 25 mm long. 4. G. spurium subsp. africanum - Inflorescences of 1 to several to many flowers, 6 6. Inflorescences several- to many-flowered, corolla large, 4-5 mm in diameter, lobes (long-) apicu­ late; leaves 3-8 mm wide. 3. G. aparinoides - Inflorescences 1- or few-flowered, corolla small, 1-1.5 mm in diameter, leaves narrower and smaller. 0.5-2 mm wide. 6. G. acrophyum 7. Inflorescences 1- or few-flowered, corolla small, 1-1.5 mm in diameter, leaves 2-12 x 0.5-2 mm. 6 G. acrophyum - Inflorescences several- to many-flowered, corolla large, (2-)2.5-4 mm in diameter, leaves 6-30 x 1-6 mm. 8 8. Fruits distinctly fleshy; flowers yellow, corolla ro­ tate; stem and leaf margins with coarse, recurved prickles, plants harshly adhesive; found in scrub and grassland 5. G. simense - Fruits dry; flowers pure white, sometimes tinged pinkish in bud, corolla slightly campanulate; stem and leaf margins without coarse prickles, con­ fined to damp or wet places. 7. G. scioanum 9. Leaves 2-12 x 0.5-2 mm, corolla small, 1—1.5 mm in diameter, inflorescences 1-or few-flowered. 6. G. acrophyum - Leaves up to 30 mm long, corolla 2—3(^4) mm in diameter, inflorescences several- to manyflowered 10 10. Leaves elliptic to linear-lanceolate, 1-4 mm wide, acute to ± obtuse at apex; flowers pure white, sometimes pinkish tinged in bud, corolla slightly campanulate; confined to damp or wet places. 7. G. scioanum - Leaves linear, 0.3-0.7 (-l) mm wide, with a long hyaline point at apex; flowers yellow, corolla ro­ tate; plants not in damp or wet places. 8. G. boreo-aethiopicum 1. G. thunbergianum Eckl. & Zeyh. (1836) - type from South Africa. Perennial with extensive rhizome. Stems straggling or procumbent, (4-) 10-45 cm long, glabrous or with spread­ ing white hairs. Leaves in whorls of 4, 3-veine.i, ovate, el­ liptic or more or less rhombic, 3-15 x 2-8 mm, more or less acute at apex, narrowed to base, glabrous or both sur­ faces and margins with spreading white hairs. Inflore­ scence usually more or less lax, broadly pyramidal to cylindrical, cymes rather many-flowered; pedicels 1-3 (-5) mm long at anthesis, slightly elongated and more or less divaricate in fruit, glabrous or hairy. Corolla greenish to yellowish, sometimes tinged reddish-purplish, 2(-3) mm in diameter. Fruit densely covered with white tuberculate, hooked hairs; mericarps more or less subglobose, each 1-1.5 mm in diameter. Fig. 141.21.1 & 2. Grassland with rocky outcrops or bush clumps, or open grassy patches in forest remnants or in ericaceous scrub; (2200-)2700-3600 m. GD GJ SU AR GG BA; south to South Africa; W Africa.

1. Leaves and stems glabrous (or nearly so). var. thunbergianum - Leaves and stems sparsely to densely covered with spreading white hairs. var. hirsutum var. thunbergianum G. natalense Rouy in Rouy & Foucard. FI. France 8: 9 (1903), in a note. Less common than var. hirsutum, probably only GD SU. Mesfin T. & Kagnew 1881; P uff et al. 810927-2/2. var. hirsutum (Sond.) Verde, in Kew Bull. 30: 326 (1975); G. rotundifolium L. var. hirsutum Sond. (1865) type from South Africa. G. dasycarpum Schweinf. (1867) - type: GD, Semien, Debra-Eski. Schimper s.n. (B holo. destroyed) G. biafrae Hiem (1877). Distribution as for species as a whole. Mesfin 670; P uff et al. 820925-3/3; Puff&Ensermu K. 821219-1/5. 2. G. chloroionanthum K. Schum. (1901) - type from Tanzania. Scrambling or climbing perennial. Stems to 1 m long, with coarse recurved prickles on the 4 angles. Leaves in whorls of 6 , 1-veined. elliptic-oblanceolate, (10—) 15—? 5 x 5-8 mm, mucronate at apex, narrowed to base, margins and midvein below with coarse recurved prickles, blades otherw ise mostly glabrous. Inflorescence 1- to 3-flowered; peduncles 20-25 mm long; true pedicels up to I mm long, mostly supported by a single bract, less than 10 mm long. Corolla pale yellow to yellowish-green, up to 2 mm in diameter, lobes more or less acute. Fruit dry. dark brown, covered with whitish-brownish hooked hairs, 1-1.5 mm long; mencarps 2-3 mm in diameter. Montane ev ergreen forest; 2600-2700 m. KF; from S Sudan and tropical E Africa to Zimbabwe, Madagascar Puff& Ensermu K. 861111-1/3. 3. G. aparinoides Forssk. (1775) - type. Yemen Forsskal (C lecto ). G. hamatum A. Rich. (1848) - types: GD, Wodjerat. Petit s.n. & Simien, Mt Selki, Schimper 671 (both P sya, K UPS isosyn.). G. aparine L. var. hamatum (A. Rich.) Hook.f in Joum. Linn. Soc. 6: 11 (1861), quoad nom. excl. specim. cit Straggling or scrambling perennial. Stems up to 50 cm long, with coarse recurved prickles on the 4 angles. Leaves in whorls of 6, 1-veined, oblanceolate, elliptic or linear-(ob)lanceolate, (8-)10-25(-30) x 3-8 mm, acu­ minate to mucronate at apex, narrowed to base, margins and midvein below with coarse, recurved prickles, blades otherwise glabrous or sometimes bristly hairy above. In­ florescence several- to many-flowered; pedicels up to 3 mm long. Corolla often tinged reddish-purplish, yellow­ ish-green, greenish or pale yellow, 4-5 mm in diameter, lobes (long) acuminate. Fruit dry, black, covered with

141. RUBIACEAE: 20. Galium

sho t, whitish, hooked hairs; mericarps 2.5-3.5 mm in diameter. Fig. 141.21.3 & 4. -iontane evergreen forest (often in clearings and at the edg: i. secondary montane scrub, grassv slopes. (1700-) 22C K3600 m. EW TU GD WU SU AR SD BA HA, tnojjcal E African mountains; SW Arabia. Puff et al. 810 '26-2/2, 820921-3/2, 820930-3/1. . \ 4. ( . spurium L. (1753) ubsp. africanum V erdeinK ew Bull. 30: 324 (1975) - type: Kenya, Milne-Redhead & Taylor 7147 (K iolo.). G. aparinoides sensu Cufod. (1949, 1965), non -"orssk. We.k. scrambling annual. Stems 15-60(-100) cm long, witl recurved prickles on the 4 angles. Leaves in whorls of x 1—4(-5) mm, distinctly acuminate at apex, nar­ row :d to base, margins and midvein below with coarsc re­ cun :d prickles, blade otherwise glabrous. Most of the flov ers/fruits solitary on unbranched peduncles arising fror main stem or branch nodes; peduncles often more or less ;:urved and up to 25 mm long in fruit; some flowers occ; sionally also in few- to several-flowered cymes. Co­ rolla white, greenish-white, greenish or yellowish-green. 1-1 ^1- 2) mm in diameter. Fmit dry, black, covered with hoo ed hairs; mericarps 2-3.5 mm in diameter. Fig. 141 .11.5-8. J lostly in much disturbed areas (grassland or bush­ land. edge of scrub, roadsides, forest edges, etc.); (850-) 100« -3000 m EW TU GD GJ WU SU AR WG IL GG SD BA HA; Somalia; from Sudan south to South Africa. Mesfin T. & Kagnew 1681; Puff et al. 811005-1/6. 820‘ 20-1/3. J obsp. spurium. with more extensive inflorescences, is native to Europe, W Asia and N Africa but lias also been intrc :iuced as a weed elsewhere. It may also be introduced to th : Flora area. e.g. GD. Puffela\. 811005-5/4; BA. Puff et al *70512-5/1.

237

ish-white, (2-)2.5-4 mm in diameter, lobes acute to shortly acuminate. Fruit fleshy, dark purple to black, gla­ brous; mericarps 2-4 mm in diameter. Fig. 141.21.9-13. Scrub (forest edge, bushland, farmland, roadsides, plantations); less often in grassland, in the shelter of rocks or, at high altitudes, in tussock grassland; (1700-) 2000-3700M 000) m. TU? GD GJ SU AR WG IL KF GG SD BA; tropical E African mountains, W Africa. P uff engadum (1979); Randia nilotica Stapf (1906); Lachnosiphonium niloticum (Stapf) Dandy (1952); Xeromphis nilotica (Stapf) Keay (1958)-type: Sudan [nr GD/GJborder], Sennar, Kotschy 400 (K. holo.). Gardenia dumetorum sensu A. Rich. (1848), non Retz. Shrub or low, several-stemmed tree up to 5 m tall: branches grey to whitish; thorns massive, woody, up to 2.5 cm long, solitary and alternate above paired, muchcontracted, leafy' short shoots. Leaves obovate. 1—L5(-6)

Figure 141.33 C A T U N A R E G A M N IL O T IC A 1 - flowering branch x 2; 2 -calyx opened x 2; 3 -c o ro lla opened x 2; 4 - fruit \ 2 No artist given and no specimens cited in orig. publ. (Repro­ duced from Andrews. FI. Angl.-Eg. Sudan fig. 159 as Lachnosiphonium niloticum.)

x 0.5—1 5(—2) cm. obtuse at apex, long-cuneate at base, glabrous to more or less pubescent. Flowers (4-)5-6merous, solitary or fasciculate, shortly pedicellate Calyx glabrous, lobes up to 3 mm long. Corolla hairy, tube somewhat longer than calyx, lobes c 5 mm long. Fruits subglobose (8-) 10-18 mm in diameter, glabrous. Fig. 141.33. Woodland or busliland, or grassland with scattered trees, 400-1700 m EE EW GD/GJ?; west to Cameroun and Nigeria. S Somalia and tropical E Africa. Fiori 436, 616; Schweinfurth & Riva 2249.

34. GARDENIA Ellis (1761), nom. cons. Shrubs or trees. Leaves decussate or sometimes in whorls of 3(4), petiolate. Stipules connate. Flowers 5-12inerous. mostly solitary or paired, tenninal on (very ) short lateral branches Calyx forming a distinct (sometimes long) lube, truncate or with distinct lobes above. Corolla often large, salver- or funnel-shaped, lobes contorted in bud. Anthers included or barely exserted; pollen in tet­ rads. Style with club-shaped or fusiform stigma, mostly

141. RUBLACEAE:34. Gardenia

nc i much exserted. Ovary l-locular, ovules numerous on 2- parietal placentas. Fruits indehiscent, often hard and leuhery. Seeds many, compressed, elliptic, embedded in fit;:; hy pulp \ large paleotropical genus with about 200 species; 3 in ne Flora area. I Semi-desert or desert shrub; leaves bullate, 7-25 x 3-12 mm. densely pubescent; corolla tube 7-15 (-20) mm long; fruits hairy, subglobose, 5-10 mm in diameter. 1. G. fiorii - Not a semi-desert or desert shrub; leaves mostly (much) larger, neither bullate nor densely pu­ bescent: corolla tube 35-100 mm long; fruits not hairy, much larger. 2 I Corolla tube 70-100 mm long, calyx lobes 9; fruits subglobose. conspicuously lenticellate, distinctly (9-)ribbed, (50-)70M 10 x (50-)60-100 mm in diameter. 2. G. volkensii - Corolla tube 35-60(-75) mm long, calyx lobes 6 or 7; fruits elongate or ± ellipsoid, neither conspicu­ ously lenticellate nor distinctly ribbed, (30-) 40-70 x 20-25(-35) mm. 3. G. ternifolia 1 G. fiorii Chiov. (1916); Randia fiorii (Chiov.) Chiov. (1932) - type from Somalia. R. sphaerocarpa K. Schum. (1904), nom. nud. S irub up to 3(-4) m tall, much and densely branched; b; rx smooth, grey. Leaves in whorls of 3(4), often c: owded on short lateral branches, bullate, 7-25 x 3-12 it m. densely pubescent. Flowers 6-merous, solitary. Ca1>x pubescent, lobes 2-5 mm long at first, elongating up to 6-9 mm in fruit. Corolla wliite, pubescent outside, tube narrowly funnel-shaped. 7-15(-20) mm long, lobes 3-12 n m long. Fruits crowned by persistent, enlarged calyx, h iry. subgiobose, 5-10 mm in diameter. 4cacia - Commiphora open scrub and bushland; 4 'i -800 in BA HA; Somalia, N & NE Kenya. Simmons 46. 196.

253

long. Fruits grey, conspicuously lenticellate, distinctly (9-)ribbed, subglobose, (50-)70-110 x (50-)60-100 mm in diameter. Fig. 141.34. Acacia - Commiphora, Terminalia - Combretum woodland, often along (seasonal) watercourses; 5001300 m. GG SD; Somalia; tropical E Africa to South Af­ rica Mesfin T. & Tewolde B.G.E. 2803; Puff & Ensermu K. 821221-4/1, 821228-1/11. Subsp. volkensii var. saundersiae (N.E.Br.) Verde., with exceptionally large calyx lobes, occurs in southern Africa. Subsp. spathulifolia (Stapf & Hutch.) Verde., with smaller fruits, is confined to Angola Zambia and Bot­ swana. 3. G. ternifolia Schumach. & Thonn. (1827) - type from Ghana. Small tree 3-6 m tall; shrubby, gnarled, only to 0.5 m tall if browsed. Leaves in whorls of 3, mostly on short branch­ lets, oblanceolate-obovate, more or less oblong or elliptic, 1—15(—18) x0.7-7.5(-l 1) cm, rounded at apex, cuneate at base, glabrous or scabrid (“sand-papery”). Flowers 6-7merous, mostly solitary. Calyx lobes 6-7 (in the Flora area), basal tubular part of calyx 5 -8 (-10) nun long, lobes mostly shorter than tube. Corolla wliite or creamy, ftiming yellowish with age, tube more or less cylindrical, 35-60 (-75) mm long, lobes 20-40(-60) mm long. Fruits brown­ ish, elongate or more or less ellipsoid. (30-)40-70 x 20-25(-35) mm. neither conspicuously lenticellate nor distinctly ribbed. A veiy common and widespread species in the Flora area and elsewhere in tropical Africa; very variable in growth form and vegetative characters. Two subspecies are recognized: 1. Plants, including youngest shoots, glabrous. subsp. ternifolia - At least youngest parts of stems and stipules sparsely to densely strigose-pubescent. subsp. jovis-tonantis subsp. ternifolia

2 G. volkensii K. Schum. (1904) subsp. volkensii var. volkensii - ty pe from Tanzania. G. somalensis Chiov. (1916). G. somalensis Chiov. var. tubicalyx Chiov. in Re­ sult. Sci Miss. Stef.-Paoli. Coll. Bot.. 93. (1916). G. volkensii K. Schum. var. somalensis (Chiov.) Cufod. Enum.: 1002 (1965). G. stu p in ica rp a Chiov. (1932). Small tree or shrub up to 4-6 m tall, untidily branched. L :aves in whorls of 3, usually crowded on short lateral b anchlets, obovate-cuneate, 2-7(-10) x 1.2—3(— 4) cm. n unded at apex. Flowers (8)9-merous, mostly solitary. C ;ilyx lobes 9, basal tubular part of calyx 4-8 mm long, It Des variable in length, in the Flora area mostly as long as o somewhat longer than the tube. Corolla white, turning y : llowish with age, tube narrowly funnel-shaped to more o less cylindrical. 70-100 mm long, lobes 15-30 mm

Deciduous open woodland, 900-1500 m. Probably confined to IL KF WG. Chaffey 932; Friis et al. 2282; Brehme s.n. subsp. jovis-tonantis (IVelw.) Verde., in Kew Bull. 34: 354 (1979). Decamerisjovis-tonantis Welw. (1859); Gardenia jovis-tonantis (Welw.) Hiem (1877) - type: Angola. Welwitsch 2573 (LISU lecto., BM K isolecto.). G. luteaFresea (1837)-type: GD,“two days N of Gondar”, Riippell s.n. (FR holo.). Woodland (more or less natural or disturbed), grassland w ith scattered trees or scrub; 900-2250 m. TU GD GJ WU SU AR WG IL KF SD BA HA. P uff & Ensermu K. 820807-3/5, 821214-1/10; 870509-6/2. A further subdivision of subsp. jovis-tonantis into the two varieties var. jovis-tonantis, with glabrous leaves, and var. goetzei (Stapf & Hutch.) Verde. (1979) |syn. G.

254

141. RUBIACEAE: 34 Gardenia, 35. Rothmannia

Figure 141.34. G A R D E N IA V O L K E N SII subsp V O L K E N SII var V O L K E N SII I flowering branch x Vy, 2 - part o f branch showing stipules x 3; 3 - juve­ nile leaf x Vy, 4 - flower bud x Vy, 5 calyx x 2 /3; 6 - longitudinal section o f corolla x 2/ 3; 7 - longitudinal section o f calyx and ovary x 2 ; 8 - stigma x 2 ;9 longitudinal section o f ovary showing ovules x 4; 10- cross section of ovary showing placentas and ovules x 4 .1 1 fruit x V y 12 - seed x 3. I & 2 from Glover el al. 2011; 3 from Rally 12 358; 4 , 6 -1 0 from Glover et al. 344; 5 collector not given in orig. publ.; 1 1 from Greenway & Kirrika 10 970. 12 from Zanzibar, collector unknown. Drawn by M E. Church. (Reproduced with permission from FI. Trop. E A fr R ubiaceae: fig. 81.)

goetzei Slapf & Hutch. (1909)), with leaves scabridly pubesccnl ("sand-papery”), does not seem worth keeping. Both plants with glabrous and “sand-papery" leaves can occur witliin single populations O'sand-papery” leaves are often found on browsed individuals). 35. ROTHMANNIA Thunb. (1776) Shrubs or trees. Leaves decussate, or rarely in whorls of 3, large, petiolate, blackish upon drying. Stipules triangular, soon falling. Flowers 5(-8)-merous, mostly solitary, often

seemingly axillary but actually terminal on short, ABBREVIATED shoots, frequently above a single leaf. Calyx more or less tubular, truncate above or with subulate to filiform lobes, splitting lengthwise or not. Co­ rolla large and showy, broadly funnel-shaped or cyiindrial below and more or less campanulate above, lobes con­ torted in bud. Anthers inserted near throat, included or lialf-exserted. Style with large, club-shaped stigma more or less exserted. Ovary 1-locular, ovules numerous or 2 parietal placentas Fruits indehiscent. with a leathery skin. Seeds embedded in a fleshy pulp.

141. RUBIACEAE: 35. Rothmannia

255

Figure 141.35. R O T H M A N N IA U RC E LLIF O R M IS: 1 - flowering branch x 14; 2 - fruit x V2; 3 - cross sec­ tion o f fruit showing laterally compressed seeds embedded in fleshy pulp x 14. 1 from E.F. Gilbert 542; 2 from Thomerson 827; 3 from M ooney 8766. Drawn by Damtew Teferra.

\ paleotropical genus of c 30 specics: American spe­ cie* sometimes included in the genus probably belong to oth r genera; 1 species in the Flora area. R. i!reel Iifo rm is (Hiern) Robyns (1947); Gardenia urcelliformis Hiem (1877); Randia ircelliformis (Hiem) Eggeling (1940) - types from Sudan and Zaire. I Gardenia riparia K. Schum. (1895); Rothmannia tparia (K. Schum.) Fagerl. (1943). Gardenia tigrina Hiem (1898). Randia stenophylla K. Krause (1909). R. spathicalyx De Wild. (1910). Rothmannia arcuata Bremek. (1956). Shr jb or small tree up to 10 m tall, much-branched; young brai :hes reddish-hairv. Leaves elliptic to ovate-elliptic (7- 9-14( -16) x 3-6(-9) cm, acuminate at apex, cuneate at t ase, glabrous, or lower surface (especially veins)

hairy; petioles up to 1 cm long, hairy. Stipules more or less triangular, with setae up to 5 mm long. Flowers 5-merous, solitary and subsessile, fragrant, opening at night. Calyx somewhat hairy outside, shaggy inside, tubular part up to 20 mm long, usually split lengthwise, filiform lobes 5-10 mm long. Corolla hairy, greenish-white with dark-brown spots outside, white with maroon spots in the throat; tube broadly funnel-shaped. 50-60 mm long, lobes up to 25 (-30) mm long, acuminate. Anthers half-exserted. Sty le with clavate stigma up to 75 mm long, stigma to 3 mm wide. Fruit ellipsoid to subglobose, 40-60 mm in diame­ ter, sometimes faintly ribbed. Seeds up to 8 mm long Fig 141.35. Forest or wooded gullies in open woodland; 1200— 2000(—2500) m. GD GJ SU AR WG IL KF SD; west to Guinea, tropical central Africa, south to Zimbabwe and Mozambique, Angola. P uff & Ensermu K. 820813-2/1, 861105-1/2, 8611074/2.

141. RUBIACEAE: 36. Oxyanthus

256

Figure 141.36. O X Y A N T H l S

SPECIOSUS subsp. STENO­ CARPUS 1 - branch with tlowering and fruiting inflore­ scence x V2, 2 - bud x I 3 flower x I; 4 - fruit x c V2. subsp. GLOBOSUS 5 - fruit x c V2. 1 from Samson Dashu 004 and Mogattyi M 13; 2 from Friis 1776; 3 from Samson Dashu 004; 4 from Mesfin T. 229-8246. 5 from Friis 4073. Drawn by Sarah Howard

6 Cl- a X a.

36. OXYANTHUS DC. (1807) Shrubs or small trees Leaves decussate, shortly petiolate Stipules triangular to ovate. Flowers 5-merous. usually white, in pseudo-axillary. few- to many-flowered cymes, inflorescences solitary at nodes. Calyx fused to form a truncate or toothed tube. Corolla salver-shaped, tube (ex­ tremely) long, narrowly cylindrical, lobes contorted in bud. elliptic to lanceolate, acuminate. Anthers and style often with club-shaped, bifid stigma exserted. Ovary (l)2-locular. numerous ovules on fleshy placentas. Fruits indehiscent. leathery, ellipsoidal, fusiform, pynfonn to globose, many-seeded A tropical African genus of c 40 species. 2 in the Flora area. I. Inflorescences many-flowered, corolla tube 35-50 mm long, calyx lobes 1-2.5 nun long; stipules tri­ angular. 1 O. speciosus

-

Inflorescences few-flowered, corolla tube 100—150 mm long, calyx lobes 2.5-4 mm long, stipules tri­ angular at base, linear to subulate above. 2. O. lepidus

1 O. speciosus DC. (1807) - type from Sierra Leone. Shrub or small tree up to 4-6 m tall. Leaves elliptic to ob­ long, (8-) 10-22 x (3—)4—10 cm. acuminate at apex, nar­ rowed to more or less rounded al base, petioles 0.5-1.5 cm long Stipules triangular, up to 12 mm long. Flowers in several- to many-flowered (pseudo-)a\iliary inflorescen­ ces, solitary at nodes. Calyx teeth 1-2.5 mm long. Corolla tube (of open flowers) 35-50 mm long, lobes more or less lanceolate, 10-20 mm long. Style with stigma exserted for up to 10 mm Fruits (sub)globose to (short-)fusifonn, 20-30 mm in diameter or 25-65 x 7-15 mm.

141. RUBIACEAE: 36. Oxyanthus, 37. Tricalysia

1 f ruits fusiform. 25-65 nun long, subsp. stenocarpus - Fruits subglobsc to globose. 20-30 mm in diame­ ter. subsp. globosus Of he five subspecies recognized in O. speciosus. only the \o keyed out above occur in the Flora area. As the distinction of these two is based on fruit shape and size onl a flowering specimen cannot always be correctly as­ sign :d to a subspecies. However, it will often be possible to make a detenni nation by reference to the place of origin of i. collection: subsp. globosus appears to be confined to the ower-lying parts of W Ethiopia (IL and KF?). while subsp. stenocarpus occurs at higher altitudes and is much mor: widely distributed (see below). ; jbsp. speciosus occurs in W Africa, subsp. gerrardii (Soid.) Bridson in South Africa and subsp. mollis (Hatch.) Bridson in E and C Africa.

257

var. unyorensis (.S’. Moore) Bridson in Kew Bull. 34: 121 (1979) O. unyorensis S. Moore (1908) - type from Uganda. O. microphyllus K. Krause (1911). Understorey of lowland rain forest; 500-1000 m. IL IL/KF, tropical E Africa, Rep. Dem. du Congo, Burundi. Chaffey 946; Friis et al. 4077. DIPLOSPORINAE Miq.

Mostly shrubby. Inflorescences axillary and paired at nodes. Flowers mostly small, 4-5(-8)-merous. Chary 2-locular. placentas axile, with one to many ovules per locule. Fruits mostly small. Seeds embedded in fleshy pla­ cental tissue or not. 37. TRICALYSIA DC. (1830)

subsp. stenocarpus (K. Schum.) Bridson in Kew Bull. 34: 116(1979), O. stenocarpus K. Schum. (1903) - type from Tanumia. Fig S41.36.l-4. Mouane evergreen forest, often in damp valleys; \3 S -2400 m. GD SU WG IL KF GG SD BA; south to Soi. l Africa Puff& Ensermu K 820819-3/2; P uffct al. 811 >07-4/1. 870510-3/2. subsp globosus Bridson in Kew Bull. 34: 115 (1979) - type from Uganda. Fig 141.36.5. Lo'a and rain forest; 1000-1200 m. IL IL/KF; Uganda. Tanzania, tropical central Africa. Friis et al. 4073, 4126.

Slirubs or small trees. Leaves decussate, shortly petiolate. Stipules cup-like, apiculate. Flowers fragrant. (4-)5-6 (-12)-merous, in paired, sessile or shortly pedunculate axillary clusters: bracteoles often distinctly cup-like. Ca­ lyx cup-like below, truncate, toothed or variously lobed above. Corolla mbe cylindrical to funnel-sliaped, lobes contorted in bud, more or less elliptic. Anthers exserted. Sty le with 2-lobed stigma exserted. Ovary 2-locular. one to many ovules per locule, embedded in placentas at­ tached to septum. Fmits drupes, crowned by persistent ca­ lyx; endocarp thin. Seeds 1-several per locule. An Afro-Madagascan genus of over 100 species; c 95 restricted to Africa; 1 in the Flora area. T. niamniamensis Hiem (1877) - type from Sudan.

2. O. lepidus S. Moore (1905) Shrub up to 5 m tall. Leaves evergreen or occasionally de­ O. ox} carpus S. Moore (1907)- type from Uganda. ciduous. elliptic or ovate. 1.5—8(—9) x l-3(-4) cm, more or less acute to rounded at apex, cuneate, rounded or cordate Shrub up to 2.5 m tall. Leaves elliptic, 4—15(—16) x at base, discolourous. with pilose domatia in angles be­ 1.5 '.5 cm, acuminate at apex, narrowed to base, glatween midvein and lateral veins; petioles 1-3 mm long. broi or finely pubescent beneath or with a few coarse Flowers sometimes appearing before the leaves, 6-merous. liai ; on nerves below; petioles c 1 mm long. Stipules tri­ in subsessile axillary clusters of 3 to several, subtended by ant t! ar at base, linear to subulate above, up to 10 mm cupular bracteoles. Calyx cup-like, 1-2 mm long, with Ion : Flowers in few- (mostly 2^)-flowered (pseudo)minute teeth. Corolla creamy wliite, tube densely hauv out­ axi ry inflorescences, solitary at nodes. Calyx teeth side, 4-6 nun long, lobes 2-4 nun long. Ov ary pubescent, 2.5-- mm long. Corolla tube (of open flowers) 100-150 with two ovules per locule. Fruits 5-7 mm in diameter, with mn long, lobes more or less lanceolate, 20-25 mm long. normally 1 hemispherical seed per locule. St> le with stigma exserted for c 5 mm. Fruits yellowish, ellipsoidal to ovoidal, 20-25 x 12 mm. : he species is divided into 2 subspecies; the 2 varie­ ties keyed out below belong to subsp. lepidus. Subsp. kig i . oensis Bridson is restricted to Tanzania. 1. ..eaves finely pubescent beneath. var. lepidus - .eaves glabrous or with a few coarse hairs. var. unyorensis

1. Leaf blades up to 8(-9) cm long, elliptic, mostly cuncatc at base; petioles up to 3 mm long. var. niamniamensis - Leaf blades up to 3(-4) cm long, ovate, mostly rounded or cordate at base; petioles c 1 mm long. var. djurensis var. niamniamensis Fig. 141.38.7 & 8.

var epidus U n f rstorey of lowland rain forest; 1050 m. KF, S Sudan, Re ' Dem. du Congo, tropical E Africa. Friis et al. 3911.

Riverine forest; 700-1400 m. IL KF GG; Sudan, tropical E Africa, Rep. Dem. du Congo. Zimbabwe. Friis et al. 2.520; Gereau 1321; Haugen 219.

141. RUBIACEAE: 37. Tricalysia, 38. Galiniera

Figure 141.37. T R IC A L Y SIA N IA M N IA M E N S IS var D J V R E N S IS : 1 - flowering branches x 1; 2 - bud x 3; 3 - flower x 3; 4 opened corolla with anthers, style and stigma x 3; 5 - leaf, upper surface >. V2; 6 —fruit x 3. var. N IA M N IA M E N S IS . 7 - leaf, upper surface x Vi; 8 - leaf, lower surface, showing domatia x Vi. 1-6 from Friis et al. 2582; 7 & 8 from Gilbert & Phillips 9001 Drawn by Sarah Howard.

var. djurensis (Hiern) Robbr. in Bull. Jard. Bot. Nat. Belg. 57: 174(1987): Tricalysia djurensis Hiem (1877) - type from Sudan.

-

Plants evergreen; flower clusters (subtended by cup-like bracteoles) on long shoots; fruits ± ses­ sile. 40. Polysphaeria

Fig. 141.37.1—6. Rocky places in woodland; 950 m. IL; S Sudan. N Rep. Dem. du Congo and Central African Republic. Friis el al. 2582.

OCTOTROPIDEAE Bedd. Hypobathreae (Miq.) Robbr. Trees or shrubs. Inflorescences axillary and paired at nodes. Corolla lobes contorted to the left. Ov ary 2-locular, placentas axile, one to many pendulous ovules per locule. Fruits mostly small, often fleshy. 1. Flowers in pedunculate corymbose cymes in the axils of evergreen long shoot leaves; corolla subrotate. 38. Galiniera - Flowers in (sub)sessile axillary clusters; corolla funnel-shaped. 2 2. Plants deciduous, flowers clustered on usually leafless short shoots; fruits pedicellate 39. Feretia

38 GALINIERA Del. (1843) Shrubs or small trees. Leaves decussate, petiolate. Stip­ ules small, deltoid to triangular. Flowers fragrant, 5-merous, in paired, axillary pedunculate corymbose cy mes on long shoots. Corolla subrotate, with very short funnel-shaped tube, lobes contorted in bud, spreading. Anthers exserted Style with bilobed, compressed and shortly pilose stigma exserted. Ovary 2-locular, two col­ lateral ovules per locule, pendulous from placenta at­ tached to near top of septum. Fruits (sub)globose, drupes, mesocarp fleshy, endocarp chartaceous. Seeds (2-)4, with ruminate endosperm. A genus of 2(1?) species, 1 in tropical Africa, 1 (doubtful) in Madagascar. G. saxifraga (Hochst.) Bridson (1988); P o u c h etia sa x ifra g a Hochst. (1842); P tych o stig m a sa x ifra g a (Hoclist.) Hochst. (1844), nom . illeg.

141. RUBIACEAE: 38. Galiniera

259

Figure 141.38. G A LIN IE R A S A X IF R A G A : 1 - flowering branch x 2/ 3; 2 - node showing stipule x 2; 3 - ovary and calyx x 4; 4 - corolla, one lobe removed x 3; 5 - stigma lobes x 4; 6 - longi­ tudinal section o f ovary showing ovules x 8; 7 —fruit x 2, 8 - seed, different views x 3; 9 - longitudi­ nal section o f seed showing rumi­ nate endosperm x 3; 10 - detail o f seed surface x 18. I~6 from Newbould & Jefford 1741; 7 10 from Ash 1474. Drawn by Mary Millar Watt. (Reproduced with permission from FI. Trop. E. Afr. R ubiaceae: fig. 119.)

- type GD, Aber. nr. Addeselam. S c h im p e r 863 (B holo. destroyed, K WU iso.). Galiniera coffeoides Del. (1843) - type: GD, be­ tween Addis and Maitalo, Ferret c£ Gallinier 23 (MPU holo.). Ilirub up to 4 111 or sometimes small tree up to 10 111 tall, often with long, down-curved branches. Leaves glossy reen above, paler green and often with reddish veins be­ low. elliptic-oblong to more or less lanceolate. (6—)8—16 etta elliptica Hochst. [in P i Schimp. Abyss . An Afro-Madagascan genus of c 20 species; 2 in tlie scct. II, 9061 (1844), nom. nud Flora area. Shrub up to 2-3 m tall. Leaves ovate-elliptic to obovate. 1. Leaves linear-lanceolate to narrowly oblong1.5-5.5 x 0.7-2(-2.5) cm, acute to rounded at both ends; lanceolate, 0 4 -1 .5 ( - 1.8) cm wide. 1. P. aethiopica petioles 1-4 mm long. Flowers subsessile at first, clus­ i - Leaves ovate to elliptic-lanceolate, up to 3 cm tered on short lateral branchlets, opening usually before wide. 2. P. parvifolia branchlets elongate; flower clusters subtended by dry. brownish ovate-lanceolate bud scales. Calyx lobes 1.5—2 nun long. Corolla white, tube and lobes often pinkish 1. P. aethiopica Verde. (1980) tinged outside, 10-15 nun long, lobes rounded. Anthers type: SD. nr. Dawa Parma River bridge. 4°47’ N, up to 8 mm long. Fruits with pedicels up to 5(-7) mm 39° 19' E, De Wilde & Gilbert 359 (K holo.. WAG iso.). ? P. parvifolia sensu Cufod. (1965), non Hiem long, often paired or solitary at lowermost nodes of newly [based on BA upper Wabc valley, Ellenbeck s.n (B produced lateral branchlets, 3-7 mm in diameter. Fig. destroyed), fide Engler(1904)] 141.39.

141. RUBIACEAE: 40. Polysphacria

261

Figure 141.40. P O L Y SP H A E R IA P A R V IF O L IA : 1 — flowering branch x 2/ 3; 2 —fruiting branch x 2/3; 3 - flower with opened corolla x 4; 4 - fruit x 2; 5 - seed x 2; 6 - cross section o f seed showing ruminations x 2; 7 - calyx x 6: 8 - node with stipule and modified leaf x 2/3 1 from Polhill & Paulo 782; 2, 4. 5. 7 & 8 from Graham I'D 1522; 3 from Faulkner 1099; 6 from Aiagogo 110. Drawn by Ann Farrer (1-5) and Sally Dawson (6-8). (Reproduced with permission from FI. Trop. E. Afr. R ubiaceae: fig. 95.)

: ! rub. branched from the base, up to 3 m tall. Leaves lint 1 -lan ceo late to narro w ly o b lo n g -la n c e o la te , ( 2 - ) 3 - 6

1 S) x 0.4—1.5(—1.8) cm, acute at apex, cuneate at base, f 1 ibrous or more or less pilose below, especially at mid-

\ :tn; petioles 2-3 mm long. Flowers fragrant. 4-merous, i clusters or (2-)4-6 at nodes. Calyx shallowly cups liped. with more or less triangular minute teeth less than 1 mm long Corolla wliite, shortly pubescent outside, tube 3 -4 mm long, lobes I.5-2.5(-3) mm long. Style with susma white, hairy, up to 8 mm long. Drupe globose. 5-9 r 1:1 in diameter, slightly didymous. Seeds brow'n, semigk bose. c 5 mm in diameter.

Mostly in riverine vegetation, often at or below high water mark; 800-1350 m. SD BA; endemic. P uff & Ensermu K. 821227-1/5; P uff el al. 870510-5/1. Collections from BA have wider leaves than from SD (ty pe locality) and leaves are hairy beneath. The species seems to be veiy close to P. p a rv ifo lia Hiem (occurring both in SW Ethiopia and S Somalia), differing primarily in its narrower leaves. P. parv ifolia Hiem (1877); - type from Zanzibar.

262

141. RUBIACEAE: 40. Polysphaeria. 41 Tarcnna. 42. Pavetta

P. parvifolia Hiem var. ? glabra Hiem in FI. Trop. Afr. 128(1877).

Tarenna boranensis Cufod. (1948) - ty pe: SD. El Banno, Corradi 2765 (FT lecto ).

Shnib or small tree, up to 4(—7) m tall. Leaves ovate to elliptic-lanceolate, (2.5-)3-8.5 x (1-) 1.5-3 cm, acute to obtuse at apex, cuneate to subcordate at base, pubescent to glabrescent on midrib below; petioles up to 2 mm long. Flowers fragrant, 4-merous, in sessile clusters of (4-) 6-10 at nodes. Calyx pubescent shortly tubular, distinctly dentate, teeth less than 1 mm long. Corolla white, (densely ) spreading pubescent outside, tube 2-A mm long, lobes 1.5-2 mm long. Style with stigma white, hairy, slightly exserted. Dmpe subglobose. 6-8 mm in diameter. Seeds brown, subglobose. 4-5 mm in diameter. Fig. 141.40.

Shrub or small tree up to 3 m tall. Leaves coriaceous, often shiny above, elliptic to broadly lanceolate. (4—)5—10 x (1 5 )2-4 cm, acute at apex, cuneate at base, mostly gla­ brous: petioles 0.5-1.5 cm long. Inflorescences usually many-flowered, often more or less corymbose, up to 60 mm in diameter Calyx lobes rounded to more or less acute, up to 1 mm long. Corolla tube 2.5—4(—5) mm long, hairy at throat, lobes as long as or slightly longer than rube. Ovary 2-locular. 2-3 ovules per locule, embedded in placenta. Fmit subglobose, (4-)5-7 mm in diameter, usu­ ally I-seeded. Seed globose, c 4 mm in diameter, with a more or less circular adaxial cavity. Fig. 141.41

Mostly in evergreen riverine vegetation; 600-920 m. IL KF SD; Sudan. Somalia. Kenya. Tanzania. Friis et al. 2519; Gereau 1250; Meyer 9009.

PAVETTEAE Dumort.

Shrubs or trees. Inflorescences tenninal on main or lateral branches or pseudo-axillary. Corolla lobes contorted to the left. Ovary 2(-7)-locular, placentas axile. one to nu­ merous ovules on fleshy placentas usually attached to sep­ tum. Fruits mostly small, drupes or berries.

Acacia - Commiphora. Combretum - Terminalia or dry Olea - Dodonaea - Terminalia woodland: degraded dry Juniperus Olea forest; often in rocky places; 1000-2100 in SU GG SD BA HA; Somalia. Uganda to Tanzania. Puff& Ensermu K. 820927-2/1. 821217-4/10, 870426-9/3. A second subspecies, subsp. a ra b ic a (Cufod.) Bridson. with nanower leaves and smaller inflorescences with fewer flowers, occurs in N Somalia and SW Arabia. 42. PAVETTA L. (1753)

1. Flowers (mostly) 5-merous, tube ± as long as lobes. 41. Tarenna - Flowers 4-merous. tube often longer than lobes. 42. Pavetta

41. TARENNA Gaertn. (1788) Shrubs or trees. Leaves decussate, often coriaceous, petiolate or sessile. Stipules ovate to triangular, falling soon Flowers mostly 5-merous, in terminal many - to fcvv-fiowercd. often more or less corymbose inflorescen­ ces. Calyx with short tube, lobed. very shortly toothed or more or less tnmcate above. Corolla whitish, tube cylin­ drical to narrowly funnel -shaped, long or short, lobes con­ torted in bud. spreading or reflexed. often as long as or longer than tube. Anthers subsessile. long, exserted. Style with club-shaped stigma long exserted Ovary 2-locular. 1-15 ovules per locule, either pendulous from, or more or less embedded in. placenta. Fruits fleshy , more or less globose, several- to 1-seeded. Seeds with a round or elon­ gated adaxial cavity . A paleotropical genus of c 180 species, the majority of which occur in Asia. 1 in the Flora area. T. graveolens (S. Moore) Bremek. (1934) subsp. graveolens. Pavetta gra\>eolens S. Moore (1907) - type from Uganda. P. ligustriodora Chiov. (1916). Tarenna edgardi Chiov. (1935, nom. event.. 1939) - types; SD. Neghelli. Za\>attari 209 & Arero. Zavattari 356 (both FT syn.).

Shrubs or small trees. Leaves decussate or occasionally in whorls of 3(4). with bacterial nodules on the blades, petiolate. Stipules connate at base, apiculatc. Flowers fra­ grant. mostly white or creamy white. 4-merous. in sessile or subsessile mam - to several- or occasionally fewflowered corymbose inflorescences terminal on (some­ times short and more or less leafless) lateral branches, flowers rarely solitary. Calyx lobes long to short and in­ distinct Corolla tube cylindrical to narrowly funnelshaped. often longer than the lanceolate to oblong lobes: lobes contorted in bud. Anthers exserted. Style with club-shaped, indistinctly 2-lobed stigma (long) exserted. Ovary 2-locular. 1(rarely 2) ovules per locule, on placenta pendulous from near top of septum. Fruits drupes, (sub) globose or more or less 2-lobed. crowned by persistent ca­ lyx. with I (rarely 2) chartaceous py renes with a circular hollow on plane, ventral surface. A palaeotropicaJ genus of c 400 species. 6 in the Flora area. 1. Leaves in whorls of 3 or 4. up to 25 x 4(-5) cm. tough, yellowish-green or yellowish-brown when dry. ' ] | 4. P. crassipes - Leaves decussate (if in whorls of 3 much smaller than above), colours in dry state not as above. 2 2. Calyx lobes nanowly triangular, linear or subulate: entire calyx at least 2 mm long, often much lon­ ger. 3 - Calyx lobes broadly tnangular (lobes about as long as wide); entire calyx less titan 2 nun long. 5 3. Basal tubular part of calyx very short and indis­ tinct not as w ide as the ovary below. 1 P. oliveriana

141. RUBIACEAE: 41. Tarenna, 42. Pavetta

263

Figure 141.41. TA R E N N A G R A V E O L E N S subsp. GRA VEOLENS: 1 - branch with in­ florescences x 1; 2 — bud x 1; 3 — flower x 1; 4 - fruit x 3. 1 & 4 from Friis et al. 6264; 2 & 3 from P u ff et al. 870927-1/10. Drawn by Sarah Howard.

-• Basal tubular part of calyx 1.5(-2) mm long, wider than the ovary below. 4 I Tubular basal part of calyx a little wider than tlie ovary below, lobes 3.5-5 mm long; only SD. 2. P. aethiopica Tubularbasal part of calyx distinctly wider than the ovary below, lobes l-4(-12) mm long. 3. P. abyssinica i\ Corolla lube 15-20 mm long, much longer than lobes; inflorescences ± head-like. 6. P. subcana - Corolla tube (4—)6—10(—12) mm long, only slightly longer than lobes; inflorescences not as above. 5. P. gardeniifolia I P. oliveriana H iern (1877); Ix o ra o liveria n a (Hiem) O. Kuntze (1 8 9 1 )- type from Tanzania. Pavetta humbertii Bremek. (1934). P. lebrunii Bremek. (1937). P. elliottii sensu Cufod. (1965), non K. Schum. & K. Krause (1907). Pavetta sp., Bremek. in Pichi-Sermolli (1951).

Shrub or small tree, 1-4 m tall; young branches pu­ bescent. Leaves elliptic, ovate-lanceolate or lanceolate, 4—15(—18) x 1.5—6(—8) cm, acute to more or less apiculate at apex, narrowed to base, pubescent to glabrous above, pubescent below, sometimes only along midvein, less commonly more or less glabrous altogether, petioles 0.3-1.5(-2.5) cm long, pubescent. Stipules triangular, c 5 mm long. Inflorescences corymbose, rather lax, 25-75 mm in diameter, terminal on sometimes leafless lateral branches; peduncles and pedicels pubescent, 3-5 mm long (up to 8 mm in fruit). Calyx pubescent, tubular basal part short and indistinct, lobes n arro w ly triangular to subulate, 2-5(-7) mm long. Corolla typically pubescent outside (occasionally more or less glabrescent), tube 15—25(—30) mm long; lobes oblong, 4-8 x 1.5-3.5 mm. Fruit shiny black, globose to more or less bilobed, often crowned by persistent calyx, 7-10 mm in diameter, more or less pubescent when young, sparsely pubescent to more or less glabrous when fully mature. Riverine vegetation; deciduous (open) woodland; rocky outcrops with bush clumps; montane scrub or forest

264

141. RUBIACEAE: 42. Pavetta

remnants. Olea - Podocarpus forest; (1150-) 1450-2450 m. EW GD GJ WU SU AR WG IL KF GG SD BA HA; tropical East Africa. Sudan, Rep. Dem du Congo, Rwanda and Burundi. P uff & Ensermu K. 820807-3/3, 820819-6/2, 821213-1/1. Leaf indumentum variable: pubescent all over (typi­ cal and most common), some pubescent only on lower surface, some hairy only on midvein below and some glabrous altogether Glabrescent forms are often also less densely pubescent in the inflorescence region. Glabrescent to glabrous plants (e.g., AR, Tewolde 2501; HA P uff et al. 820922-1/14; SU, Mesfin et al. 1181) could be referred to “var. denudata (Bremek.) Bridson (1978)' [= P. denudata Bremek. (1949); P. cineraseens (A. Rich.) Chiov. var. glabrescens Chiov., nom. nud., based on Negri 2421 (FT) & P. ehiovendae Pic.Serm., nom. nud., based on Negri 1004 & 2421 (FT), both from SU, Lake ZwaiJ. The variety is, however, not formally upheld, primarily because both the “typical" pubescent and the glabrescent morphs can occur within a single population. Pavetta abyssinica “var. cinerascens A. Rich" [nom illeg.\ P. abyssinica sensu A. Rich. (1848). non Fresen., var. “cinerescens” A. Rich. (1848) - type: GD or TU, Choho, Ouartin Dillon & Petit s.n. (P holo.); = P. cinerascens (A. Rich.) Chiov. (1911), non Bremek. (1934)] should, as already suggested by Pichi-Sermolli in Ricerche Bot. 7,1, Miss. Stud. Lago Tana: 148 (1951). be placed in P. oliveriana. “cinerascens” approaches “var. denudata" and more or less links P. oliveriana and P abyssinica [e.g.. GD. Gilbert & Getachew 3021; GJ Puff et al. 811008-2/3]. 2. P. aethiopica Bremek. (1956) type: SD, Mega Mt. Bally 9397 (K holo., EA iso). Shrub, 2-3 m tall. Leaves elliptic to narrowly obovate. 5-9 x 3-5 cm, shortly acuminate at apex, more or less cuneate at base, glabrous above except for midrib, (sparsely) pubescent below, petioles up to 1 cm long, pu­ bescent. Stipules lanceolate, papery, soon falling, up to 10 mm long. Inflorescences corymbose, rather compact, 30-40 mm in diameter, terminal on lateral branches: pedi­ cels 2-5 mm long, glabrous. Calyx with tubular basal part c 1.5 mm long, lobes linear. 3.5-5 mm long, glabrous or ciliate. Corolla glabrous outside, tube 15 mm long; lobes lanceolate-oblong, 8 x 3 mm. Fruit shiny black (sub) globose, crowned by persistent calyx, 7 mm in diameter, sparsely puberulous or with a few longer hairs. Montane scrub, c 2050 m. SD; Kenya. Bally 9397. In Ethiopia only known from the type collection. Very closely allied to P. abyssinica and possibly scarcely dis­ tinct. 3. P. abyssinica Fresen. (1837); Ixora abyssinica (Fresen.) Oliv. (1873) - type: GD, between Halei and Temben, Riippell s.n. (FR holo.).

Figure 141.42. PA V E TTA A B Y S S IN IC A var. A B Y SSIN IC A : 1 - flowering branchlet x 2 - fruiting branchlet x V y ,3 - leaf x 2,s; 4 - flower x 3/-i. 5 - longitudinal section o f flower x 114. 1, 4 & 5 from Bally 8814; 2 & 3 from Verdcourt 1822. Drawn by M E. Church. (Reproduced with permission from Dale & Green way. 1961: Kenya Trees & Shrubs: fig. 86.)

Shrub or small tree up to 5 m tall; young branches gla­ brous or finely pubescent. Leaves elliptic, oblongianceolate or oblanceolate, 4—15(—18) x 1.2—6.5(—8) cm acute to more or less acuminate at apex, narrowed to base, glabrous above and below' or occasionally pubescent be­ low, especially along midvein; petioles 2-10 mm long. Stipules truncate, more or less papery, soon falling. 5-10 mm long. Inflorescences corymbose, rather compact and more or less head-like, up to 50 mm in diameter, terminal on sometimes leafless lateral branches: peduncles and pedicels glabrous or finely pubescent. 2-3 mm long (up to 10 mm in fruit). Calyx glabrous or finely pubescent, tubu­ lar basal part distinctly wider than ovary below, 1.5(-2) mm long, lobes more or less subulate. 1—4(—12) mm long. Corolla glabrous outside, tube 15—25(—37) mm long; lobes oblong, 5-8 x 2-2.5 mm. Fruit shiny black, more or less globose, often crowned bv persistent calyx, (5—)7—10 mm in diameter, glabrous. Bridson (1978) in Kew Bull. 32: 634 divides the spe­ cies into 7 varieties, 3 of which occur in the Flora area (one variety, var cinerascens, is excluded; see P. oliveriana, above).

141. RUBIACEAE: 42. Pavetta

1. Calyx lobes 7-12 mm long. - Calyx lobes 1-4 mm long.

var. brem ekam piana 2

2. Inflorescence branches, ovary and calyx mostly (yellowish) pubescent or haiiv; leaves mostly at least a little hairy. var. dolichosiphon - Ovary and calyx mostly glabrous. var. abyssinica var. abyssinica Pa\>etta abyssinica sensu A. Rich. (1848), non Fresen., var. glabra A. Rich. (1848), nom. illeg. type: TU, Mt Koubi, Schimper 353 (K holo., BM W WU iso.). P. ellenbeckii K. Schum. (1903) - type: HA Gara Mulata, Ellenbeck 519 (B holo. destroyed). ?P. erlangeri Bremek. (1934) - type. HA, Gara Mulata, Ellenbeck 603 (B holo. destroyed). P kenyensis Bremek. (1934). P. maitlandii Bremek. (1934). P. silvicola Bremek. (1934). P. trichotropis Bremek. (1949).

265

Shrub or small gnarled tree up to 5 m tall: stems to 15 cm in diameter; bark corky, yellow. Leaves in whorls of 3 or 4 (in the Flora area), thickish, yellowish-green or yellow­ ish-brown when dry, lanceolate to oblanceolate, (6-) 10-25 x (l-)1.5-4(-5) cm. rounded to more or less acute at apex, narrowed to base, glabrous; petioles short. Stip­ ules triangular, c 5 mm long. Inflorescences cory mbose, many-flowered, up to 120 mm in diameter, terminal on short leafless lateral branches; peduncles and pedicels (3—)5—10 mm long (up to 15 mm in fmit). Calyx glabrous, 1-1.5 mm long, truncate to slightly dentate. Corolla gla­ brous outside, tube 10-15 mm long; lobes oblong. 4—6 x 1.5-2.5 mm. Fmit shiny black, more or less globose, crowned by persistent calyx, 5-7 mm in diameter, gla­ brous. Deciduous w'oodland; 900-1500 m. KF GG SD: w'est to W Africa, south to Zambia, Malawi and Mozambique. Gereau 1333; Puff&Ensermu 821214-2/1, -3/2.

Somewhat obscure; seems to be known from the type collection (possibly from GD or TU) only; no material seen.

5. P. gardeniifolia A Rich. (1848) var. gardeniifolia: Ixora gardeniifolia (A. Rich.) O. Kuntze (1891) type: GD, Choata in Semien, Schimper 1141 (P lecto., K WU isolecto.). Pavetta gardeniifolia A. Rich. var. breviflora Vatke in Osterr. Bot. Zeitschr. 25: 231 (1875) - types: EW Habab, Hildebrandt 495 (B syn. destroyed) & ISoinalia, without locality] 889 (B syn. destroyed). P. gardeniifolia A. Rich. var. laxiflora K. Schum. in Pflanzenw. Ost. Afr. C.: 389 (1895). P. gardeniifolia A. Rich. “var. angustata A. Rich " sensu Annari in Pirotta. FI. Eritrea 1: 153 (1904). P. adelensis Del. (1846), nom. nud. P. assimilis Sond. (1865); Ixora assimilis (Sond.) O. Kuntze (1891). Pax’etta assimilis Sond. var. brevituba-glabra Bremek. in Ann. Transv. Mus. 13: 210 (1929). P. assimilis Sond. var. glabra Bremek. loc. cit., nom. superfl. P. krauseana K. Krause (1912). P. saxicola K. Krause (1912). P. hochstetteri Bremek. var. glaberrima Bremek. in FeddesRep. 37: 182 (1934)-type: TU, Schahaganc in Memsah. Schimper 210 (K holo., WU iso.). P. hochstetteri Bremek. var. mollirama Bremek. loc. cit.: 183 - type: SD Galla, Bidduma, Ruspoli & Riva 122 (FT holo.). P. hochstetteri Bremek. var. graciliflora Bremek. in Feddes Rep. 47: 95 (1939). P. petraea Bremek. (1939). P. termitaria Bremek. var. glabra Bremek. loc. cit.: 94. P.fossorum Bremek. (1949). P. somaliensis Bremek. (1949).

4. P. crassipes K. Schum. (1895) - type from Kenya. P. barteri Dawe (1906). P. utilis Hua (1907). P. crassipes K. Schum. var. major De Wild. PI. Bequaert. 2: 293 (1923).

Shmb or small tree up to 3 m tall; branches mostly gla­ brous; bark yellowish-grey. Leaves decussate or occa­ sionally in whorls of 3, yellowish(-brown) when dry, obovate, elliptic, lanceolate or occasionally more or less spathulate, (1.5—)2.5—10 x (0 .5 -)l-4 cm, acute to rounded at apex, gradually narrowed to base, glabrous or. seldom, pubescent; petioles 5-10 mm long. Stipules triangular to

Fig. 141.42. Evergreen (montane) forest, Podocarpus or Juniperus forest; bush clumps; riverine vegetation; (1350—) 1700-2850 in. EW TU GD GJ WU SU AR WG KF SD HA; south to Tanzania. Puff et al. 811001-1/3; Puff & Ensermu K. 820818-7/7; P uff Sc Sileshi N. 881209-1/10. Bridson (loc. cit.) pointed out that atypical luxuriant forms with rather large leaves occurring in undergrowth of rain forest (1350-1400 m, KF; Gilbert 2005, Mooney 9185, 9225) may belong to a separate taxon. var. dolichosiphon (Bremek.) Bridson in Kew Bull. 32: 636(1978); P. dolichosiphon Bremek. (1934) - type: TU?, Etchelicote, Ouartin Dillon & Petit 132 (K holo.. P iso.). P. eritreensis Bremek. (1934) - type: EW Assaorta. Pappi 5214 (FT holo ). P. cinerascens sensu Bremek. (1934), non (A. Rich.) Chiov. Riverine vegetation, scrub; 1600-2700 in. EW TU? GD: endemic. Baldrati 225, 505; Chiovenda 2503 bis. var. bremekam piana (Pic.-Ser.) Bridson in Kew Bull. 32:637(1978); P. bremekampiana?ic.-Ser. (1951) —type: "Ethio­ pia”, O uartin D illo n & P e tit s.n. (P holo.).

266

141. RUBIACEAE: 42. Pavetta. 43. Cofiea

subtruncate, 2-5 mm long. Inflorescences coiymbose, rather many-flowered, lax to more or less congested, up to 50 mm in diameter, terminal on sometimes very short leafless lateral branches; peduncles up to 10 mm long, pedicels 2—7(—12) mm long (up to 15 mm in fruit). Calyx glabrous, tubular below, shallowly toothed or with short but broad triangular lobes above, altogether up to 1 mm long. Corolla yellowish-green to dull yellow, glabrous outside, tube (4—)6—10(—12) mm long; lobes lanceolate to obovate, 4-8 x 1.5—2.5(—3) mm. Fruit shiny, black, more or less globose, crowned by persistent calyx, (4-)5-8 mm in diameter, glabrous.

2-5 mm long. Inflorescences more or less head-like, many-flowered, up to 20 nun in diameter, terminal on mostly very short leaf-less lateral branches; peduncles ab­ sent orvery short in flower, pedicels 1-2 mm long (up to 8 mm in fruit), glabrous or finely pubescent. Calyx glabrous or haiiy on margins, tubular below, with short but broad triangular lobes above, altogether less than 1-2 mm long. Corolla white or creamy-white, glabrous outside, tube 15-20 mm long, lobes ovate-lanceolate, 3-5 x 1.5-2 ram. Fruit shiny black crowned by persistent calyx, more or less globose, 4-8 mm in diameter, or more or less bilobed and up to 12 mm wide, mostly glabrous.

Combretum - Terminalia and other woodland, ever­ green bushland, in bush clumps in much disturbed sites, often associated with rocky places (granite, limestone); 800-2100 m. EE EW TU SU KF GG SD BA HA; Soma­ lia, south to South Africa, west to Togo. P uff & Ensermu K. 861030-1/2; P uffei al. 810916-2/10, 870428-2/5.

Bushland and woodland, often along streams; 730-1500 m. EE EW GG SD, Sudan, Rep. Dem. du Congo, Central African Republic, tropical E Africa. Puff iflora Afz. (1815) - type from Sierra Leone. Canthium afzelianum Hiem (1877), nom. now for Pa\>ettapar\>if1ora Afz., non C. par\>iflorum Lam., p.p Plectronia vulgaris K. Schum. (1895); Canthium vulgare (K. Schum.) Bullock (1932). C. golungense Hiem (1898); Plectronia golungensis (Hiem) K. Schum. (1900). Canthium golungense Hiem var. parviflorum S. Moore in Journ. Linn. Soc. 37: 161 (1905). Plectronia brieyi De Wild. (1914). Canthium giordanii Chiov. (1941) - types: WG. Galla Sidama, Saio. Giordano 2450 & 2539 (FT syn ). Tree up to 10 m tall, often single-stemmed and with more or less horizontally arranged branches; bark grey. Leaves decussate, evergreen, ovate to oblong-ovate, 5-15 x 2.5-5.5 cm. acuminate at apex, narrowed to base, gla­ brous. glossy above, diying green to brownish; petioles up to 1 cm long. Stipules broad-based, subulate above, up to 6 mm long. Flowers fragrant 4-merous, «in pedunculate many-flowered cymes: peduncles up to 15 mm long. Ca­ lyx cup-like, not distinctly toothed. Corolla creamy while to greenish, tube up to 1 mm long, lobes reflexed, longer than tube. Drupe 2-lobed. 4-6 x 8-10 pun. or with only I developed pyrene and more or less globose, c 6 mm in di­ ameter. Pyrene strongly curved, sides of lateral groove touching.

1.5-4(-4.5) cm, more or less acute at apex, narrowed to base, glabrous, glossy above, drying green to brownish; petioles up to 5 mm long. Flowers fragrant, 4-merous. in (sub)sessile, more or less umbel-like, paired, axillary, many-flowered inflorescences; pedicels 5—8(—10) nun long. Calyx minute, lobes shortly triangular. Corolla (creamy) white, tube 2.5-3.5 mm long, lobes reflexed. as long as or longer than tube. Drupe black, subglobose to more or less 2-lobed. 4-6 mm in diameter, or oblong up to 8 mm wide Py rene slightly curved. Fig. 141 48. Acacia or Termmalia - Combretum woodland. Olea Juniperus forest, montane sc mb and forest; not uncom­ monly associated with rocky areas; 900-2600 m. EE EW TU GD WU SU AR GG SD BA HA; Somalia, Rep Dem du Congo, Rwanda Burundi, south to Malawi, SW Ara­ bia. Puffei al. 821002-1/9, 870424-10/1, 870509-5/3. A second subspccies. subsp. occidentalis Bridson. with shorter pedicels and winged angles on young steins, occurs in W Africa. 49. PYROSTRIA Commers. exJuss. (1789) Canthium sensu auctt. div. le g. Cufod. (1965)]. p.p., non Lam. Plectronia sensu auctt. div. p.p., non L.

Shrubs up to medium-sized trees. Leaves decussate, usu­ ally glabrous, mostly petiolate. Stipules triangular at base. Flowers unisexual (in the Flora area) or bisexual. 4-5merous. in pedunculate umbels; in dioecious species in­ florescences often dimorphic (female flowers often soli­ tary. male inflorescence with more flowers); paired, connate bracts surrounding inflorescence in bud Calyx reduced to a nm, sometimes dentate or unequally lobed Corolla tube shorter or longer than lobes, throat densely covered with hairs. Anthers attached to throat with short filaments. Style with more or less capitatc. lobed stig­ matic knob exserted. Ovary 2-locular (in the Flora area; Riverine forest or deciduous woodland; 1050-1750 up to 10-locular elsewhere). 1 ovule per locule pendulous from placenta attached to near top of septum; in dioecious m. WG IL KF GG SD; west to West Africa, southwest to Zambia and Angola. P uff & Ensermu K. 820812-1/4, species, male flowers with rudimentary ovary, style and 821212-6/18. 861107-5/1. stigmatic knob Fruits drupes, lobed; pyrenes (thinly) 2 more subspccies, subsp. rubrocostata (Robyns) woody. Bridson and subsp melanophengos (Bullock) Bridson A paleotropical genus with 14 African species, many centred in E and S Central Africa, are recognized, differ­ of which were formerly associated with Canthium; 1 spe­ ing in stipule, leaf and/or fruit characters. cies in tlie Flora area 2. P. schimperiana (A. Rich.) Bridson (1985) subsp. schimperiana: Canthium schimperianum A. Rich. (1848); Plectronia schimperiana (A. Rich.) Vatke (1876) types: TU. nr. Mayc-Gouagoua, Quartin Dillon s.n. & Mt Selcuda | ScholodaJ. Quartin Dillon s.n., Schimper 328 (all P syn.). Phallaria schimperi Hochst. (1841). nom. nud Canthium lucidum R Br. (1814), nom. nud. C. myrtifolium S. Moore (1907). Plectronia angiensis De Wild. (1925). Shrub or small tree up to 6 ni tall Leaves decussate, ever­ green, elliptic lo ovate-lanceolate, (2—)2.5—8(—10) x (1-)

P, phyllanthoidea {Baill.) Bridson (1987); Canthium phyllanthoideum Baill. (1878); Rytigynia phyllanthoidea (Baill.) Bullock (1932) type from Kenya. Plectronia bogosensis Martelli (1886); Canthium bogosense (Martelli) Penzig (1893) - type: EW. Keren and Luglia. Beccari 155 (FT holo.). Dioecious shrub or small tree up to 5 m tall; older indiv iduals often with leaves crowded on short shoots; browsed individuals vers irregularly and densely branched. Leaves frequently slightly dimorphic (long shoot leaves laiger than short shoot leaves), obovate. oblanceolate. oblong-

275

141. RUBIACEAE: 49. Pvrostria. 50. Fadogia

Figure 141.49. P Y R O ST R IA P H YLL A N T H O ID E A : 1 - fruiting branch x Vi; 2 - bud x 3; 3 - female flower x 5; 4 - rudimentary ovary and calyx o f male flower (co­ rolla fallen off), subtended by connate bracts x 5; 5 —drupe x 2. 1 fro m /3H$ret al. 870425-811A; 2 & 3 from P u ff ct al. 870501-211; 4 from P u ff ct al. 870427-1/1; 5 from P u ff et al. 870427-1/8. Drawn by Sarah Howard.

lptic to linear-oblong, (0.25-)0.4-3.5(-6) x (0.15-) i 2—1.5 cm, usually rounded at apex, cuneate to more or ;s rounded at base, tough, thickish and more or less suclent, venation more or less invisible; petioles short. ] bwers 4-merous, subtended by a pair of bracts 1-2.5 n n long, in male plants in groups of few to solitary', in fe­ ll lie plants solitary; pedicels up to 1 mm long. Calyx with t oth 0.5-1 mm long. Corolla tube more or less cylindric ii. 2-3 mm long, lobes spreading to reflexed. as long as or somewhat shorter than tube. Style with stigmatic knob e serted fo re 1 nun in female plants (male plants: rudii mtary style and stigma present but included). Ovary 2I :ular. Dmpe on short pedicel with paired bracts, 2-lobed, r ore or less heart-shaped, 5-15 x 6-11 mm; pyrenes 2, r ore or less kidney-shaped in side view. Fig. 141.49. Mostly associated with rocky places in woodland or b i>hland areas, sometimes in woodland on sandy flats; 1 *50-1800 m. EE? EW AR SD BA HA; N Somalia, I :nya and Tanzania; Saudi Arabia. P uff et al. 8704271 1. 870512-2/2; Tewolde B.G.E. 2503. Leaves extremely variable in shape and size; forms v ith unusually long leaves (up to 6 cm) (on long shoots) a near to be confined to EW. In many areas virtually all plants are heavily browsed a id apparently a favourite source of food for camels.

50. FADOGIA Schweinf (1868) Mostly subshrubs with several to many stems from a sub­ terranean woody base. Leaves in vvhoris of 3-5, glabrous or hairy, subsessile. Stipules small, with a narrowly trian­ gular seta between leaf bases. Flowers 5(-8)-merous, in axillary, paired, pedunculate, few- to several-flowered cy mes. Calyx truncate above or shortly lobed. Corolla tube straight or slightly curved, funnel-shaped to cylindri­ cal, haiiy within, lobes valvate in bud, reflexed, acute to shortly apiculate. Anthers inserted at throat. Style with crown- or hat-like stigmatic knob as long as corolla tube or exserted. Ovary' 3-5-Iocular, I ovule per locule pendu­ lous from placenta attached to top of septum. Fruits drupes, crowned by persistent calyx, with 3-5 (or fewer due to abortion) pyrenes. An African genus of 40-50 closely related spccies; 1 in the Flora area.

F. cienkowskii Schweinf. (1868); Canthium cienkowskii (Schweinf.) Roberty (1954) - type: Sudan [nr. WG border], Fesoglu, nr. Fadoga, Cienkowski 159 (W holo.). Vangueria tristis K. Schum. (1903); Fadogia tristis (K. Schum.) Robyns (1928). Fadogia katangensis De Wild. (1913).

276

141. RUBIACEAE: 50 ladogia, 51 Meyna

Figure 141.50. FAD O GIA C IE N K O W SK II 1 - branch with axillary inflorescences x '/i; 21 woody rootstock with bases o f aerul stems x Vi; 3 - leaf, upper surface x 14; 4 leaf, underside detail x 15. 5 —bud x 3, 6 - flower x 3: 7 - corolla opened out to show stamens x 3; 8 - developing drupe with calyx lobes x 3; 9 - mature drupe x 1'/j. 1 & 4 -7 from M oonev 7730; 2 from P u ffe i al 820810-1/1; 3, 8 & 9 from M ooney 7749. Drawn by Sarah Howard

Subshrub; several erect, often unbranched stems up to 40(-50) cm tall from a more or less massive subterranean woody base. Leaves in whorls of 3(4), elliptic to ovate. (2-)3-8.5 x (0.8—)1.5—3.5 cm, cuspidate to rounded at apex, cuneate to rounded at base, markedly discolourous, very dark and hispid above, golden-brown and densely velvety beneath. Flowers 5-merous, 2-6 in shortly pedunculate axillary clusters. Calyx lobes up to 2 mm long, triangular-subulate. Corolla hairy outside, green­ ish-yellow to whitish, tube broadly funnel-shaped to cy­ lindrical, 2.5-4 mm long, lobes lanceolate, acute to apiculate, 2-3 mm long. Crown-like stigmatic knob 3(4)lobed, up to 1 mm long. Ovary 3(4)-locular. Drupe dark blue to blackish, globose to ellipsoidal, 5-8 mm long, with 3 (or fewer due to abortion) pyrenes. Fig. 141.50. Grassland, wooded grassland or woodland; 12001600 m. WG: west to Mali, south to Zimbabwe (? and South Africa). Angola Mooney 7730; Puff & Ensermu K. 820809-4/5, 820810-1/1. Flora area material belongs to var. cienkowskii. The somewhat dubious var. lanceolata Robyns, with leaf

blades up to only 1 cm wide, is recorded from Cameroun. Rep Dem. du Congo and Tanzania.

51. MEYNA Roxb. ex Link (1820) Shrubs or small trees, mostly with thorny branches. Leaves often crowded on short shoots, borne below thorns, decussate, petiolate. Stipules small. Flowers 4- or 5-merous. small, pedicellate, few- to several, clustered on short shoots. Calyx small, linear to subulate. Corolla tube cylindrical, throat hairy inside; lobes valvate in bud, small, recurved, often apendiculate. Anthers inserted at throat Style with crown-like stigmatic knob exserted. Ovary 4-5-locular, 1 ovule per locule pendulous from pla­ centa attached to top of septum. Fruits drupes, crowned by persistent calyx, w ith 4 or 5 (or due to abortion fewer) py­ renes. A small Asiatic genus with only one species in Africa mid the Comoro Islands. The species are poorly delimited and it is possible that the Flora area species is but a variant of the Indian M spinosa Link.

277

141. RUBIACEAE: 51. Meyna, 52. Rytigynia

Figure 141.51. M E YN A T E T R /i P H YLLA subsp T E T R A P H Y L L A : 1 - young branch with thorns x V&; 2 - portion o f older, flowering branch, corollas already fallen x 12/s; 3 - flower x 5; 4 —drupe x V/i. 1 from Jensen et al. X2/3; 2 from Turner 227; 3 & 4 from Haugen 208. Drawn by Sarah Howard.

M. tctraphylla (Schweinf ex Hiem) Robyns (1928) subsp tctraphylla; Vangueria tetraphylla Schweinf. ex Hiem (1877); I' Canthium tetraphyllum (Schweinf. ex Hiem) Baill 1 (1878) - type from Sudan. Shrub up lo 4 111 tall, usually much and irregularly branched, sometimes subscandent; older branchlets usu■Uy dark grey, lenticellate. glabrous, mostly armed with paired, straight, stout thorns, up to 20 nun long, borne above leaf- and/or flower-bearing short shoots; younger :iranclilets reddish, puberulous, often without short • hoots, occasionally without thorns. Leaves decussate but :iften seemingly in whorls of 4 on short shoots, (ob)ovate :o more or less elliptic. 2.5-4(-5) x 1,5-3(-4) cm. more or less obtuse at apex, cuneate at base, more or less hairy to iensely pubescent, margins often ciliolate; petioles up to :>i-8) mm long. Flowers appearing before the leaves, on iihort. mostly hairy pedicels; buds sparsely to densely hairy. Calyx lobes 0.5 mm long. Corolla yellowish-green. 1-5 111111 long. Drupe subglobose, slightly ribbed. 10-15 mm in diameter, with 4 or 5 (or fewer due to abortion) pyrenes. Fig. 141.51. Woodland, often associated with thickets in gullies; 550-1500 m. IL GG; S Sudan. N Uganda. S Somalia.

Friis et al. 2437; Haugen 208; P uff & Ensermu K. 861103-2/3. A second subspecies, subsp. comorensis (Robyns) Verde. (1982). with glabrous buds and typically glabrous leaves, occurs in Kenya, Tanzania and on the Comoro Is­ lands; it may not be worth upholding. (See also comments following the genus description.) 52. RYTIGYNIA Blume (1850) Shnibs or small trees, unarmed (in the Flora area); branches usually with lenticcls. Leaves decussate, rarely also in whorls of 3, glabrous or hairy. with domatia in axils of veins, shortly petiolate. Stipules with linear trian­ gular appendages; base of stipules villous inside. Flowers 5-merous. axillary. several to few in axillary , paired, ses­ sile clusters or shortly pedunculate umbel-like cymes. Ca­ lyx truncate above or with distinct teeth. Corolla tube cylindrical, lobes valvate in bud, acute to apiculate al apex. Anthers (shortly) exserted. Style with crown-like or capitate stigmatic knob exserted. Ovary 2-locular [in the Flora area; 3-5(6)-locular elsewhere]. 1 ovule per locule. pendulous from placenta attached to near top of septum. Fruits drupes, mostly 2-lobed and with 2 one-seeded py­ renes.

278

141. RUBIACEAE: 52. Rytigynia

Figure 141.52. R Y T IG Y N IA N EG LEC TA var. N E G L E C T A . 1 - fruiting branch x */4; 2 - part o f stem showing lentieels, stipule and bases o f petioles x 2; 3 - leaf, lower surface showing domatia x V2; 4 - bud x 2V3; 5 - flower x 2Vi, 6 - calyx, ovar>, style and stigma x 2 Vi, 7 - drupe x 2 V i. 1 & 2 from M acLachlan 2125; 3 from P u ff et al. 870423-2/4; 4 - 6 M esfm T. 6879; 7 from de Wilde & D e Wilde-Duyfjes 6723. Drawn by Sarah Howard.

An Afro-Madagascan genus of 60-70 species; I in the Flora area. Not to be confused with Canthium and allied genera which, in the Flora area, neither have apiculate corolla lobes nor stipules which are villous inside.

long. Calyx minute, more or less truncate above or indis­ tinctly toothed. Corolla greenish-yellow, yellowish or creamy-white, tube 1.5-2 mm long, with a ring of hairs in­ side, lobes as long as tube, more or less apiculate. Dmpe black, mostly 2-lobed, 7-10 x 6-9 mm. Forest, forest edge, montane scrub, woodland; river­ ine vegetation; (1200-)1650-2400(-2600) m.

R. neglecta (Iliern) Robyns (1928); Canthium neglectum Hiem (1877) - type: GD. nr. Gaffet. Schimper 1106 (K holo.. BM E iso.) [p.p.]. Shrub up to 3 m tall, often with more or less horizontal decussate branches; younger parts distinctly lenticellate. Leaves decussate, rarely also in whorls of 3, oblong- or ovate-elliptic. (3—)5—11 (—13) x (1.2—)2—6(—7.5) cm. acu­ minate at apex, narrowed or more or less rounded at base, glabrous (save for domatia) or pubescent to more or less velvety; petioles up to 6 mm long. Stipules apiculate, up to 8(—10) mm long, densely (yellow'ish-)white villous in­ side. Flow'ers several, subumbellate or fasciculate in leaf axils, sometimes appearing before or with the leaves; pedicels up to 3 mm at first elongating up to 8(—10) mm in fruit: inflorescences sometimes on peduncles up to 2 mm

I Leaf blades quite glabrous (save for domatia). var. neglecta - Leaf blades pubescent to ± velvety beneath. var. vatkeana var. neglecta Fig. 141.52. GD GJ SU AR WG IL KF SD BA HA; Sudan. N Uganda and N Kenva. P uff & Ensermu K. 820809-1/1, 8208181/3. Puff et al. 820924-2/1. Plants found in open habitats (bush clumps, etc.) often have relatively larger, tougher and lighter green leaves titan those found in shadier localities.

141. RUBIACEAE: 52. Rytigynia, 53. Vangueria

an vatkeana (Hiem) Verde, in Kew Bull. 42: 158 (1987); Canthium vatkeanum Hiem (1877) - type: GD, Repp [Reb| valley, Gerra-Abuna-Tecla-Haimanot, Schimper 1130 (K holo., BM iso.). Rytigynia kiwuensis sensu Cufod. (1948, 1949, 1965), non (K. Krause) Robyns. .}D WG GG SD BA; endemic or possibly also in Uganda. Haugen 392; Mooney 7785; Puff & Ensermu K. 320809-1/2. 53. VANGUERIA Juss. (1789) Shrubs or small trees, unarmed. Leaves decussate, dccidu:>us. glabrous or pubescent, petiolate. Stipules broadly tri­ angular at base, narrowly triangular to linear above, villous inside. Flowers 5-merous, usually many in axillary, paired, pedunculate, often lax dichasial cymes; nracts and bracteoles small to minute. Calyx lobes trian­ gular, linear or elongated and more or less band-like. Co­ rolla frequently greenish, tube cylindrical or campanulate, inside with a ring of deflexed hairs, lobes acute to apicu­ late, spreading to recurved. Anthers attached to throat with short filaments, usually shortly exserted. Style with stigmatic knob slightly exserted; stigmatic knob more or less cylindrical. Ovary 5-locular, 1 ovule per locule, pen­ dulous from placenta attached to near top of septum. Fruits drupes, large and globose, with 5 (or occasionally Iewer by abortion) woody pyrenes. An Afro-Madagascan genus of c 15(?) species (taxa. in part, ill-defined); some species cultivated in tropical re­ gions because of their edible fmits; 3 species in the Flora area. 1. Calyx lobes narrowly triangular, 0.5—1,5(—2.5) nun long; corolla lobes acute, mostly shorter than the corolla tube; buds 5-7 mm long; inflorescences ± lax. 1. V. madagascariensis - Calyx lobes ± band-like. (1,5-)3-7 mm long, often curved; corolla lobes distinctly apiculate. as long as to (much) longer than the corolla tube; buds 10—12 mm long; inflorescences ± congested. 2 2. Young parts and leaves glabrous. 2. V. apiculata -r Young parts and leaves rusty- or golden-brown pubescent. 3. V. volkensii 1. V. madagascariensis Gmel. (1791) - type from Madagascar. Shrub or several-stemincd small tree up to 5(—15) m tall; oung pans glabrous or golden-brown pubesccnt. Leaves "iroadly elliptic, ovate-lanceolate or lanceolate, (4-)6.520 x (1.5—)2.5—9 cm. acute to shortly acuminate at apex, narrowed to rounded at base, glabrous or golden-brow n pubescent; petioles 0.5-1 cm long. Stipules haiiy inside, and (on younger parts) often covered with resinous excrelon, triangular at base, with a narrowly lanceolate to lin­ ear lobe, up to 12 mm long, above. Inflorescences ypically rather many-flowered and more or less lax, up to i0 mm long, branched 3 to 4 times; flowers on the

279

branches arranged in zig-zag fashion; pedicels 1-2 mm long (those of terminal flowers longer, thickened and elongated in fruit). Calyx lobes narrowly triangular, 0.5—1.5(—2.5) nun long, often spreading or recurved. Co­ rolla greenish-yellow, yellowish or creamy-white, tube wide, broadly cylindrical to more or less funnel-shaped, 3-4 mm long, lobes mostly shorter than tube, acute; bud 5-7 mm long. Mature fruit globose, c 30 mm in diameter, mostly only one fruit per inflorescence; pyrenes 15-20 mm long. 1. Plants glabrous or nearly so. var. madagascariensis - Plants golden-brown pubescent. var. abyssinica var. madagascariensis Vavanga chinensis Rohr (1792). Va\>anga edulis Vahl (1792), based on Vavanga chinensis, nom. illeg.\ Canthium edule (Vahl) Baill. (1880), nom. illeg.', Vangueria edulis (Vahl) Vahl (1794), nom. illeg. V. edulis Lam. (1792. 1812). V. commersoni Jacq. (1797). V cymosa Gaertn.f. (1805). V. acutiloba Robyns (1928). V floribunda Robyns (1928). V. venosa Hochst. ex Robyns (1928) non Sond., nom. illeg. - type: TU Dieladjeranne, Schimper 653 (P holo., K UPS iso.). Canthium maleolens Chiov. (1939) - type: SD, Moyale, Cufodontis 731 (FT holo.). Vangueria robynsii Tennant (1968). Mostly associated with rocky outcrops in Acacia Commiphora and other woodland or in Juniperus forest; often al base of large boulders; 1000-1600 m. EE/EW EW TU GD SU GG SD BA; west to Nigeria, tropical East Africa south to South Africa (Transvaal); occasionally cultivated elsewhere in the tropics. Beccari 136; Puff el al. 870429-1/2. 870509-1/1. var. abyssinica (A. Rich.) Puff, stat. nov.; Vangueria abyssinica A. Rich., in Tent. FI. Abyss. 353-354 (1848) - type: TU, Tchelatchekanne (Djeladjeranne], Takazze River, Quartin Dillon s.n (P holo.). ? V. barnimiana Schweinf. (1862). V. edulis Lam. var. pubescens Fiori in Agric. Col. I t a l 5, Suppl. 140 (1912) - type: EE from Ghinda to Dongollo, Fiori 434 (FT holo.). “ K. venosa Robyns var. pubescens (Fiori) Robyns (1928)” sensu Cufod. (1965). V. infausta sensu Cufod. (1965). non Burch. (1824). Riverine vegetation; 450-1300 m. EE EW TU. possibly endemic. Baldrati 227. 3687; Tellini 895. 2. V. apiculata K. Schum. (1895) - type from Kenya. V longicalyx Robyns (1928). Small tree or shrub, 3-10 m tall, several-stemmed; young parts glabrous. Leaves elliptic- or ovate-lanceolate to lan­ ceolate, (3-)4.5-20(-23) x 2-8.5(-12) cm. acute to acu­

280

141. RUBIACEAE 53 Vangucna

Figure 141.53. VAN G U E R IA A P1C U LATA : 1 - shoot with flowering and fruiting inflorescence and veg­ etative branch x 2^3; 2 — bud x 3'/3; 3 - flower x V h . 4 - sta­ men x 4'/a; 5 - style and stigma x 614; 6 - drupe (shrunk, show­ ing outlines o f pyrenes within) x 2/ 3. 1-3 from Zenna Abebe 1109; 4 & 5 from Burger 175 :; 6 from Thomerson 727. Drawn by Sarah Howard.

minate at apex, narrow ed to more or less rounded at base, glabrous, petioles 0.5-1 cm long. Stipule haiiy within, tri­ angular at base, with a more or less linear lobe, up to 10 mm long, above. Inflorescences more or less congested and mostly rather many-flowered, 20-40 mm in diameter, more or less clearly pedunculate; peduncles up to 10(—15) mm long, pedicels 1-3 mm long (elongating in fmit). Ca­ lyx lobes more or less band-like, (2—)3—7 mm long, often curved. Corolla greenish-white, greenish or yellowish, glabrous outside, tube densely hairy inside, 3—5 mm long, lobes (much) longer than tube, (long) apiculate; bud 10-12 mm long, with distinct apiculate appendages above. Mature fruit globose, up to 30 mm in diameter, pyrenes 10-15 mm long. Fig. 141.53. Riverine forest or scrub, Podocarpus forest or (ever­ green) montane scrub; 1200-2100 m. GD WG IL GG SD BA HA; Sudan, Somalia, tropical E Africa south to Zim­ babwe, Rwanda Rep. Dem. du Congo. Burger 1751; Mesfm 4960; P uff & Ensermu K. 820813-1/2.

Plants growing in sun-exposed open habitats tend to have lighter green and tougher leaves than plants occur­ ring elsewhere (e.g., HA, P uffstal. 820922-1/1; Ensermu K 1386). 3 V. volkensii A.'. Schum. (1895) var. volkensii - type from Tanzania. V. linearisepala K. Schum. (1903). V. bicolor K. Schum. var. crassiramis K. Schum. (1905). Shrub or small tree up to 5 m tall, youngest parts rusty- or golden-brown hairy. Leaves ovate- or elliptic-lanceolate to lanceolate, (2—)4—17 x (1—)3—8 cm, acute to acuminate at apex, narrowed to more or less rounded at base, golden-brown pubescent, pubescence denser below and on veins; petioles hairy-, 3-15 mm long. Stipule hairy within, triangular at base, with a more or less linear lobe, 3-8 mm long, above. Inflorescences more or less con­ gested. 15-25 mm in diameter, several- to rather mam -

141. RUBIACEAE: 53. Vangueria, 54. Pentanisia

281

Figure 141.54. P E N T A N IS IA O URANO GYNE 1 - habit x Vy, 2 - node showing stipule x 2; 3 - calyx x 10: 4 - disc x 10; 5 & 6 —longitudinal section o f short- and long-styled flower x 3; 7 — style with stigma lobes x 6; 8 - longi­ tudinal section o f ovary x 20; 9 — fruit x 6; 10 - cross section o f fruit x 6; 11 - seed, different views x 10. 1 & 5 from Richards 23360; 2 from Dyson Hudson 19; 3 , 4, 6 -8 , 11 from Lewis 5989; 9 & 10 from Sym es 573A. Drawn by Diane Bridson. (Reproduced with permission from FI. Trop. E .Afr. Rubiaceae: fig. 18.)

f nvered, more or less clearly pedunculate, peduncles up to 10 mm long, pedicels 1-3 mm long (elongating and thickening in fruit); inflorescence branches pubescent. C ;iiyx lobes more or less band-like, (1.5—)4—5 mm long, often curved. Corolla hairy outside (at least the lobes), greenish to whitish- or yellowish-green, tube hairy inside, 4 -5 nun long, lobes as long as, or longer than, tube, apicukrne; bud c 10 mm long, with distinct apiculate append­ ages above Fruit globose, up to 25 mm in diameter (larger v hen fully mature?); pyrenes 12-18 nun long.

tropical E Africa, Rwanda, Rep. Dem. du Congo. P uff et al. 820922-1/1, -1/12; Sebsebe D. & Tewolde B.G.E. 905.

Rocky outcrops with bush clumps; rocky areas in J miperus forest or in Combretum - Terminalia-Acacia v oodland, 1250-2300 m. GD SU SD BA HA; Sudan,

Herbaceous plants or sometimes more or less shrubby. Inflorescences terminal. Flowers heterostylous. Ovary 2-5-locular; 1 pendulous ovule per locule. Fruits either in-

Two more varieties, distinguished by glabrous corol­ las, occur in Uganda [var. fyffei (Robyns) Verde.] and Tanzania [var. kyimbilensis (Robyns) Verde.].

KNOXIEAE Hook.f.

Puff & Robbrecht (1989) in Bot. Jahrb. Sysl. 110: 511-558.

141. RUBIACEAE: 54. Pentanisia

282 I

dehiscent and hard, or drapes, or splitting into two indehiscent mencarps. 54. PENTANISIA Harv. (1842) Perennial heit>s, subshrabs or dwarf shrubs. Leaves decussate, mostly (sub)sessile. Stipular sheaths usually with several setae. Flowers 5-merous, heterostylous. mostly in terminal, many-flowered, subcapitate inflores­ cences, usually somewhat elongated and more or less spike-like in fruit. Calyx lobes unequal, 1-3 often en­ larged, foiiaceous, the others small to minute. Corolla tube narrowly cylindrical, throat densely hairy, lobes valvate in bud. ovate to oblong. Anthers included in slightly widened uppermost part of tube (long-styled flowers) or exserted. Style with 2-5 filiform stigma lobes included (short-styled flowers) or exserted. Ovary 2-5locular, 1 ovule per locule, pendulous from placenta at­ tached to near top of septum. Fruits crowned by persistent calyx, mostly more or less subglobose, slightly lobed or ribbed, dry. indehiscent, hard and more or less woody (or, not in Flora area, indehiscent more or less succulent or spongy and with hard endocarp or tardily dehiscent into mericarps). Seeds small, somewhat compressed. An Afro-Madagascan genus of 16 species; 2 in the Flora area. 1. Corolla tube 7-20(-23) mm long, corolla bright blue; enlarged calyx lobes up to 8(—10) mm long. 1. P. ouranogyne - Corolla tube 25-45 mm long, corolla pale blue, whitish(-blue) or white; enlarged calyx lobes up to 15(—20) mm long. 2. P. longituba 1. P. ouranogyne S. Moore (1880); P. ouranogyne S. Moore var. glabrifolia Cufod. in Nuov. Giorn. Bot. Ital. n.s. 55: 87 (1948) —type. SD. El Dire, Corradi 2731-2733 (FT syn). Perennial herb, up to 30 cm tall, with several to mam branched or unbranched, mostly hairy stems from a some­ what woody, branched rootstock. Leaves decussate but often seemingly in larger numbers at nodes due to leafy, much-contracted short shoots, lanceolate to linear-

lanceolate, (2-)2.5-6(-10) x (0.2-)0.3-1.5(-2) cm, most­ ly hairy; petioles up to 5 mm long. Stipular sheaths with several setae up to 5 mm long. Inflorescences manyflowered, sometimes branched, 10-20 mm in diameter, on peduncles up to 50 mm long. Enlarged calyx lobes up to 8(—10) x 2 mm, other lobes minute. Corolla bright blue, mbe 7-20(-23) mm long, lobes narrowly lanceolate to ovate. 2-4 x 0.5-2 mm. Ovary mostly 3-locular, h a m . Fruits more or less globose, 2.5-4 mm in diameter, slightly furrowed, hairy, very hard and more or less wood) . Fig. 141.54. Acacia - Commiphora (- Combretum) woodland or bushland. grassland often with Acacia drepanolobium; (800-) 1400-1900 m. SU AR GG SD BA HA; Somalia, south to S Tanzania. Ensermu K. 1274; P uff et al. 870426-7/6. 870513-5/2. 2 P. longituba (F ranch.) Oliv. (1888); Knoxia (Pentanisia) longituba Franch. (1882) type from Somalia. Pentanisia pentasiana Mattei (1908). Perennial herb, up to 60 cm tall with several unbranched or branched stems from a somewhat woody branched rootstock; stems glabrous and shiny below, hairy above. Leaves decussate but often seemingly in larger numbers at nodes due to leafy, much-contracted short shoots, lanceo­ late to linear-lanceolate, 3-7.5(-8) x 0.2—1(—1.5) cm, hairy or glabrescent; petioles up to 7 mm long. Stipular sheaths with several setae up to 3 mm long. Inflorescences many-flowered, compact, 10-20 mm in diameter, on peduncles up to 35(-45) mm long. Enlarged calyx lobes up to 15(—20) x 2 5 mm. other lobes minute. Corolla pale blue, whitish (-blue) or white, tube 25-45 mm long, lobes oblong, 5-7 (-8) x 1-2 mm. Ovary mostly 3-locular, hairy Fruits subglobose, 4-6 mm in diameter, slightly furrowed, hairy, very hard and more or less woody. Acacia - Commiphora wood- or bushland, grassland, often also on rocky hillsides, 900-1400 m. GG SD HA; Somalia N Kenya. P u ff & Ensermu K. 821220-1/2; P uff et al. 870424-9/i, 870425-6/1. Very closely allied to the previous species. Hybrids may be expected in areas where both species occur.

283

142. CAPRIFOLIACEAE by Ensermu Kelbessa* Vericourt, Caprifoliaceae in FI. Trop. E. Afr. 4 pp. (1968); Fukuoka, Mem. Fac. Sci. Kyoto Univ., ser. Biol. 6: 15-58 (19' 2); D ’Arcy, 180. Caprifoliaceae in FI. Panama, Part IX. Ann. Missouri Bot. Gard. 60: 155-161 (1973).

Shrubs, herbs or lianas, seldom small trees. Leaves simple or pinnately compound, opposite, or rarely alternate. Stipules present or absent. Flowers in cymes, panicles or umbels, or rarely solitary, borne terminally or in the axils o f leaves. Bracts present or sometimes absent; bracteoles usually present. Flowers usually bisexual, regular or irreguiiar. Calyx fused to the ovary, with 4-5 or rarely 3 lobes. Corolla often irregular and 2-lipped, or regular, with 4-5 .ual or unisexual, usually zygomorphic, 5-merous. Calyx often small or obsolete, lobes forming a pappus, u;;..ally toothed. Corolla funnel-shaped or tubular, sometimes saccate or spurred below, limb 3-5-fid. Stamens 1 4, epipetalous, alternating with the corolla lobes. Ovary inferior, carpels 3 and 3-locular, but only one locule ft "lie. Fruit a one-seeded achene, often with a persistent, hair-like or w ing-like, calyx at the top. Seed pendulous. About 13 genera, 4 o f which (C entranthus, Fedia, Valeriana and Valerianella) occur in Africa, and about 4' ’i) species, occurring m ostly in the tem perate region o f the northern hem isphere; 1 genus and 1 species in the F ora area.

VALERIANELLA Mill. (1754) A .nual herbs with dichasial branching. Basal leaves entiiu n d iv id e d or dentate, sometimes forming a rosette; st< m leaves often toothed, sometimes entire, joined at the b; se. Flowers bisexual, in terminal partial inflorescences. C. l\x often short or obsolete at anthesis, entire or 3-6to thed. Corolla funnel-shaped, narrowly campanulate or tu ular, attenuate at the base, with 5 slightly unequal lo es, bluish or pinkish. Stamens 3, inserted towards the t o : of the corolla-tube. Style shortly 3-fid at the apex. A hene oblanceolate to spathulate, glabrous to pubescent, or .'-seeded. 3-locular with the 2 abaxial locules empty.

Afro-alpine moorland, montane forest, grassland and as a weed in cultivated areas; (2200-)2900-4100 m. TU GD AR BA; Kenya, the Mediterranean region. Mooney 9602A; Hedberg 5528; Friis et al. 5656.

About 80 species, natives of temperate and subtropical co intries of the northern hemisphere, most numerous in so it hem Europe; 1 species in the Flora area.

V. m icrocarpa Lois. (1810); V. dentata (L.) Poll. var. microcarpa (Lois.) Fiori, iRel. Monogr. Agr.-col. 58:40 (1940)- type: France, . Robert s.n. (?P holo.).

F

V. abyssinica Fresen. (1836) - type: GD. Simen, fiuppell s.n. (FR holo.). V dentata sensu Hiem. (1877), non (L.) Poll.

At nual herb 5-25(-50) cm high, erect or ascending to al­ most spreading. Leaves sessile (0.5-) 1-6.5 x 0.2-1.1 cm. lo wer ones spathulate. entire, upper oblong, dentate, uppe m ost bract-like and linear. Inflorescence a compound di hasium with bisexual, sessile flowers. Calyx-limb obscire, oblique and entire. Corolla funnel-shaped, 1-1.8 m n long, pale pink to wliite, with 5 lobes, 2 of which are sn .iller than the others. Stamens 3, included or slightly ex icrted. Style club-shaped to shortly 3-lobed. Achene o\ nd, 1-2 x 0.5-1 mm, hispid to pubescent (glabrous on th< abaxial face), brown, sterile, loculi reduced to ribs. Fig. 143.1. * v niversity o f Nairobi, Department o f Botany, Chiromo Campus, P.O. Box 30197. Nairobi, Kenya.

Figure 143.1. V A L E R IA N E L L A M ICRO CARPA: 1 - habit x 2/3; 2 - part o f inflorescence x 4'/2; 3 - flower x 16; 4 - corolla, opened out to show stamens x 16; 5 - stamen x 20; 6 - ovary and style x 16; 7 - fruit x 10; 8 fruit in transverse section x 10; 9 — seed x 10. 1 from Leaky & Evans 144; 2 -9 from G. Taylor 1436. Drawn by I leather Wood. (Reproduced with permission from FI. Trop. E. Afr. V alerianaceae: fig. 3.)

286

144. D1PSACACEAE by I. Hedberg* and 0 . Hedberg* Cufodontis, E n u m 1028-1030 (1965); Napper. Dipsacaceae in FI. Trop. E. Afr: 11 pp. (1968); M.J. Cannon & J ?M . Cannon, Dipsacaceae in FI. Zamb. 7(1): 77—85 (1983); Verlaque, in Rev. Gen. Bot. 91: 81-121 (1984), Rev. Cytol. 3iol. Veg. 8(2): 117-168 (1985), 8(3): 171-243 (1985) and 9(1): 5-72. 9(2): 97-176 (1986).

Annual or perennial herbs, sometimes suffrutescent. Stems erect. Leaves opposite, without stipules, entire or dissected, often heterophyllous. Inflorescence one or several dense, cymose heads with convex receptacles, which are naked or provided with bracts, each subtending a single flower. Flowers sessile, bisexual, zygomorphic with the ovary enclosed in a tubular involucel, formed by the fusion o f 2 or 4 bracts, often with a mem­ braneous cupular or spreading lobed limb. Calyx small, irregularly 4- or 5-lobed, sometimes with long bristles. Corolla united, irregularly 4- or 5-lobed with spreading lobes Stamens 2-4, alternating with the corolla lobes; anthers exserted. Ovary inferior and unilocular with a single ovule; style filiform; stigma entire or bifid i*ruit an achene surrounded by the involucel and crowned by the persistent calyx. A small and primarily temperate family o f 8 genera and about 250 species occurring in Eurasia and Africa: 3 genera and 3 species in the Flora area. Key to genera 1. Calyx without bristles; corolla cream or white. 1. Dipsacus - Calyx with bristles, corolla lilac, mauve or pink. 2 2. Calyx bristles naked, sometimes scabrid; recepta­ cle bracts present. 2. Scabiosa - Calyx bristles plumose; receptacle bracts absent. 3. Pterocephaius 1. DIPSACUS L. (1753) I. & O. Hedberg in Bot. Not. 129: 383-389 (1977). Erect, perennial herbs, with aculeate, pubescent or rarely smooth stems. Leaves pinnatipartite or entire, the upper ones sometimes united at the base. Inflorescences termi­ nal and lateral. Heads involucrate with one or two rows of foliaceous, often spine-tipped bracts of variable length. Receptacle bracts acute, leathery and usually spinetipped, usually shorter than the involucral bracts. Involucel (with or without a cup-shaped limb) developing 8 longitudinal furrows. Calyx cup-shaped, small, undulate or minutely toothed, usually pubescent or pilose. Corolla 4-lobed, with unequal lobes. Stamens 4, exserted. Stigma entire or rarely shortly bifid. 15 species in Eurasia, tropical Africa and Sri Lanka; 1 in the Flora area. D. pinnatifidus Steud. ex A. Rich. (1848) type: SD. Mt Aber near Tschenausa, Schimper 11:665 (P lecto.. K S isolecto.). D. appendiculatus Steud. ex A. Rich. (1848) type: SD, Mt Aber near Tschenausa, Schimper II: 865 (P holo., K iso.). D. schimperi A. Br. (1867) - type: cultivated in Berlin (not seen).

* Dept, o f System atic Botany, Uppsala University, SE-752 36 Uppsala. Sweden.

N o rbyvS gen

18D.

D. setosus Hiem (1867) - type: Ethiopia. Chire. Quartin-Dillon s.n. (K holo.). D. pinnatifidus var. integrifolius Engl., in Bot. Jahrb. Syst. 19 Beibl. 47: 49 (1894). D. bequaertii De Wild. (1922). D. kigesiensis Good (1924). D. eremocephalus Pic.Seim (1951) - type: GD, Simien, Mt Buahit, Pichi-Sermolli 2619 (FI holo., K photo). Cephalaria acaulis Steud. ex A. Rich. (1848); Simenia acaulis (Steud. ex A. Rich.) Szabo (1940); Dipsacus acaulis (Steud. ex A. Rich.) Napper (1968) type: GD. Simien, Mt Buahit, Schimper 563 (P holo., K UPS iso.). Perennial, with stout rhizome. Stem erect, from 2 cm to 3 m tall, usually angular and scabrid but occasionally terete and smooth. Basal leaves up to 25 cm long, petiolate. Stem leaves sessile, lanceolate, pinnatipartite, 4-20 cm long, toothed or entire, usually glabrous Ter­ minal inflorescences of one to several globose heads, 1.5-4 cm in diameter. Involucral bracts spreading, foliaceous, as long as or shorter than head. Receptacle bracts leathery, variable in size and shape, spine-tipped or obtuse. Involucel pubescent, up to 4 mm long in fruit. Calyx 1-1.5 mm long, shallowly 4-lobed, pubescent. Corolla white or cream, 6-15 mm long, pubescent on the outside. Fig. 144.1. Ericaceous scrub and grassland; 1980-4100 m. GD GJ SU AR KF SD BA HA; Cameroun, Sudan, Uganda, Kenya, Tanzania, Rwanda, Burundi, Rep. Deni du Congo. Hedberg & Getachew A. 5397; Hedberg 5592; Friis et al. 87.

2. SCABIOSA L. (1753) Annual or perennial herbs, glabrous or pubescent, often heterophyllous Leaves opposite, simple or deeply pin­ nate, often in basal rosettes. Inflorescence terminal. Flower heads long-pedunculate, involucrate; involucral

144. DIPSACACEAE: 1. Dipsacus, 2. Scabiosa

287

Figure 144.1. D IP SA C U S P IN N A TIH D U S : 1 - habit x V2. 2 - head showing bract variation x 2/3, 3 involucel and flower x 6; 4 - fruit with persistent calyx x 6. J from Napier 664; 2 & 3 From M ooney 8305; 4 from Gillett 14703. Drawn by Derek Erasmus. (Reproduced with permission from FI. Trop. E. Afr. Dipsacaceac: fig. ].)

D.E.

bracts herbaceous, in 1-3 rows. Receptacle hemispherical to o lindrical. receptacle bracts linear-lanceolate. Involucel smooth or 8-fumowed throughout its length or towards the :op only, expanded above into an orbicular or funnelshaped. scarious. many-veined corona. Calyx cup-shaped with 5 teeth, each ending in a long scabrid or barbed bnsue; brisUes subequal or equal. Corolla with a short tubs and 5 unequal lobes. Marginal flowers usually con^derably longer than tlie central ones. Stamens 4. Stigma oblique, entire.

About 100 species widespread throughout the temper­ ate regions o f Europe and Asia; also occumng in tropical and southern Africa; 1 species in the Flora area. S. columbaria L. (1753) - type from Sweden. •S', columbaria var. longibracteolata Chiov in Ann di Bot. 9(l):55-85 (1911) Peremual, 6-70 cm tall, erect with thick rootstock and branched, pubescent stems, markedly heterophyllous.

288

144 DIPSACACEAE 2. Scab\osa. 3. IHeroccphalus

Figure 144.2. SC A B IO SA COLU M ­ B A R IA 1 - habit x Vi; 2 flower x 3; 3 fruit with persistent calyx x 3. 4 fruiting flower head x Vi. All from Saunders Davies s.n. Drawn by Marga­ ret Tebbs. (Reproduced with permis­ sion from FI. Zamb. 7(1) Dipsacaireae: Tab. 12.)

Basal leaves sessile, entire or rarely lobed in the lower part, the upper pan with crenate or toothed margin, gla­ brous or pubcscent. Lower stem-leaves entire or pinnalifid. rarely bipinnate: upper ones smaller, deeply pinnatipartite or simply pinnate. Flower heads 1.5-3(-4) cm in diameter Involucral bracts lanceolate, acute, at least half the length of the marginal flowers. Bracts of the receptacle linear-lanceolate, acute. 3-5 mm long, pu­ bescent. hrvolucel pubescent, becoming 8-furrowed and (including the limb) c 3.5-5 mm long in fruit Calyx pilose, with 1.5-6 mm long bristles elongating at matu­ rity. Corolla lilac, mauve, or rarely pinkish. 4.5-18 mm * Instilut fur I orslentomologie. Forslpalhologie und Forstschul/. Univcrsital lur Bodcnkultur. Masenaucrstr. 38. A -1190 Wien. Austria. ** iiotanischer se - with prominent nerves, ribs, or veins, net-i eined - when the sm aller veins are interconnected to form unet-likc (reticulate) pattern.

net venation - closed venation in which the veins branch and loin.

nei l.ral flower - a flower without sexual parts, without func­ tioning stamens or pistils.

nocturnal - active at night, as in flowers that open at night; the opposite o f diurnal.

not'

the place on a stem where a leaf or bud is formed; a thick­ ened area on a stem-like organ where other parts are attached «'T where the organ will later break in two.

nocule - a small node or thickening; especially used in roots with thickened swellings in which nitrogen-fixing bacteria lave.

nu< . flower - a flower without a perianth, naked or abhlamydeous.

numerous - used for parts o f a flower which are 15 or more in number, see also indefinite.

nut a fruit with a hard outer f overing that does not split open vthcn ripe, m ostly comparatively large and with one or two •.ceds; see also nutlet and achene.

nutlet - a small nut. ob- a prefix meaning ‘opposite’, ‘inverse’, or ‘against’. obcl avate - attached at the thicker end. obci irdate - with a broad 2-lobed apex and a narrowed base, as !he leaflets in species o f Oxalis.

obc iplostemonous - having two whorls o f stamens (usually

301

oligo- - a prefix meaning ‘few -’, as in oligocarpus: with few fruits.

opaque - something that does not allow light to pass through, a surface that is dull and not lustrous or shiny.

Operculate - opening by a lid or cover; covered with a cap. operculum - a lid, cap or covering that comes off as a single unit.

opposite - a term used for two leaves or two branches that arise from the same node on the opposite sides o f the stem; a term also used for organs that arise opposite each other or when one arises at the base o f another, as stamens opposite a petal or sepal.

o rb icu la r - a flat structure with an almost circular outline.

organism - any individual living thing. ortho- - a prefix meaning ‘straight’, ‘upright’, or ‘true’ or ‘cor­ rect’.

ostiole - an opening or pore. oval - broadly elliptic or having an egg-shaped outline, usually widest at the middle.

ovary - that part o f the pistil which contains the ovules within one or more locules and which will produce the fruit if polli­ nation (and fertilization) takes place, all angiosperms have ovaries.

ovate - a Hat structure which is egg-shaped in outline with the broadest part near the base and the narrow part near the apex; the opposite o f obovate; compare elliptic and oblong.

ovoid - egg-shaped with the broadest part near the base and the narrow part near the apex; the solid or 3-dimensional form o f ovate.

ovule - an organ which contains the embryo sac and the egg cell

rtvice as many as the petals) with the outer stamens opposite tile petals.

within the locule o f the ovary (in angiosperms) or borne on fertile scales (in gymnosperms); after fertilization develops into a seed.

Obi ;riceolate- with the shape o f the end o f a lance or spear but

palmate - with three or more parts attached to a single point and

v.ith the narrow end towards the base; the inverse o f lanceoute.

Obi lie - broadly elliptic with the long dimension perpendicular 0 the axis o f the organ.

obl iiate - no choice; generally used for organisms which can 1nly exist as parasites.

obl.ijue - a leaf-base in which the two sides are unequal, as in rtegonia.

obi ng - a plane shape longer than broad with nearly parallel

radiating outward, as the fingers o f an open hand radiating outwards from the palm o f the hand; as in palmate venation, or in palmately compound leaves.

palm atifld - with palmately arranged lobes, the leaf being di­ vided more than halfway to the single point from which the nerves radiate.

palmatilobcd - with palmately arranged lobes, the leaf being divided halfway or less to the single point from which the nerves radiate; compare with palmatifld.

i ides, almost rectangular in outline but with rounded ends ..nd with the length two or three times the width.

palm atipartite - palmately divided almost to the centre or

obo rate - a plane shape with an egg-shaped outline but with the

palmatisect - palmately divided to the centre or midrib. pandurate - a rounded plane figure with a slender portion near

(iroadest part near the apex and the narrow side near the lase.

obo void - egg-shaped but with the broadest part near the apex

midrib.

the centre and with two broad ends one o f which is broader than the other; the shape o f a body o f a violin.

- r»d the narrow part near the base; the solid or 3-dimensional orm o f obovate.

panicle - an inflorescence with an indeterminate axis (that con­

obptriform - shape like a water pot. obsolete - wanting or rudimentary. obtuse - with a blunt or rounded end or the margins o f the tip

tinues to grow and does not end in a flower) and many side branches each o f which bears two or more flowers. This term is often used for a branched inflorescence which is dif­ ficult to classify into any o f the other more precise types such as raceme, cyme, etc.

palming an angle o f more than 50°; compare acute.

OCr -a (ochrea) - a stipular growth that sheaths the stem near Hie

leaf-base, as in Polygonaceae.

papilionaceous - flowers that resemble the flower type o f the subfamily Papilionoideae o f the Fabaceae (Leguminosae).

OCtii- - a prefix meaning ‘eight’. odd -pinnate - a pinnate leaf with an odd number (3, 5 ,7 , etc.)

papillate - covered with many minute rounded gland-like

't leaflets, with a single terminal leaflet; the same as impariinnate; compare with even-pinnate and paripinnate.

parallel venation - generally used for veins that are parallel

structures or papillae; compare with muricate. with each other and with the margin of the leaf; sometimes

GLOSSARY OF BOTANICAL TERMS

302

used for secondary veins that are parallel with each other but not with the midrib or the margin, as in Musa and Ensete.

parasite - a plant (or animal) that lives upon another plant (or animal) and takes nourishment from it; compare epiphyte

parietal placentation - when the ovules are attached to the in­ ner surface o f the peripheral or outside wall o f the ovary, or the outer wall o f the locule.

paripinnate - a pinnate leaf with an even number (2, 4, 6, 8, etc.) of leaflets and without a single leaflet at the end; the same as even-pinnate; compare with imparipinnate, oddpinnate.

parted - divided almost to the base or to the midvein. parthenogenesis - the development o f a female gamete into a new individual without fertilization.

partite - the same as parted, also used as a suffix showing the number of parts in a structure, for example tripartite (3 parts).

patent - spreading or open peat - a deposit o f incompletely decomposed plant material with little or no soil, dark brown or black with a high carbon content; in Europe, mostly made up o f the moss Sphagnum

pectinate - divided to form many parallel parts like the teeth of a comb.

pedate - used for leaves which are palmately divided, with each division two-cleft, as in some Passifloraceae.

pedicel - the stalk o f a single flower within an inflorescence or group o f flowers (also used for the stalk o f a solitary flower).

perigynous - arising from a cup-like or tubular structure around the ovary, surrounding the ovary but not at its base nor united to it

periphery - along the margin or on the outer wall, persistent - remaining attached to the plant and not falling of' |>erulate - with a covering o f protective scales, as in many buds.

petal - a flat and usually broad part o f the inner whorl o f stenle appendages in the flower that together arc called the corolla, different from the outer whorl (sepals) and often brightly coloured.

petaloid - like a petal in colour and form; used tor bracts (as in Bougainvillaea), sepals and stamens,

petiole - the stalk o f a leaf on which the blade is borne, petiolule - the stalk o f a leaflet. ptianerophyte a plant with the perennating organs borne high above ground level,

phenotype - the visible, or chemical, or biologically detect­ able, manifestation o f the genotype produced as a consequencc o f g m y th and development,

phloem - the part o f the vascular system made up o f living cells that function primarily in the conduction o f tood, the inner bark.

photosynthesis • the process through which green plants make sugar from carbondioxide and water using sunlight energy captured by the green pigment chlorophyll,

phototaxis - the movement o f a whole organism in response to light

peduncle - the stalk that bears an inflorescence consisting o f

phototropism - a change in growth direction in response to

two or more flowers; the flowers may themselves each have a stalk (the pedicel) or be without a stalk.

phyllary - a bract-like part o f the involucre which subtends the

pellucid - clear and translucent, bright when viewed against the light.

peltate - with the stalk attached near the centre o f a more or less rounded shape and not at the edge, as in a peltate leaf.

pendulous - hanging down or drooping. penicillate' with a tuft o f hairs, often shaped into a point like a pencil.

penninerved - with pinnate nervation. penta- - a prefix meaning 'fiv e-’. pentam erous - with five parts, or with sets o f five parts, some­ times with multiples o f five (5.10.15. etc.).

pepo - a fleshy fruit with hard outer rind and without septa or separate chambers within, as in the Cucurbitaceae.

perennating - surviving from one growing season to the next; a structure or organ which enables a plant to survive a non-growing season. perennial - living for three or more growing seasons.

perfect flower - a bisexual flower with stamens and pistil functional.

perfoliate - when the leaf-base grows around the stem and the stem appears to have grown through the leaf.

light flower-hcads in the Compositae and some other families.

phyllode - a flattened leaf-stalk (petiole) or leaf-rachis with the form and function o f a leaf, as in some o f the species o f.lcacia introduced from Australia, compare with cladode

phyllotaxy - the arrangement o f leaves on the stem. phytogeny - the evolutionary history o f an organism or group o f organisms

pilose - with a loose covering o f soft long simple hairs. pinna - the primary division o f a pinnate leaf which can be a leaflet in simple pinnate leaves, or can be divided again into pinnules in a bipinnate leaf,

pinnate - when a compound leaf has its leaflets borne along an extension (the rachis) o f the leaf-stalk (petiole) or when the leaflets are borne on divisions or branches o f the rachis; leaves can be bipinnate, tripinnate, etc.; compare palmate.

pinnately - in pinnate fashion. pinnatifid - with the margin divided more than halfway to the midvein or centre and forming pinnate lobes.

pinnatilobed - with the margin divided to about half the dis­ tance to the midvein or centre and forming pinnate lobes.

pinnatipartite - with the leaf divided alm ost to the midvein or centre and forming pinnate lobes,

peri - a prefix meaning "around5. perianth - the outer sterile whorls or envelopes o f a flower,

pinnatisect - with the leaf divided to the midvein or centre and

made up of identical perianth segments, or by two different kinds o f perianth segments; sepals (calyx) or petals (co­ rolla).

pinnule - the second or third branching o f a compound pinnate

pericarp - the wall o f the ripened ovary or fruit (between the

pistil - the individual female structure that contains the ovule

locules and the outer surface); it may be o f one or as many as three layers (see exocarp, mesocarp, and endoearp).

and will produce the seeds. It is made up of one or several carpels and is usually divided into the following parts:

forming pinnate lobes, leaf; the branches or divisions that arise from the primary rachis o f a pinnate leaf; compare pinna,

GLOSSARY OF BOTANICAL TERMS

ovary, style, and stigma. A flower may have one pistil or many pistils. p i tilla te - oitcn used for flowers with female parts but no male parts and. therefore, unisexual; a female flower. pi: :illo d e - a reduced non-functioning pistil that does not pro­ duce seeds; often present in functionally male flowers. pii h - the soft spongy tissue found in the centre o f many stems. pii ted - marked with small depressions.

placenta - the part o f the ovarv on which the ovules are at­ tached pliicentation - the arrangement o f the ovules in the ovary. p lcom o rp h ism - having more than one form or shape. plicate - folded, with the edges folded together like a fan. plum ose - v\ith tufted or feather-like hairs or feather-like bris­ tles. pD:.‘u m a t 0 p h 0 r e s - a ir passages; more often used for the verti­ cal wooden projections from the roots o f some trees growing in very wet situations, as in swamps or mangrove forma­ tions. These woody projections rise above the water level and are believed to function in bringing air to the roots.

po I- a dry fruit that opens when ripe; often used as synonvmous with legume. po Ion - the powder-like grains produced in the anthers that will produce the male gametes necessary in fertilization. The pollen will germinate on the stigmatic surface o f the stvle (in angiosperms) or in the micropyle (in gymnosperms) and produces the pollen tube. The pollen tube grows into the m u le and releases the male gametes where thev can reach and fuse with the female gametes; the pollen-grain is homolDgous with a microspore. pollen-grain - see pollen. p o lle n -sa c - the chamber in which the pollen is produced after it eiosis has taken place; the theca (pi. thecae) o f the anther. polliinarium - pollinia joined by a pair o f translator arms to the central corpusculum.

303

pome - a fleshy fruit with a soft outer covering and seeds borne within stiff central partitions, as in the fruits o f M alus (apple) and Casimoroa (Rutaceae).

pore - a small, usually round opening, as in anthers opening by pores in the Ericaceae.

posterior - towards the axis, on or nearest the axis (adaxial or ventral); the opposite o f anterior.

precocious - developing early; used for flowers that develop before the leaves.

prickle - a small sharp outgrowth from the bark or surface. procumbent - lying along the ground. pro parte (proparte) - often abbreviated p.p ., meaning ‘in part only’.

prophyll - a much reduced leaf or bract. precocious - developing early; used for flowers that develop before the leaves.

procumbent - with stems that lie on the ground. proliferous - with adventitious buds on the leaves or in the (lowers that are capable o f producing new plants.

propagule - a short, lateral shoot by which certain plants are propagated; any offshoot.

prostrate - lying flat on the ground. protandrous - when the anthers discharge pollen before the stigma (in the same flower) is receptive; sometimes spelt proterandrous.

proto- - a prefix meaning ‘first’ or ‘original’. protogvnous (proterogynous) - when the stigma is recep­ tive before the anthers (in the same flower) have discharged their pollen.

proximal - nearer to the place o f attachment; the opposite o f distal.

pruinose - covered with a whitish wax or very fine powder pseudo- - prefix meaning ‘false’. pseudowhorl - Where leaves arise so close together around the stem that they appear to be in a ring.

pollination - the act of taking pollen from the anther and bring­ ing it to the receptive stigmatic part o f a style or to an ovule; (his can be carried out by different agents.

puberulous (puberulent) - covered with very short fine hairs

pollinium - a body comprised o f all the pollen-grains o f an anther-loculus; a pollen mass.

pubescent - with a covering o f soft hairs. pulverulent - covered with a powder; see also pruinose and

poly- - a prefix meaning ‘m any-’ or ‘m uch-’. polyadelphous (p o lyad clp h u s) - with three or more groups of united stamens. pol • and ro us - with many (more than 15) stamens. pol carpic - bearing fruit many times, as opposed to monoearpic. p o l' gam odioecious - dioecious but with some bisexual flow­ ers or with some flowers o f the opposite sex. pol i»amom onoecious - monoecious but with a few bisexual flowers. pol \ gam ous ■when a plant or different plants o f the same spe­ cies have both bisexual and unisexual flowers. po lvgyn o u s - with many pistils. pol vm orp h o us - with several or many forms, variable. pol ''petalous - with petals that are separate and free from each Uher p o h phyletic - evolved from more than one ancestral group. polysepalous - with sepals that are separate and free from each other.

or slightly hairy.

glaucous.

pulvinus (pu Ivin ate) - an enlargement or swelling, shaped like a cushion or pad. round and flattened, usually found at the base of the leaf or petiole.

punctate - marked with dots or glands or very small depres­ sions 1 mm or less in diameter.

punctiform - covered in small dots or depressions 1 mm or less in diameter.

pungent - ending in a sharp stiff point; with a strong smell or taste.

pustule (pustulate) - many small elevations like pimples or blisters.

pyrene- a nutlet or kernel; the ‘sto n e’ o f a drupe or similar fruit. pvriform (piriform) - shaped like a pear (Pynis)\ a solid shape with a broad apex (away from the point o f attachment) narrowing to a wide neck at or below the middle.

pyrophytic - a plant able to tolerate fire or needing fire to stim­ ulate flowering.

pyxis - a capsular fruit in which the top comes off as a lid. quadr- or quadri- - a prefix meaning ‘lour’.

GLOSSARY OF BOTANICAL TERMS

304

racem e - an indeterminate inflorescence in which the flowers

rhipidium - a cymose inflorescence with branches alternating

are borne along a single axis with the uppermost the youn­ gest, each flower with a stalk o f about the same length, com­ pare with cyme

from one side o f the vertical axis to the other; normally flat­ tened in one plane and fan-shaped.

racemose (racem ous) - arranged like a raceme. rachilla (rhachilla) - a small axis or rachis; the axis o f pinnules in compound pinnate leaves. The central axis o f the spikelct in the Poaceae (Grammeae) and Cyperaceae.

rachis (rhachis) - the axis o f a compound leaf or frond; the axis o f an inflorescence

radiaJ sym m etry - symmetrical about a central axis; when the structure is divided longitudinally along any axis, the two halves are mirror images o f each other.

radiate (radiating) - spreading outward from a point. radical leaves - leaves that an seso close to the base o f the stem that they appear to come from the top o f the root.

random b ran ch in g - branches which arise without any rela­ tionship to subtending leaves.

rank - a vertical row; when 2-ranked leaves arc in two vertical rows; a general term to denote a level in the taxonomic hier­ archy (variety, subspecies, species, genus, family, etc.).

raphe - the part o f the stalk o f the ovule (funicle) that is united or coherent to the outer wall o f the ovule and forms a ridge on the surface o f the seeds.

raphides - needle-shaped crystals found within cells o f plants. recaulescent - adnated to the stem or pedicel. receptacle - the axis or the central body o f the flower on which the flower parts (sepals, petals, stamens, and pistil) are borne

recurved - curved backwards, reduced - not properly developed or undeveloped, reflexed - bent downwards or backwards, bent abruptly back­ wards towards the base,

regular - (of flowers) radially symmetrical or actinomorphic, a flower that can be divided into two equal halves by two or more vertical sections (some authors consider regular syn­ onymous with symmetrical and include both bilateral and radial symmetry), compare with irregular, asymmetric, and zygomorphic.

reniform - kidney-shaped, as the seed o f many Fabaceae (Leguminosae), and some leaves.

repand - with an uneven or wavy margin, not as uneven as sinuate, compare with undulate.

repent - stems that creep along the ground and root at the nodes. replum - a septum within the ovary formed by the placenta, as in the Brassicaceae (Cruciferae) where the replum forms a frame to which the seeds are attached when the two outer valves fall away,

resin (resinous) - a solid or semisolid substance produced by plants from special canals or ducts found within the plant, not soluble in water and often sticky and aromatic

reticulate - with many interconnections as in a net. applied to a surface being marked by a network o f fine lines or ridges

reticulate venation - see net-venation. retrorse - bent abruptly backwards or downwards. retuse - With a rounded tip that has a small notch at the centre revolute - with the margin or tip rolled or bent backwards to­ wards the centre.

rhachis - synonymous with rachis. rheophyte - plant growing in flowing water.

rhizome - a root-like stem on or beneath the ground with roots growing downwards and leaves and shoots upwards, differ­ ing from a true root in the presence o f buds, leaves, or scales; especially robust rhizomes are often called rootstocks.

rib - a primary vein or prominently raised vein or nerve. riparian - growing in or at the edge o f rivers and streams root - the underground portion o f the plant that never produces leaves and thus lacks nodes and intemodes; with a special anatomy.

root cap - a cap of tissue over the root apex. root hair - a hair-like outgrowth o f an epidermal cell which ab­ sorbs water and minerals, found on young actively growing roots.

rootstock - a short, vertical, underground stem, bearing roots. rosette - a cluster o f parts in a circular form; often used for leaves produced at ground level.

rostrate - with a beak or beak-like projection, rosulate - when the leaves are in a circle or in a rosette, rotate - a calyx or corolla, or other organ, with a very short tube and spreading lobes,

rotund - with a shape between orbicular and broadly elliptic, rudimentary - incompletely developed, rufous - rusty or brownish red. rugose - with wrinkles or grooves on the surface, ruminate - very uneven, looking as chewed. runcinate - a margin in which the lobes or teeth point back­ wards towards the base.

runner - a slender stem running along the ground and rooting at the nodes, soc also stolon.

rupturing - breaking open irregularly. saccate - bag-shaped or sack-shapcd, pouched. sack (sac) - a pouch or bag-1 ike structure. sagittate - arrow shaped; the base has two acute lobes that point backwards to the base o f the petiole.

samara - a one-seeded nut-like fruit with a wing, saponins - a toxic, soap-like group o f compounds which is present in many plants.

saprophyte - a plant that obtains its nourishment from dead or­ ganic matter and usually docs not possess chlorophyll

sarcocarp - a fleshy covering to a fruit sarmentose - with long slender runners. scaberulous - minutely scabrous. scabrid (scabrous) - rough to touch, usually caused by the presence of very short stiff hairs which point backwards to the line o f growth.

scale - any small flat and thin structure like a flattened hair or very' small leaf, often triangular in shape.

scale leaf - small, usually adpressed as in Erica (Ericaceac) or much reduced as in Tamarix (Tamaricaceae).

scandent - a general term for climbing. scape - a flower stalk (a pcdicel or. more often, a peduncle) without leaves that arises from the ground, as in Crinum (Amaryllidaceae).

scarious (scariose) - thin and dry. not green.

GLOSSARY OF BOTANICAL TERMS

SCI i/o c a r p - a dry fruit breaking up into one-seeded nut-like parts (each part called a mericarp), as in many genera in the Malvaceae and in Apiaceae (Umbelliferae). sci id io id a l - a compressed and impoverished panicle resulting in a pseudo-umbellate appearance.

SCt rpioid - a cymose inflorescence curvcd to one side and boiled like the tail on a scorpion; see also helicoid cyme.

scram b ler - a plant which usually climbs with the help o f iiooks. thorns or tendrils so that it spreads out over or through its support.

scrctiform - pouch-shaped.

305

silique (siliqua) - the long slender fruit o f some Brassicaceae (Cruciferae), more than two times as long as broad; divided into two compartments by a thin partition on which the seeds are borne and from which the two outer valves separate.

simple fruit - derived from a single carpel or syncarpous ovary.

simple leaf - with only a single blade, the opposite o f a com­ pound leaf.

sinuate (sinuose) - when the margin is uneven or wavy by turning inwards or outwards but not deeply enough to be lobed.

Scru b - more or less dense vegetation of small shrubs (up to j-bout 3 m high).

sinus - a rounded depression between two projecting lobes or

sec.I nd - one-sided; as when branches, leaves or flowers are all

siphon (siphonaceous) - a tube (tubular), soboliferous - bearing vigorous shoots, solitary - one alone and without companions or similar struc­

attached along one side o f an axis.

see J - the ripened ovule containing the embryo with or without additional storage tissue. S eed lin g - the young plant which develops from a germinated -ted.

segm ent - a division or part o f an organ, s e n i - - a prefix meaning 'h a lf .

sen* i-parasite - a plant which can parasitize another plant but s also capable o f growing by itself. S ensu - Latin meaning ‘in the sense o f .

sepal - a flat part o f the outer whorl o f sterile appendages that j 'otects the flower in bud, often green or dull in colour; the •:pals together are called the calyx.

sepsiloid - a structure which is similar to a sepal. sepi I - partitions; plural o f septum. Sept ute - divided by one or more partitions or septa. sept cidal - opening at or along the partition (or septum) or ationum botanicarum iconibus Op. Bot - Opera Botanica op cit. - 'opera citato’ Latin for ‘in the work already cited’ Opera Bot. Belg. - Opera botanica belgica orth. var. - orthographic variant Osterr. Bot. Zeitschr. - Osterreichische Botanische Zeitschrift, Wien

311

OXF - Herbarium, Department of Botany, University of Oxford, England P - Museum National d’Histoire Naturelle, Laboratoire de Phanerogamie, Paris, France p. - page pp. - pages p.p. - ‘pro parte’ Latin ‘for in part’; used to show that only a part of the group as circumscribed by the original au­ thor is being used by the later author PAL - Erbario Siculo and Erbario Generale, Palermo, It­ aly PAT - Laboratoire d ’Ethnobotanique, Museum National d ’Histoire Naturelle, Paris, France P-LA - Lamarck Collection, Paris, France PI. - plate Planzenfam. - Die naturlichen Pflanzenfamilien Planzenw. Ost. Afr. - Die Pflanzenwelt Afrikas (J), Ostafrica zwischen Abbaja-See undKenia PI. Bequaert. - Plantae Bequaertianae PI. Bog. - FJorula Bogosensis PI. Syst. Evol. - Plant systematics and evolution P.O.A.C. - Die Pflanzenwelt Ost-afrikas, C PRE - National Herbarium, Botanical Research Institute, Pretoria, South Africa Prodr, - ‘prodromus’ Latin for ‘forerunner’, a preliminary work which should be followed by a more complete one Prodr. - Prodromus systematis naturalis regni vegetabilis of De Candolle quoad nom. excl. specim. cit. - ‘quoad nomen exclusive specimen citatum (specimina citata)’ Latin for ‘con­ cerning the name but not the specimen quoted’ R. - river R - Herbario, Departamento de Botanica, Museu Nacional, Universidade Federal do Rio de Janeiro Racc. Bot. Miss. Consol, Kenya - Raccolta botaniche (embryophyta diploidalia) fatta dai missionari della consolata nel Kenya Rar. PI. Afr. - J. Burmanni rariarum Africanarum plantarum Decas Rel. Monogr. Agr.-col. - Relazioni e monografie agrario-coloniali, Firenze. Result. Sci. Miss. Stef-paoli Coll. Bot. - Resultati Scientifici della Missione Stefanini-Paoli nella Soma­ lia ltaliana Rev. Bot. Appl. Agric. Trop. - Revue international de botanique appliquee et d ’agriculture tropicale Rev. Gen. PI. - Revisio generum plantarum RO - Erbario dell’Istituto Botanico dell’Universita di Roma, Roma. Italy S - Herbarium, Swedish Museum of Natural History (Naturhistoriska riksmuseet), Stockholm, Sweden S - south S. Afr. J. /Joum. Bot. - South African Journal o f Botany Sect. - Section of a genus s. lat. - 'sensu lato' Latin for 'in a broad sense’ s.n. - ‘sine numero’ Latin for ‘without a number’ SD - Sidamo region, Ethiopia SE - southeast

I

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ABBREVIATIONS

sect. - section sensu - Latin for in the sense o f ’ SINET: Eth. J. Sci. - SINET: An Ethiopian Journal o f Sci­ ence sp. - species (singular) spp. - species (plural) SRGH - National Herbarium and Botanic Garden. Cause­ way, Harare. Zimbabwe stat. nov. - status novus’ Latin for 'new rank' Stap. und Klein. - Stapelieen und Kleinien STR - Institut de Botamque de 1’Universite Louis Pasteur, Strasbourg, France SU - Shewa region, above and to the west of the 1000 m contour, Ethiopia subgen. - subgenus subsp. - subspecies (singular) subspp. - subspecies (plural) Suppl. - supplement Svensk. Bot. Tidskr. - Svensk Botanisk Tidskrift SW - southwest Symb. Bot. Upsal. —Symbolae Botanicae Upsaliensis syn. - sy nonym t./tab. - tabula’ Latin for ‘illustration’ TCD - Herbarium. School of Botany , Trinity College, Dublin. Ireland Tent. FI. Abyss. - Tentamen Florae Abyssinicae TO - Herbarium, Museum Botanicum Horti Taurinensis. c/o Istituto ed Orto Botanico dell’ Universita, Torino. Italy Trans. Linn. Soc. (Lond.) - Transactions o f the Linnaean Society o f London

Trop. - tropical TU - Tigray region, Ethiopia TUB - Herbarium, Institut fiir Biologie I. Tubingen, Ger­ many Univ. Calif. Publ. Bot. - University o f California Publica­ tions in Botany UPS - The Herbarium, University of Uppsala, Uppsala. Sweden US - United States National Herbarium, Smithsonian In­ stitution, Washington, U.S.A. var - variety Verh. Kon. Ned. Akad. Wetensch. Afd. Natuurk. Verhandelingen der koninklijke nederlandsche akademie van wetenschappen: afdeeling natuurkunde, Amsterdam vol - volume of a published work which appears in more than one part W - Naturhistorisches Museum, Wien, Austria W - west WAG - Herbarium Vadense, Laboratory for Plant Taxon­ omy and Plant Geography, Netherlands Wag Agnc. Univ Papers - Wagenmgen Agricultural University Papers WG - Welega region, Ethiopia WIR - Herbarium, The All-Union Institute of Plant Indus­ try, St Petersberg (Leningrad), Russia WU - Welo region. Ethiopia WU - Herbarium Inst, fiir Botanik, Universitat Wien. Austria Z - Institut fiir systematische Botanik der Universitai Zurich, Zurich, Switzerland ZSS - Herbarium Stadtische Sukkulentensammiung. Zurich. Switzerland

INDEX TO SCIENTIFIC NAMES *COKANTHERAG. D on. 87 Acokanthera abyssinica K. Schum., 87 Acokanthera deflersii Schweinf. ex Lewin. 88 var. africana Markgr.. 88 acokanthera schimperi (A. DC.) Schw einf.. 87.8 8 •\DENIUM Roem. & Schult.. 88 Adenium coelaneum Stapf. 88 .Adenium honghel A. DC., 88 Adenium obesum (Forssk.) Roem. & Schult.. 88. 89 mdeniurn sotttalense Ball', f . 88 var. caudatipelalum Chiov.. 88 var. crispum Chiov.. 88 Adenium speciosum Fenzl. 88 . 1denium iricholepis Chiov.. 88 . \dina galpinii Oliv.. 240 . Idina lasiantha K. Schum.. 240 var. parviflora Hochr.. 240 Adina microcephala (Del.) Hiem. 240 var galpinii (Oliv.) Hiern. 240 Adina ruhrostipulata K. Schum.. 242 var discolor Chiov., 242 Afrocoffea Bridson. 268 Afrosison gallabatensis Schweinf. cx Wolff. 2 AGARISTA D. Don ex G. Don. 46. 47 AGATHISANTHEMUM Klotzsch. 215 Agarista salicifolia (Comm, ex Lam.) Hook. f . 48 Agathisanthemum globosum (/I. Rich.) Bremek.. 215. 216 var subglabrum Bremek.. 215 Agauria (DC.). Hook.f.. 4 6 .4 7 Agauria salicifolia (Comm, ex Lam.) Hook.f. cx. Oliv.. 48 AGROCHARIS Hochst.. 11 Agrocharis incognita (Norman) Heyw. & Jury. 12. 13 Agrocharis m elanantha Hochst.. 13 \g ro ch aris pedunculata (Bak. f ) Heyw. & Jury, 12 ALEPIDEA De la Roche. 7 1lepidea congesta Schlecht. & H. W olff. 7 Hep idea longifolia E. Mcv. cx Steud.. 7 Alepidea longifolia Diimmer.. 7 subsp coarclata (Diimmer) Weim., 7 subsp propinqua (Diimmer) W eim.. 7 subsp swynnertonii (Diimmer) W cim ..7 Alepidea peduncularis Steud. ex A. Rich.. 7. 8 var fischeri Engl.. 7 Alepidea propinqua Diimmer. 7 ALSTONIA R. Br.. 88 Alstonia boonei De Wild.. 89 AMMI L.. 25 Ammi copticum L.. 20 Ammi majus L.. 25. 26 Ammi pauciradialum Hochst. ex A. Rich.. 25 Ammi visnaga (L.) Lam ..2 5 Andromeda salicifolia Comtn. cx Lam.. 48 ANETHUM L.. 35 Anethum foeniculum L., 34 Anethum graveolens L.. 35. 36 Anethum sowa Fleming, 35 Angolluma R. Munster, 185 Angolluma abayensis (M.G. Gilbert) Plowes. 187 Angolluma baldratii (W hite & Sloane) Plowes. 187

Angolluma circes (M.G. Gilbert) Plowes, 187 Angolluma foetida (M.G. Gilbert) Plowes, 186 Angolluma gemugofana (M.G. Gilbert) Plowes, 186 Angolluma gilbertii Plowes, 186 Angnlluma laticorona (M.G. Gilbert) Plowes, 186 Angolluma ogadensis (M.G. Gilbert) Plowes, 186 Angolluma sacculata (N.E. Br.) Plowes, 189 Angolluma sprengeri (N.E. Br.) Plowes, 185 “Angolluma’'sudanica Plowes, 187 Angolluma vihratilis (B ruce & Bally) Plowes, 190 Aningeria Aubrev. & Pellegr.. 60 Aningeria adolfi-friederici (Engl.) Robyns & Gilbert, 61 subsp. adolfi-friedcrici, 61 subsp. australis J.H. I lemsl., 61 subsp. floecosa J.H. Hcmsl.. 61 subsp. kcniensis (R.E. Fries) J.H. Hemsl., 61 subsp. usambarensis J.H. Hemsl.. 61 Aningeria altissima (A. Chev.) Aubrev. & Pellegr., 63 Annesorhiza abyssinica A. Br., 18 ANTHOCLEISTA Afzel. ex R. B r . 70 Anthocleista niamniamensis Gilg, 70 Anthocleista schweinfurthii G ilg, 70, 71 ANTHOSPERMEAE Cham. & Schlechtd.. 231 ANTHOSPERMUM L., 231 Anthospermurn ferrugineum Eckl. & Zeyh., 232 Anthospermurn hedyotideum Sond., 232 Anthospermurn herbaccum L .f, 232. 233 var. villosicarpum Verde., 234 Anthospermurn hirsutum sensu A. Rich, non DC., 233 Anthospermurn lanceolatum Thunb.. 232 Anthospermurn m ildbraedii K. Krause. 232 Anthospermurn muriculalum Hochst. ex A. Rich.. 232 Anthospermurn pachyrrhizum H iem , 233 Anthospermurn unisetum Hochst., 233 Anthospermurn villosicarpum (Verde.) Puff. 234 ANTHRISCUS Pers., 14 Anthriscus africana Hook.f., 25 Anthriscus sylvestris (L.) H o f f m 14 var. abyssinica A . Rich., 14 var. nemorosa (M. Bieb.) Trautv., 15 var. sylvestris. 14 ANTIRHEOIDEAE R a f. 268 Antura (without epithet) Forssk., 90

APIACEAE. 1 APIOIDEAE. I APIUM L.. 20

Apium graveolens L.. 20 var. dulce (Miller) DC.. 20 var. rapaceum (M iller) DC., 20 Apium nodiflorum (L.) Lag., 20, 21 APOCYNACEAE. 87 Apteranthes tessellata Decne.. 179 Arbulocarpus Tennant. 224 Arbulocarpus erythraeae (Lanza) Cufod., 228 Arbulocarpus sphaerostigma (Hochst.) Tennant, 228 A S C L E P IA D A C E A E , 99 ASCLEPIADOIDEAE. 100, 115 Asclepias abyssinica (Decne.) N.E. Br.. 124 Asclepias albida N.E. Br.. 124

3

314

INDEX TO SCIENTIFIC NAMES

Asclepias angustata (K. Schum.) N.E. Br., 131 Asclepias coarctata S. Moore. 121 Asclepias cordata Forssk.. 134 Asclepias crinita (G. Bcrtol.) N E. B r. 123 Asclepias daemia Forssk.. 133 Asclepias denticulata Schltr., 123 A sclepiasflavida N.E. Br. 124 Asclepiasfluvial ilis A . Chcv.. 121 Asclepias forskaolii Schult.. 138 Asclepias frutieosa L.. 123 Asclepias geminata Roxb.. 153 Asclepias gigantea, 119 Asclepias glaberrima (Oliv.) Schltr.. 121 Asclepias glabra Miller. 123 Asclepias integra N.E. Fir.. 126 Asclepias laniflora Forssk.. 121 . Isclepias laniflora Del. non Forssk.. 121 Asclepias laurenliana (Dewevrc) N.E. Br., 131 Asclepias leuchotricha Schltr.. 131 Asclepias leucocarpa Schltr.. 126 Asclepias lineolata sensu Hutch. & Dalziel non Decne., 126 Asclepias lilocarpa Chiov.. 126 Asclepias macrantha sensu Bullock, non Hochst ex Oliv . 130 Asclepias macrantha Hochst. ex Oliv.. 130 Asclepias negri Chiov.. 126 Asclepias nivalis Forssk.. 138 Asclepias pedunculata (Decnc.) Dandy, 130 Asclepias phillipsiae N.E. Br.. 124 Asclepias procera Ait.. 119 Asclepiaspubiseta N.E. Br.. 124 Asclepias rhacodes (K Schum.) N.E. Br.. 130 Asclepias riva Iis S Moore. 121 Asclepias robusla (A. Rich.) N.E. Br.. 127 Asclepias salicifolia Salisb.. 123 Asclepias semiltinala (A Rich ) N.E. Br.. 123 Asclepias selosa Forssk.. 124 . tsclepias spiralis Forssk . 138 Asclepias stenophylla A Gray. 126 Asclepir.sstipilacea Forssk . 148 Asclepias undulata L.. 131 Asclepias trvirensis S. Moore. 130 Asclepias welwitscliii (Britt. & Rendle) Britt. & Rendle ex N.E. Br.j 131 ASPIDOGLOSSUM E. M ey.. 1 17 Aspidoglossum elliotii (Schltr.) Kupicha, 1 19 Aspidoglossum inten upturn (E. Mey.) Bullock. 118. I 19 Aspidoglossum kulsii Cufod.. 117 Aspidoglossum masaicum (N.E. Br.) Kupicha. 117 Aspidoglossum sp. J de Wilde 4921. I 19 Astephanus schimperi Vatke. 137 Bassia parkii G. Don. 57 BERULA K o ch . 3 1 Berula crecta (Hudson) Coville. 31. 32 Berula thunbergii (DC.) W olff. 31 Bidaria (Endl.) Dccne . 153 Binectaria laurifolia Forssk . 55 Blaeria condensata 1lochst e \ A Rich., non Erica condensala Benth . 47 Blaeria guguensis PicSerm . & I lein.. 47 Blaeria L.. 46 Blaeria spicata Hochst ex A Rich . non Erica spicala Thunb . 47 Blaeria tenuipilosa Engl ex Aim & Fries. 47 BLYTTIA A m .. 138

Blyttia arabica Arn., 138 Blyttia fruticulosum (Decne.) D. V Field. 138. NO Blyttia spiralis (Forssk.) D.V. F ield & J.R.I. Wood. 138, 140 Borreria G.F.W. Mey., 224 Borreria chaetocephala (DC.) Flcppcr. 225 Borreria compacta (Hiem) K. Schum., 225 Borreria hebecarpa Hochst. ex A. Rich., 225 '*Borreria neglecta A Rich., 225 Borreria paludosa I lepper. 225 Borreria prm ceae K. Schum.. 227 Borreria pus ilia (W all.) D C . 225 Borreria quadrisulcata Bremek.. 225 Borreria repens DC,, 226 Borreria rhodesica Suesseng.. 228 Borreria ruelliae sensu auctt. mult., non (DC.) FI. Thoms.. 228 Bfjrreria scabra sensu Cufod. p.p. non (K. Schum. & Thonn ) K Schum.. 228 'Borreria somatic a K. Schum., 228 Borreria squarrosa Schinz. 228 Borreria stachydea (I )C.) Hutch & Dalz.. 228 Borreria slolzii K Krause. 228 Borreria ienuiflora Chiov.. 224 Boucerosia Wight & Am., 170 Boucerosia acutangula (Decne.) Decne.. 174 Boucerosia cylindrnu Brongn., 179 Boucerosia edulis Edgew.. 174 Boucerosia penicillata Defiers. 176 Boucerosia russelumu Brongn.. 174 Boucerosiasocotrana Balf. f , 176 Boucerosia stocksiana Boiss., 174 Boucerosia lombuctuensis A. Cheval.. 174 B RA C H Y STELM A R. Br . 169 Brachystelma. 160 Brachystelma asmarense Chiov.. 169 Brachystelma elhpnca A Rich.. 169 Brachystelma johnstomi N.E. Br . 169 Brachystelma lineare A. Rich.. 169 Brachystelma s p M & S. Gilbert 2 147. 169 BREONADIA R id sd . 239 Breonadia microcepliala (Del.) Ridsdalc. 240 Breonadia salicina W ahl) Hepper A Wood. 240 var galpinii (Oliv.) Hepper & Wood. 240 BUCKOLLIA Venter & R.L. Verh.. 102 Buckoliia tomentosa (E.A Bruce) I enter & R.L. Verh.. 103. l()-i Buckollia volubilis ^Schltr.) Venter & R.L. Verh . 103. / 0-4 BUDDLEJA L., 70 Buddleja cordata fl.B K . 1 1. 72 Buddleja davidii Francli., 72 Buddleja madagascariensis Lam . 72 Buddleja polystachya Fresen.. 73 var parvifolia Marquand. 73 Buddleja rufa Fresen., 73 Buddleja saltiana Steud.. 73 BUPLEURUM L.. 19 Bupleurum lancifolium Hornem.. 19. 20 Bupleurum subovatum Link, 20 var heterophyllum (Link) Wolff. 20 Butvrospermum Kotschy. 55. 57 Butyrospermum niloficum Kotschy. 57 Butvrospermum paradoxum (Gaertn. f ) I lepper. 57 subsp nilolicum (Kotschy) Hepper. 57 Butyrospermum parkii (G. Don) Kotschy. 5n var nilolicum (Kotschy) Engl.. 57

INDEX TO SCIENTIFIC NAMES

ubsp. niloticum (Kotschy) J.H. Hemsl.. 57 Cat iirys abyssinica Hochst. ex A. Rich.. 33 CA LOTROPIS R .B r.. 119 Caloptropis inflexa Chiov., 1 19 Calotropis procera (Ait.) A i t . f . 1 19. 120 CamUropissyriaca Woods.. 1 19 Cat panolea mildbraedii Gilg & Schellenb.. 81 CA '‘THIUM Lam., 270 Canthium sensu auctt. div. p.p.. non Lam.. 271. 273. 274 Canihium afzelianum Hiern. 274 Ca< 'iiium bogosense (Martel li) Penzig, 274 Ca* ' iium charadophilum (K. Krause) Bullock. 272 C an’ lium cienkowskii (Schweinf.) Roberty. 275 Canihium crassum sensu Chiov. non Schweinf. ex Hiern. 270 Ca thium edule (Vahl) Baill.. 279 Ca thium gentilii (De Wild.) Evrard, 273 Ca thium giordanii Chiov.. 274 Canthium golungense Hiern. 274 var. parviflorum S. Moore. 274 Canthium gueinzii Sond.. 272 Ca ithium hispidum sensu Cufod. non Benth.. 272 Ca t thium lactescens Hiern. 270. 27] Ca tbium lucidum R. Br.. 274 Canthium maleolens Chiov.. 279 Ca ithium myrtifolium S. Moore. 274 Ca ithium neglectum Hiern. 278 Ca ithium oligocarpum Hiem . 270 ubsp. angustifolium Bridson. 271 ■ubsp. capium (Bullock) Bridson. 271 •ubsp. friesiorum Bridson. 271 subsp. intermedium Bridson. 271 ubsp. oligocarpum. 270. 271 Ca Kthium parviflorum Lam. p.p.. 274 Cti nhium pin llanthoideum Baill.. 274 Ct lithium pseudosetiflorum Bridson. 2 7 1 Cc ithium ruwenzoriense Bullock. 270 Cc uhium scabrosum Bullock. 272 Cc 'ithium schimperianum A. Rich.. 274 CMlthium setijlorum sensu Cufod. non Hiern.. 271 Ct anchum fruticulosum Decne., 138 Cyitanchum gerrardii (Harv.) Liede. 145 CyHiinchum gonoloboides Schltr.. 142 Cynamehum hastaturn Pers.. 156 Cy/ anchum hastifolium K. Schum.. 147 Cynanchum hastifolium N.E. Br., 147 Cyn ; nchum heteromorphum Vatke. 147 Cyt anchum heierophyllum Del.. 155 Cyt anchum holstii (K. Schum.) K. Schum.. 146 Cynanchum lanceolalum Poir.. 156 Cyt anchum lancifolium Schum. & Thonn., 156 Cyi nchum Icnewtonii Liede. 145, 146 Cynanchum mensense Schweinf. ex K. Schum.. 146 Cynanchum pleianthum K. Schum.. 147 Cyti mchum pyrotechnicum Forssk., 155 Cyt anchum roseum Chiov.. 135 Cyt mchum sarcostemmatoides K. Schum., 145 Cyt inchum sarcostemmoides K. Schum., 145 Cyt nchum scabrum Schum. & Thonn., 156 Cyt nchum schimperi Hochst.. 151 Gyn nchum senegalense Sieb. ex Decne.. 153 Cyt. mchum somaliense (N.E. Br.) N.E. Br., 142 Cyt nchum subvolubile Schum. &. Thonn., 153 Cyh mchum tetrapterum sensu auctt. non (Turcz.) R.A. Dy er, 145 ( V’ mchum trifurcatum Schltr., 142 Cyt nchum validum N.E. fir. non Cynanchum alatum Wight • Am ., 136 D ai'm a aethiopica Decne., 133 Dai 'ina angolensis Decne., 133

317

Daemia cordata (Forssk.) R. Br. ex Schult., 134 Daemia cordifolia (Retz.) K. Schum., 133 Daemia extensa (Jacq.) R. Br. ex Schult., 133 D aem iaforskalii Schult., 133 Daemia iticana Decne., 134 Daemia tomentosa (L.) Pomel, 134 DAUCUS L., 10 D aucus abyssinicus C.A. Mey., 11 Daucus abyssinicus Hochst. ex A. Rich, non C.A. Mey.. 11 Daucus carota L.. 11,1 2 var. abyssinica A. Braun, 11 Daucus hochstetteri Braun ex Drude. 1 I Daucus melananthos Steud.. 13 Daucus visnaga L., 25 D ecam erisjovis-tonantis Welw., 253 Deringa A dans., 25 Desmidorchis Ehrenb., 170 Desmidorchis acutangula Decne., 174 Desmidorchis edithae (N.E. Br.) Plowes. 176 Desmidorchis penicillata (Defiers) Plowes, 176 Desmidorchis retrospiciens Ehrenb., 174 Desmidorchis speciosa (N.E. Br.) Plowes, 176 DIODIA L.. 223 Diodia benguellensis Hiern, 228 Diodia breviseta Benth., 223 Diodia pilosa Schumach. & Thonn., 223 Diodia sarmentosa S>v., 223 Diodia scattdens sensu Cufod. non Sw.. 223 Diodia senensis Klotzsch, 228 Diodia stipulosa S. Moore, 227 DIOSPYROS L., 49 Diospyros abyssinica (Hiern) F. White, 50, 52 subsp. abyssinica, 50 subsp. chapmaniorum F. White, 50 Diospyros kaki L., 49 Diospyros mespiiiformis Hochst. ex A. DC.. 5 0 .5 2 Diospyros scabra (Chiov.) Cufod., 51. 52 DIPLOLOPHIUM Turcz.. 32 Diplolophium abyssinicum (Hochst. ex A. Rich.) Benth., 33 Diplolophium africanum Turcz.. 33. 34 DIPLOSPORINEAE M iq., 257 DIPLOSTIGMA K. Schum ., 139 Diplostigma canescens K. Schum., 139, 141 DIPSACACEAE, 286 DIPSACUS L.. 286 Dipsacus acaulis (Steud. ex A. Rich.) Napper, 286 Dipsacus appendiculatus Steud. ex A. Rich.. 286 Dipsacus bequaertii De Wild. 286 Dipsacus eremocephalus Pich.Serm., 286 Dipsacus kigesiensis Good, 286 Dipsacus pinnatifidus Steud. ex A. Rich.. 286, 287 var. integrifolius Engl., 286 Dipsacus schimperi A. Br., 286 Dipsacus setosus Hiern, 286 Dirichletia Klotzsch. 207 Dirichletia asperula K. Schum.. 208 Dirichletia borziana Mattei. 208 Dirichletia ellenbeckii K. Schum .. 208 Dirichletia giumbensis Chiov.. 208 Dirichletia glaucescens H iem. 207 Dirichletia macranlha K. Schum., 208 Dirichletia paolii Chiov., 208 DREGEA E .M ey.. 151

318

INDEX TO SCIENTIFIC NAMES

Dregea abyssinica (Hochst.) K. Schum., 151 Dregea africana (Decne.) Martelli, 151 Dregea rubicunda Fliern non K. Schum., 151 Dregea rubicunda K. Schum., 151,1 5 2 Dregea schimperi (Decne.) Bullock, 151 Dregea stelostigma (K. Schum.) Bullock. 152 Durieua abyssinica Boiss. & Reuter, 11 DU VALIA Haw., 190 Duvalia somalensis Lavranos, 190 Duvalia sulcata N.E. Br., 190 subsp. seminuda (Lavranos) Meve. 190 subsp. somalensis (Lavranos) M eve, 190 subsp. sulcata, 190 Duvaucellia tenuis S. Bowd.. 209 EBENACEAE. 49 ECHIDNOPSIS HOOK. F ., 178 Echidnopsis cereiformis H o o k .f, 179. 180 var. brunnea Berger, 179 var. obscure Berger, 179 Echidnopsis chrysantha Lavranos, 180 Echidnopsis dam m anniana Sprenger, 179, 180 Echidnopsis golathi Schweinf. ex Defiers, 176 Echidnopsis jacksonii Plowes, 181. 183 Echidnopsis lavraniana Plowes, 181 Echidnopsis malum (Lavranos) Bruyns, 182 Echidnopsis montana (Dyer & Bruce) Bally, 179, 181 Echidnopsis nubica N.E. B r , 179 Echidnopsis planiflora Bally, 179, 182 Echidnopsis scutcllata (Defiers) Berger, 180 subsp. planiflora (Bally) Bruyns. 179 Echidnopsis sharpei White & Sloane, 181, 182 Echidnopsis somalensis N.E. Br., 179 Echidnopsis lessellata (Decne.) K. Schum., 179 Echidnopsis urceolata B ally , 181, 183 Edgeworthia Falc. non Meisncr. 58 Edgeworthia buxifolia Falc., 60 EDITHCOLEA N.E. Br.. 184 Edithcolea grandis N.E. Br., 184 EMBELIA Burnt, / . , 64, 68 Embelia schimperi Vatke, 68,6 9 ERICA L„ 46 Erica acrophya Fresen.. 46 Erica arborea L., 46, 47 Erica tenuipilosa (jEngl, ex Aim & Fries) Cheek. A l subsp. spicata (Hochst. A. Rich.) Cheek, 47 subsp tenuipilosa, 47 Erica trim era (Engl.) Beentje, 46. 47 ERICACEAE. 46 Eryngium pedunculare (A. Rich.) Koso-Poljansky. 7 ERYTHROSELINUM Chiov., 43 Erythroselinum atropurpureum (Hochst. ex .4. Rich.) Chiov., 43, 44 EUCLEA M urr , 49, 51 Euclea divinorum H iem , 52 Euclea divinorum Hiem., 52 subsp. kenyensis (Fries) Dale & Greenway. 52 Euclea kellau Hochst.. 51 Euclea kenyensis Fries, 52 Euclea lanceolata auct., non E. Mey. ex A. IX)., 52 Euclea latidens Stapf, 51 Euclea microcarpa Giirke, 51 Euclea neghellensis Cufod., 51 Euclea racemosa M urr., 51

subsp. raccmosa, 52 subsp. schimperi (A. DC.) White, 51, 52 Euclea schimperi (A. DC.) Dandy, 51 Euphorbia vimmalis L., 148 FA DOG IA S ch w ein f, 275 Fadogia cienkowskii S ch w ein f, 275, 276 var. cienkowskii, 276 var. lanceolata Robyns, 216 Fadogia kalangensis De W ild., 275 Fadogia tristis (K. Schum.) Robyns, 275 FERETIA Del., 260 Feretia ?canthioides Hiem, 260 Feretia apodanthera Del., 260 subsp. apodanthera. 260 subsp. keniensis Bridson. 260 subsp. tanzaniensis Bridson, 260 FERULA L., 37 Ferula abyssinica Hochst. ex A. Rich., 37 Ferula communis L., 37, 39 Ferula erythraeae Schweinf. ex Penzig, 37 FOENICULUM M iller, 33 Foeniculum capillaceum Gilib., 34 Foeniculum officinale All., 34 Foeniculum vulgare. M iller, 34, 35 Franchetella arborescens (Spreng.) Kuntze, 18 var. abyssinica (A. Rich.) K untze, 18 GALINIERA D el., 258 Galiniera coffeoides Del., 259 Galiniera saxifraga (Hochst.) Bridson, 258, 259 GALIUM L .,2 3 5 Galium abyssinicum Chiov., 237 Galium acrophyum Hochst. ex Chiov., 237 Galium apanne L., 236 var. hamatum (A. Rich.) Hook.f., 236 Galium aparinoides Forssk., 236, 238 Galium aparinoides sensu Cufod. non Forssk., 237 Galium biafrae Fliem, 236 Galium boreo-aethiopicum Puff, 238 Galium chloroionanthum K. Schum ., 236 Galium dasycarpum Schweinf., 236 Galium hamatum A. Rich., 236 Galium hochstetteri Pic.Serm., 237 Galium homblei De Wild.. 237 Galium latum De Wild., 237 Galium mollugo sensu Cufod. non L., 238 Galium natalense Rouy in Rouy & Foucard, 236 Galium rotundifolium L., 236 var. hirsutum Sond., 236 Galium ruwenzoriense (Cort.) Chiov., 237 Galium scabrellum sensu Cufod. non K. Schum., 237 Galium scioanum Chiov., 237 var. glahrum Brenan, 237, 238 var. latum (De Wild.) Verde., 237 var. latum, 238 var. scioanum, 238 Galium sidamense Chiov., 237 Galium simense Fresen., 237, 238 Galium simense A . Rich, non Fresen., 237 Galium spurium L., 237 subsp. africanum Verde., 237, 238 subsp. spurium, 237 Galium thunbergianum Eckl. & Zeyh., 236, 238 var. hirsutum (Sond.) Verde., 236

INDEX TO SCIENTIFIC NAMES

var. th u n b erg ian u m , 236 Gah um verum sensu Cufod. non L.. 238 G ARDENIA Ellis, 252 Gar t’nia dumetorum sensu A. Rich, non Retz., 252 Gar ienia fiorii Chiov., 253

Gan ;nia goetzei Stapf & Hutch., 253 Gar ieniajovis-ionantis (Welw.) Hiern, 253 Gar ienia longistyla (DC.) Hook., 252 Gardenia lutea Fresen., 253 Gar ienia paleacea A. Rich., 252 Gar ienia riparia K. Schum., 255 Gar ienia somalensis Chiov., 253 var. tubicalyx Chiov.. 253 Gar ienia stupinicarpa Chiov., 253 Gar Ienia ternifolia Schumach. sp. jovis-tonantis ( W elw .) Verde., 253 var. goet'ei (Stapf & Hutch.) Verde., 253 var. jovis-tonantis, 253 s >sp ternifolia. 253 Gar Ienia tigrina Hiern, 255 Gar ienia urcelliformis Hiern, 255 G at Ienia volkensii K. Schum., 253 var somalensis (Chiov.) Cufod., 253 subsp. spathuli folia (Stapf & Hutch.) Verde.. 253 si t>sp. volkensii Verde., 253 var. saundersiae (N.E. Br.), 253 var volkensii, 253, 254 G A R D E N IE A E D C , 250 G A R D EN IIN A E DC’., 251 GE i PHI LAD. Don, 201 Geophila reniformis D. Don, 202 Get phila repens (L.) /. M. Johnston, 201, 202 GeMahila uniflora Hiem, 202 Gilgui Candida Pax. 1 15 G LO SSON EM A Decne . 1 15 Glakronema affine N.E. Br.. ! 17 G la so n e m a hoveanum (Decne.) Decne., 1 17 Glaktonema echinatum Hochst. ex Di Capua, 1 17 GlaH ' onema elliotii Schltr., 115 Glostonema erlangeri K. Schum., 1 17 Glo >onema hispidum Hutch. & Bruce. 117 Glo ' onema linearis (Fenzl) Decne., 157 GlojBKtnema ntacrosepalum Chiov., 1 15 G lotsonem a revoilii Tranche 115, 116 G ldtfonem a rivaei K. Schum.. 1 15 G lossonema 'h ru p p ii Oliv.. 1 17 G O M PH O C A R PU S R. Br., 121 Goi ip h o carp u s abyssinicus Decne.. 124 Gomphocarpus abyssinicits Hochst . 124 Gomphocarpus anguslalus Hochst., 131 Gomphocarpus bisacculatus Oliv., 126 Gomphocarpus brasiliensis Fourn.. 123 Gomphocarpus cornutus Decne., 123 Gomphocarpus crinilus G. Bertol., 123 Get iphocarpus drepanostephanus Hochst.. 127 GormUiocarpusfrutescens E. Mey.. 123 G om phocarpus fruticosus (L.) Ait. f , 123 var. angustissimus Engl., 124 var. purpureus Schweinf., 124 s.asp. flavidus (N.E. Br.) Goyder, 124 d.Dsp. setosus (Forssk.) Goyder, 124 ( to, phocarpus glaherrimus Oliv., 121 Go ii phocarpus integer (N.E. Br.) Bullock, 126

I

319

Gomphocarpus longipes Oliv., 130 Gomphocarpus pedunculatus Decne., 130 G om phocarpus pniilipsiae (N.E. Br.) Goyder, 124 G omphocarpusphysocarpus sensu Bullock p.p., 123 G om phocarpus p u rp u rascen s A. Rich., 124, 125 Gomphocarpus robustus A. Rich., 127 G om phocarpus sem ilunatus A. Rich., 123 Gomphocarpus setosus (Forssk.) N.E. Br. ex Schult., 124 G om phocarpus stenophylius Oliv. (1875), non Oliv. (1887), 126 Gomphocarpus verticillatus Turcz., 124 G O N G R O N EM A Decne., 153 G ongronem a angolense (N.E. Br.) Bullock, 153, 154 Grumilea bussei K. Schum. & K. Krause, 199 Grurnilea capensis (Eckl.) Sond., 199 Grumilea e/liottii K. Schum. & K. Krause, 199 Grumilea riparia K. Schum. & K. Krause, 199 G Y M N A N TH ER A E, 102 GYM NEM A R. Br.. 153 Gymnema subgen. Gongronema Endl., 153 Gymnem afruticulosum Hochst., 153 Gymnema humile Decne., 153 Gymnema macrocarpum A Rich.. 152 Gymnema m kenii Harv., 153 Gymnema par\’ifolium Oliv., 1 13 Gymnema rufescens Decne , 153 Gymnema subvolubile (Schum. & Thonn.) Decne., 153 Gymnema sylvestre (Retz.) R. Br. ex Schult., 153 Gymnosciadium pimpinelloides Hochst.. 28 Gymnosciadium pusillum Pich.Scrm., 28 HALLEA J.-F. Leroy, 241 Hailea rubrostipulata (K Schum.) J.-F. Leroy, 242, 243 HAPLOSCIADIUM Hochst., 16 Haplosciadium abyssinicum Hochst., 17 H ED Y O TID EA E Cham. A Schlechtd., 202 Hedyotis (Kohautia) abyssinia Hochst ex A. Rich., 21 1 Hedyotis aspera Heync ex Roth. 212 Hedyotis aspera sensu auctt. div.. non Heyne cx Roth., 211 Hedyotis caespitosa (Schnizl.) W alp., 210 Hedyotis corymbosa (L.) Lam., 220 Hedyotis dichotoma A. Rich, non Roth, 220 Hedyotis globosa Hochst. ex A. Rich., 215 Hedyotis goreensis DC.. 219 Hedyotis grandiflora (DC.) A. Rich., 209 Hedyotis herbacea L., 220 Hedyotis heynii (G. Don) Sond.. 220 Hedyotis lancifolia Schumach.. 219 var. brevipes (Bremek ) Fosberg. 219 Hedyotis linearis (DC.) Steud.. 221 Hedyotis macrophylla (DC.) A. Rich . 217 Hedyotis monanthos A Rich.. 219 Hedyotis pentandra Schumach. ssinicum Hochst. ex D C., 85 var amplifolium Fieri, 85 var. gratissimum (Defiers) Di Capua. 86 | ?Jasminum basilei Chiov., 84 i ?Jasminum bogosense Becc. ex M artelli, 84 Jasminum dichotomum Vahl. 84 Jasminum eminii auct., 84 Jasminum flonb u n d u m R.Br. ex Fresen., 86 forma decipiens Di Capua. 86 var. decipiens (Di Capua) Fiori. 86 var. steudneri (Schweinf. ex Bak.) Gilg & Schellenb., 86 Jasminum fluminense Veil., 86 subsp. fluminense. 86 subsp. gratissim um (Defiers) P.S.Green, 86 subsp. m auntianum sensu Cufod.. 86 Jasm inum gran d iflo ru m L., 86 subsp. floribundum (R.Br ex Fresen.) P.S.Green, 86

subsp. grandiflorum, 86

I

I

■ '

,

I i :

, i

Jasminum gratissimum Defiers, 86 Jasminum humile L.. 84 JasOminum humile sensu Engl, non L.. 86 Jasminum matinIdae Chiov.. 84 Jasminum officinale L., 84 Jasminum schimperi Vatke. 84 Jasminum st*n%Pax. 86 Jasminum steudneri Schweinf. ex Bak.. 86 Jasminum streptopus E M e y .. 84 Jasminum tetten&f sensu Martelli non Klotzch.. 86 KANAHIA R. Br.. 121 Kanahia carlsbergiana F ield et al., 121 Kanahia consimilis N.E. Br.. 121 Kanahia deli lei Kotschv ex Decne., 121 Kanahia forssk alii Decne.. 121 Kanahia glaberrima (Oliv.) N.E. Br., 121 Kanahia kanrtph Schult.. 121 Kanahia laniflora (Forssk.) R. Br., 121,1 2 2 Kanahia mortroi S. Moore, 121 Karamyschewia hedyotoides Fisch. & Mev.. 222 KEETIA E P. Phillips , 271 Keetia gueinzii (Sond.) Bridson, 212 Keetia transvaalensis E. P. Phillips p.p.. 272 K eetia zan zib arica (Klolzsch) Bridson, 273 subsp corm-licjiides (De W ild.) Bridson. 273 subsp. gentilii(D t' Wild.) Bridson. 272. 273 subsp zanzibarica. 273 Kellaua schimperi A . DC., 51 Knaxia (P entw nsfa) longituba F ranch., 282 KN O XIEA E H o o k.f. 281 K O H AU TIA O t^m . & Schlechtd.. 209 K OH AU TIA Chfim & Schlechtd subgen KOHAUTIA. 209 KOHAUTIA C/iIm. Schlechtd. subgen. PACHYSTIGM A. 209 Kohautia araZucafHochst. ex Boiss , 210 K ohautia a sp e r» (//e v « t’ ex Roth) Brem ek.. 212 Kohautia caespiitosa S ch n izi, 210 var. amaniensis (K. Krause) Bremek.. 210 var dclagoetisis sensu Cufod., 210 var. eritrevnsis Bremek.. 210 var hispidula Bremek.. 210 var kitalienlis Verde., 210

INDEX TO SCIENTIFIC NAMES

var. ramosior Bremek., 210 var. schimperi (C. PresI) Bremek., 210 Konautia caespitosa S chn izi, 210 subsp. amtf niensis (K. Krause) Aiantell ex Puff, 210 *jlbsp. brachvloba (Sond.) Mantel I. 210 ■ibsp. caespitosa. 210, 212 KonButia coccinea Royle, 211. 212 Kohautia gracillima Bremek., 210 Konautia grandiflora D C , 209 Kohautia lasiocarpa Klotzsch, 21 1 var. eritreensis Bremek., 211 Kohiiutia nocliflora Hochst., 209 Koh.iutia pappii Bremek., 210 Koh ftutia platyphylla (K. Schum.) Brem ek., 211, 212 Koh lutiapleiocaulis Bremek., 211 Kohautia quartiniana (A. Rich.) Bremek., 209 Kohautia schimperi Steud. & Hochst. ex Hiern, 210 Kohautia senegalensis Cham. & Schlechtd., 209 TKohautia somaliensis Bremek., 210 Kol nutia subverticillata (K . S ch u m .)M a n tell,2 \ 1 sufcsp eritreensis (Bremek.) M antell ex P u f f 2 \ 1 sufcsp. subverticillata, 2 11 Kol uutia tenuis (S. Bowd.) Mabberley, 209 Kur . j Hochst. & Steud., 245 Kur .afloribunda Hochst. & Steud., 245 Lacnnopyhs congesta (R. Br. ex Fresen.) C.A. Smith, 74 Lacnnopylis oppositifolia Hochst., 74 Lachnopylis ternifolia Hochst., 74 Laduiosiphonturn Hochst., 252 Laci \K>siphomum niloticum (Stapf) Dandy, 252 L a g tn n th u s abyssinicus Hochst. ex Benth. & Hook., 119 Lag( n n th u s interruptus E. Mey., 119 LAP DOLPHIA Beauv., 92 Lan lolphia buchananii (H a llf.) S ta p f 92, 93 Lantcbiphia comorensis (Boj.) K. Schum., 93 var florida (Benth.) K. Schum., 93 Landolphia florida Benth. 93 var. leiantha Oliv., 93 Lasi stelma somalense Schlecht., 171 LEF liBVREA A. R ich .,4 \ Lefebvrea abyssinica A. Rich., 42 var. angustisecta Engl., 42 Lefe \^ea stuhlmannii Engl., 42 LEP T \D EN IA R. Br., 155 Leptadenia abyssinica Decne., 155 Lepi itdenia arborea (Forssk.) Schw ein f, 155 Lept denia cla\ipes S. Moore, 155 Lephtuenia deli lii Decne., 155 Lepbi-denia ephedriformis Defiers, 106 LepUidenia forskalii G. Don., 155 Leptadenia gracilis Decne., 155 Leptadenia hastata (Pers.) Decne., 156 Lepti jenia jacquem ontiana Decne., 155 Leptadenia lancifolia (Schum. & Thonn.) Decne., 156 ''ar. scabra Decne., 156 Leptm enia pyrotechnica (Forssk.) Decne., 155 Leptadenia spartium Wight. & Arn., 155 Leptadenia visciformis Vatke, 104 Linociera Sw., 81 Linociera giordanii Chiov., 81 Linociera johnsonii sensu Cufod., 81 Linoc era latipetala M.Taylor, 81 Lithot aulon Bally not Meneghini, 183

321

Lochnera rosea (L.) Rchb. ex Endl., 92 Logania capensis Eckl., 199 LOGAN1ACEAE, 70 LONICERA L., 283 Iconic era flexuosa Thunb., 283 Lonicera halliana Dipp., 283 Lonicera hildebrandtiana Coll. & Hemsl., 283 Lonicera japonica Thunb., 283, 284 var. halliana (Dipp.) Nichols., 284 Maba abyssinica Hiern, 50 Maba scabra Chiov., 51 MACROSPHYRA H o o k.f, 251 M acrosphyra longistyla (DC.) H iern, 251, 252 Macrosphyra paleacea (A. Rich.) K. Schum., 252 M adoriusprocerus (Ait.) Kuntze, 119 MAESA F o rssk, 64 Maesa lanceolata Forssk., 64, 65 var. serralifolia A. Rich., 64 Maesa picta Hochst., 64 M alabaila abyssinica Boiss., 45 Malabaila atropurpurea (A. Rich.) Vatke, 44 M alabaila elgonensis W olff, 45 Malabaila rivae Engl., 45 Malacantha alnifolia (Bak.) Pierre, 61 M alacanthaferrugineo-tom entosa (Engl.) Engl., 61 Malacantha Pierre, 60 Manettia lanceolata (Forssk.) Vahl. 205 MANILKARA Adans., 57 M anilkara butugi Chiov., 57, 59 Manilkara multinervis (Bak.) Dubard, 58 Manilkara multinervis auct., non (Bak.) Dubard, 57 Manilkara sulcata (Engl.) Dubard, 58 Marsdenia R.Br., 151 Marsdenia auctt. p.p., non R. Br., 153 Marsdenia abyssinica (Hochst.) Schltr., 151 Marsdenia angolensis N.E. Br., 153 Marsdenia gondarensis Chiov., 153 Marsdenia rubicunda (K. Schum.) N.E. Br., 151 Marsdenia schimperi Decne., 151 M arsdenia spissa Moore, 151 Marsdenia stefaninii Chiov., 152 M arsdenia stelostigma K. Schum., 152 Mayapea welwitschii Knobl., 81 MEYNA Roxb. ex Link, 276 Meyna spinosa Link, 276 Meyna tetraphylla (Schweinf. ex H iem ) Robyns, 277 subsp. comorensis (Robyns) Verde., 277 subsp. tetraphylla, 277 M icroloma pyrotechnicum (Forssk.) Spreng., 155 Microstephanus N.E. Br., 137 MIMUSOPS L , 54 Mimusops djurensis Engl., 55 M im usopsfragrans (Bak.) Engl., 55 Mimusops kummel A DC., 55, 56 ?Mimusops degan auct., non Chiov., 55 Mimusops laurifolia (Forssk.) F riis, 55, 56 Mimusops schimperi A . Rich., 55 Mimusops stenosepala Chiov., 55 MITRACARPUS Zucc., 229 M itracarpus ampliatus Hochst., 228 Mitracarpus dregeanus Sond. (“Mitracarpum ” dregeanus), 228 M itracarpus hirtus (L.) DC., 229, 230 M itracarpus scaber Zucc., 229

322

INDEX TO SCIENTIFIC NAMES

Mitracarpus senegalensis DC., 230 Mitracarpus sphaerostigma Hochst., 228 Mitracarpus verticillatus (Schumach. & Thonn.) Vatke, 230 Mitracarpus villosus (Sw.) DC, 229 Mitragyna sensu Cufod. non Korth., 241 Mitragyria rubrostipulata (K. Schum.) Havil., 242 Mitragyna stipulosa sensu Cufod. non (DC.) O. Kuntze. 242 Monolluma Plowes, 170 Monotheca A. DC., 58 Monotheca buxifolia (Falc.) A. DC., 60 Monotheca mascatensis A. DC.. 60 Motandra erlangeri K. Schum., 92 MUSSAENDA L., 245 Mussaenda abyssinia Chiov., 246 Mussaenda arcuata Poir., 246, 248 var. parviflora S. Moore, 246 var. pubescens W ernham, 246 M ussaendaflava (Verde.) Bakh.f., 247 Mussaenda luteola Del., 247 Mussaenda luteola Hochst., 206 Mussaenda schimperi Hochst., 203 MYRSINACEAE. 64 MYRSINE L., 65 Myrsine africana L., 66 Myrsine melanophloeos ( L ) R. Br., 67 Myrsine simensis Hochst. ex DC.. 67 Nathusia alata Hochst., 82 Nauclea L., 241 Nauclea esculenta (Sabine) Merr., 241 Nauclea latifolia Smith. 241 Nauclea microcephala Del., 240 NAUCLEEAE AYn/., 239 NERIUM L., 97 Nerium indicum Mill.. 97 Nerium obesum Forssk., 88 Nerium oleander L., 97 Nerium salicinum Vahl, 240 Neurocarpaea lanceolata (Forssk.) R. Br., 205 var. stenostygma Chiov.. 205 Neurocarpaea quadranguiare (Rendle) Rendle. 213 Neurocarpaea thomsonii (Scott Elliot) S. Moore, 203 NUXIA Lam., 14 Nuxia congesta R.Br ex Fresen., 14 Nuxia dentata R.Br. ex Benth.. 74 Nuxia oppositifolia (Hochst.) Benth., 74. 75 OENANTHE L., 31 Oenanthe palustris (Chiov.) Norman, 31, 33 OCTOTROPIDEAE Redd.. 258 Odontanthera linearis (Fenzl) Mabberley. 157 Odontanthera reniformis sensu Mabberley non Wight, 115 Oenanthe procumbens (Wolff) Norman, 31. 33 Oenanthe ruwenzoriensis Norman. 31 Oenanthe uhligii (Wolff) Norman, 31 OLDENLANDIA L., 218 Oldenlandia L. sect. Conostomium Stapf, 212 Oldenlandia abyssinia (Hochst. ex A. Rich.) Hiem, 211 Oldenlandia aemulans Bremek.. 221 Oldenlandia amaniensis K. Krause, 210 Oldenlandia aspera (Heyne ex Roth) DC., 212 Oldenlandia aspera sensu auctt. div.. non (Heyne ex Roth) DC., 211 Oldenlandia caespitosa (Benth.) Hiem, 221 var. major Bremek.. 221

var. subpedunculata (O. Kuntze) Bremek , 221

Oldenlandia capensis L .f, 222 var. capensis, 222 var. pleiosepala Bremek., 222 Oldenlandia capensis sensu Chiov. non L.f., 221 Oldenlandia coccinea (Royle) Hook.f., 211 Oldenlandia commutata Cufod., 210 Oldenlandia corymbosa L., 220 var caespitosa (Benth.) Verde., 221, 222 var. corymbosa, 221, 222 var. linearis (DC.) Verde., 221 var. nana (Bremek.) Verde., 221 var. subpedunculata O. Kuntze, 221 Oldenlandia corymbosa sensu Chiov. non L .f, 221 Oldenlandia corymbosa sensu Cufod. p.p. non L., 221 Oldenlandia cyanea Dintcr, 212 Oldenlandia debeerstii De Wild. & Thonner Dur., 211 Oldenlandia (Conostomium) dolichantha Stapf, 214 Oldenlandia fasciculata Hiem, 214 Oldenlandia fastigiata Bremek., 221 var. fastigiata. 221, 222 var. longifolia Bremek., 221 var. pscudopcntodon Verde.. 221 var. somala (Bremek.) Verde., 221 Oldenlandia globosa (A. Rich.) Hiern, 215 Oldenlandia goreensis (D C .) Sum merh., 219, 222 var. tnchoclada Bremek., 219 Oldenlandia grandiflora (DC.) Hiem, 209 Oldenlandia hedyotoides (Fisch. & Mey .) Boiss., 222 Oldenlandia herbacea (L.) Roxb., 220 var. caespitosa Benth.. 221 var.flaccida Bremek., 220 var. herbacea. 220. 222 var. holstii (K. Schum.) Bremek., 220 Oldenlandia heynii G. Don, 220 Oldenlandia holstii K. Schum., 220 Oldenlandia ichth yo d erm a Cufod., 220 Oldenlandia lancifolia (Schumach.) D C ., 219 var. brevipes Bremek., 219, 220 var. scabridula Brem ek., 219, 222 Oldenlandia leclercii A. Chev., 212 Oldenlandia linearis DC., 221 var. nana Bremek., 221 Oldenlandia longituba B cck.214 Oldenlandia macrodonta Bak., 2 1 1 Oldenlandia macrophylla DC., 217 Oldenlandia megistosiphon K. Schum., 213 Oldenlandia monanthos (A. Rich.) Hiern, 219, 222 Oldenlandia noctiflora Hiern, 209 Oldenlandia pentandra (Schumach. & Thonn.) DC., 217 Oldenlandia platyphylla K. Schum., 211 Oldenlandia rhodesiana S. Moore, 212 Oldenlandia rhynchotheca K. Schum., 214 Oldenlandia roseiflora K. Schum. & K. Krause. 219 Oldenlandia rotata Bak., 214 Oldenlandia saganensis Cufod., 210 Oldenlandia schimperi (C. Presl) T. Anderson, 210 Oldenlandia schimperi sensu Chiov. non (C. Presl) T. Anderson , 210 Oldenlandia schweinfurthii A. Terr., 210 Oldenlandia senegalensis (Cham. & Schlechtd.) H iem , 209 Oldenlandia somala Bremek., 221 Oldenlandia strumosa (A. Rich.) H iem , 212

INDEX TO SCIENTIFIC NAMES

nlandia subverticillata K. Schum., 2 11 nlandia wauensis Schweinf. ex Hiern, 218 Oldenlandia violacea K. Schum., 219 C LEA L., 79 Ciea africana Mill., 79 O lea capensis L., 80 subsp. capensis, 81 *ubsp. hochstetteri (Bak. ) Friis & P.S.Green, 80 subsp. m acro carp a (C.A. Wright.) Verde., 80 subsp. welwitschii (Knobl.) Friis & P.S.Green, 81 Olea chrysophylla Lam., 79 Olea cuspidata Wall, ex G.Don, 79 Oiea europaea L., 79 subsp. africana (Mill.) P.S.Green, 79 subsp. cuspidata (Wall, ex G.Don) C if, 79. 80 subsp.europaea, 80 O ita hochstetteri Bak., 80 O k a lauri/olia sensu Richard non Lam., 80 Oiea macrocarpa C.A. Wright, 80 O l;a mildbraedii (Gilg & Schellenb.) Knobl., 81 Olea mussolinii Chiov., 81 O k a schimperi Gand.. 79 O ita somaliensis Bak., 79 Olea subtrinervata Chiov., 79 Olea welwitschii (Knobl.) G ilg & Schellenb.. 81 OLEACEAE. 79 ONCINOTIS 92 Oncinotis chirindica S. M oore, 92 C icinotis inandensis Wood & Evans. 92 inotis natalensis Stapf, 92 Oncinotis oblanceolata Engl., 92 Oncinotis tenuiloba Stapf. 92, 94 Ophiorrhiza lanceolata Forssk., 205 Orbea abayensis (M.G. Gilbert) Bruyns, 187 Ort ea baldratii (White & Sloane) Bruyns, 187 Orbea dummeri (N.E. Br.) Bruyns, 187 subsp. circes (M.G. Gilbert) Bruyns, 187 Orbea gemugofana? (M.G. Gilbert) Bruyns, 186 Orbea laticorona (M.G. Gilbert) Bruyns. 186 Orbea sprengeri (N.E. Br) Bruyns, 185 subsp. foetida (M.G. Gi Ibert) Bruyns, 186 ubsp. ogadensis (M.G. Gilbert) Bruyns, 186 Ch^ea subterranea (Brucc & Bally) Bruyns. 187 O ’ bea succulata (N.E. Br.) Plowes, 189 0*~ea vibratilis (Bruce & Bally) Bruyns, 190 CfceopsisL.C. Leach, 170 OI IEOSCHI1MPERELLA Rauschert. 23,24 Oreoschimperella verrucosa (A. Rich.) Rauschert. 24 Otomeria ?heterophylla K. Schum., 214 O I OMERIA Benth., 206 Otomeria blomniaertii Dc W ild., 206 O nm eria m adiensis Oliv., 206 Otomeria ocuiata S. M oore, 206, 207 O '[YANTHUS DC., 256 O yanthus lepidus S. M oore, 257 var. lepidus, 257 var. unyorensis (S. M oore) Bridson. 257 Oy\'anthus microphyllus K. Krause, 257 O 'vanthus oxycarpus S. Moore, 257 O anthus speciosus D C., 256 iubsp. gerrardii (Sond.) Bridson, 257 subsp. globosus Bridson. 256. 257 ;ubsp. mollis (Hutch.) Bridson, 257

323

subsp. speciosus, 257 subsp. stenocarpus (K. Schum .) Bridson, 256, 257 Oxyanthus stenocarpus K. Schum., 257 Oxyanthus unyorensis S. Moore, 257 Oxyanthus villosus G. Don, 252 OXYSTELMA/? Br., 149 Oxvstelma alpini Decne., 150 Oxystelma bornouense R. Br., 149, 150 Oxvstelma esculentum ( L .f ) Schult., 150 var. alpini (Decne.) N.E. Br., 150 Oxystelma secamone (L.) Karsten, 150 Oxystelma senegalense Decne., 149 PACHYCARPUS E. Mey., 126 Pachycarpus bisacculatus (Oliv.) Goyder Pachycarpus bullockii Cavaco, 126 Pachycarpus corniculalus Hochst., 130 Pachycarpus lineolatus sensu Bullock non Decne., 126 Pachycarpus petherickianus (Oliv.) Goyder. 128 Pachycarpus robustus (A. Rich.) Bullock, 127 Pachycarpus schumannii Chiov., 128 Pachycarpus terminans Fiori. 128 Pachycarpus verrucosa (A. Rich.) Vatke, 24 var. simensis (A . Rich.) Vatke, 24 P A C H Y C Y M B IU M Leach, 185 Pachycymbium Leach, 170 Pachycymbium abayense (M.G. Gilbert) M.G. Gilbert, 187,189 Pachycymbium baldratii (White & Sloane) M.G. Gilbert, 187 subsp. baldratii, 187 subsp. subterraneum (Bruce & Bally) M.G. Gilbert, 187, 188.189 Pachycymbium circes (M.G. Gilbert) M.G. Gilbert, 187, 188 Pachycymbium dummeri (N.E. Br.) M.G. Gilbert, 189 Pachycymbium gemugofanum (M.G. Gilbert) M.G. Gilbert, 1 8 6 .188.189 Pachycymbium gilbertii (Plowes) M.G. Gilbert, 1 8 6 ,189 Pachycymbium laticoronum (M.G. Gilbert) M.G. Gilbert, 186, 188,189 Pachycymbium sacculatum (N.E. Br.) M.G. Gilbert, 188, 189 Pachycymbium sprengeri (N.E. Br.) M.G. Gilbert, 185 subsp. foetidum (M.G. Gilbert) M.G. Gilbert, 186 subsp. ogadense (M.Gilbert) M.G. Gilbert, 186 subsp. sprengeri, 1 8 6 ,188,189 Pachycymbium vibratilis (Bruce & Bally) M.G. Gilbert, 188, 190 Pacouria comorensis (Boj.) Roberty, 93 Pacouria florida (Benth.) Hiem, 93 PAEDERIA L., 231 Paederia petrophila K. Schum., 231 Paederia pospischilii K. Schum ., 231, 232 PAEDERIEAE DC., 230 Pastinaca atropurpurea Steud. ex A. Rich.. 44 PAVETTA L., 262 Pavetta abyssinica Fresen., 264 var. abyssinica. 264, 265 var brem ekam piana (Pic.Serm.) Bridson, 265 var. cinerascens A. Rich., 264, 265 var. dolichosiphon (Bremek.) Bridson, 265 Pcn’etta abyssinica sensu A. Rich, non Fresen., 264 var. “cinerescens" A. Rich., 264 var. glabra A. Rich., 265 Pavetta adelensis Del., 265 Pavetta aethiopica Bremek., 264 Pavetta albertina S. Moore, 266 Pavetta assimilis Sond., 265

324

INDEX TO SCIENTIFIC NAMES

var. brevituba-glabra Bremek., 265 var. glabra Bremek., 265 Pavetta barteri Da we. 265 Pavetta bremekampiana Pic.Serm., 265 Pavetta chiovendae Pic.Serm., 264 Pavetta cinerascens (A. Rich.) Chiov. non Bremek., 264 var. glabrescens Chiov., 264 Pavetta cinerascens sensu Bremek. non (A. Rich.) Chiov., 265 Pavetta crassipes K. Schum., 265 var. major De Wild., 265 Pavetta denudata Bremek., 264 Pavetta dolichosiphon Bremek., 265 Pavetta ellenbeckii K. Schum., 265 Pavetta elliottii sensu Cufod. non K. Schum. & K. Krause, 263 Pavetta elliptica Hochst., 260 Pavetta eritreensis Bremek., 265 ?Pavetta erlangeri Bremek., 265 Pavetta fossorum Bremek., 265 Pavetta gardeniifolia A. Rich., 265 “var. angustata A. Rich.” sensu Armari , 265 var. breviflora Vatke, 265 var. gardeniifolia, 265 var. laxiflora K. Schum., 265 var. tongijlora Vatke, 266 var. subtomentosa K. Schum., 266 Pavetta graveolens S. Moore, 262 Pavetta grumosa S. M oore, 266 Pavetta hochstetteri Bremek., 265 var. glaberrima Bremek., 265 var. graciliflora, 265 var. mollirama Bremek., 265 Pavetta humbertii Bremek., 263 Pavetta Icabarensis Bremek., 266 Pavetta kenyensis Bremek., 265 Pavetta kerenensis Becc., 266 Pavetta kotschyana Cufod., 266 Pavetta krauseana K. Krause, 265 ?Pavetta erlangeri Bremek., 265 Pavetta lebrunii Bremek., 263 Pavetta ligustriodora Chiov., 262 Pavetta maitlandii Bremek., 265 Pavetta murleensis Cufod., 266 var. glabrescens Cufod., 266 Pavetta oliveriana H iem , 263 var. denudata (Bremek.) Bridson, 264 Pavetta parviflora A fz ., 274 Pavetta petraea Bremek., 265 Pavetta rudolphina Cufod., 266 var. robusta Cufod., 266 Pavetta saxicola K. Krause, 265 Pavetta sihicola Bremek., 265 Pavetta somaliensis Bremek., 265 Pavetta sp. Bremek., 263 Pavetta subcana H iem , 266 var. longiflora {Vatke) Bridson, 266 var. subcana, 266 Pavetta termitaria Bremek., 265 var. glabra Bremek., 265 Pavetta trichotropis Bremek., 265 Pavetta unguiculata Bremek., 266 Pavetta utilis Hua, 265 PAVETTEAE Dumort., 262 PENTANISIA H arv., 282

Pentanisia longituba (F ranch.) Oliv., 282 Pentanisia ouranogyne S. M oore, 281, 282 var. glabri/oJia Cufod., 282 Pentanisia pentasiana Mattei, 282 PENTANOPSIS Rendle, 214 Pentanopsis fragrans R endle, 214 PENTARRHINUM E. M ey., 139 Pentarrhinum abyssinicum Decne., 141 subsp. abyssinicum. 142 Pentarrhinum balense (Liede) Liede, 1 4 2 ,144 Pentarrhinum fasciculatum K. Schum., 138 Pentarrhinum gonoloboides (Schltr.) Liede, 142 Pentarrhinum insipidum E. M ey, 142, 143 Pentarrhinum somaliense (N.E. B r.) Liede, 142 Pentas ainsworthii Scott Elliot, 205 PENTAS Benth., 202 PENTAS Benth subgcn. LONGIFLORA Verde., 203 PENTAS Benth. subgen. PENTAS Verde., 203 PENTAS Benth. subgen. VIGNALDIOPSIS Verde., 203 Pentas caffensis Chiov., 205 Pentas cam ea sensu Cufod. non Benth., 206 Pentas coccinea sensu Chiov., 205 Pentas coerulea Chiov., 206 Pentas concinna K. Schum ., 204 Pentas concinna sensu Chiov. non K. Schum., 206 Pentas decora S M oore, 204 var. decora, 205 var. lasiocarpa Verde., 205 var. triangularis {De Wild.) Verde., 204 Pentas globifera Hutch., 204 Pentas lanceolata (Forssk.) Deflers, 205 forma velutma Verde., 205 var. membranacea Verde., 205 subsp. lanceolata, 205 var. stenostygma (Chiov.) Cufod., 205 subsp. quartiniana (A. Rich.) Verde., 206 var. leucaster (K. Krause) Verde., 206 var. nemorosa (Chiov.) Verde., 206 Pentas leucaster K K rause, 206 Pentas longiflora sensu Cufod., non Oliv., 205 Pentas modesta Bak., 211 Pentas mombassana Oliv., 205 Pentas parvifolia H ie m , 205 var. nemorosa Chiov., 206 Pentas quadrangulare Rendle, 213 Pentas quartiniana (A. Rich.) Oliv., 206 Pentas schimperi Engl., 203 Pentas schimperiana (A. Rich.) Vatke, 203 subsp. schimperiana, 203, 204 subsp. occidentalis (Hook.f.) Verde., 203 Pentas schweinfurthii Scott Elliot, 205 Pentas tenuis Verde., 203 Pentas thomsonu Scott Elliot, 203 Pentas triangularis De W ild., 204 Pentas verruculosa Chiov., 206 Pentas verticillata Scott Elliot, 204 var. pubescens S. M oore, 204 PENTATROPIS/?. Br. ex Wight & A m ., 137 Pentatropis cynanchoides R. Br. ex N.E. Br., 138 Pentatropis fasciculatum (K. Schum.) N.E. Br., 138 Pentatropis hoyoides K. Schum, 138 Pentatropis nivalis (J F . Gmel.) D .V F ield & J.R.I. Wood, 138. 139

INDEX TO SCIENTIFIC NAMES

P 'Htatropis rigida Chiov., 138 Pmttodon abyssinicus Hochst.. 217 PENTODON Hochst., 216 P ntodon pentandrus (Schum ach. & Thonn.) Vatke, 217 var. minor Bremek.. 218 var. pentandrus, 217 P RGULARIA L., 133 P rgularia daemia (Forssk.) Chiov., 133, 134 P 'rgularia extensa (Jacq.) N.E. Br., 133 P rgularia tomentosa L., 134 P nanthostelm a Baill., 144

p : r i p l o c a l , 103 P 'nploca aphylla sensu Chiov.. 106 Periploca ephedriformis (D cflcrs) Schwcinf. ex Defiers, 106 P n p lo c a esculenta L. f., 150 P riploca linearifolia Quart.-Dill. -L T-t, ®*-w. £-y, 7i-z, Tr-z, h-k, 'fi-k. The representation of the diphthong fourth forms is given as follows:

f id e l s

in

their

fJL bwa, 3; -cwa, 3,-dwaA -fwa, 7,-gwa, ^-hwa,*L-jwa, U.-kwa, '9,-kwa, 4,-lwa, 9-nwa. £ nwa, *-qwa.