WAGENINGEN AGRICULTURAL UNIVERSITY PAPERS 96-I(1996)

Acontributio n toth elea f anatomy andtaxonom yo f Apocynaceaei n Africa

- The leaf anatomy of Apocynaceae in East Africa - A monograph of Pleiocarpinae (Series of revisions of Apocynaceae XLI)

Elizabeth Omino

Wageningen MMAgricultura l University

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CIP-gegevensKoninklijk e Bibliotheek,De nHaa g

Omino,Elizabet h

Acontributio n toth elea f anatomy andtaxonom y of Apocynaceae inAfric a :Th e leaf anatomyo f Apocynaceae inEas tAfric a : Amonograp ho f Pleiocarpinae (Serieso frevision so fApocynacea eXLI )/ Elizabet h Omino.- Leide n: Backhuys Publishers.- Dl .- (Wageninge n AgriculturalUniversit ypaper s ; 96-1) Alsopublishe da sthesi sLandbouwuniversitei tWageningen , 1996- Wit hindex ,ref . - Withsummar yi nDutch . ISBN90-73348-53- 6 NUGI835 Subject headings:lea f anatomy/ Apocynacea e ; taxonomy. TABLE OF CONTENTS

General Summary i Samenvatting 2 i Leaf anatomy of Apocynaceae in East Africa 3 i.i Summary 3 1.2 Introduction 3 1-3 Material and Methods 4 1-4 Observations 4 1-5 Characteristics of the genera io 5.1 Acokanthera G.Do n 10 5.2 Adenium R.Br . 11 5.3 .A/o/iaThouars 12 5.4 Ancylobotrys Pierre 13 5.5 Baissea A.D C 14 5.6 Carissa L 15 5.7 Carvalhoa K.Schu m 16 5.8 Clitandra Benth 17 5.9 Dictyophleba Pierre 18 5.10 DiplorhynchusWelw. ex Fie. & Hiern 18 5.11 Funtumia Stapf 19 5.12 Holarrhena R.Br. 20 5.13 Hunteria Roxb 21 5.14 Landolphia P.Beau v 22 5.15 Mascarenhasia A.D C 23 5.16 Motandra A.DC 24 5.17 Oncinotis Benth 25 5.18 Picralima Pierre 26 5.19 Pleiocarpa Benth 26 5.20 Rauvolfia L 28 5.21 Saba (Pichon) Pichon 29 5.22 Schizozygia Baill 30 5.23 Stephanostema K.Schu m 30 5.24 Strophantus DC 31 5.25 Tabernaemontana L 32 5.26 VoacangaThouars 33 1.6 General discussion 34 1-7 Conclusions 37 1.8 Figures and tables 42 1-9 Anatomy references 76 2 A monograph of Pleiocarpinae (Apocynaceae-Plumerioideae-Carisseae); Series of revisions of Apocynaceae XLI 81 2.1 Summary 81 2.2 Introduction 8i 2.3 General part 82 2.3.1 History of the subtribe Pleiocarpinae 82 2.3.2 History of the genera Hunteria Roxb.,Pleiocarpa Benth. and Picralima Pierre 82 2.3.3 Geography and distribution 83 2.3.4 Morphology 83 2.3.5 General taxonomiereference s 86 2.4 Taxonomiepar t 87 2.4.1 The subtribe Pleiocarpinae 87 2.4.2 Key to the genera of the subtribe Pleiocarpinae 87 2.5 Hunteria Roxb 88 2.5.1 Hunteria ballayi Hua 90 2.5.2 Hunteria camerunensis K.Schum . ex Hall 93 2.5.3 Hunteria congolana Pichon 96 2.5.4 Hunteria densiflora Pichon 99 2.5.5 Hunteria ghanensis Hall & Leeuwenberg 102 2.5.6 Hunteria hexaloba (Pichon) Omino comb, nov 104 2.5.7 Hunteria macrosiphon Omino sp.nov . 107 2.5.8 Hunteria myriantha Omino sp.no v 109 2.5.9 Hunteria oxyantha Omino sp.no v 112 2.5.10 Hunteria simii (Stapf) H.Huber 115 2.5.11 Hunteria umbellata (K.Schum. ) Hall 118 2.5.12 Hunteria zeylanica (Retz.) Gard. exThw 122 2.6 Picralima Pierre 128 2.6.1 Picralima nitida (Stapf) Th. & H.Dur. 128 2.7 Pleiocarpa Benth 134 2.7.1 Pleiocarpa bicarpellata Stapf 135 2.7.2 Pleiocarpa brevistyla Omino sp.no v 139 2.7.3 Pleiocarpa mutica Benth 142 2.7.4 Pleiocarpa pyenantha (K.Schum. ) Stapf 146 2.7.5 Pleiocarpa rostrata Benth 152 2.8 Doubtful species 156 2.9 Excluded species 157 2.10 Index of exsiccatae 158

Index of scientific names 169 Curriculum Vitae 177 Acknowledgements 177

vi GENERAL SUMMARY

Thispublicatio n comprisestw omai nparts . Parton e(Chapte r i) dealswit hth elea f anatomyo fApocynacea ei nEas t Africa, wherethi sfamil y isrepresente db y7 7 speciesi n3 0genera . Outo fthese ,3 7specie s in2 6gener aar eexamine dan dthes eexhibi ta wid erang eo f anatomicalvariations . Many speciesca nb eseparate do nth ebasi so f theirlea f anatomyan dman ygener a are confirmed to be closely related.Th e most outstanding anatomical characters includeth edorsiventra lleaves ,presenc eo flaticifers ,an dbicollatera lvascula rbun ­ dleswit hintraxylar y phloem. Parttw o(Chapte r2 )i sa monograp ho fth eapocynaceou ssubtrib ePleiocarpinae - Plumeriodeae-Carisseae, which comprises of three genera, Hunteria(1 2 spp.); Pleiocarpa( 5 spp.)an dPicralima ( 1 sp.).Th esubtrib ePleiocarpina ei srestricte dt o Africa, althoughon especies ,Hunteria zeylanica, extend st oAsia .I ti seasil ydistin ­ guishedfro mth eothe rmember so fth etrib eCarissea eb yth epresenc eo fcompletel y apocarpous ovaries, with 2-5 carpels, which are a great exception in the family Apocynaceae. Thegenu sPicralima wa sfirst describe db yPierr ei n189 6an dha ssinc eremaine d monotypic.It sonl yspecies ,Picralima nitida, i sno tonl yexploite dfo rit swood ,bu t also used medicinally in its present distribution area and employed as well as an arrowan da fish poison . In this publication, three new species are described in the genusHunteria (Roxburgh, 1814, 1824)an don ene wcombinatio n ismade . Thegenu sPleiocarpa (Bentham, 1876)ha sprove dt ob ea ver ydifficul t subject duet oinsufficien t collections andlac ko f sufficient information from thefield. A t present it has 5 species and one doubtful species of which the collections are all eithersteril eo ronl ybearin gfruits . Onene w speciesi sdescribed . Parallelt oth einvestigation s reported herea taxonomi etreatmen t wasprepare d forth eFlor ao f TropicalEas tAfrica .Thi scontribution ,coverin gal lth ementione d 30gener aan d7 7 species,wil lb esubmitte dt oth eeditor si nKe wshortly .

WageningenAgricultural UniversityPapers 96-1 (1996) SAMENVATTING

Dezepublicati ebestaa tvoornamelij k uittwe edelen . Hoofdstuk i behandeltd ebladanatomi e vanvertegenwoordiger s vand ei nOos t Afrikavoorkomend eApocynaceae ,ee nfamili edi edaa r7 7soorte ni n3 0gener atelt . Hiervan werden 37 soorten van 36gener abestudeer d en deze vertonen een grote variatieaa nanatomisch ekenmerkstaten .Vee lsoorte nkunne nworde nonderscheide n aand ehan dva nd ekenmerke nva nd ebladanatomie ,e nd enauw everwantschappe n tussen vele genera kunnen er mee worden bevestigd. De meest opvallende anatomische kenmerken zijn de dorsiventrale bladeren, de aanwezigheid van melksapvaten, enbicollateral evaatbundel sme tintraxylai r floeem. Hoofdstuk 2i s een monografie van de subtribus Pleiocarpinae, welke uit drie generabestaat : Hunteria (met 12 soorten); Pleiocarpa (5soorten )e nPicralima (1 soort). De subtribus Pleiocarpinae van de onderfamilie Plumerioideae, tribus Carisseae,i svoornamelij k beperktto tAfrika , maaréé n soort,Hunteria zeylanica, komt ook in Azië voor. De subtribus verschilt duidelijk van andere in de tribus Carisseae door de aanwezigheid vangehee l apocarpe vruchtbeginsels met 2to t5 vruchtbladen,hetgee n eengrot euitzonderin g isi n defamili e Apocynaceae. Het genus Picralimawer d door Pierre in 1896 beschreven en is sindsdien monotypischgebleven .D eenig esoort ,Picralima nitida, word tnie tallee nvanweg e hethou tgeëxploiteerd ,maa roo kmedicinaa lbenu te nbovendie nzowe lvoo rpijlgi f alsvoo r visvergift. InHunteria (Roxburgh , 1814,1824)worde ndri enieuw esoorte nbeschreven ,e n eri see nnieuw ecombinati egemaakt . Hetgenu sPleiocarpa (Bentham , 1876)blee kee nzee rlasti gonderwerp ,doorda t erweini gmateriaa lverzamel di se nveldgegeven snauwelijk sbeschikbaa rwaren .E r worden nu vijf soorten onderscheiden en er is een probleemgeval waarvan de belegexemplarenó f sterielware nó f alleenvruchte ndragen .E ri séé nnieuw esoor t beschreven. Tijdens het onderzoek dathie r wordtgepubliceer d werd ookee n taxonomische bewerkingvoorberei dvoo rd eFlor ava nTropisc hOos tAfrika .Dez ebijdrag edi ed e genoemde 30 genera en 77 soorten omvat, zal binnenkort aan deredacti e van de Florao f TropicalEas tAfric a inKe wworde n aangeboden.

WageningenAgricultural UniversityPapers 96-1 (1996) 1 LEAFANATOM YO FAPOCYNACEA EI N EASTAFRIC A

i.i SUMMARY

A study has been made of 37 species in 26gener a of the Apocynaceae in East Af­ rica.Th e anatomical characters found tob e useful in distinguishing taxa are:pres ­ ence and absence of hairs/domatia, hair types and position on the leaf, cuticular architecture (SEM), stomatal outline, and occasionally density,thei r leaf venation; in transverse section: petiole/midrib outline, shape of vascular bundle, phloem ar­ rangement, presence or absence of fibres in the petiole, and occasionally spongy mesophyll arrangement and presence or absence of foliar sclerids, arrangement of laticifers atth e midrib,sclerenchymatou s marginal veins andpresenc e of papillose cells at the margin.Th e present study confirms a close relationship between the Pleiocarpinae (Hunteria, Picralima and Pleiocarpa); Motandra and Oncinotis; Mascarenhasia and Funtumia; Carissa, and Acokanthera and Tabernaemontana, VoacangaSchizozygia and Carvalhoa. Themos t outstanding anatomical characters areth edorsiventra l leaves;presenc eo f lalicifers; typically bicollateral vascular bundle in petiole and leaf and presence of intraxylary phloem.

1.2 INTRODUCTION

In East Africa, the plant family Apocynaceae is represented by 77 species in 30 general (Table 3). The family is usually subdivided into two subfamilies, Plumerioideae andApocynoideae .Th esubfamilie s arefurthe r subdividedint otribe s and subtribes (Table 3).Th e earliest work on the general leaf anatomy was by Solereder (1899, 1908) followed by Poulsen (1917) andMetcalf e & Chalk (1950). Other work on leaf anatomy includes Sayeed-ud-Din (1941) on Lochnera pusilla (now Catharanthus pusilla); Holm (1910) on Apocynum cannabinum; Ballard (1926),o ntrichome si nApocynum, Ngan(1965 )o nth egenu s Wrightia;Newcomb e & Patel (1966) on Voacanga schweinfurthii (now V.africana); Fjell (1983) on Allamanda neriifolia (nowA . schottii), Thevetiaperuviana and Wincaminor; Araujo etal., (1984) onPlumeria rubra. Epidermal and venation studies have been carried out by Chandra et al., (1969, 1972); Kapoor et al., (1969); Sharma et al., (1970); Kapoor & Mitra (1979). Extensive stomatal studies inWes t African species have been carried out by Nyawuame & Gill (1991), other work on stomata include Karatela & Gill (1983).Apar t from the stomatal studies inWes t African species of Apocynaceae and a few remarks on some African genera by Metcalfe & Chalk (1950), leaf anatomy especially of theEas t African species is hitherto unknown. Inth epresen tpape rlea f anatomical charactersar edescribe di ndetai lfo r 26o f the

WageningenAgricultural University Papers 96-1 (1996) 3 30 genera in East Africa and their effect on the current taxonomierelationship s in the family is discussed atth e end of the chapter. All scientific authorities of the species names havebee n omitted, thesear e given in the final index.

1.3 MATERIAL AND METHODS

All living material was obtained from the field in Kenya, fixed in FAA and stored in 70% alcohol. Material from Uganda andTanzani a was obtained from Kew and East African herbaria (see appendix 1). Dried leaves were boiled for several minutes and stored in 70% alcohol. Petiole and leaves were sectioned atth eapproximat e midpointusin g aReicher t sliding mi­ crotome, cleared with parazone, and mounted ineuparo l after dehydration through an alcohol series.Technica l methods for cuticular preparations, clearing leaves, stomatalcount san dmeasurement sfo rlamin athicknes sar eadapte dfro mWilkinso n (1989).

1.4 OBSERVATIONS

Characteristics of the family Apocynaceae as a whole:

Leaves: opposite, rarely alternate or bunches set close together as in Adenium obesum,o rwhorle da si nRauvolfia, Pleiocarpa andAlstonia; petiolate ,rarel y sessile asi n Voacangaaf ricana, entire ;textur evarie sfro mmembranaceou s asi n Carvalhoa campanulata to coriaceous as in Carissa and Acokanthera, to succulent as in Adenium obesum; sizerange s from 0.8-3.6 x 0.3-1.4c m in Strophanthus mirabilis to i.8-50(-70) x 0.8-19 cm in Rauvolfia coffra; petiole with colletors and axils of secondary veinswit hdomati aa si nMotandra, Baissea andOncinotis; blad eobovat e to linear as in Adenium obesum to elliptic as in Clitandra cymulosa; completely glabrous as inHunteria to densely short-pubescent asi nStrophanthus hypoleucos.

Leaf venation: According to the terminology proposed by Hickey (1979) for the architecture of dicotyledonous leaves, all the species studied have camptodromous leaves,a sth e secondary veins dono tterminat e atth emargin .Th e venation is either brochidodromous with secondariesjoine d together in a series of prominent arches (Figs:22A, B) or eucamptodromous with secondaries connected to superadjacent veinsb y aserie so f cross-veins withoutformin g prominentmargina lloo p(Fi g22c) .

Leaf surface Hairs:presen to nth epetiol ea si nDictyophleba, Ancylobotrys andCarissa (Fig . iC, F); on both surfaces of midrib as in Baissea myrtifolia and Holarrhena pubescens (Fig. 11,B) ;o n adaxial midrib as in Carissa tetramera oro n abaxial leaf surface as in H. pubescens and Stephanostema stenocarpum. Hairs simple, uniseriate, 1-20-

4 WageningenAgricultural UniversityPapers 96-1 (1996) celled;thin-walled , short asi nMascarenhasia arborescens (Fig. iE); thick walled, long and straight as inHolarrhenapubescens (Fig. iB); thick-walled, curved, with very short cells asi nDictyophleba lucida (Fig. iF);thick-walled , hooked, but with longerbasa l cells and shortertermina l cells asi nBaissea myrtifolia (Fig. il). Glan­ dular hairs with three basal cells have been observed in the petiole of Motandra guineensis (Fig. iD).Tuft s of hairs (domatia) are present in the axils of secondary veins e.g. in Baissea myrtifolia, with long curled hairs (Fig. 3E) and in Oncinotis tenuiloba with short straight hairs (Fig. 3F).

Cuticularsurface: smootha si nDiplorhynchus condylocarpon (Fig.2A )an dCarissa tetramera (Fig. 5C); with wings of striae across subsidiary cells as inT. stapfiana (Fig. 5B); with concentric rings of striae over subsidiary cells as in Strophantus courmontii (Fig. 4C); with ridges extending from the stomatal poles as in Stephanostema stenocarpum (Fig. 5A); with striatums confined to each cell as in Schizozygia coffaeiodes (Fig. 2D) or striate over the whole surface as in Tabemaemontana stapfiana (Fig. 2C).

Stomata: distribution hypostomatic in almost all species examined except in Stro- phanthus courmontii (Fig.6A) ,Schizozygia coffaeiodes, Voacangaafricana andV. thouarsii where it is amphistomatic and in Carissa edulis where a few stomata overly the adaxial midrib.I n this case the stomata found on the adaxial surface are very few and larger than the ones on the abaxial surface (Fig. 6A,B) .

Stomata outline:broadl y elliptica si n Carissatetramera (Fig.5C) ;narrowl y elliptic as in Rauvolfia mannii (Fig. 7G);circula r as inAncylobotrys tayloris (Fig. 5E).

Densityper mm2:(Tabl e2) :lowes ti nTabemaemontana pachysiphon (31-56-69 )an d highest in Dictyophleba lucida (700-825-925).

Stomatal size:(Tabl e2) :rangin gfro m 14.6-16.6u, mlon gi nLandolphia watsoniana to 26.4-36.5 [imlon g inTabemaemontana stapfiana.

Subsidiary cells:i n most cases the subsidiary cells werereadil y distinguished from adjacent epidermal cells by their specially thin cuticle; paracytic (Fig. 7D, F, G), sometimes extending over one or both poles (laterocytic) as m Carvalhoa campanulata, Holarrhena pubescens (Fig. 6D) and Stephanostema stenocarpum; usually with one or two pairs of subsidiary cells, rarely 3 pairs as in Acokanthera oppositifolia (Fig. 6F); occasionally paracytic cells subdivided by radiating walls on one or both sides as in Strophanthus courmontii (Fig. 6B); anomocytic and anisocytic as in Hunteria congolana; anomocytic with 5-8 cells around the stoma as in Clitandra cymulosa and 3-6 cells as in Dictyophleba lucida (Fig. 6H); cyclocytic or cyclocytic-tetracytic with 3-4 subsidiary cells as in Pleiocarpa (fig 7B), with 7-8 subsidiary cells asi n Carissa edulis;tetracyti c with 4 subsidiary cells as in A. oppositifolia (Fig. 6F); occasionally complex as in A. oppositifolia. Pri­ mary stomata observed in several species of most genera (Fig.4C) ;doubl e stomata

WageningenAgricultural UniversityPapers 96-1 (1996) 5 present in several species (Fig. 6B); single guard cell observed in Holarrhena pubescens.

Anticlinal walls:(Tabl e 2),± straight onbot h sides as inAcokanthera oppositifolia (Fig. 6E, F); very undulate on both sides as in Baissea myrtifolia (Fig. 7C, D); slightly undulate onbot h sides asi nRauvolfia mannii (Fig.6E ,F) ;adaxia l straight, abaxial undulate asi n Oncinotis tenuiloba andRauvolfia mombasiana (Fig.7G ,H) .

T.S.(Transverse Section) of leaf lamina (Figs. 13, 14)

Thickness: (Table 1),rangin g from 82-110 urn inBaissea myrifolia to 384-458 um Acokanthera oppositifolia.

Cuticle and outerpericlinal walls: (Fig. 8,9,) the outer cuticle is of even thickness alongth e lamina (Fig. 8A, K,W ) and of eventhicknes s to crenate atth emidri b and margin (Fig.9H ,I ,J , K,L ,T) ;generall y thicker atth emidri ban dmargi n in allspe ­ cies examined (Fig. 9A, B, G, M, P,T) ; ranging from 1u m in Carvalhoa campanulata to9.4-4 8 urn inAcokanthera oppositifolia.

Epidermal cells:(Fi g8,9) ,mor eo rles srectangula ri nmos to fth especie s examined (Fig. 8 A, B, O, S,T) ;Tabl e 1,t o almost square or taller than wide as in Adenium obesum whichha sth elarges tepiderma lcell s (upt o68. 4 x 61.8um )(fig . 8C);adax ­ ial epidermal cells generally larger than abaxial (Fig. 8C,D) ,occasionall y of ± the same size asi nDiplorhynchus condylocarpon; usually smaller,talle rtha n wide and with an obtuse apex at the midrib inAdenium obesum (Figs.9C ,D ) and Oncinotis tenuiloba;wit ha nacut eape xa si nMascarenhasia arborescens (Figs.9I ,J) .Adaxia l epidermal cells 1-layeredi nmos t species;2-layere d atth e midrib and margin as in Mascarenhasia arboresens (Fig.9S) ;intermittentl y 2-layered withlarge rcell si nth e secondlaye r inAncylobotrys tayloris (Fig. 8E);irregularl y 2-3-layered, sometimes with tangential divisions in Funtumia elastica andTabernaemontana pachysiphon (Figs. 8M,U ,W 18F);abaxia l mostly i-layered; 1-2-layered asi nF elastica andT. stapfiana (Figs. 8N, X).

Stomata T.S.: (Figs. 10 & n), Stomata level with the cuticular surface as in Oncinotis tenuiloba (Fig. 11D),Adenium obesum or Alafia microstylis; raised on abaxial surface as in Strophantus courmontii andTabernaemontana pachysiphon (Figs. 1iF ,H) ;sunke n onadaxia l surface asi nStrophanthus courmontii and abaxial surface Voacangathouarsii (Figs. 1iE , J); with a very thick cuticular ledge as in Acokanthera oppositifolia andPicralima nitida (Figs. 10A, 11A) ;wit hsmalle rcutic ­ ular ledges in several other species (Figs. 1iB , C, 1,10B, C, F,H , I).

Mesophyll: (Table 2; Fig. 12), bifacial in all the species examined except in Saba comorensis (Fig. 12A)an d wherei ti s homogenous (thepalisad e and spongy tissue cannot be distinguished). In some species likeAcokanthera oppositifolia there are 2-3 layers of poorly developed palisade-like cells above the lower epidermis (Fig.

6 WageningenAgricultural UniversityPapers 96-1 (1996) I2E), 3-rows in Carissa edulis, 2-rows in Clitandra cymulosa (tendency to be isobilateral).

Palisade tissue: (Table i), the palisade tissue consists of 1-4 layers of regularly arrangedcell selongate da trigh tangle st oth eadaxia lepidermis ;i tma yexten dt oth e midrib as in Tabernaemontana elegans (Fig. 15F) or leave a gap as in Saba comorensis (Fig 15D);i ti s i-layeredi nHolarrhena pubescens (Fig. 12B), Baissea myrtifolia andCarvalhoa campanulata; 2-layere da si nClitandra cymulosa and3-4 - layered asi nStmphanthus courmontii, Tabernaemontana elegans (Fig. 12D) andZ stapfiana. Inth e multiseriatepalisad etissu eth e outermost cells areth elongest , the innermost the shortest; palisade cells are characteristically short in Stmphanthus courmontii and several times longer than wide as in Carissa tetramera (Fig. 16B), Rauvolfia caffra,Landolphia buchananii and Funtumia elastica (Fig. 17E).

Spongy mesophyll cells: (Fig. 12), appear less regular; transversely elongated and compact asi nHunteria congolana andH. pubescens (Fig. 12B),o rles s compact as inTabernaemontana elegans (Fig. 12D) except in Saba comorensis where all the cells are vertically elongated and very compact (homogenous) (Fig. 12 A) and in Ancylobotrys tayloris where they are also more or less elongated perpendicular to the leaf surface, but very lacunate (Fig. 12C), and inDiplorhychus condylocarpon wherethe yar eles slacunat e(Fig . 12F)an di nAdenium obesum wherethe yar emor e or less irregularly arranged.

Minor veins:embedde d in most of the species examined, collateral, with xylem on adaxial and phloem on abaxial side; xylem vessels usually surrounded by thick- walled fibres as in Pleiocarpa bicarpellata (Fig. 16F), the larger veins of Acokanthera oppositifolia are bicollateral; transcurrent as in Diplorhynchus condylocarpon (Fig. 12F),Ancylobotryspetersiana andSaba comorenis (Fig. 12A) withgirder stoward son eo rbot hepidermi s(Figs .1 2A ,F) .Girder san dbundl esheat h cells composed of thin- tothick-walle d parenchyma cells (Fig. 16F).

Midrib: (outlines illustrated in Figs. 13& 14).

Thickness: thickest inTabernaemontana stapfiana (2013-2141 um) and thinnest in Baissea myrtifolia (201-238 urn); adaxial outline ± straight as in Motandra guineensis (Fig. 14B)an dBaissea myrtifolia (Fig.13F) ;prominen ta si n Holarrhena pubescens andTabernaemontana pachysiphon (Figs. 13N, 14N, 15B); abaxial ± straightasia Ancylobotrys taylorisandPleiocarpapycnantha (Figs.13E ,14E ,15E) , veryprominen ta si nTabernaemontana elegans(Fig . 15F);outlin e± straigh to nbot h sides as in Diplorhynchus condylocarpon and Landolphia watsoniana (Figs. 13L, R).

Vascular bundle bicollateral, usually consisting of one median crescentic bundle (Figs. 13, 14) but may have small accessory bundles as in Pleiocarpa pycnantha (14E);usuall y inth efor m of ashallo w arca si n Carissa edulis (Fig. 13G)althoug h

WageningenAgricultural UniversityPapers 96-1 (1996) 7 it may rarely be crenate as in Pleiocarpa bicarpellata (Fig. 14D), comparative­ ly deep as in Mascarenhasia arborescens (Fig. 14A) and v-shaped as in Tabemaemontana (Figs. 14M, N, O); ends of bundle in contact with one another, forming a cylindrical bundle as inLandolphia and Clitandra(Figs . 13J, Q),i n this caseth eto ppar to fth ebundl ei spredominantl y composedo ffibres (Fig . 15C);wit h a lower crescentic bundle and five smaller ones at the top as in Saba comorensis (Fig.15D ); i nClitandra cymulosa th eoute rphloe mi ssurrounde db ylignifie d tissue.

Phloem: generally surrounds the xylema s in (Figs. 13,14), but may extend across the bundle as in Ancylobotrys, Tabemaemontana and Landolphia (Figs. 13D, E, 14N, O); may also form islands of phloem in the middle of the bundle as in Dictyophleba lucida andTabemaemontana elegans (Figs. 13K, 14M, 15F);th e is­ landso fphloe mma yalternat ewit hthick-walle dmucilaginou sfibres a si n Oncinotis tenuiloba (Fig. 16A) and Motandra guineensis.

Fibres:mucilaginou sfibres usuall ypresen ti ngroup so f 1-20immediatel ybelo wth e vascular bundle (Fig. 16F), may extend to the abaxial side which usually has less fibres. Lignified, thick-walled fibres present on the adaxial side of the bundle in Carissa tetramera.

Laticifers: (Fig. 17)Th e laticifers are typically branching tubes with thin, smooth walls and no dividing transverse walls (Fig. 17B, C, E).Th e degree of branching varies from species to species and may be very branched as in Funtumia elastica, (Fig.17E )t oles sbranche da si nLandolphia. Thelaticifer s aremostl ynarrowe rtha n the surrounding cells (6-13 um diam.) but may be very wide as in Carissa and Acokanthera (32 um diam.) andbecom e increasingly narrower towards the epider­ mis (Fig. i7B).They ramify throughout the mesophyll, commonly following the veins (Fig. 17A, D) and ramify outwards towards the epidermis as in Adenium obesum (Fig. 17B) or towards the palisade as in Holarrhenapubescens. Also ob­ servedadjacen t toth einne rpericlina lwall so fth eadaxia lepidermis ,abov eth epali ­ sade tissue as in Strophanthus courmontii and Hunteria zeylanica; present in the groundtissu eo fth epetiol eramifyin g towardsth eepidermi s asi nH. pubescens and Ancylobotrys petersiana (Fig 17C).

Crystals: (Fig. 18), clustered, solitary, small prismatic or rhomboidal crystals of calciumoxalat erecorded .Clustere dcrystal soccu ri nlarg eidioblast sa si n Hunteria zeylanica, mainlyi nth epalisad e tissue(Fig . 17E, 18A);i nenlarge d cells inth efirst layero fth espong ymesophyl la si nHolarrhena pubescens ori nth eentir emesophyl l asi nFuntumia elastica and Oncinotis tenuiloba (Fig. 18B);i nlayer so f 2-4 crystals above the minor veins asi n Rauvolfia caffra and R. mombasiana (Fig. 18E); in the ground tissue of the midrib especially in Schizozygia coffaeoides and Funtumia elastica. Smaller crystals common in the layer above the lower epidermis in most species. Solitary crystals occasional in both epidermis of Pleiocarpa bicarpellata and Hunteria; solitary and crystals of irregular shape frequently abundant in the in thephloe mo fmos to fth especie sinvestigate dan di nth egroun dtissu eo fth emidri b

8 WageningenAgricultural UniversityPapers 96-1 (1996) of Motandra guineensis alternating with clustered crystals; along the veins in Dictyophleba lucida (Fig. 18D).

Margin: (Fig. 19) usually straight as in Acokanthera oppositifolia (Fig. 19A) to revolute asi nSchizozygia coffaeoides (19B); slightly revolute (downcurved) at the tip with arounde d apex asi n Carissatetramera (Fig. 19C).Epiderma l cellsma y be small as in Diplorhynchus condylocarpon and Baissea myrtifolia (Fig. 9Q); ex­ tremely large and with an obtuse apex asi nAdenium obesum (Fig.9N , 19D), acute asi n Carissa edulis to long, pointed and papillose as inAcokanthera oppositifolia (Fig. 9M, 19A). Marginal vein with a well developed sclerenchymatous bundle sheath of several layers in Carissa tetramera (Fig. 19C) and A. oppositifolia, and Dictyophleba lucida (Fig. 19C).

Petiole: (Figs.2 0 & 21), Outline varies from ridged onth e abaxial and canaliculate on adaxial asi n Funtumia elastica (Fig. 20G) or straight on adaxial asi n Adenium obesum (Fig.20B )t osubglobos ea si nSchizozygia coffaeoides (Fig.2 iD )t ocircula r asm Dictyophleba lucida (Fig. 20F).Vascula rbundl ebicollateral ,o fon e crescentic median bundle which may be shallow as in Adenium (Fig. 20B); deep and with incurvededge sa si nHunteria andPleiocarpa (Figs.20H ,21C )t oalmos tcylindrica l asi nAncylobotrys, Dictyophleba orMotandra (Fig.20C ,F ,2 1A) ;t ocylindrica l but flattened onth eadaxia l sidet ofor ma semi-circl ea si n Clitandracymulosa and Saba comorensis (Fig. 20D, 2iB) .Vascula r strand strongly supported in the pericyclic region by thick-walled mucilaginous fibres inHunteria, Pleiocarpa (Fig. 16C)an d Picralima. Cluster crystals present in the ground tissue of the petiole in Funtumia elastica, Mascarenhasia arborescens (Fig. 18C), Schizozygia coffaeoides and in Motandra guineensis, styloids also present.

YOUNG STEM (Transverse section): The stem of 20 species of Apocynaceae was examined. All had the similartyp e of arrangement incros s section (Figs.22E ,F) ,wit hth eepidermi soutermost ; followed byth ecortex ,sometime swit hsolitar yo rgroupe dston ecells ;the nth epericycl ewit h white, sometimes mucilaginous, unlignified fibres, mostly in groups; followed by the phloem; then the xylem which forms a continous cylinder traversed by narrow rays; all the species have an intraxylary phloem at the margin af the pith; pith fre­ quently containing sclerosed elements as in Acokanthera, Carissa, Hunteria, Landolphia, Picralima, Pleiocarpa and Tabemaemontana (Figs.2 2D ,F) .Accord ­ ingt o Mecalfe and Chalk (1950), no pericyclic fibres were seen Rhazya orientalis (Dene.)A .DC .Amsonia orientalis Dcne.Latifers arepresen ti nal lth especie sexam ­ inedan di sgenerall y situatedi nth eprimar y cortex,pericycle ,phloe man dpit h (Fig. 17F).

WageningenAgricultural UniversityPapers 96-1 (1996) 1.5 CHARACTERISTICS OF THE GENERA

1.5.1 Acokanthera G.Don

(Figs. 3D, 6E, F, 8A, B, 9A, B, M, 10A, M, 12E, 13A, 17A, 19A, 20A;Table s 1, 2, 3;Appendi x 1)

Agenu so f 5species ,mainl ytree so rshrub sfro mYeme nt oSout hAfrica . Iti srepre ­ sented by three species in East Africa. A. oppositifolia is examined here.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM): more or less smooth but very uneven on both surfaces. STOMATA (Table 2):hypostomatic ; paracytic, with 2-3 pairs of subsidiary cells and cyclocytic-teracytic, sometimes with very complex subsidiary cells Figs. 6E,F) .outlin e broadly elliptic to circular and rounded at the poles, with wide rims and a round aperture; subsidiary cells obscured by thick peristomatal rims; size 15-22 x 11.5-16 [im; density per mm2 650-719. EPIDER­ MAL CELL OUTLINE: indistinctly visible as grooves. ANTICLINAL WALLS (LM): straight to slightly curved on both sides (Figs. 6E,F) .

T.S. lamina THICKNESS (Table 2) 384-458 urn thick. CUTICLE: adaxial 19-26.6 um thick, abaxial9.4-1 5 umthick ,ver ythic ka tth emidrib ,eve nthicke ra tth emargi na t22-4 8 um. ADAXIAL EPIDERMAL CELLS: 1-layered,rectangular , 11-19 x 19-40 urn, much smaller at the midrib. ABAXIAL EPIDERMAL CELLS: i-layered, 11.4- 10 x 57-28.5 (im,muc h smallertha n adaxial,eve n smaller atth emidrib ,triangula r and acute at the apex. STOMATA (T.S.) (Fig. 10A): with a very thick cuticular ledgean dwid eaperture .MESOPHYLL :bifacial ; palisade extendst oth emidrib ,i n 2(-3) rows, with 1-2layer s of poorly developed palisade on the abaxial side (Fig. 12E);spong ymesophyl lwit hsomewha thorizontall y elongatedcells ,wit hampl eai r spaces. MINOR VEINS: small veins collateral with xylem on the adaxial and phloemo nth eabaxia l side;large rvein sbicollateral ,wit ha lo to ffibre s onth elatera l and abaxial side of the xylem tissue. MIDRIB (Fig. 13A;Tabl e 1): 677-732 um thick; abaxial outline almost flat to slightly curved;abaxia l with a slightridge. Vas ­ cular bundle bicollateral, forming a shallow arc, with 6-10mucilaginou s fibres on theadaxia l side,xyle mtissu ewit h alo to f thick-walled,xyle mfibre s onth e adaxial and lateral sides (Fig. 17A). Collenchyma on the adaxial periphery of the midrib. MARGIN: straight,wit h amargina lvei n surrounded with severallayer s of scleren- chymatous tissue; cuticle very thick; epidermal cells papillose, pointed (Fig. 19A) CRYSTALS:cluste rcrystal s (druses)presen t inth epalisad e layer,especiall y inth e 2nd row and the layer immediately above the lower epidermis. LATICIFERS: al­ ways accompany the veins, quite prominent at the midrib (Fig. 17A), branching throughout the mesophyll towards the adaxial or abaxial epidermis, becoming in­ creasingly narrower atth epoin t of branching.

10 WageningenAgricultural UniversityPapers 96-1 (1996) T.S. petiole Vascular bundle bicollateral, forms a relatively deep arc with incurved edges (Fig. 20A), with small accessory bundles predominantly surrounded with thick-walled fibres; Laticifers observed along the veins asi n themidrib ; ground tissue predomi­ nantly parenchymatous, lignified, with pits.

Comments: Thisspecie sexibit svariou sxeromorphi c characters sucha scoriaceou sleaves ,thic k cuticle and outer epidermal cell walls as well as a tendency towards being isobilateral.Th e stomata has a very thick cuticular ledge.

1.5.2 Adenium Roem. & Schult.

(Figs. 8C,D , 9C D, N, ioB, 13B, 17B, 19B, 20B;Table s 1& 2, 3;Appendi x 1)

A genus of 6species ,mainl y succulent shrubs ortree s upt o 5m talli n Africa from Senegali nth eWes tt oSouth Africa . Representedb yAdenium obesum inEas t Africa which extendst o Socotra andth eextrem e southern part of Arabia, andi s examined here.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM):adaxia l surface smooth, abaxial striateove rth ewhol esurface .STOMATA :hypostomati can dparacytic ,occasionall y subsidiary cells subdivided by radiating walls on one or both sides or sometimes cyclocytic-staurocytic with 3-4 cells around the stomata; outline elliptic; size 23- 33.5 x 19-22.5 [im; density per mm2 150-188. EPIDERMAL CELL OUTLINE (SEM): adaxial with cuticular ridges over anticlinal walls, smooth within the cells. ANTICLINAL WALLS (LM): adaxial straight angular, abaxial straight to very slightly curved.

T.S. Lamina THICKNESS (Table 1,Fig . 13B):403-43 9 um.CUTICLE :thickes t onadaxia l epi­ dermis and leaf margin, thinnest on lower epidermis. ADAXIAL EPIDERMAL CELLS (Fig. 8C): regular, i-layered, very large, 43.7-68.4 x 28.5-61.8 urn, taller thanwide ,smalle ra tth emidrib .ABAXIA LEPIDERMA LCELL S (Fig.8D) :irreg ­ ular, i-layered, 19-28.5 x 15.2-57 um,widertha ntall .STOMAT A(TS. )(Fig .10B) : slightlyraise dan dwit ha wid eaperture .MESOPHYLL :bifacial ;palisad elayer s 1-2 , does not extend to the midrib; spongy mesophyll cells more or less irregularly ar­ ranged and quite compact. MINOR VEINS: collateral, with a parenchymatous sheath.MIDRI B (Fig. 13B; Tabl e 1) : 1245-126 3u mthick ,wit ha prominen t abaxial ridge and a slight adaxial projection.Vascula rbundl e crescent-shaped, bicollateral; ground tissue of midrib collenchymatous towards the periphery, especially on the abaxial side.MARGIN :slightl yrevolut ea tth etip ,wit hcollenchym aa tth eacut etip ; epidermal cellstalle rtha n widewit ha rounde d apex (Figs.9N , 19D).CRYSTALS : none seen. LATICIFERS (Fig. 17B): many, branched throughout the mesophyll;

WageningenAgricultural University Papers 96-1 (1996) 11 alongth eveins ;branchin gbelo wth epalisad ean depidermis ;betwee nth epalisad e andepiderma lcell san dtoward sth eabaxia lo radaxia lepidermis .

T.S.petiole Vascularbundl ebicollateral ,formin g ashallo w arc,wit haccessor ybundle sa tth e wings (20B) ;collenchym a cellsa tth eperiphery ; epidermalcell sver ysmall .

Comments:th eleave sexamine dhav en ohairs ,bu tth eleave so fthi sgenu sar eusu ­ ally glabrous to hairy.Nyawuam e &Gil l (1991) recorded anomocytic stomatai n Adeniumobesum wit hstraigh tanticlina lwall so nbot hlea fsurface s andth eagenou s typeo f stomatal development.

1.5.3 AlafiaThouars

(Fig.90 , 13C;Tabl e 1,2 ;Appendi x 1)

Agenu so f2 3lianescen tspecie si nAfrica . Representedb y5 speciesi nEas t Africa. A. microstylisi sexamine dhere .

Leafsurface HAIRS: absent. CUTICULAR SURFACE (SEM): smooth. STOMATA: hypostomatic;paracytic ;outlin eelliptic , size 18-25x 19-20urn ;densit y permm 2 140-150.EPIDERMA LCEL LOUTLIN E(SEM) :undulat eANTICLINA LWALL S (LM): bothundulate .

T.S. lamina THICKNESS (Table 1)256-30 2 ujn. CUTICLE: 1.9-2.9ur nthic k on both sides, 3.8-4.8Ji mthic karoun dth emidrib .ADAXIA LEPIDERMA L CELLS: i-layered, 13.3-18x 13.3-34.2um ,rectangular ,2-layere da tth emidri ban dlea fmargin ,thick - walled. ABAXIAL EPIDERMAL CELLS: i-layered, 8.6-12.4x 13.3-32.3 um, narrower. STOMATA (T.S.):mor e or less level with the surface. MESOPHYLL: bifacial;palisad e 15-36x 4-1 3fim , in i-2(-3)rows ,extend st oth emidrib ;spong y mesophyll with somewhat horizontally elongated cells, not compact. MINOR VEINS: embedded, collateral, with xylem surrounded with thick-walled fibres, phloemabaxial ,wit ha noute rparenchym a sheath;large rmino rvein sbicollateral . MIDRIB(Tabl ei;Fig .13C) :402-43 9[i mthick ;adaxia loutlin ewit ha sligh tdepres ­ sion, abaxial slightly projecting.Vascula r bundle bicollateral forming a shallow crescenticar cwit hisland so fmucilaginou sfibres below .Collenchym apresen tonl y onth eadaxia lperiphery .MARGIN :straight ,epiderma lcell ssmalle r(Fig .90 ); cuti n thick.CRYSTALS : afe wlarg eclustere dcrystal si nth emesophyll ,smalle rone so n thero wabov elowe repidermis ;a fe wrhombohedr aan dman ysmal lclustere dcrys ­ talsi nth ephloem .LATICIFERS :accompan y theveins ;branchin gbelo wth epali ­ sade;betwee n epidermalcell stoward sbot hepidermis .

12 WageningenAgricultural UniversityPapers 96-1 (1996) T.S. petiole Vascular bundle bicollateral.

Comments: N yawuam e& Gil l( 199 1) reporte dparacyti c stomata,wit hsinuou santi ­ clinalwall so nbot hsurface san dth eeumesogenou sontogeneti ctyp eo fdevelopmen t in Alafia barteri, A. multifiora and A. landolphioides (as A. scandens). Stomatal sizes ranged from 23.2-25.8 x 13.6-19.8 [im.

1.5.4 Ancylobotrys Pierre

(Figs. iC, 4D, 5E, 8E, F,9E , F, 10C, D, 12C, 13D , E, 15E, 20C;Table s 1& 2,3 ; Appendix 1)

A genus of 7 species,mainl y lianas or sarmentose shrubs restricted to Africa, with A. petersiana spreading toth eComor o Islands andMadagascar . Iti srepresente d by three species in E. Africa, A. petersiana andA. tayloris are observed here.

Leaf surface HAIRS:presen t on thepetiol eo f both species,2-7-celled ,thin-walle d inboth . CU- TICULAR SURFACE (SEM): adaxial of A. tayloris with small bumps, abaxial smooth but with thick cuticular ridges over subsidiary cells (Fig. 5E) . STOMATA (Fig. 4D, 5E): hypostomatic, anomocytic in A. petersiana, outline circular in A. tayloris and with a round aperture. EPIDERMAL CELL OUTLINE (SEM): not visible on adaxial surface inA. tayloris. ANTICLINALWALL S (LM):ver y undu­ late on both sides inA. petersiana, undulate on both sides inA. tayloris.

T.S. Lamina THICKNESS (Table 1): 229-300 u,mi nA. tayloris, 293-320 fim inA. petersiana. CUTICLE: thicker in A. tayloris, thickest on adaxial midrib in both. ADAXIAL EPIDERMALCELL S (Figs.8E ,9E) :intermittentl y 2-layeredi nA .tayloris, second layerlarger ,thick-walled ;1 -layere di nA .petersiana, 2-3-layeredtoward sth emargi n andth emidrib ;oute rpericlina l wallo f thefirs t layer very thick inboth . ABAXIAL EPIDERMAL CELLS (Figs.8F ,9F) : i-layeredi nbot hbu twit h 1-2 layers of short, compact cells above the lower epidermis. STOMATA (T.S.) (Figs. 10C, D):wit h a wider aperture inA . tayloris. MESOPHYLL: bifacial in both, but with a tendency towardsbein gisobilatera l inboth ;palisad e 2(-3)-layered inboth ,doe sno texten dt o the midrib; spongy mesophyll composed of more or less vertically elongated cells with a lot of air spaces (Fig. 12C). MINORVEINS : embedded or transcurrent and bicollaterali nth elarge rvein si nbot hspecies .MIDRI B (Figs. I3D,E;Tabl e 1)549- 586 u,mi nA .tayloris, 714-750 uini nA .petersiana; outline almost straight onbot h sides in A. tayloris; adaxial gently curved, abaxial prominently ridged in A. petersiana. Vascular bundle bicollateral, forms a shallow arc in A. tayloris and a deeper one inA .petersiana; phloem extends across the vascular bundle and forms a semi-circle above iti nbot h species (Figs. 13D,E , 15E)MARGIN : slightly revo- lute in both, with an acute tip in A. tayloris. CRYSTALS: solitary crystals in the

WageningenAgricultural UniversityPapers 96-1 (1996) 13 phloem and a few in the ground tissue below vascular bundle in A. petersiana. LATICIFERS:alon gveins ;branchin gi nth epalisad etoward sbot hepidermis ;belo w palisadetissu ean depidermi si nbot hspecies .

T.S.petiole Endso fvascula rbundl ealmos ti ncontac twit hon eanothe rformin ga nalmos tcylin ­ dricalstran dwit haccessor ybundl ei nbot h(Fig .2 0C) ;laticifer sbranchin gbetwee n cellsi nth egroun dtissue . Comments: Both species have a very thick cuticle and outer epidermal cells andbot h have a tendency towards being isobilateral. A. taylorisca n be distinguished from A. petersianab yit slarge rstomat awit ha wider ,circula raperture ;2-layere depidermi s anda midri bwhic hi salmos tstraigh to nbot hsides .

1.5.5 BaisseaA.DC .

(Figs, il, 3E,7C , D,8G ,H ,9Q , 13 F;Table s 1,2 ,3 ; Appendix1 )

Agenu s of 18mainl y lianescent species,mainl y climbers or scandent shrubsre ­ strictedt oAfrica . Representedb yfive specie si nEas tAfrica .B. myrtifolia i sexam ­ inedhere .

Leafsurface HAIRS: occuro nadaxia lsurfac eo fmidri ban do npetiole ;multicellular ,uniseriate , sometimeshooke dwit hlonge rbasa lan dshorte rapica lcell s(Fig .il) ;domati apres ­ enti nth eaxil so fsecondar yvein s(Fig .3E) .CUTICULA RSURFAC E(SEM) :mor e orles s smoothbu t with uneven surface onbot h sides.STOMAT A (Table 2;Figs . 7C,D):hypostomatican dparacytic ;outlin eelliptic ;sizei5-i9. 5 x8-1 3ujn ;densit y permm 2150-200 .EPIDERMA LCEL LOUTLIN E(SEM) :no tvisible .ANTICLI ­ NALWALL S(LM) :ver yundulat eo nbot h surfaces (Figs.7C , D).

T.S. lamina THICKNESS(Tabl e1) :82-11 0|im .CUTICLE :1- 3 jimthick ,thickes ta tth emidrib . ADAXIALEPIDERMA LCELL S(Fig .8G) :i-layered ,rectangular ,wit ha nirregu ­ lari noutline .ABAXIA LEPIDERMA LCELL S(Fig .8H) : 1-layered,rectangular , witha nirregula routline .STOMAT A(T.S.) :slightl yraised .MESOPHYLL :bifacial ; palisade1 -layered ,extend st oth emidrib ;spong ymesophyl lwit hmor eo rles strans ­ verselyelongate dcells .MINO RVEINS :wit ha parenchym asheath . MIDRIB(Fig . 13F): 201-238u mthick ;outlin ealmos flatt o nadaxia lsurface ;abaxia lprominentl y ridged.Vascula rbundl ebicollateral ,formin g ashallo warc .Groun dtissu eo fmidri b collenchymatoustoward sth eperiphery . MARGIN:slightl y revolute.CRYSTALS : cluster crystals present in the palisade layer; cluster and solitary crystals in the spongymesophyll .LATICIFERS :alon gth eveins ;belo wth epalisade ;branchin gi n themesophyl ltoward sbot hepidermises .

14 WageningenAgricultural UniversityPapers 96-1 (1996) T.S. petiole

Vascular bundle u-shaped, bicollateral.

1.5.6 CarissaL.

(Figs. 4F, 5C,9G , H, 10E,F , 13G,H , P, 16B, 17D;Table s 1& 2,3 ;Appendi x 1) A genuso f about 20specie s mainly shrubso r smalltree si n Africa, Asiaan d Mada­ gascar. Represented by three species inEas t Africa, C.edulis and C.tetramera are investigated here.

Leaf surface

HAIRS:ver y few on adaxial midrib of C. tetramera ando n petiole of both species; unicellular, simple and thin-walled. CUTICULAR SURFACE (SEM): smooth on both sides in C. tetramera and on abaxial surface of C. edulis. STOMATA: hypostomatic but in C.edulis afe w stomata found on abaxial surface overlying the midrib and on adaxial surface of petiole; cyclocytic in C. edulis, paracytic in C. tetramera; outline broadly elliptic in both, and with atruncat e polei n C. tetramera (Fig.5C) ;siz elarge r in C.edulis at 29-36 x 23-32 ujn, sunken with anarro w aper­ ture (Fig. 4F);densit y per mm2 500-572 in C. tetramera, 66-176 mm2 in C. edulis. EPIDERMAL CELL OUTLINE (SEM): outline obscurely visible on both sides. ANTICLINAL WALLS (LM) adaxial ± straight, angular in C. tetramera, abaxial straight to slightly curved; straight to slightly curved on both surfaces in C. edulis.

T.S. lamina THICKNESS (Table 1): 247-302 [xmi n C. edulis; 366-412 [im in C. tetramera. CUTICLE: thicker in C. tetramera on the adaxial midrib at 9-20 um. ADAXIAL EPIDERMALCELL S (9G): i-layeredi nboth ,large ri nC .tetramera, triangula ran d acutea tth eape xi nC. edulis.ABAXIA LEPIDERMA LCELL S(Fig .9H) : 1-layere d inboth ,rectangula r inC. tetramera, smalleran dtriangula r inC. edulis. STOMATA (T.S.) (Figs. 10E, F): raised, with a smaller aperture in C. edulis. MESOPHYLL: bifacial in both species, C. edulis with a tendency towards being isobilateral; pali­ sade cells 2(-3)-layered, characteristically long, extends up to the midrib in both species; spongy mesophyll with transversely elongated cells in C. edulis; C. tetramera with shorter cells with arms; many transversely elongated thick-walled scleridsi nth epalisad e andmesophyl l of C. tetramera (Fig. 16B) .MINO RVEINS : smaller ones collateral with aparenchym a sheath; larger ones in C.tetramera with an outer parenchyma sheath and an inner sclerenchymatous sheath.MIDRI B (Fig. 13 G, H): almost fiat on adaxial surface in both species, more ridged on abaxial surface inboth .Vascula r bundlebicollatera l andformin g a shallow arc in both spe­ cies, withthick-walle d lignified fibres above and below thebundl e inC. tetramera. Collenchyma only on abaxial periphery of midrib. CRYSTALS:cluste r crystals in the mesophyll of both; solitary crystals in the phloem and mesophyll of C. tetramera. LATICIFERS:alon g veins in the midrib (Fig. 17D) and mesophyll; be-

WageningenAgricultural UniversityPapers 96-1 (1996) 15 tween epidermal cells;belo w palisade;toward s the upper and lower epidermisi n bothspecies ;widt hdecreasin ga tth epoin to fbranching .MARGIN :revolute in bot h species,ape x bluntly obtuse inC. edulis androunde d inC. tetramera (Fig. 19C); epidermal cellstriangula rwit h acuteape x inC. edulis (Fig.9P) ;margina l bundle surroundedb y alarg eamoun to f sclerenchyma inC. tetramera (Fig.19C) .

T.S.petiole Vascular bundle forming a deeper and slightly incurved bundle in C. tetramera; epidermalcell ssmall ,triangular ;laticifer s observedalon gth evein si nth emidrib .

Comments:C. tetramera iseasil y distinguishedb yth efolia r scleridsi nth emeso - phyllo fth eleave s(Fig . 16B)an dit svenatio nwit ha stron gmargina lvei n(Fig . 22 A).Th e stomata are smaller in C. tetramera which also has a higher density. Nyawuame& Gil l( 199 1) reporte dth eparacyti ctyp eo fstomat awit hstraigh tanticli ­ nalwall so nbot hside san deumesogenou styp eo fstomata ldevelopmen ti nC. edulis and sizeo f 30.1x 17.4 um.

1.5.7 CarvalhoaK . Schum.

(Figs.2E ,5F ,81 ,J , 10G, 13I;Table s 1,2 ,3 ;Appendi x1 )

Amonotypi cgenu srestricte dt oEaster nan dSout hEaster nAfrica .I nEas tAfric ai t isrestricte dt oKeny aan dTanzania .Th eonl yspecies ,C. campanulata,i sexamine d inth epresen tstudy .

Leafsurface HAIRS:absent .CUTICULA R SURFACE(SEM) :adaxia l with striations overth e whole surface, striations crossing cell outlines (Fig. 2E);abaxia l with concentric rings striaeove rsubsidiar ycell san dwit hcuticula rridges o nth eres to fth e surface (Fig.5F) .STOMAT A(Fig .5F ;Tabl e2) :hypostomatic ;paracyti cwit hon esubsid ­ iary cell on either side, subsidiary cells extending over oneo rbot hpoles ;outlin e elliptic;siz e20-2 5x J3-i9M™ ;densit ype rmm 281-106-150 .EPIDERMA LCEL L OUTLINE (SEM): adaxial with cuticular ridges over anticlinal walls (Fig. 2E), abaxial with irregular cuticular ridges. ANTICLINAL WALLS (LM): adaxial straightt oslightl y curved, abaxialundulate .

T.S. lamina THICKNESS (Table 1; Fig 81, J): 156-220 um. CUTICLE: very thin, c. 1urn . ADAXIAL EPIDERMAL CELLS (Fig. 81): regular, 1-layered, wider than tall, collenchymatousa tth emidrib .ABAXIA L(Fig .8J) : i-layered.STOMAT A(T.S. ) (Fig. 10G):slightl yraised . MESOPHYLL:bifacial ;palisade , i-layered,extend st o the midrib; spongy mesophyll with short transversely elongated cells. MINOR VEINS: collateral,wit hxyle mo nadaxia lside .MIDRI B(Fig .13I ;Tabl e 1):1409 - 1446 um thick; adaxial ridge small, less prominent, abaxial more prominent, u- shaped.Vascula rbundl ebicollateral ,formin g adee parc ;collenchym ai n2- 3layer s

16 WageningenAgricultural UniversityPapers 96-1 (1996) on both surfaces. MARGIN: revolute, with 2-3 layers of collenchymatous cells; epidermal cells subglobose. CRYSTALS: cluster crystals in the palisade layer. LATICIFERS: along the veins; in the mesophyll and among parenchyma cells of ground tissue.

T.S. petiole Vascular bundle bicollateral, u-shaped, with accessory bundles on the wings.

NOTES:Th e petioles and leaves of C.campanulata sometimes have some pubes­ cence.

1.5.8 Clitandra Benth.

(Figs. 8K, L, 9R, 13J, 20D;Table s 1& 2, 3;Appendi x 1)

A monotypic genus restricted in Africa from Guinea toTanzania . C. cymulosa, a large liana, is examined.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM): STOMATA: hypostomatic; anomocytic with 5-8 cells around the stomata; outline broadly elliptic to rounded; size 17-23x i5-i9nm;densit ypermm 2288-340 .EPIDERMA L CELLOUTLINE: not visible. ANTICLINAL WALLS (LM): more or less straight to very slightly curved on both sides.

T.S.lamina THICKNESS (Table 1): 192-247(im .CUTICLE :thickes ta tth emargin .ADAXIA L EPIDERMAL CELLS (Fig. 8K):± square, smaller atth e midrib.ABAXIA L EPI­ DERMALCELL S(Fig .8L) :rectangula ri nshape ,muc hsmalle ra tth emidrib .STO ­ MATA (T.S.):slightl y sunken stop.MESOPHYLL : bifacial, but with two layers of vertically elongate shortpalisade-lik e cells above abaxial epidermis;palisad e layer in 2rows ,doe s not extend toth e midrib; spongy mesophyll with horizontally elon­ gated cells, not compact. MINOR VEINS: smaller ones embedded, with a paren­ chyma sheath; larger ones transcurrent towards the adaxial side. MIDRIB: 640- 659 urn thick; outline more or less straight on adaxial side, prominent on abaxial side,acute .Vascula rbundl ebicollateral ,formin g anova lshape .MARGIN : straight. CRYSTALS: none seen. LATICIFERS: branching in the mesophyll and palisade layers towards both epidermis.

T.S. petiole Vascular bundle bicollateral, cylindrical and flattened on adaxial side forming a semi-circle (Fig. 20D).

WageningenAgricultural UniversityPapers 96-1 (1996) 17 1.5.9 Dictyophleba Pierre

(Figs. iF, 4A, 6G,H , 13K, 18D , 19C, 20F;Table s 1,2 , 3;Appendi x 1)

Agenu s of 5 species in Africa, all lianas with tendrils.Onl y one species,D. lucida, extends to East Africa: Tanzania and Kenya.

Leaf surface HAIRS (Fig. iF):presen t onth epetiole ,u pt o 25-celled,cell s short. CUTICULAR SURFACE (SEM) (Fig. 4A): smooth. STOMATA (Fig. 6 G, H;Tabl e 2):hyposto - matic and anomocyticwit h 3-6cell s aroundth eguard cells ;outlin ealmos t circular; size 14.6-16.6 x 14.6-16.6 um;densit ype rmm 2700-825-925 .EPIDERMA LCEL L OUTLINE (SEM),cel loutline sobscurel y visiblea swid egroove s(Fig .4A) .ANTI ­ CLINALWALL S (LM): undulate on both surfaces.

T.S. lamina THICKNESS (Table 1): 183-311 H™.CUTICLE : very thin, 1-3urn . ADAXIAL EPIDERMAL CELLS: i-layered, much larger than abaxial ones.ABAXIA L EPI­ DERMAL CELLS: i-layered, rectangular in outline. STOMATA (T.S.): raised. MESOPHYLL: bifacial; palisade layer in 2-rows, extends to the midrib; spongy mesophyll with vertically elongated, lacunate cells.MINO RVEINS : transcurrent, with outer parenchyma sheath and inner sclerenchymatous sheath; bundle sheath extensionstoward sbot hepidermis .MIDRI B (Fig. 13K):714-78 7ur nthick ; slightly concaveo n adaxial surface; abaxial ridge prominent, with anirregula routline .Vas ­ cularbundl eformin g awid eu-shape ,bicollateral ,wit hisland so f phloemwithi nth e u-shape (Fig. 13K).Collenchym a in 3-4 layers on both sides of the midrib. MAR­ GIN (Fig. 19C): revolute with an obtuse tip, marginal vein with sclerenchyma. CRYSTALS: solitary crystals arranged in a neat row along the veins in the meso­ phyll.LATICIFERS :man y laticifers seenespeciall y branching from the mesophyll to the palisade and to the upper epidermis and margin.

T.S. petiole Outline of petiole circular; ends of vascular bundle almost in contact with one an­ otherformin g analmos tcylindrica lstran dwit haccessor ybundle sa tth ewing so fth e petiole (Fig. 20F).

1.5.10 DiplorhynchusWelw .e x Fie. & Hiern

(Figs. 2A, 5D, 9K, L, 12F, 13L;Table s 1, 2, 3;Appendi x 1)

A monotypic genus with its only species,D. condylocarpon, restricted to Africa, which hasbee n examined.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM): adaxial smooth with very fine

18 WageningenAgricultural UniversityPapers 96-1 (1996) striatums,abaxia lsmoot h(Figs .2A ,5D) .STOMAT A(Tabl e2) :hypostomatic ;type ; outlinebroadl y elliptic withperistomata lrims (Fig . 5D);siz e 16-18x 14-16mm ; densitype rmm 2260-340 ;EPIDERMA LCEL LOUTLIN E(SEM) :no tvisible .AN ­ TICLINALWALL S(LM) :slightl y undulate.

T.S. lamina THICKNESS (Table 1):256-27 5 urn.CUTICLE :2.9-5. 7l« no nth euppe repider ­ mis;1.9-5. 7ur ni nth elowe repidermis ;thickes ta tth emargin .ADAXIA LEPIDER ­ MALCELLS : i-layered,rectangula ri nshape ,7.6-15. 2x 7.6-28. 5urn ,smalles ta t themidrib ,triangular . MESOPHYLL (Fig. 12F):bifacial ; palisadei n 2(-3 )rows , very narrow; mesophyll vertically elongated. MINORVEINS : transcurrent with bundlesheat hextension sreachin gtoward sbot hepidermises .MIDRI B: 439-45 8ur n thick;outlin emor eo rles sstraigh to nbot hsides .Vascula rbundl ebicollateral ,form ­ inga wid ear cwit hphloe macros sth ear can dman yfibres belo wth evascula rbun ­ dle.MARGIN :mor eo rles s straight withmuc hcollenchym a atth etip ;epiderma l cellsnarrow ,small ,acut ea tth eapex .

T.S. petiole Vascularbundl ebicollateral .

Comments: theleave sar enormall yglabrou st opubescent ;petiole sar eglabrescen t topuberulent .

1.5.11 FuntumiaStap f

(Figs.8M ,N , 13M, 17E,20G ;Table s 1,2 ,3 ;Appendi x 1)

Agenu so ftw otre especie srestricte dt oAfrica . F. elasticawa savailabl efo rstudy .

Leafsurface HAIRS: absent. CUTICULAR SURFACE (SEM): rough. STOMATA: hypo­ stomatic;paracytic ;outlin e narrowly elliptic; size 19-25x 10-15M™ ; density per mm2300-320 .EPIDERMA L CELLOUTLINE :completel y obscured.ANTICLI ­ NALWALLS :straigh tt oslightl y undulateo nbot hsides .

T.S. Lamina THICKNESS (Table 1): 311-366(im . CUTICLE: thickest on adaxial midrib. ADAXIALEPIDERMA LCELL S(Fig .8M ,9I) :irregularl y2-3-layered ,sometime s withpericlina l divisions, thick-walled, 7.6-24.7x 13.3-24.7 urn.ABAXIA LEPI ­ DERMAL CELLS (Fig. 8 N): 1-2-layered, thick-walled, smaller, 7-11.4x 10- 20(im ,eve nsmalle raroun dmidrib ,acut ea tth eapex .STOMAT A(T.S.) :leve lwit h the surface. MESOPHYLL:bifacial ; palisade extends toth e midrib,i n 2-3-rows, characteristicallylong ;spong ymesophyl lwit htransversel yelongate dcell swit hai r spaces.MINO RVEINS :embedded ,large rone sbicollateral ,smalle rone scollateral , witha parenchym asheath .MIDRI B(Fig .13M) : 586-768ur nthick ;adaxia loutlin e

WageningenAgricultural UniversityPapers 96-1 (1996) 19 almostflat, abaxia lridged ;Vascula rbundl ebicollateral ,formin g awid eshallo warc ; pockets of phloem interspersed with mucilaginous fibres above the arc, few fibres below it.Groun dtissu e collenchymatous towardsth eperiphery .MARGI N revolute with two layers of epidermal cells and collenchymatous cells.CRYSTALS : many clustercrystal spresen ti nth emesophyl lan dlowe rcorte xo f midriban dlarge rmino r veins. LATICIFERS(Fig. 17E): branching freely in the mesophyll and along the veins; branching just below the epidermis and palisade tissue into the epidermal cells.

T.S. petiole The vascular bundle forms a deep arc with adaxial phloem extending across the bundle and has three accessory bundles on each side of the wings; numerous clus­ tered crystals in the ground tissue of the petiole; outline deeply canaliculate on adaxial side (Fig. 20G).

Comments: Leaf may have afe w hairs on midrib.

1.5.12 Holarrhena R.Br.

(Figs. IB, 2F,4E , 6C,D , 12B, 13N, 15B)

A genus of 4 species in Africa, Southern Continental Asia and Sri Lanka. Repre­ sented by only one species,H. pubescens, confined to Kenya andTanzania , which is examined here.

Leaf surface HAIRS (Figs. iB, 4E):presen t onpetiole ,abaxia l leaf surface ando nbot h surfaces ofth emidri ban da tth emargin ; 1-2 (-7)-celled, thebasa lcell sar eshorte ran dtermi ­ nal cells longer and may be hooked. CUTTCULAR SURFACE (SEM) (Fig. 2F): abaxial striate; abaxial smooth. STOMATA (Figs. 6C, D;Tabl e 2): hypostomatic; paracytic with subsidiary cells extending over the poles, a few anisocytic stomata also seen; double stomata and stomata with single guard cell seen;outlin e rounded toelliptic ,slightl yraised ;wit hdoubl erims; apertur ewide ,elliptic ;siz e 17-31 x 14- 19 [im; density per mm2 250-289-312. EPIDERMAL CELL OUT LINE (SEM): with cuticular ridges over anticlinal walls (Fig. 2F).ANTICLINA LWALL S (LM): ± straight, angular on both surfaces.

T.S. lamina THICKNESS: 201-220 urn. CUTICLE: 1-3 um thick. ADAXIAL EPIDERMAL CELLS: 1-layered, at some points seen dividing periclinally, taller than wide or moreo rles ssquare .ABAXIA LEPIDERMA LCELL S: 1 -layered ,rectangular ,talle r than wide at the midrib. STOMATA (T.S.): slightly raised. MESOPHYLL (12B): bifacial; palisade layer 1-layered , extends to the midrib, but leaves a gap; spongy mesophyll cellstransversel y elongated, very compact.MINO RVEINS : embedded, with athin-walle dparenchym a sheath;olde r ones transcurrent towards adaxial epi-

20 WageningenAgricultural UniversityPapers 96-1 (1996) dermis.MIDRI B(Figs .13N , isB;Tablei):970-i02 5 |xmthick ;adaxia loutlin ewit h asmal lridge, abaxia lmor eprominent ,u-shaped .Vascula rbundl eformin ga shallo w arc.Collenchym ao nth eperipher yo fbot hridges. MARGIN :revolut ea tth eextrem e tipo rmor eo rles s straight.CRYSTALS :cluste rcrystal s in large subglobose cells immediatelybelo wth epalisad elayer .LATTCIFERS :i nth emesophyl lalon gveins ; belowth epalisad eeventuall y endingi nth euppe ro rlowe repidermis .

T.S. petiole: Vascular bundle forms a very wide and a shallow u-shape, with small accessory bundles;man ylaticifer ssee nbetwee ncortica lparenchym acell so fth epetiol eeven ­ tually endingi nth eepidermis .

Comments: the stomata are also reported to be paracytic in H. pubescensb y Nyawuame &Gil l (1991)an di nH. antidysenterica (=H. pubescens) byChandr a etal. ,(1969) ,wit heumesogenou styp eo fstomata ldevelopmen t(Nyawuam e& Gil l 1991).

1.5.13 HunteriaRoxb .

(Figs.3A , 18A, 10H ,1,13O ,P ,20H ,22A ;Tabl e 1,2 ,3 ; Appendix1 )

Agenu so f 12specie srestricte d toAfrica , onlyH. zeylanica extendst oSout han d South East Asia up to Sumatra. Represented by two species in East Africa, H. congolana andH. zeylanica, bothexamine dhere .

Leafsurface: HAIRS: absent. CUTICULAR SURFACE (SEM): smooth on both sides in H. congolana, smoothwit hfin e striatumsi nH. zeylanica. STOMATA: hypostomatic; anomocytican danisocyti ci nH. congolana, complex-cyclocytic withsun ksubsid ­ iarycell si nH. zeylanica(Fig .3 A);outlin ebroadl y elliptict ocircular ;large ri nH. zeylanica;densit ype rmm 2greate ri n H.zeylanica a t294-350 .EPIDERMA LCEL L OUTLINE:no t visible in both.ANTICLINA LWALL S(LM) :bot h walls straight tover y slightly curvedi nbot hspecies .

T.S. lamina Width183-22 0i nH. congolana,an d266-34 8\im i nH. zeylanica.CUTICLE :thick ­ esta tth emidri ban dmargi ni nbot hspecies . ADAXIALEPIDERMA LCELLS :1 - layeredi nbot h species,mor eo rles srectangula r inH. congolana, tallertha nwid e inH. zeylanica. ABAXIALEPIDERMA L CELLS: i-layeredi nboth ,rectangular , largeri nH. zeylanica; epiderma lcell spapillos ewit hacut eape xaroun dmidri bi nH. congolana. STOMATA(T.S. )(10H ,I) :raise di nbot hspecies .MESOPHYLL :bifa ­ cial,H. zeylanicawit htw olayer so fshor tpalisade-lik ecell sabov eth elowe repider ­ mis;palisad e 1-2-layeredi nH. congolana,extend st oth emidrib ;2-3(-4)-layere di n H.zeylanica; spongymesophyl lwit hshor thorizontall y elongatedcells ,ver ycom ­ pacti nbot hspecies .MINO RVEINS : embedded,wit ha parenchym asheath ,xyle m

WageningenAgricultural UniversityPapers 96-1 (1996) 21 surroundedb yfibres. MIDRI B( 13O,P ;Tabl e i):9i5-95i [imthic ki nH. zeylanica, 403-439 [im thick in H. congolana; adaxial outline more or less straight, abaxial ridged in both.Vascula r bundle bicollateral, forming a wide shallow arc in both species;mucilaginou s fibres present on lower sidei n H. congolana. Ground tissue collenchymatousespeciall y onth eabaxia l side.MARGIN :slightl yrevolut ei nboth , epidermal cells taller than wide, with acute apex in H. congolana. CRYSTALS: cluster crystals in the palisade layer in large subglobose idioblasts in both species, idioblastsextremel ylarg ei nH. zeylanica (Fig.18A) ;mesophyl lwit hcluste rcrystal s in enlarged cellsi nH. zeylanica; H. congolana withnumerou s drusesi nth e ground tissueo f midriban dmesophyl l andman y solitary crystals inth ephloe m (Fig. 18F); solitary crystals in both epidermis. LATICIFERS: common adjacent to the lower periclinalwall so fth eadaxia lepidermis ,an dbetwee npalisad ecell si nH. zeylanica; along veins of midrib in H. congolana.

T.S. petiole Vascular bundle bicollateral; forms a deep u-shape with incurved margins, in both species; strongly supported in the pericyclic region by thick-walled mucilaginous fibres in groups of 2-35 scattered sclerosed elements in the ground tissue of H. zeylanica (Fig.ióE).

Comments: anisocytic stomata with straight anticlinal walls on both surfaces and hemimesogenous typeo f stomatal development havebee nreporte d inH. umbellata by Nyawuame & Gill (1991), size 18.7 x 14.6 (im.

1.5.14 Landolphia R Beauv.

(Figs. 13Q , R, 15C, 20E ;Tabl e 1, 2, 3;Appendi x 1)

Agenu so f 51specie si nAfric a andca .8 specie sar eendemi ct oMadagascar .Repre ­ sented by 7 species in East Africa and two are examined here.

Leaf surface

HAIRS:absent . CUTICULAR SURFACE (SEM): smooth on abaxial epidermis in L watsoniana. STOMATA:hypostomati can danomocyti cinL . watsoniana; outline circular in L. watsoniana, elliptic inL. buchananii; size 15-18x15-16 [im; density permm 2500-62 5i nL. watsoniana. EPIDERMAL CELLOUTLINE :obscurel y vi­ sible as groves. ANTICLINAL WALLS (LM): undulate on both surfaces in both species.

T.S. lamina THICKNESS: 256-311 [im in L. buchananii, 201-256 [im thick in L. watsoniana. CUTICLE: thickest at the margin in both species. ADAXIAL EPIDERMAL CELLS: 1-layeredo nbot h species,rectangular , largeri nL. buchananii. ABAXIAL EPIDERMALCELLS :1-layered ,rectangular ,smalle rinL .watsoniana. STOMATA

22 WageningenAgricultural UniversityPapers 96-1 (1996) (T.S.): slightly raised in L. buchananii; slightly sunk in L. watsoniana. MESO­ PHYLL: bifacial; palisade i-2-layered in L watsoniana, extends to the midrib; 2(-3)-layered with much longer cells in L. buchananii, does not extend to midrib; spongymesophyl lcell shorizontall yelongate di nboth .MINO RVEINS : embedded, with outer parenchyma sheath and inner sclerenchyma sheath; larger ones transcurrenttoward sbot hepidermis .MIDRI B(Figs .13Q ,R) :adaxia lslighd yprom ­ inent in L. watsoniana, more or less straight in L. buchananii; abaxial more or less straight inL. watsoniana, ridged inL. buchananii.Vascula rbundl e forming acylin ­ drical strand inL. buchananii, flattened onth eadaxia l side (Figs. 13Q, 15C)an d an almost cylindrical strand in L. watsoniana (Fig. 13R).MARGIN : straight in both. CRYSTALS:L. watsoniana with afe w cluster crystals above minor veins and soli­ tary crystalsi n theparenchym a sheath and alongth eveins ;L. buchananii with soli­ tary crystals in the ground tissue of midrib and in the 2nd layer of palisade, small cluster crystals in the palisade.LATTCIFERS :toward s upper and lower epidermis; along veins;betwee n palisade cells and in the mesophyll.

T.S. petiole Vascularbundl ecylindrical ,formin g anova lshap ei nL. watsoniana (13E) ,flattened adaxially in L. buchananii; ground tissue with solitary crystals in L. buchananii.

Comments:anomocyti cstomat awit hsinuou santiclina lwall so nabaxia lsurfac e and agenous type of stomatal development reported in L. calabrica and L. dulcis var. barteri (=L. dulcis) and paracytic stomata in L. owariensis with sinuous anticlinal walls on both surfaces and eumesogenous type of stomatal development by Nyawuame& Gill (1991).Landolphia buchananii maysometime shav eleave swit h pubescence.

1.5.15 Mascarenhasia A. DC.

Figs. iE, 9I,J , S, 14A, 18C;Tabl e 1,2 , 3;Appendi x 1)

Agenu so f 8species ,al ldistribute di nMadagascar ,on eo f whichals ooccur si nEas t Africa: M. arborescens.

Leaf surface HAIRS (Fig. iE): simple,unicellula r hairs on abaxialpetiole .CUTICULA R SUR­ FACE (SEM):rough . STOMATA: hypostomatic, elliptic,outline ; size 21-31 x 10- 18 um; density per mm2 200-213; EPIDERMAL CELL OUTLINE (SEM): ob­ scured. ANTICLINALWALLS : abaxial straight, adaxial undulate.

T.S. lamina THICKNESS (Table 1):211-25 6um .CUTICLE :thickes ta tth emidri ba tth emidri b and margin. ADAXIAL EPIDERMAL CELLS: irregularly 1-2-layered, almost squaret orectangular ; smallera tth emidrib ,talle rtha nwid ewit ha nacut eape x(Fig . 9I). ABAXIAL EPIDERMAL CELLS: i-layered, rectangular; smaller around the

WageningenAgricultural UniversityPapers 96-1 (1996) 23 midriban dwit ha pointe dape x(Fig .9J) .MESOPHYLL :bifacial ;palisad e1-2-lay - ered, extends to the midrib; spongy mesophyll with transversely elongated cells, compact.MINO RVEINS :embedded ,collatera l witha noute rparenchym aan da n inner sclerenchymatous sheath. MIDRIB (Fig. 14A): 750-787 umthick ; adaxial outline almost straight, abaxial prominentlyridged. Vascula r bundle bicollateral, forming adee parc ,wit hisland so fphloe mabov eth eu-shape ;mucilaginou s fibres belowth ebundle ,a fe wabov eit .Collenchym ai n3- 4layer so nth eperiphery .MAR ­ GIN:revolute ,epidermi s2-layered ,collenchymatous .CRYSTALS : clustercrystal s in the mesophyll and ground tissue of midrib; solitary crystals in the phloem. LATICIFERS:branche dfreel y inth emesophyll ;alon gth eveins ;belo wth epalisad e andepiderma lcell stoward sbot hepidermis .

T.S.petiole: Vascularbundl ebicollateral , forms au-shape , withnumerou scluste rcrystal sdis ­ persed inth ecortex .

1.5.16 MotandraA .DC .

(Figs. iD, H, i4B;Table 1,2 ,3 ;Appendi x 1)

Agenu so f 3specie sconfine d toAfrica .M. guineensisi sexamine dhere .

Leafsurface

HAIRS:presen to nth epetiol ean dmidrib ;simple ,thick-walled ,1-5-celled ,wit ha n acute apex (Fig. iH); few glandular hairs alsoobserve d onth epetiole , withthre e shortbasa l cell (Fig. iD). CUTICULAR SURFACE(SEM) :smoot h onbot h sur­ faces.STOMATA : hypostomaticanomocytic ;type ;outlin eelliptic ,raised ;siz e15 - 19x9-15um ;densit ype rmm 270-160 .EPIDERMA LCEL LOUTLINE :featureles s onbot h sides.ANTICLINA LWALLS :adaxia l straightt o slightly curved,abaxia l undulate.

T.S. lamina

THICKNESS (Table 1): 99-110 um thick. CUTICLE: adaxial 1.9-3.8 urn thick; abaxial 1.9-2.9um , thickest at the midrib.ADAXIA L EPIDERMAL CELLS: 1- layered, 9.5-13.3x8.6-25.7 urn. ABAXIAL EPIDERMAL CELLS: i-layered, smaller,6.7-9. 5x 7.6-22. 8um.MESOPHYLL :palisad ei(-2)-layered;transversel y elongated,compact .MINO RVEINS :embedde dMIDRI B(14B) :513-54 9u mthick ; adaxial outline straight, abaxial prominently ridged.Vascula r bundlebicollateral , forminga dee par cwit hmucilaginou sfibre sintersperse dwit hphloe mwithi nth ear c andman ypocket so f mucilaginous fibresbelo wth evascula rbundle . CRYSTALS: numerous solitary crystals inth e ground tissue below the vascular bundle andi n enlargedcell si nth epalisad elayer ;man ysolitar y crystalsi nth ephloem .

24 WageningenAgricultural UniversityPapers 96-1 (1996) 7^5. petiole: Vascularbundl ebicollateral ,ova l shaped, endso f vascularbundl e almost in contact with one another; phloem in the middle of the oval shape; numerous solitary and clustered crystals in the cortical region.

Comments: anomocytic stomata with sinuous anticlinal walls on both surfaces and the agenous type of stomatal development have been reported in M. guineensis by Nyawuame & Gill (1991).

1.5.17 Oncinotis Benth.

(Figs. iG, 3C,3F ,4C , 11D, 14C, 16A, 18A;Tabl e 1, 2, 3;Appendi x 1)

Agenu s of 7lianescen t species confined toAfrica . O. tenuiloba hasbee n inspected here.

Leaf surface HAIRS: present on adaxial surface of petiole; with short, thick-walled basal cells, terminal celllonge rwit ha nacut eape x (Fig. iG) andi nth eaxil so f secondary veins as domatia (Fig. 3F). CUTICULAR SURFACE (SEM) (Fig. 3C): smooth on both surfaces. STOMATA:hypostomati c andparacytic ;outlin e elliptic, slightly sunken; rimver y narrow; aperture elliptic,wide ;siz e 15-22.5 x 10-14 ^m; density per mm2 369-439;primar ystomat aobserved .EPIDERMA LCEL LOUTLIN E(SEM) :articu ­ lar ridges over anticlinal walls on both surfaces. ANTICLINAL WALLS (LM): adaxial straight to slightly curved, angular; abaxial undulate.

T.S. lamina THICKNESS (Table 2) 147-183 urn. CUTICLE: very thin. ADAXIAL EPIDER­ MAL CELLS: i-layered, 2-layered towards the midrib, rectangular in outline. ABAXIALEPIDERMA LCELLS :smaller ,rectangula rt osquare .STOMAT A(T.S. ) (Fig. 1iD) : more orles s level withth e surface. MESOPHYLL: bifacial; palisade in 1- 2rows ;spong ymesophyl lcell scompact ,transversel yelongated .MINO RVEINS : embedded, collateral, with a parenchyma sheath. MIDRIB: abaxial prominently ridged, adaxial almost flat. Vascular bundle bicollateral, forming a deep arc with many fibres below and above interspersed with phloem tissue (Fig. 16A). Ground tissue collenchymatous at the periphery. CRYSTALS:numerou s cluster crystals in thegroun dtissu eo fmidri ban dspong ymesophyll ;solitar ycrystal san dsmal lcluste r crystals in the phloem.LATICIFERS : occur along veins;branchin g below the epi­ dermis and palisade towards both epidermis. MARGIN: slightly revolute.

T.S. petiole Vascular bundle bicollateral, deeply curved with phloem in the middle of the arc; numerous crystals in the ground tissue (Fig. 18A).

WageningenAgricultural UniversityPapers 96-1 (1996) 25 Comments: paracytic stomata have also been reported in O.gracilis and O. nitida, withth eeumesogenou styp eo f stomataldevelopmen tan danomocyti c stomatawit h the agenous type of development in O. glabrata by Nyawuame & Gill (1991).

1.5.18 Picralima Pierre

(Fig. 11A;Tabl e 1,3 ;Appendi x 1) A monotypic genus restricted to Africa.Th e only species P.nitida is examined.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM): rough. STOMATA: hypo- stomatic;anisocytic . Outline elliptic; size ;densit y permm 2. EPIDERMAL CELL OUTLINE: not visible. ANTICLINALWALLS : more or less straight.

T.S. lamina THICKNESS:366-40 3urn .CUTICLE :thickes ta tth euppe rmidri bregion .ADAX - IAL EPIDERMAL CELLS: i-layered, rectangular in shape, 13.3-19 x 13.3- 25.7 um.ABAXIALEPIDERMALCELLS:rectangular,io.5-i3.3 x 11.4-28.5um, 1-layered, smallest around the midrib and with an acute apex. STOMATA (T.S.) (Fig. 11 A).MESOPHYLL :bifacial ;palisad edoe sno texten dt oth emidrib ,(1-)2-3 - layered; spongy mesophyll withhorizontall y elongated cells with ample air spaces. MINORVEINS : with aparenchym a sheath.MIDRIB : 1208-1190 um; adaxial out­ lineslightl yconcave ,abaxia lprominentl yridged, acute ;vascularbundl ebicollateral , forming a deep arc, incurved at the edges, with many islands of fibres below the bundle.Groun dtissu ecollenchymatou s towardsth eperiphery .MARGIN :revolute , epidermalcell s smallerwit ha littl ecollenchyma .CRYSTALS :druse si nlarg eidio - blasts in palisade layer; numerous druses in the mesophyll especially in the layer above the lower epidermis; a few in the cortex below the vascular bundle, fewer above.

T.S. petiole Vascular bundle bicollateral, forms au-shape , with many fibres below the vascular bundle.

Comments: anomocytic stomata, with straight anticlinal walls on both sides and agenous type of stomata have been reported in P. nitida by Nyawuame & Gill (I99I)-

1.5.19 Pleiocarpa Benth.

(Figs.7A , B ,9T , 1iB , C, 14D,E , 16C,E ,F , 21B, 21C;Tabl e 1,2, 3;Appendi x 1)

Agenu so f 5specie srestricte dt oAfrica , mainlytree so rshrubs .Represente di nEas t Africa by two species, P. bicarpellata and P. pycnantha, and both are examined here.

26 WageningenAgricultural UniversityPapers 96-1 (1996) Leaf surface HAIRS:absent .CUTICULA R SURFACE (SEM):adaxia lroug h inP. bicarpellata, smooth inP. pycnantha; abaxial smooth inboth . STOMATA: hypostomatic in both species; cyclocytic with 3-4 cells around the stomata (Fig. 7A); outline broadly elliptic; slightly raised with sunken subsidiary cells in both; aperture slit like with narrowrim si nboth ;o f± th esam esiz e 16-23x 14-20urn ;densit ype rmm 2133-153 in P. bicarpellata, 200-238 in P. pycnantha. EPIDERMAL CELL OUTLINE: vaguely visible as wide ridges in P. bicarpellata and as round mounds in P. pycnantha. ANTICLINALWALL S (LM):bot h walls more or less straight to very slightly curved in P.pycnantha (Fig. 7A), abaxial slightly undulate in P. bicar­ pellata.

T.S. lamina THICKNESS (Table 1): 293-348 urn in P. pycnantha and 275-329^ in P. bicarpellata. CUTICLE:thickes t atth emargi nan dmidri bi nbot hspecies .ADAX ­ IALEPIDERMA LCELL S: 1 -layered ,rectangula ri nbot hspecies ,smalle ran dtalle r than wide atth e midrib.ABAXIA L EPIDERMAL CELLS: 1-layered, rectangular in both species, smaller and taller than wide atth e midrib. STOMATA (T.S.) (Fig. 1iB , C): slightly raised. MESOPHYLL (Table 1):bifacial ; palisade layer in 2 (-3) rows, extends to the midrib in both species;mesophyl l with transversely elongated cellsi nbot h species.MINO RVEIN S (Fig. 16F):embedde d andwit ha parenchym a sheath inbot h species.MIDRIB :622-64 1 urnthic k inP. bicarpellata, 750-767 um thick inP. pycnantha, adaxial outline almost flat inP. bicarpellata, prominent inP. pycnantha, abaxial outline slightly prominent in both species.Vascula r bundle bicollateral, forming analmos t straight arc with accessory bundles inP. pycnantha and a crenate one in P. bicarpellata; abaxial side of vascular bundle with a few pocketso f mucilaginous fibres andver yfe w ornon eo nth eadaxia l side.MARGIN : more or less straight to slightly revolute at the tip;epiderma l cells taller than wide and papillose in P. pycnantha; marginal vein with a well developed sclerenchyma- tousbundl esheat hi nP. pycnantha. CRYSTALS:abaxia lan dadaxia lepidermi s with solitary crystalsi nboth ;phloe mwit h solitary crystals andmesophyl lwit h clustered crystals in P. pycnantha; palisade with clustered crystals in enlarged cells in both. LATICIFERS:branchin gi nth emesophyl lan dals oalon gth evein si nboth ;immedi ­ ately below thepalisad e tissue.

T.S. petiole Vascular bundle bicollateral, forming a deep u-shape with incurved ends of bundles (Fig. 21C); strongly supported by thick-walled islands of mucilaginous fibres in groups of 2-35 in both (Fig. 16C); P. bicarpellata with accessory bun­ dles in the wings of petiole also supported with mucilaginous fibres adaxially; P.pycnantha with thick-walled, sclerosed elements scattered in the ground tis­ sue (Fig. 16E).

Comments: in my study of stomata of 3 species of Pleiocarpa, all were found to have cyclocytic-staurocytic typeo f stomata (Fig.7B) .O nth e contrary, Nyawuame

WageningenAgricultural UniversityPapers 96-1 (1996) 27 & Gill (1991) record anomocytic stomata in P.mutica, P. pycnantha var. tubicina (now P. pycnantha) and P.talbotii (now P. rostratd).

1.5.20 RauvolfiaL.

(Figs. 7E,F ,G , H, 8 0, P, 14F,G , H, 15A, 18E,Table s 1,2 , 3;Appendi x 1)

Apantropica l genuso f about 60species ,7 restricte dt oAfric a and 3t o Madagascar. Represented by 5specie s inEas tAfric a and 3o f them areexamine d here:R. caffra, R. mannii, and R. mombasiana.

Leaf surface HAIRS:absent .CUTICULA R SURFACE(SEM) :wit hfine stria eacros s subsidiary cells and over the whole surface on both sides in R. mannii. STOMATA (Figs.7F , G): hypostomatic and paracytic in all, sometimes with subsidiary cells subdivided by radiating walls onon eo rbot h sides;outlin eelliptic ;apertur e long, slit-like; size ranges from 19-32 x 10-18 [im;densit y per mm2 125-138-175 in Rauvolfia mom­ basiana. EPIDERMAL CELL OUTLINE (SEM): cuticular ridges over anticlinal walls on both surfaces in R. mannii. ANTICLINAL WALLS (LM) (Figs. 7E, H): adaxial straight in R. mombasiana, undulate on both sides in the other two species and on abaxial side of R. mombasiana.

T.S. lamina THICKNESS (Table 1): 165-183 um thick in R. mannii, 201-275 urn thick in R. mombasiana and 183-238 um thick in R. caffra. CUTICLE: thickest atth e margin R.manii. ADAXIA LEPIDERMA LCELL S(Fig .8O ): large rtha nabaxial ,rectangu ­ lar, 1-layered,2-layere d atth emargi n withpericlina l divisions atsom epoint s in R. caffra, much smaller around atth e midrib area. ABAXIAL EPIDERMAL CELLS (Fig. 8P): i-layered in all species, smaller. MESOPHYLL: bifacial in all species; palisade layer in a single row in R. mannii and R. caffra; in 1 (-2) rows in R. mombasiana, extendst oth emidri b in allbu tleave s a smallgap ;spong y mesophyll with somewhat horizontally elongated cells,no t socompact . MINORVEINS : em­ bedded, with a parenchyma sheath. MIDRIB (Figs. 14F,G , H): 567-604 urn thick inR. caffra; 695-714 urnthic k in R. mannii, 714-732 um thick inR. mombasiana; outline more or less straight in R. mombasiana on both sides, with a small abaxial ridge and a prominent adaxial ridge in R. mannii, with ridges on both sides in R. caffra. (Fig. 14F).Vascula rbundl ebicollateral ,form s ashallo war ci nal lspecie sbu t widest inR. mombasiana (Fig. 15A).Groun d tissue collenchymatous for 3-4 layers onbot h sides.MARGIN : more or less straight to very slightly curved in R. mannii and revolute in the other two; epidermal cells 2-layered in R. caffra, thick-walled. CRYSTALS: in subglobose idioblasts in the palisade in R. mannii and R. mombasiana; 2-4 druses present above minorvein s in a singleo r doublefile i n the palisade tissue inR. caffra and R. mombasiana (Fig. 18E);clustere d crystals in the groundtissu eo f midribi nR. mombasiana. LATICIFERS:alon gveins ;i nth emeso ­ phyll and towards both epidermis.

28 WageningenAgricultural UniversityPapers 96-1 (1996) T.S. petiole Vascular bundle bicollateral and forms a shallow arc in both but wider in R. mombasiana; with accessory bundles in R. mannii and R. mombasiana. Clustered crystals present in ground tissue and solitary crystals in thephloe m parenchyma of R. mombasiana. Ground tissue collenchymatous towards the periphery.

Comments: paracytic stomata are characteristic of the genus. Also reported in R. vomitoria,b yNyawuam e& Gil l( 199 1) .Eumesogenou san deumesoperigenou s type of stomatal ontogeny havebee n reported in R. coffra and R. mannii respectively by Karatela (1983) and Gill (1988).

1.5.21 Saba (Pichon) Pichon

(Figs. 4B, 8B, 8Q, R, 10J, 12A, 14I, 15D, 21B;Table s 1, 2;Appendi x 1)

A genus of three species restricted toAfrica . Only one species,Saba comorensis is represented in East Africa and is examined here.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM): smooth. STOMATA (Fig.4B) : hypostomatic andanomocytic ;outlin ebroadl y elliptict ocircular , sunken;rim sver y narrow, aperture short, circular; size 13-18x11-16 [im;densit y perm m2 850-856. EPIDERMAL CELL OUTLINE (SEM): cell outlines visible as round mounds. ANTICLINALWALLS : straight to slightly curved on both sides.

T.S. lamina THICKNESS (Table 1): 165-183 um.CUTICLE :thickes t on adaxial midrib at 10- 15 (im. ADAXIAL EPIDERMAL CELLS (Fig. 8Q): i-layered, taller than wide. ABAXIAL EPIDERMAL CELLS: i-layered, rectangular. STOMATA (T.S.) (Fig. 10J): slightly sunken: MESOPHYLL (Fig. 12A):homogenou s with 5-6 (-7) layers of vertically elongated cells, taller than wide, longest in the first row. MINOR VEINS: smaller veins collateral, embedded, larger ones transcurrent towards the adaxialepidermis ;girder swit hlignifiedparenchym a cells.MIDRI B (Fig. 14I;Tabl e 1): 878-924 [imthick ; adaxial outline ridged, more prominently ridged on abaxial surface.Vascula rbundl ebicollatera l andform s anova lshap ewit ha shallo w abaxial bundle and 5-6 adaxial bundles (Figs. 15D, 14I). Ground tissue with 3-4 layers of collenchyma onth eperiphery .MARGIN :mor eo rles s straight, witha n acute apex; epidermal cells taller than wide but with a rounded apex. CRYSTALS: a few styloids present in the mesophyll. LATICIFERS: a few noted, branching between palisade cells.

T.S. petiole Vascular bundle bicollateral, circular, but flattened at the top.

WageningenAgricultural UniversityPapers 96-1 (1996) 29 1.5.22 Schizozygia Baill.

(Figs. 2D, 14J, 19B, 21D;Table s 1, 2, 3;Appendi x 1)

A genus in Central and East Africa and Comoro Islands.Th e only species S. coffaeoides is investigated here.

Leaf surface HAIRS:absent . CUTICULAR SURFACE (SEM) (Fig.2D) :adaxia l with striations confined to antepericlinal wall of each cell; abaxial more or less smooth. STO- MATA: hypostomatic; paracytic; outline elliptic; rim narrow, aperture slit like, sometimes closed; size 27-31 x 16.6-20.3 H*m;densit y per mm2 63-106. EPIDER­ MALCEL LOUTLINE :adaxia l surface withthick ,wid eridges o fcuticl eove ranti ­ clinal walls (Fig. 2D). ANTICLINAL WALLS (SEM): adaxial straight, abaxial straight to slightly curved.

T.S.lamina THICKNESS (Table 1): 256-275 [imthick . CUTICLE: not very thick. ADAXIAL EPIDERMALCELLS : 1-layered,2-3-layere dtoward sth emidrib ,large rtha n abax­ ial. ABAXIAL EPIDERMAL CELLS: 1-layered, wider than tall. MESOPHYLL: bifacial; palisade in 1-2 rows,extend s toth e midrib; spongy mesophyll with some­ what transversely elongated cells but with many air spaces.MINO RVEINS : with aparenchymatou s sheath.MIDRI B (Fig. 14J): 1437-1473u mthick ;adaxia l outline very slight,abaxia l veryprominent .Vascula rbundl ebicollateral , forming au-shap e with a few pockets of fibres on the lower side, fewer above.Collenchyma , adaxial ridge with 5-6 layers, abaxial with 3-4 layers. MARGIN: revolute. CRYSTALS: present in enlarged cells in the palisade layer and in the cortex of the midrib. LATTCIFERS: along veins;branchin g in the mesophyll towards both epidermis.

T.S. petiole Vascularbundl ebicollateral ,form s awid eu-shape d arc(Fig .21D) ;wit h crystals in the ground tissue and collenchyma on the periphery.

Comments:Ca nb edistinguishe dfro mTabernaemontanab y cuticularornamentatio n

1.5.23 Stephanostema K. Schum.

(Figs. 1A , 5A, 8S,T 14K;Tabl e 1, 2,3;Appendi x 1)

A monotypic genus known from only one locality inTanzania .Th e only species 5. stenocarpum is examined here.

Leaf surface HAIRS: present on abaxial epidermis near the midrib and on the petiole; simple, uniseriate,wit h 1-5 cells,thick-walle d (Fig. 1A ) CUTICULAR SURFACE (SEM):

30 WageningenAgricultural UniversityPapers 96-1 (1996) abaxial withthic kridges extendin g from thepole ,acros s subsidiary cellsan d onth e rest of the surface (Fig.5A) . STOMATA: hypostomatic; paracytic, with subsidiary cellsove rth epoles ;outlin eelliptic ;siz e 15-21 x 9-13 um;densit ype rmm 2132-264. EPIDERMAL CELL OUTLINE: with wide ridges over anticlinal walls on both surfaces. ANTICLINAL WALLS (LM): adaxial slightly undulate, abaxial more undulate.

T.S. lamina THICKNESS:229-25 6urn .CUTICLE :thickes ta tth emidrib .ADAXIA LEPIDER ­ MAL CELLS (Fig. 8S):rectangular , i-layered, 19-28.5 x 32.3-72 urn. ABAXIAL EPIDERMAL CELLS (Fig. 8T): i-layered, smaller, 15.2-20.9 x 24.7-49.4 urn. MESOPHYLL:bifacial ; palisade extendst oth emidrib , 1-3-layered;spong y meso- phyll with transversely elongated cells, not compact. MINORVEINS : embedded, witha parenchym asheath .MIDRI B (Fig.14K) :787-823. 5ur nthick ;adaxia loutlin e with a slight projection, abaxial prominently ridged. Vascular bundle bicollateral, with a shallow arc.Collenchym a with 5-6 layers on adaxial ridge and 2-3 layers on abaxial side.MARGIN : straight. CRYSTALS: none observed.

T.S. petiole

Vascular bundle bicollateral, forms au-shape . Ground tissue collenchymatous.

1.5.24 Strophantus DC.

(Figs. 4C, 6A, B, 11E,F , 14L;Table s 1,2, 3;Appendi x 1) Strophanthusi sa genu so f 38specie soccurrin gi nAfric a includingMadagasca r and SouthAsia .I ti srepresente d by 11specie si nEas t Africa. S. courmontii is examined here.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM) (Fig. 4C): abaxial surface with concentric rings of striae around subsidiary cells, finer striae on theres t of the sur­ face; adaxial surface with wide ridges. STOMATA: amphistomatic (Fig. 6A); paracytic with a few anomocytic-actinocytic stomata (Fig. 6B); outline elliptic, slightly raised; rim wide;apertur e narrow, slit-like; size 24-33 x ll-29 um; density permm 2 63-144.EPIDERMA L CELL OUTLINE:wit h wide ridges over anticlinal walls. ANTICLINAL WALLS (LM): adaxial straight to slightly curved, abaxial undulate.

T.S. lamina THICKNESS: 192-238 urn thick. CUTICLE: 1-2 urn. ADAXIAL EPIDERMAL CELLS: i-layered, taller than wide. ABAXIAL EPIDERMAL CELLS: i-layered rectangulart osquar esmalle ra tth emidrib .STOMAT A(T.S. )(Figs . 1iE ,F) : abaxial epidermis raised, adaxial sunk, larger. MESOPHYLL: bifacial; palisade layer in 3 (-4) rows, extends to the midrib but leaves a very small gap; spongy mesophyll

WageningenAgricultural University Papers 96-1 (1996) 3I with somewhat horizontally elongated cells,bu t not compact. MINOR VEINS: embedded, with a parenchyma sheath; larger ones transcurrent towards adaxial side. MIDRIB; 970-988 um thick, abaxial outline with a slight ridge, adaxial prominently ridged (Fig. 14L).Vascula rbundl e bicollateral, forms a shallow median crescentic arc,wit h thin-walled mucilaginous fibres above and be­ low thebundle .Collenchym a 3-4 layers onbot h sides.MARGIN : slightly revolute at the tip. CRYSTALS: clustered and solitary crystals in the ground tissue of the abaxial ridge. LATICIFERS: common adjacent to the inner periclinal walls of the epidermis;betwee n thepalisad e cells and along the veins.

T.S.petiole Vascular bundle bicollateral, forms a curved arc; numerous crystals in the ground tissue especially below the vascularbundle ;thin-walle d mucilaginous fibres above and below the bundle. Collenchyma on the periphery.

Comments: stomata reported to be paracytic in S. barteri and S. gracilis, with eumesogenousdevelopmen tan danomocyti ci nS. hispidus (als oamphistomatic )an d S.preussii, with agenous type of development by Nyawuame & Gill (1991).

1.5.25 Tabernaemontana L.

(Figs. 2B, C, 3B, 5B, 8U,V,W , X, 11G, H, I, 12D, 14M, N, O, P, 15F, 16D, 17F, 18F, 21E, F;Tabl e 1,2 , 3;Appendi x 1)

Tabernaemontana is a genus of about 99 species, 18 in Africa, 15i n Madagascar, 1 on the Mascarene Islands, 21 in Asia, Oceania and Australia and 44 inTropica l America. In East Africa it is represented by 5 species, four of which are examined here:T. elegans,T. pachysiphon, T. stapfiana andT. ventricosa.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM) (Fig. 3B, 5B): abaxial surface withwing so f striaeacros s slightly sunkensubsidiar y cellsi nT. stapfiana, otherwise withthic k ridges on theres t of the surface; afe w striae across subsidiary cells of T. elegans, thicker ones from the poles and the rest of surface striate; striate over the whole surface andacros sth epole si nT. pachysiphon; adaxial surface withver yfine striae inT. elegans, striate inT. stapfiana andT. pachysiphon (Figs. 2B, C). STO­ MATA: hypostomatic and paracytic with 1o r 2 pairs on both sides in all species examined; outline elliptic in all, slightly raised; rims very narrow inT. elegans and T. pachysiphon; narrow in the others; aperture very wide inT. elegans; size largest inT. stapfiana; density per mm2 highest inT. stapfiana at 99-123, lowest inT. pachysiphon at31-69 .EPIDERMA LCEL LOUTLIN E(SEM) :adaxia lindistinc ta s very shallow grooves inT. elegans, abaxial obscure in all.ANTICLINA LWALLS : adaxial straight, angular in all;abaxia l straight to slightly curved; slightly undulate inT. elegans.

32 WageningenAgricultural UniversityPapers 96-1 (1996) T.S. lamina THICKNESS: 165-384 um, thickest inT. stapfiana at 329-384 um. CUTI- CLE:thickest at the upper midrib in almost all. ADAXIAL EPIDERMAL CELLS (Figs. 8U,W): i-layered in7 ? elegans and7 ! ventricosa, towards the margin or midrib, 2-layered; irregularly 2(-3)-layered inT. stapfiana andZ pachysiphon. ABAXIAL EPIDERMAL CELLS (Figs. 8V, X): i-layered inT. ventricosa; 1- layered inZ elegans except at some points near the margin, 2-layered; irregularly 2-layered inT. pachysiphon andZ stapfiana. STOMATA (T.S.) (Figs. 11G, H, I): with a wide aperture inT. elegans, a narrow one inT. pachysiphon and with a cuticular ridge inT. stapfiana. MESOPHYLL: bifacial in all species; palisade 2-3(-4)-layered inT. stapfiana andT. elegans (Fig. 12D); i-2(-3)-layered inZ pachysiphon andT. ventricosa, extends to the midrib in all species; spongy meso- phyll cells transversely elongated with numerous air spaces in all species. MI­ NORVEINS : collateral, embedded and with a parenchyma sheath in all; each supported by one large sclerid inZ pachysiphon (Fig. 16D).MIDRI B (Figs. 14M, N, 0,P): 604-2141 |j,mthick , thickest inZ pachysiphon at 2013-2141 um; midrib outline forms a slight adaxial ridge and a prominent abaxial ridge in all species. Vascular bundle deep v-shaped in all species; islands of phloem in the middle of the vascular bundle inZ elegans (Figs. 14M, 15F): phloem extending across the bundle on the adaxial side in the rest of the species (Figs. 14N, O, P). Ground tissue of midrib collenchymatous towards the periphery. MARGIN: revolute inZ ventricosa andZ pachysiphon; slightly revolute inZ elegans. CRYSTALS:Z pachysiphon with many single crystals in lower epidermis and with star-like crystals in the palisade layer and in large cells in the layer immediately below it, few in spongy mesophyll;Z stapfiana with many small star-like crystals in palisade layer and phloem parenchyma, very large star-like crystals in the layer above the lower epidermis and in the ground tissue of midrib;Z ventricosa with clustered crystals in enlarged cells ofth e mesophyll, in the ground tissue of mid­ rib and in both epidermis. LATICIFERS: in the spongy mesophyll; below the palisade and epidermis; between palisade and epidermal cells and towards both epidermis.

T.S. petiole Vascular bundle deep u-shaped inZ elegans andZ stapfiana, numerous crystals in the u-shape and some in the cortical region.

Comments: paracytic stomata are characteristic of this genus. Stomata are alsore ­ portedt ob eparacyti ci nZ eglandulosa,T. pachysiphon andZ pendulifiora, and with eumesogenoustyp eo fstomata ldevelopmen tb yN yawuam e& Gil l( 199 1) .Hair sar e sometimes present onZ elegans leaves.

1.5.26 VoacangaThouars.

(Figs. 11J, 14Q, R, 22C;Table s 1, 2, 3;Appendi x 1)

WageningenAgricultural UniversityPapers 96-1 (1996) 33 A genus of 12 species, 7 of which occur in Africa, and 5 in Asia.Tw o species are examined here:V. africana andV. thouarsii.

Leaf surface HAIRS: absent. CUTICULAR SURFACE (SEM): with striae across subsidiary cells. STOMATA: amphistomatic; outline elliptic; size larger in V. thouarsii; den­ sity per mm2;EPIDERMA L CELL OUTLINE (SEM):no t visible. ANTICLINAL WALLS (LM): adaxial straight, abaxial undulate.

T.S. lamina TfflCKNESS:CUTICI^:thickestmKa^canaADAXIALEPIDERMAL CELLS: 1-layered, 2-layered at the margin in V. thouarsii. ABAXIAL EPIDERMAL CELLS: i-layered, smaller. STOMATA (T.S.) (Fig. 11J) : sunken in the abaxial epidermis of V. thouarsii MESOPHYLL: bifacial; palisade in 1-2-layered in V. africana, 2-3(-4)-layered inV. thouarsii, extends to the midrib; spongy mesophyll with somewhat horizontally elongated cells.MINO RVEINS : embedded. MIDRIB (Figs. 14Q,R) :thicke r inV. thouarsii at476-55 0 umthick ; outlinecurve d on abax­ ial, ridged on adaxial side.Vascula r bundle bicollateral, v-shaped in V. africana, wider inV. thouarsii. Ground tissue collenchymatous towards adaxial side. MAR­ GIN: revolute on both sides.

T.S. petiole Vascular bundle bicollateral, forms a shallow \ inV. thouarsii and a deeper one in V. africana.

Comments: paracyti c stomatahav ebee nreporte db yN yawuam e& Gill ( 199 1) i nV. africana, with hemimesogenous type of development.

1.6 GENERAL DISCUSSION

Leaves The results of the present study and the information from previous studies carried outb y various authors onth elea f anatomy ofApocynaceae species clearly indicate thatth eleave s aredorsiventra l (bifacial) excepti na fe w caseswher ethe y havebee n described as isobilateral such as in Wrightia saligna (Ngan 1965), Aspidosperma quebracho andNerium oleander (Metcalfe and Chalk 1950).Th eleave s of Nerium oleander are not always isobilateral as shown by Esau (1965) in Fig. 16.2, a fact whichcoul db econtrolle db yth eenvironment .Som eleave shav ea tendenc ytoward s beingisobilatera l e.g.i nth egener a Carissa,Acokanthera andAncylobotrys. Allth e species examined in these genera show xeromorphic characteristics such as thick cuticle (Figs. 8A, 12C), papillose marginal cells (Fig. 19A) and marginal veins strongly supportedb ysclerenchymatou s cells(Fig . 19C).Saba comorensis ha sbee n shown to have homogenous leaves in the present study (Fig. 12A).

34 WagemngenAgricultural UniversityPapers 96-1 (1996) Hairs Five basic types are recognised: (i)Unicellular :ma yb eonl y 10[i mlon g asi nCarissa t o24 7 (imlon gi n Holarrhena pubescens, or even up to 850 um long as reported inAllamanda violacea (now A. blanchetii) by Chandra et al. (1972). (ii) Multicellular: vary from 57 [im in Ancylobotrys to 304 Jimi n Dictyophleba but longer lengths of 800 um have been recordedi nIchnocarpusfrutescens byChandr ae tal .(1972) .(iii )Multicellula rhair s with compressed basal cells embedded inth eepidermis :reporte d inEchites peltata (nowPeltastes peltata) byMetcalf e andChal k (1950)(iv )Variousl ybranche d hairs, shaped like antlers: recorded in Oncinotis hirta by Solereder (1908) and de Kruif (1985),als orecorde di nO. gracilisan dO. tomentella (deKrui f 1985)(v) :Glandula r hairs: present in Motandra guineensis (Fig. iC) and Clitandra cymulosa (Leeuwenberg & Berndsen 1988).

Cuticular ornamentation Apocynaceae leavessho wa wid erang eo fvariatio no fth ecuticula rsurfac ewhic hca n besmoot ha si nCarissa tetramerat ostriat eove rth ewhol esurfac e asi n Holarrhena pubescens. Tabemaemontana stapfianaca nb edistinguishe dfrom Z pachysiphon by itswholl ystriat ecuticl e(Compar eFig s2 B& 2C) .Kure r( 1917) ,a ssee ni nWilkinso n (!979).wa sabl et odistinguis hadulterant sfro mofficia l leaveso fcertai n compounds based onpresence , absence orposito n of striaean d granules onth e leaf. According toPaganell iCappellett i (1975),afte r his study ofAtropa belladona underth es.e.m , cuticular striatums areo fgrea tusefulnes s fordiagnosti cpurpose s andenabl eon et o distinguish even small samples of species from adulterants.

Stomata Stomata arerecorde d on both surfaces in only 5 of the 37 species studied. Stomata on both leaf surfaces have also been reported in Lochnera pusilla (now Catharanthus pusillus by Sayeed-ud-Din (1941) and Chandra et al. (1972); Ervatamia coronaria (nowTabemaemontana divaricata) by Kapoor et al. (1969); Plumeria rubra, Wrightia tinctoria, Wrightia tomentosa (now W. arborea) by Chandrae tal .( 1969) ;Wrightia saligna andWrightia laevis b yNga n( 1965 ); Carissa hirsuta (now C.spinarum), by Kapoor &Mitr a (1979);Landolphia owariensis, L. dulcis var.barteri (nowL. dulcis), Strophanthus hispidus andIsonema bulcholzii by Nyawuame and Gill (1991).

Subsidiary cells May not always fit exactly in the types described byWilkinso n (1979) and I have indeedexperience dsom edifficult y indescribin gsom earrangement si n Acokanthera oppositifolia, Hunteria zeylanica and Carissa tetramera. Jansen and Baas (1973), Baas (1975)an dHarto g andBaa s (1979)hav euse dth eter m "complex" for several intermediatetype slik ecomplex-anisocyti can dcomplex-laterocytic .I ti sals osome ­ timesdifficul t todecid ewhethe rsurroundin gcell sar etrul yanomocyti co rsomewha t anisocytic (Wilkinson 1979) and this problem is experienced in Strophanthus cormontii.

WageningenAgricultural UniversityPapers ç6-l (1996) 35 Stomatal size Ranges from 13 ^1111-36.5 um long (Table 2). An extensive survey of stomata of WestAfrica nApocynaceae byNyawuam ean dGil l(1991) ,resulte di nstomata l sizes between 16-35.5u mlong .O nth econtra y Chandra etal .(1972 )recorde d unusually large sizes inAllamanda violacea (nowA. macrocarpa) (15-45 urnlong) , Carissa grandiflora (nowA .blanchetii) (25-45 (Amlong ) and Catharanthuspusillus (25-68 [imlong) .Size sar edependen to npositio no fstomat ao nth elea f surface andwhethe r the sizes of primary stomata are recorded or not.

Anticlinal walls Differences in waviness between sun and shade leaves were first reported by Areshong (1897) and confirmed by Anheisser (1900).The y found that undulations are consistently more pronounced in shadeleave s and this fact is confirmed by nu­ merous other investigators.Accordin g toHaberland t (1934)an dWatso n (1942) the epidermal cell wallundulatio n isaffecte d byligh tintensit y whichinhibit sth egene s for waviness. Baissea myrtifolia has very undulate walls on both sides, a fact ex­ plained by its habitat in the forest understorey which lacks light. On the contrary, Ancylobotrys petersiana also has very undulate walls, although the specimen ana­ lysed was from aver y dry forest and noti n a shaded area. On the contrary, Sharma and Dunn (1968) found that undulations were always markedly reduced in xeric habitats in Kalanchoë fedschenkoi.

Stomatal abnormalities Single guard cells are observed in Holarrhena pubescens and were also reported in Trachelospermumjasminoides by Chandra et al. (1972). Aborted stomata and sto­ matawit hsingl eguar dcell sar ereporte di nAllamanda cathartica, Vincarosea (no w Catharanthus roseus) wadThevetiaperuviana by Nyawuame & Gill (1991). Single guard cells and giant stomata with aborted guard cells are reported in Rauvolfia manniiandR. caffra (Karatellaan dGill 1983).Gian tstomat awer erecorde di nman y species, as in Strophanthus courmontii, Carvalhoa campanulata etc, also frequent inRauvolfia mannii (Karatella and Gill 1983).

Lamina thickness The lamina is thickest in Adenium obesum, Acokanthera oppositifolia, Carissa tetramera,Picralima nitidaaadTabernaemontanapachysiphon. T.pachysiphon has a thick lamina because it has very large leaves.Adenium obesum has a very thick lamina because of the succulent nature of the leaves andth e other three species al­ ways have thick coriaceous leaves due toth e nature of the environment in Carissa tetramera, but the habitat of A. oppositifolia and P. nitida are quite diversified. Baissea myrtifolia has a very thin lamina (82-110 urn) because it is usually found growingi nth efores t understorey whereligh ti sminimal .Th eres to fth especie shav e alamin a thickness below 350 urn and cannotb e separated on thebasi s of this char­ acter.

36 WageningenAgricultural UniversityPapers 96-1 (1996) 1.7 CONCLUSIONS

Thepresen t study carried out onth e leaf anatomy of Apocynaceae is not very con­ clusivea sonl y3 7o f the7 7 speciesi nEas t Africa wereexamined .Th eresults ,how ­ ever, show that the leaves exhibit a wide range of variability in anatomical charac­ ters,som eo f whichca nb econfidentl y usedt odistinguis hth egener ao rspecies .Th e present conclusions arebase d on limited material available for examination and in situations whereI fel t moremateria l ofth esam e speciesshoul db eexamined ,I hav e indicated soan dinten dt od os oi nfuture .Liste dbelo w aresom eo fth elea f anatomi­ cal characters which are either characteristic to the family or genera or useful in distinguishing the taxa. Conclusions about close relationships are also mentioned and the most outstanding anatomical characters are listed.

Leaves aretypicall y dorsiventral (bifacial) excepti nver y few instances wherethe y may be isobilateral depending on their environment as observed in Nerium oleander (Metcalfe &Chal k 1950;Esa u 1965)o r have atendenc y towards being isobilateral as observed in some xeric species.Trul y isobilateral leaves in Apocynaceae have been recorded by Ngan (1965), in Wrightia saligna.

Hairs are simple, rarely glandular and uniseriate, rarely branched. May be uni- or muticellular, sometimes with compressed basal cells.Th e number of cells and the size is variable.Th e hairs of both Baissea myrtifolia and Oncinotis tenuiloba are terete (Figs. 3E, F), whereas those of Holarrhena pubescens are flattened with a widened base (Fig. 4E).Mor e specimens should be examined under s.e.m.t o find out whether the hairs are always terete or not.

Cuticular surface highly variable and in many instances diagnostic. More material need tob e exam­ ined as striations may be of great taxonomie value in some groups. (Stace (1965): genus Macropteranthes (Combretaceae); Wilkinson (1971): Anacardiaceae; van Staveren &Baa s (1973):Icacinaceae ) orma y showconsiderabl evariatio ni n degree of development in somegroup s as seen inth egenu s Combretum(Stac e 1965),o ri n Pistacia terebinthus, which shows a wide range of cuticular ornamentation from coarsely striate to smooth (Wilkinson 1971).

Stomata havebee nreporte d onadaxia lsurfac e inonl y 10gener aof Apocynaceae. Inthi scas e they are usually larger and scattered (Figs.6A , B; Chandra et al. 1969) and appear tob e of notaxonomi e value.Onl y Wrightia saligna which is endemic to Australia canb esai dt ob etrul y amphistomatic asstomat aar ereporte dt ooccu ri nequa lnum ­ bers on both surfaces by Ngan (1965).

WageningenAgricultural UniversityPapers 96-1 (1996) 37 Outline variable from narrowly to broadly elliptic to circular. Can be used to distinguish most species, especially if the width of the rim and type of aperture are also taken into consideration, e.g. the narrow rim and wide, elliptic aperture of Tabernaemontana elegans compared to the wider rim and circular aperture of Ancylobotrys tayloris (Figs. 3B,5E) .

Density per mm This isinversel y proportional to sizean d is only useful inextrem e cases. Generally higher in the lianas and not useful in the intermediate cases.

Stomatal size rarely exceeds 45 fim.Thos e species considered to have large stomata show mea­ surements between 30-36 u,m.

Subsidiary cells four basic types of subsidiary cells are recognised in the family Apocynaceae: (i) paracytic, sometimes laterocytic: very common. Characteristic of the genera Rauvolfia, Tabernaemontana and Voacanga. (ii) anomocytic, fairly common and may sometimes occur together with anisocytic type in the same species; anisocytic typeo f stomata wasneve rrecorde d on itsow n inth epresen t study,(iii ) Cyclocytic: ratheruncommo n excepti nPleiocarpa, CarissaaaAAcokanthera. (iv)Actinocytic : very rare but seen in Strophanthus courmontii in the present study and recorded in Catharanthuspusillus by Chandra et al. (1972). According to Poulsen (1917) sto­ mata are surrounded by several rings of radially arranged subsidiary cells in Acokanthera spectabilis (nowA. oblongifolia).

Anticlinal walls a situation was neverfoun d whereth e abaxial walls are straight andth e adaxial un­ dulate, it was always vice versa: abaxial walls undulate, adaxial straight or both walls straighto rbot hundulat e (seeTabl e2) .Onl yusefu l inextrem e cases and more specimens should be examined as some variaton in wall undulation of different leaves of the samespecie s ist ob eexpected ,accordin gt oWilkinso n (1979) andthi s fact was not investigated in the present paper.

Lamina thickness has notprove d tob ereliabl e in distinguishing the species asi ti s also dependent on the environment, texture and sizeo f the leaves.Thirty-on e of the 37examine d spe­ cies have a lamina thickness below 350 um and cannot be easily separated.

Cuticle and outerpericlinal walls only useful inextrem e casesan dquit edependen t onth eenvironmen t andag eo f the leaf.Mor e specimens from different localities need to be examined.

38 WageningenAgricultural UniversityPapers 96-1 (1996) Epidermal cells sizean dshap eusefu l indistinguishin g speciese.g .th elarg emor eo rles ssquar ecell s of Adenium obesum from the small rectangular shaped ones of Baissea myrtifolia. 1-2layere d species also easily distinguished from the i layered species.

Stomata t.s. useful in distinguishing species with thick cuticular ledges (Acokanthera and Picrcdima), or in distingushing those with wide apertures (Tabernaemomtana elegans) and sunken (Voacangathouarsii) or raised stomata.

Palisade cells continuity or discontinuity of chlorenchyma across the adaxial side of the midrib vascular tissue is too variable to be useful.

Spongy mesophyll very compact inHunteria, Saba andHolarrhena andver y lacunate in Ancylobotrys and Dictyophleba, not markedly different in the rest. More specimens need to be examined.

Minor veins embedded or transcurrent, not very useful in distinguishing species, except in Dictyophleba, Diplorhynchus, Ancylobotrys and Saba where they are transcurrent and easily distinguishable.

Midrib outline and thickness can be substantially used to distinguish taxa (see Figs. 13, 14)

Vascularbundle typically bicollateral inpetiol e and leaves of all specimens examined.Th e shapeo f the bundle is of some diagnostic value, i.e. the v-shaped bundles of the tribe Tabernaemontaneae from the cylindricalbundle s of Landolphia and Clitandra.

Phloem arrangement inth e midrib and petiole can be used in distinguishing some species.

Laticifers recorded in all the species examined.Variabl e in diameter, position and degree of branching.

Crystals calcium oxalate, abundant in all parts of the leaf and petiole; druses are the most common, followed by solitary crystals inth ephloem .N ocrystal s wereobserve d in Adenium obesum, Clitandra cymulosa andStephanostema stenocarpum.

WageningenAgricultural University Papers 96-1 (1996) 39 Leaf margin has notprove d tob ea usefu l character indistinguishin g mosto f the species,excep t inextrem e cases.Th elea f margin inApocynaceae isusuall y entire,althoug h itma y be straight to revolute on section, with an obtuse to almost rounded apex (Figs. 19 A-D).Th e rest of the species did not show very marked differences.

Petiole outline, size and shape of vascular bundle can be used to distinguish species (see Figs. 20, 21)

A.Th e following species can be easily distinguished from all the others:

1. Acokanthera oppositifolia: by itsthickl y coriaceousleave swit ha ver ythic k cuti­ cle and outer periclinal walls; leaf margin with papillose cells (Fig. 16A); stomata in transverse section with a very thick cuticular ledge (Fig. 10A)

2.Baissea myrtifolia mdAncylobotryspetersiana: byth epresenc eo f very undulate anticlinalwall so nbot hlea f surfaces andfro m eachothe rb yparacyti c stomatai nth e former and anomocytic inth elatter . More specimens from different localities need to be exmined to ascertain that the anticlinal walls are always undulate.

3. Carissa tetramera:b y thelea f mesophyll which has numerous sclerids;th e mar­ ginal vein which is surrounded by a large amount of schlerenchymatous tissue and the leaves which have one strong marginal vein (Fig. 22A) as opposed to all the otherspecie s studied,whic har eeucamptodromou so rhav ea submargina lvei n (Fig. 22 A,D) .

4. Dictyophleba lucida:b y its hairs withnumerous , short cells;lea f margin acutely obtuse with schlerenchymatous marginal vein and solitary crystals neatly arranged along the vascular bundle in the mesophyll.

5. Oncinotis hirta:steril e specimens canb e easily distinguished from allothe r East African Apocynaceae species by presence of many branched hairs in the leaf and petiole.

6.Tabemaemontana pachysiphon: can be distinguished from others species by its minorvein stha tar esupporte dwit hon elarg esclerid .Mor especimen sfro m different localities need to be examined.

B.The present survey confirms a close relationship between the following:

(i)Th e genera Carissa and Acokanthera Both comprise the subtribe Carissinae (Table 3) and have coriaceous leaves with laticifers prominently following the midrib vascular bundle, a situation never

40 WageningenAgricultural University Papers96-1 (1996) occuring so clearly in all the other species (Figs. 14 A & D). Both genera have leaves with atendenc y towards being isobilateral.

(ii)Motandra and Oncinotis Bothhav ephloe m aboveth emidri bbundle ,clearl y interspersed with mucilaginous fibres (Fig. 16A)

(iii)Dictyophleba and Ancylobotrys Although thethre e species studiedfro m theabov egener a were from three different localitiesi nth eKenya ncoast ,the y allexhibite dth esam etyp eo f spongymesophyl l arrangement which is highly lacunate (Fig. 12C).

(iv)Mascarenhasia and Funtumia Both have characteristically long palisade cells; similar types of vascular bundle arrangement; repeatedly branched laticifers and similar type of vascular bundle in the petiole with many druses in the ground tissue.

(v) Hunteria, Picralima and Pleiocarpa These three genera comprise the subtribe Pleiocarpinae and have 2-5 completely apocarpous carpels; the petiolar vascular bundle is strongly supported in the pericyclic region by mucilaginous fibres with very thick walls (fig. 16C).Thi s is veryeviden ti nm ypresen t studyo fth ePleiocarpina ewher eal lth ete nspecie sexam ­ inedexhibite dthi scharacter .Th epalisad elaye rals oha slarg e subglobose idioblasts with druses.

(vi) Clitandra and Landolphia Both have a cylindrical vascular bundle in the petiole and midrib and a spongy mesophyll with horizontally elongated cells.

C.The most outstanding anatomical characters include:

The dorsiventral leaves, presence of laticifers, bicollateral vascular bundle and intraxylary phloem.

Thepresen tinvestigatio nwa scarrie dou ta sa preliminar y surveyo fth elea f anatomy ofApocynaceae inEas t Africa.Th eresult s showmor e diversity inanatomica l char­ acters inth eleave s than inth e stem.Th eresult s alsoconfir m that many species can be identified using leaf venation, surface characters or leaf transverse section. It is hoped that allth e information gathered will stimulate further research.

WageningenAgricultural UniversityPapers 96-1 (1996) 41 1.8 FIGURES AND TABLES

42 WageningenAgricultural UniversityPapers 96-1 (1996) Figure i. A-I. Hairs.A .Stephanostema stenocarpum; adaxial epidermis,curve d 5-celled.B .Holarrhena pubescens; adaxial epidermis, unicellular, straight. C.Ancylobotrys petersiana; petiole, short hairs. D. Motandra guineensis; petiole, glandular hairs. E. Mascarenhasia arborescens; petiole, unicellular, thin walled.F .Dictyophleba lucida;petiole ,cell s small,numerous .G .Oncinotis tenuiloba; petiole ,shor tbasa l cells,termina lcell slonger .H .Motandra guineensis; midrib ,straigh thair ,shor tbasa lcell san dlon gtermi ­ nal cells.I .Baissea myrtifolia; adaxial midrib, basal cells long, terminal cells short, hooked at the apex. A-I = mag. x 25.

WageningenAgricultural UniversityPapers 96-x (1996) 43 Figure 2. A-F. Adaxial leaf surfaces. A. Diplorhynchuscondylocarpon; smoot h surface with very fine striae.B .Tabemaemontana elegans; surface with fine striae,cel l outline indistinct as shallow groves.C . T. stapfiana;whol e cuticle striate, obscuring cell outlines. D. Schizozygiacoffaeiodes; ridges o f cuticle over anticlinal walls; striatumsconfine d to antepericlinal wall of each cell. E. Carvalhoa campanulata; cuticular ridges over anticlinal walls, striatums over cell outlines. F. Holarrhena pubescens; cuticular ridgesove r undulate, anticlinal walls, striatums crossing cell outlines.Th e cells exhibit some inward collapse of the outer periclinal walls, due to drying. A, E = mag. x iooo; B = mag. x 1250; C = mag. x 2000;D =mag . x 1500;F = mag . x 2060.

44 WageningenAgricultural UniversityPapers 96-1 (1996) Figure3 .A-F .Abaxia llea fsurfaces .A .Hunteria zeylanica; surfacesmooth ,stom acircular ,slightl yraise d withsunke n subsidiary cells.B .Tabernaemontana elegans; wide,ellipti c aperturean dnarro wrims ,por e closed,striation sacros ssubsidiar ycell san dridges extendin ga tpola rends .C .Oncinotis tenuiloba; stoma with paracytic subsidiary cells, surface smooth. D. Acokantheraoppositifolia; circular stoma, round aperturean dwid erims .E .Baissea myrtifolia; domatiai nth eaxil so f secondary veins,hair slong ,curled , extending to the midrib. F. Oncinotistenuiloba; domati a in the axils of secondary veins, hairs straight erect A = mag. x 3080; B = mag. x 1640; C = mag. x 2730; D = mag.x 2800; E = mag. x 101; F = mag. x 185.

WageningenAgricultural UniversityPapers 96-1 (1996) 45 Figure4 .A-F .Abaxia l leaf surfaces. A.Dictyophleba lucida;stomat a and subsidiary cellsraised , surface smooth, with shallow grooves over cell outlines. B. Saba comorensis; sunken stomata; cells outlines visible astin ymounds .C .Strophantus courmontii;primar y stomaa tto p right, concentric rings of striae around subsidiary cells. D.Ancylobotrys petersiana; circular stoma, pore narrow, slit, cuticular surface ornamented.E .Holarrhenapubescens; flattened multicellularhair swit ha wid ebase ,taperin ga tth eapex ; whole cuticle striate. F.Carissa edulis;broadl y elliptic stomata, sunken; ridges over subsidiary cells and partly obscuring stomata. A, B = mag. x 2000;C , D = mag. x 1000;E = mag. x 366;F = mag. x 1520.

46 WageningenAgricultural University Papers96-1 (1996) Fig 5. A-F. Abaxial leaf surfaces and stomata. A. Stephanostemastenocarpum; thic k cuticularridges extend fromth epole s andacros s subsidiary cells. B.Tabernaemontcma stapfiana; wingso f striae across slightly sunken subsidiary cells;C .Carissa tetramera; broadly elliptic stomawit hwid erims ,truncat ea t thepoles ,surfac e smooth.D .Diplorhynchus condylocarpon; broadl yellipti cstom awit hnarro wpor ean d peristomatalrims. E .Ancylobotrys tayloris; circular stomawit h a circular aperture,ridges ove r sunken subsidiary cells. F. Carvalhoa campanulata; concentricrings o f striae around subsidiary cells, stomata narrowlyelliptic ,cuticula rsurfac eridged, otherwis esmooth .A ,B ,D ,E = mag . x 2000;C = mag . x 3500; F= mag . x tooo.

WageningenAgricultural UniversityPapers 96-1 (1996) 47 Figure6 A-H.Adaxia l and Abaxial leaf surfaces showingstomat a and anticlinal walls.A,C,E,G ,adaxial ; B,D,F,H, abaxial. A,B- Strophanthus courmontii; amphistomatic. C,D. Holarrhena pubescent; with paracytic cells extending over the poles (laterocytic). E,F. Acokanthera oppositifolia; paracytic and cyclocytic-tetracytic. G,H. Dictyophleba lucida. anomocytic stomata.T = trichome, A-H = mag. x 200.

48 WageningenAgricultural UniversityPapers 96-1 (1996) Figure7 A-H .Adaxia l andAbaxia l leaf surfaces showing stomataan d anticlinal walls.A,C,E,H ,adaxial ; B,D,F,G,abaxial .A.B .Pleiocarpapycnantha; cylocytican dtetracytic .C,D .Baissea myrtifolia; paracyti c stomata, anticlinal walls very undulate on both sides. E,F. Rauvolfia mannii; paracytic, anticlinal walls undulate onbot h sides.G,H . R. mombasiana;abaxia l undulate, adaxial straight. A-H = mag. x 200.

WageningenAgricultural UniversityPapers 96-1 (1996) 49 3 M iOC*30CG BDH—rr-r-i NI

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Figure8 A-X .Adaxia lan dabaxia lepidermi so f variousApocynaceae .A,C,E,G,I,K,M,0,Q,S,U,W , adax- ial. B,D,F,H,J,L,N,P,R,T,V,X, abaxial. A.B. Acokanthera oppositifolia. C,D. Adenium obesum. E,F. Ancylobotrys tayloris. G,H. Baissea myrtifolia. IJ. Carvalhoa campanulata.K,L . Clitandracymulosa. M,N.Funtumia elastica. O.P.Rauvolfia caffra. Q,R .Saè acomorensis. S,T .Stephanostema stenocarpum. U,V.Tabemaemontana pachysiphon. W,X .7 ? stapfiana. A-X = mag.x 25 .

50 WageningenAgricultural UniversityPapers 96-1 (1996) böQoa

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Figure 9 A-T. Adaxial andabaxia l epidermis. A, C, E, G, I, K,adaxia l midrib. B, D, F,H , J, L, abaxial midrib. M-T, margin. A.B. Acokanthera oppositifolia. C,D. Adenium obesum. E,F. Ancylobotrys petersiana. G,H.Carissa edulis. IJ .Mascarenhasia arborescens. K,L.Diplorhynchus condylocarpon. M . Acokanthera oppositifolia. N. Adenium obesum. O. A/o/ïa microstylis. P. Carissa edulis Q. Baissea myrtifolia. R. Clitandra cymulosa. S. Mascarenhasia arborescens. T. Pleiocarpa pycnantha. A-T = mag. x 25.

WageningenAgricultural University Papers 96-1 (1996) 51 B

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Figure io. A-J. Stomata in transverse section. A. Acokanlhera oppositifolia; thick cuticular ledge. B. Adenium obesum; slightly raised. C. Ancylobotrys petersiana; narrower aperture. D.A. tayloris; wider aperture. E. Carrisa tetramera: wider aperture. F. C. edulis; very narrow aperture. G. Carvalhoa campanulata; more or less level with the surface. H. Hunteria congolana; stoma raised. I. H. zeylanica. J.Saba comorensis;stom a sunken. A-J= mag. x400 .

52 WageningenAgricultural UniversityPapers 96-1 (1996) B

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Figure n. Stomata in transverse section. A. Picralima nitida; thick cuticular ledge. B. Pleiocarpa pycnantha; C.P. bicarpellata D. Oncinotistenuiloba; mor e orles s level with the surface. E, F.Strophan- thuscourmontii; E .adaxial ,ver ywid eaperture ;F .abaxia lraised .G .Tabemaemontana elegans;ver ywid e aperture. H.T. pachysiphon; narrow aperture. I.T. stapflana;J .Voacanga thouarsii; abaxial, sunken sto­ mata. A-J = mag. x 400.

WageningenAgricultural UniversityPapers 96-1 (1996) 53 Figure 12. A-F. Mesophyll. A. Saba comorensis; homogenous. B. Holarrhena pubescens; 1-layered palisade, transversely elongated spongy mesophyll cells, compact. C. Ancylobotrys tayloris; vertically elongated spongy mesophyll cells, lacunate, note thick cuticle and outer periclinal walls. D. Tabernaemontana elegans; 3-4 layered palisade, transversely elongated spongy mesophyll cells, less compact. E. Acokanthera oppositifolia; mesophyll with tendency towards being isobilateral. F. Diplorhynchus condylocarpon; vertically elongated spongy mesophyll cells, less lacunate with transcurrent veins towards both epidermis. A, B = mag. x 200; C = mag. x 312; D = mag, x 125; E = mag. x 160; F = mag. x 312.

54 WageningenAgricultural UniversityPapers 96-1 (1996) Figure 13.A-S .Midri boutline :A .Acokanthera oppositifolia. B .Adenium obesum. C.Alafia microstylis. D. Ancylobotryspetersiana. E.A. tayloris. F.Baissea myrtifolia. G. Carissa edulis. H. C. tetramera. I. Carvalhoa campanulata. J. Clitandra cymulosa. K. Dictyophleba lucida. L. Diplorhynchus condylocarpon.M .Funtumia elastica. N .Holarrhenapubescens. O .Hunteria congolana. P .// . zeylanica. Q.Landolphia buchananii. R.L . watsoniana.A- R= mag. x 25.

WageningenAgricultural UniversityPapers 96-1 (1996) 55 Figure 14. A-S. Midrib outline. A. Mascarenhasia arborescens. B. Motandra guineensis. C. Oncinotis tenuiloba. D. Pleiocarpa bicarpellata. E. P. pycnantha. F. Rauvolfia coffra. G. R. mannii. H. R. mombasiana. I. Saba comorensis. J. Schizozygia coffaeiodes. K. Stephanostema stenocarpum. L. &/w- phanthus courmontii. M.Tabernaemontana elegans. N.7 ! pachysiphon. O.T .stapfiana. P.7" . ventricosa. Q. V^iocan^aafricana. R. Kthouarsii. A-R = mag. x 25.

56 WageningenAgricultural UniversityPapers 96-1 (1996) Figure 15. A-F.Midrib .A .Rauvolfia mombasiana; midriboutlin e straighto nbot hsides ,vascula rbundl e shallow arc. B. Holarrhenapubescens; midrib outline, 1-layered palisade. C. Landolphia buchananii; vascular bundle with schrenchymatous tissue on adaxial side. D. Saba comorensis. E. Ancylobotrys tayloris; phloemextend s acrossth eto po f bundle. F.Tabernaemontana elegans; phloem formsisland si n the middle of the bundle, palisade extends to the midrib (arrow). A, B, D, F = mag. x 50; C, E = mag. xi 25.

WageningenAgricultural UniversityPapers 96-1 (1996) 57 Figure 16. A-F.Fibres . A. Oncinotis tenuiloba; Mucilaginous fibres above vascular bundle interspersed with phloem. B. Carissatetramera; sclerid s among palisade and spongy palisade tissue. C. Pleiocarpa bicarpellata; petiolar vascular bundle strongly supported by mucilaginous fibres. D.Tabemaemontana pachysiphon; seiend supporting each minor bundle; cork wart. E.Pleiocarpa pyenantha; sclerosed ele­ ments in ground tissue of petiole; mucilaginous fibres along petiolar vascular bundle. F. Pleiocarpa bicarpellata; embedded and collateral minor bundle with outer parenchyma sheath and xylem vessels surrounded with fibres. A, E = mag. x 125; B,C , D, F = mag. x 312.

58 WageningenAgricultural University Papers96-1 (1996) fj|(IJt tilt»ite rtP.ill -êÈ

Figure 17. A-E. Laticifers. A.Acokanthera oppositifolia;Laticifer s along the midribveins . B. Adenium obesum;laticife r branching below epidermis. C.Ancylobotrys petersiana; laticifers in ground tissue of petiole. D. Carissa edulis;laticifer s along midribveins . E.Funtumia elastica;laticife rbranchin g inth e palisade; crystals in mesophyll. F.Tabernaemontana elegans, two laticifers along the stem. A, D= mag. x 125; B =mag . x 312; C = mag. x 500; E= mag. x 200; F = mag. x 160.

WageningenAgricultural University Papers 96-1 (1996) 59 %*•*: %m~&-' m m%ßi

*>•• * ito % <Ï& "-....

Figure 18. A-D. Crystals. A. Hunteria zeylanica; idioblast with cluster crystal (druse) in palisade. B. Oncinotistenuiloba; cluste rcrystal si nspong ymesophyll .C .Mascarenhasia arborescens; cluste rcrystal s in the ground tissue petiole. D. Dictyophleba lucida; solitary cystals along veins in the mesophyll E. Rauvolfiamombasiana. cluste r crystals (druses) in two layers above minor vein. F. Tabemaemontana stapflana; crystalsi npalisade ;2-layere depidermis .A = mag , x 787;B , F= mag . x 312;C = mag . x 320; D, E = mag. x 400.

60 WageningenAgricultural UniversityPapers 96-1 (1996) Figure19 .A-D .Lea fmargin :A .Acokanthera oppositifolia; straigh tmargi nwit hpapillos eepiderma lcells . B. Schizozygia coffaeiodes;revolut e leaf margin. C. Carissa tetramera; marginal vein with layers of schlerenchymatous tissue. D.Adenium obesum;obtusel y acute margin, slightlyrevolut e atth etip .A , B, C = mag. x 125; D = mag. x 250.

WageningenAgricultural UniversityPapers 96-1 (1996) 61 Figure 20. A-I. Petiole outline. A. Acokanthera oppositifolia. B. Adeniumobesum. C. Ancylobotrys petersiana. D. Clitandra cymulosa. E. Landolphia watsoniana. F. Dictyophleba lucida. G. Funtumia elastica. H.Hunteria zeylanica. I .Mascarenhasia arboresens. A-D =mag . x 25.

62 WageningenAgricultural UniversityPapers 96-1 (1996) Figure 21. A-F.Petiol e outlines. A. Oncinotistenuiloba. B . Sabacomorensis. C .Pleiocarpa pycnantha. D. Schizozygia coffaeiodes.E .Tabernaemontana elegans.F .T. pachysiphon.A- F= mag . x 25.

WageningenAgricultural UniversityPapers 96-1 (1996) 63 Figure 22. A-F. A. Carissa tetramera;lea f venation with a very strong marginal vein. B. Pleiocarpa pycnantha;lea fvenatio nwit ha submargina lvein .C . Voacangaafricana; lea f venationwithou tprominen t marginalloop s(eucamptodromous) . D.Tabemaemontana elegans; stem(L.S.) ,sclerose delemen ti npith . E.Carvalhoa campamdata; stem,(T.S.) ,t osho warrangemen to fcells .F . Hunteriazeylanica; stem ,(T.S.) , to show arrangement of cells, note the sclerosed elements in the pith. A, B, C, D = mag. x 160; E = mag. x 125; F= mag. x 100.

64 WageningenAgricultural UniversityPapers 96-1 (1996) Table I.Lea f anatomical characters of someEas tAfrica n Apocynaceaespecies .

Species Cuticle size Epidermal cells, size in Palisade, num­ Midrib Lamina in um [im. bero f rows and thickness thickness sizes of 1/2ro w (Urn) (um)

a i. i9-26.6a 11-19X i9-40 2-3 rows 677-732 384-458 b 1 Acokanthera 9-4-i5b n.4-i9X5.7-28.5 32.3-59 x 8-13 2 oppositifolia 21-28.5" 12.4-22.8 x 11.4-27* 30-44 X6-12 2I-23.8y n.4-i7X2.9-i5.2y 22-48z 30.4-40 X5.7-H.42

a 2. i-9-9-5a 43.7-68.4 x 28.5-61.8 1-2row s 1245- 403-439 1 Adenium 1.9-3.8" 19-28.5 x i5-57b 51-67 X6-13 1263 2 obesum 1-9-5-7" 12.4-20.9 x 7.6-17" 32-38 x 10-15 y i-9-5-7y 133-19 x 9-5-i8 z 5-7-9-5z 22.8-36.6 x 9.5-26.6

3- i.9-2.9a 13.3-18 x 13.3-34.2" 1-2(-3 ) rows 402-439 256-302 1 Alalia i.9-2.9b 8.6-12.4 x i3-3-34-2b 15-29 x 4-13 2 microstylis 3.8-4.8" 9.5-11.4x9.5-17" 27-36 x 6-13 y i.9-2.9y 5-7-9-5 x 3-8-7-6 z z 2.9-4.8 7.6-13 x 7.6-9.5

a 4- 3-7-5a 5-10 x 7-33 1-2(-3 ) rows 714-750 293-320 b 1 Ancylobotrys 3-6b 3.5-5 x 9-i9 29-45 x 7-15 petersiana 5-8" 6-8 x 5-9" 28-42 x 8-152 3-6-5y 4-6 x 5-9y 7-9 x 4-8z

5- 6-1 la 4-8 x 8-24" 2-(3) rows 549-586 229-300 b 1 Ancylobotrys 3-9b 3-5 x 9-23 30-92 x 6-13 3 tayloris 2-6" 5-7 x 5-8-5* 19-27 x 6-13 y 4-4-5y 3-6 x 4-7 Z 3-5-5-5* 7-IIX4-7

6. ia 4.5-8 X 5-20a 1 row 201-238 82-110 Baissea ib 4-6 x 5-28b 15-28 x 4-71 myrtifolia 3* 13-19x5-16" 3y 6-12 X4-I3y z 2Z 3.8-7.6 x 7.6-9.5

a 7- 8-i3a 9-12 x i3-27 2 (-3) rows 439-458 247-302 b 1 Carissa 4-5b 7-11 X5-I2 40-86 x 6-15 edulis 8-17" 10-18 x 6-17" 25-35 x 6-I02 3-5" 8-11 xs^ z 3-8z 9-15 x 5-io

WageningenAgricultural UniversityPapers 96-1 (1996) 65 Species Cuticle size Epidermal cells, size in Palisade, num­ Midrib Lamina in (im um. bero f rows and thickness thickness sizes of 1/2ro w (um) (um)

8. I2-l6* 13-17 x 13-25* 2 (-3) rows 494-549 366-412 b Carissa 3-6" 5-12 x 7-24 45-65 x 7-I51 tetrameta 9-20" 13-19 x 14-27" 30-50 x 5-152 3-9X 4-6 x 4-8* 8-11 X7-20" i3-i5z

a 9- i 14-21 x 16-38* 1 row 1409- 156-220 b b Carvalhoa i 10-20 XII-29 25-35 x 8-14' 1446 campanulata i" 12-14x10-16" iy 10-12 x 8-i4y iz 10-13 x 7-16"

10. 1-2* 8-13x8-15* 2 rows 640-659 192-247 b 1 Clitandra I9b 5.7-9.5 X II.5-22.8 25-32 X8-I2 cymulosa 1-2.8" 7.6-11.5x3.8-15.2" 17-23 X8-I2 2 1-3.8» 5-7-9-5 x 3-8-9-5'' 7.6-11.5 x 3.8-11.4"

ii. l-2a 10-12 X7-20* 2 rows 714-787 183-311 Dictyophleba 2-2.5b 4-6 x 7-i4b 32-42x6-13.5' lucida 2-3" 8-12x3-11" 23-31 x 10-212 y 2-3 4-7 x 3-7y 3-10 x 3-5"

12. 2-9-5-7a 8.6-17 x 9-5-32.3" 2-3 rows 439-458 256-275 b 1 Diplorhynchus i-9-5.7b 7.6-15.2 x 7.6-28.5 34-42 x 6-7.5 condylocarpon 3.8-7-6" 7.6-9.5 x 2.9-15.2" 21-36 x 38.7s 3.8-7.6" 7.6-12.4 x 3.8-7.& 7.6-10.5" 12-14x4-5"

13- 3-8-5.7a 7.6-24.7 x 13.3-24.7* 2-3 rows 586-768 311-366 Funtumia 1.9-3.8" 7-11.4 X I0-20b 55-80x6-10' elastica 4.5-7" 7-12 x 5-10" 44-67 x 8-152 3.8-5-7" 8-12 xS^* 3.8-6" 9.5-19 x 38-13"

a 14- l-2 12-26 x 7-26* 1 row 970-1025 201-220 Holarrhena I-2b 10-24 x n-30b 30-37 x 5-9' pubescens 1-2" 16-21 x 8-17" I" 12-20 x io-i7y 2-3* 18-22 X9-I8"

66 WageningenAgricultural UniversityPapers 96-1 (1996) Species Cuticle size Epidermal cells, size in Palisade, num­ Midrib Lamina in um um. bero f rows and thickness thickness sizes of 1/2ro w (um) (um)

a a 15- 8-9 8-I2X 7-20 1-2row s 403-439 183-220 1 Hunteria 4-7b 5-8x8-18" 14-26 x 5-9 congolana 10-15 x 10-20" 8-13 x 6-82 I0-I4y 7-12 x 5-9" z 13-15' 7-11 X4-n

a a 16. 9-io 12-19 x 6-20 2-3 rows 915-951 266-348 1 Hunteria 5-7b 10-12.8 x 9-29" 16-31 x 5-9 zeylanica 10-13.5" 16-17 x 15-16" 14-30 x 5-102 IO-I2.5y 14-16 x n-i5y Z 9-i3z IO-I5 X9-I2

17- 2-3-5a 6-9 x 4.8-18" 1-2 (-3) rows 586-622 256-311 1 Landolphia i-3b 6-8 x 9-25" 57-90 x 7-10 buchananii 2-5-5-5" 7-10 x 4-5-14-5" y 3-5-5y 4-9 X 2-8 z 5-5-9* 6-9 x 3.5-1I

18. 4-5-8a 3-5-5 x 8.5-i8a 1-2row s 586-604 201-256 1 Landolphia 2-5-3b 3-5 x 6-15" 25-30 x 6-12 2 watsoniana 4-5-5" 5-7.5 x 12-15" 24-27 x 11-19 3-5-5y 4-5x6-8" 2 5-7-5* 6-8.5 X6-I0

19- 1-9-3-8" 17-24.7 x 13.3-34.2" 1-2row s 750-787 211-256 Mascarenhasia 1.9-3.8" 19-24.7 x 20.9-38" 50-60 x 9.5-151 3 arborescens 3-8-5-7" 133-19 x 5-7-9-5" 29-38 x 9.5-19 i-9-5-7y 7.6-13.3 x 3.8-10.5" 3-8-5-7* 13-21 x 9.5-13.3*

20. i-9-3-8a 9.5-13.3 x 8.6-25.7" 1-2row s 513-549 99-110 Motandra i.9-2.9b 6-7-9-5 x 7.6-22.8" 20-30 x 7-121 guinensis 1.9-3.8" 3.8-11.4x2.9-8.6" 15-20 x 8-132 i.9-2.9y 4-10.5 x 2.6-7y z I-9-3-82 4-5 x 6-io

21. I-2a 11-17X 13-22" 1-2row s 723-732 147-183 Oncinotis 1-2» 4-7 x 7-i6b 25-40 x 5-91 tenuiloba I-2" 4-9 x 3-9" 24-35 x 5-122 I-2y 6-8 x 6-ioy 1-2.5* 5-8 x 5-9z

22. 5-7-9-5a 13.3-19 x 13.3-25.7" 1-2(-3 ) rows 1190- 366-403 b b 1 Picralima 3-8-6.7 I0.5-I3-3 x n.4-28.5 15-45 x 8-14 1208 nitida 7-11.4" 13.3-29 x 9.5-20.9" 15-30 x 8-152 3.8-7.6" 9.5-17.1x5.7-13.3" 9-24.6* 8-25 x 4-16.5 z

WageningenAgricultural UniversityPapers 96-1 (1996) 67 Species Cuticle size Epidermal cells, size in Palisade, num­ Midrib Lamina in (im |im. ber of rows and thickness thickness sizes of 1/2ro w (jim) (|im)

23- 3-5-5a 20.9-30.4 x 19-51.3" 2 (-3) rows 622-641 275-329 Pleiocarpa 3-4b 17-24.7 x 19-57" 15-27x9-15' bicaipellata 8-10" 15.2-28.5 x 13.3-30.4" 15-20 X9-152 5-6" 19-36 x i3.3-30.4y 6-ioz 19-30.4 x 9.5-19"

24. 3-5-6" 15.2-28.5 x 26.6-58.9" 2 (-3) rows 768-787 293-348 1 Pleiocarpa 3-4b 15.2-24.7 x i9-55b 10-31 X8-15 pycnantha 8-10" 26.6-36 x 19-30.4" 10-15 X9-142 io-i3y 19-34 x i5.2-28.5y IO-IIz 19-28.5 x 7.6-19*

25- 1-9-3-8" 11.4-20.9 x 19-41" 1 row 567-604 183-238 b b Rauvolfia o.9-3.8 3.3-15.2 X7.6-34.2 67-93 x 9-5-251 caffra 1.9-2.9" 11.4-17.1 X7.6-11.4" i-i.9y 7.6-11.4 x3.8-7.6y 1.9-3.8* 9.5-13.3 x 5.7-1i.4 z

26. I-2a 16-21 x 3.5-38" 1 row 695-714 165-183 Rauvolfia Ib 6-13 x 5-26b 30-40 xi 1.5- manmi 1.8-3" 9-12 x 5-7.5" if i.5-3y 6.5-12.5 x6-n.8y z i-9-4z U.3-I4xi5.5-I9

27. I-I-5* 23-30 x 17.5-38" 1-2row s 714-732 201-275 Rauvolfia ib 5.5-9.8 X 4-20b 58-67 x 8.5-141 mombasiana 1-3" 10-15.5X 6.5-15" I-2y 4-9.5 x 3-6.9" i-3z 5-10x5.5-13.3*

28. 4-6" 15.2-20.9 x 8.6-24.7" homogenous 878-924 165-183 Saba 2-4b 9.5-19 x i3.3-34b 38-42x7-10' comorensis 10-15" 19-26.5 x 6.5-23" 14-20 x6-82 3-4" 17-28.5 x 9.5-i5.2y 4-6-5* 17-24.7 x 9-5-i9z

29. 2-3a 22-29 x 18-40" 1-2row s 1437- 256-275 b 1 Schizozygia i-i.5b 15-19 x i6-40 31-38 X6-15 1473 coffaeiodes 1-1.8" 6-13 X6-12" 17-31 x 7-92 2.6-4'' 5-11.8 X3-I2y 1-2* 7.6-17.1 XI14-28.5*

30. 1-2" 19-28.5 x 32.3-72" 1-3row s 787-824 229-256 Stephanostema Ib 15.2-20.9 x 24.7-494b 40-67 x 8-13.5' stenocarpum 1-3" 4.5-10 x 2-9" 23-38 x 9.5-152 y 1-3" 7.5-10 x 6-8 z 1-2.5* 7.6-17 x n-28

68 WageningenAgricultural UniversityPapers 96-1 (1996) Species Cuticle size Epidermal cells, size in Palisade, num­ Midrib Lamina in um urn. bero f rows and thickness thickness sizes of 1/2ro w (um) (um)

a 31- i 20-33x11-33" 3 (-4) rows 970-988 192-238 Strophanthus ib 19-28 x i5-35b 9-26 x 8-101 courmotii 1" 15.2-26.6 x 7.6-24.7" 10-15 x 5-102 2y 15.2-20.9 x i3.3-24.7y 15.2-24.7 x 9.5-20.92

32- 2-5-4-5a 14-34 x 23-41" 2-3 (-4) rows 1281- 266-311 Tabernaemontana 2-5-3-2b 6-36 x 24-43b 14-44x6.5-111 1318 2 elegans 2.5-3-5'1 9-11 x 6.5-13" 10-35 x 6-I2 2.5-3-5y 4.5-11x2.5-8.5" z 2-5-4-5z 7-11 x5.5-io

33- 3-5a 15-26 x 11-29" 1-2 (-3) rows 2031- 329-403 Tabernaemontana I-2b 12-23 x 9-25b 25-39 x 8-121 2105 pachysiphon 7.2-8" 11-14x8-11" 19-26x8-112 y 3-5" 8-13 x 6-<).5 7-8z 7-11 X5.5-ioz

34- 3-7" 6-25 x 9-35" 2-3 (-4) rows 2013- 329-384 b 1 Tabernaemontana 3-4-5b 8-18 x i6-27 31-40 x 9-14 2141 2 stapfiana 5-6.5x 11.5-14.5x5-11-5 * 22-28 x 10-13 5-6" 5-iox3-i3y z 7-5-9z 9-10 x 7-i3-5

35- 1-9-3-8" 19.5-2.8 x 11.4-32" 1-2(-3 ) rows 604-641 165-220 b Tabernaemontana i-9b 11.4-15X i5X26.6 25-35 x 9-141 ventricosa 3-8-7-6" 9.5-19 x 7.6-17" 22-30 x 20-142 3-8-5-7" 7.6-13.3 x 5.7-9.5" 2 3-8-5-7z 13.3-20.9 x 7.6-I5.2

36. 1.9-3.8» 11.4-13.3 x 15-44" 2-3 (-4) rows 439-476 146-220 b 1 Voacanga i.9-4-3b 5-18 x 9-36 21-25 X8-13 africana 3.8-6.5" 11-17X 10-20" y 1.9-3.8" 7-i3X4-8 z 3-8-5-7 11-22 X9-i8

37- i-9-3-8a 13.3-19x11.4-30.5" 2-3 (-4) rows 476-550 238-293 Voacanga i-9b 5-7-19 x 9-5-36" 29-44 x 9-5-I91 thouarsü 1.9-3.8" 11.4-17 x 11.4-22.8" 1.9-3-8" 7.6-13.3 x 3.8-9.5y 2-5-3-8z 11.4-22.8 X9.5-I9Z

"=uppe r epdermis;b= lowe repidermis ;x =uppe repidermis ,midrib; y =lowe repidermis ,midrib ; z= margin ;' =first ro w ofpalisade ;2 =secon dro w ofpalisade .

WageningenAgricultural UniversityPapers 96-1 (1996) 69 Table2 .Stomata lsize ,densit yan dtyp eo f someEas tAfrica n Apocynaceae.

Species Stomatal Stomatal Densitype r Subsidiary Anticlinal Anticlinal lengthi n widthi nu m mm cells, surface walls walls urn adaxial abaxial

i. 15-19-22 n.5-13-16 650-663-719 Hypostomatic ±straigh t ±straigh t Acokanthera cyclocytican d angular angular oppositifolia paracytic

2. 23-30-33.5 19-20-22.5 150-169-188 hypostomatic straight, slightly Adenium cyclocytican d angular curved obesum staurocytic

3- 18-20-25 19-20-20 140-150 hypostomatic undulate undulate Alalia paracytic microstylis

4- 21-23-25 20-21-23.5 400-500 hypostomatic very very Ancylobotrys amomocytic undulate undulate petersiana

5- 27-29-31 24-27-29 200-300 anomocytic undulate undulate Ancylobotrys tayloris

6. 15-16-19.5 8-11-13 150-172-200 hypostomatic very very Baissea paracytic undulate undulate myrtifolia

7- 29-30-36 23-28-32 66-132-176 hypostomatic straightt o straightt o Carissa cyclocytic slightly slightly edulis curved curved

8. 17-19.5-22 I4-I7-I9-5 418-534-572 hypostomatic straight straightt o Carissa paracytic angular slightly tetramera curved

9- 20-21-25 13-15-19 81-150-220 hypostomatic straightt o undulate Carvalhoa paracytic slightly campanulata curved

10. 17-20-23 15-16-19 288-340 hypostomatic ±straigh t straightt o Clitandra anomocytic toslightl y slightly cymulosa curved curved

ii. 14.6-16.6 14.6-16.6 700-825-925 hypostomatic slightly slightly Dictyophleba anomocytic undulate undulate lucida

70 WageningenAgricultural UniversityPapers 96-1 (1996) Species Stomatal Stomatal Densitype r Subsidiary Anticlinal Anticlinal lengthi n widthi nu m mm cells, surface walls walls um adaxial abaxial

12. 16-18 14-16 260-340 hypostomatic straight straight Diplorhynchus paracytic condylocarpon

13- 19-22-25 10-13-15 300-320 hypostomatic straightt o slightly Funtumia paracytic slightly undulate elastica undulate

14- 17-26-31 14-16-19 250-289-312 hypostomatic straight slightly Holarrhena paracytic angular curved pubescens anomocytic angular

15- 17-19-22 16-17-19 266-280 hypostomatic straightt o slightly Hunteria anomocytic very curved congolana anisocytic slightly curved

16. 21-24-25.7 22-24.7 294-338-350 hypostomatic straightt o slightly Hunteria anomocytic slightly curved zeylanica anisocytic curved

17- 15-18 13-15-16 400-450 hypostomatic undulate undulate Landolphia anomocytic buchananii

i8. 15-16 13-15-16 500-525-625 hypostomatic undulate undulate Landolphia anomocytic watsoniana

19- 21-25-31 10-15-18 200-219-213 hypostomatic slightly slightly Mascarenhasia paracytic undulate undulate arborescens

20. 15-18-19 9-10-15 168-200-370 hypostomatic slightly undulate Motandra anomocytic curved guineensis

21. 15-21-22.5 10-12.5-14 369-406-439 hypostomatic slightly undulate Oncinotis paracytic curved tenuiloba angular

22. 16-20-22 14-15-20 133-148-153 hypostomatic slightly angular Pleiocarpa cyclocytic curved slightly bicarpellata undulate

WageningenAgricultural UniversityPapers 96-1 (1996) 71 Species Stomatal Stomatal Density per Subsidiary Anticlinal Anticlinal length in widthi nur n mm cells, surface walls walls urn adaxial abaxial

23. 16-20.8-23 15-16.6-19 200-219-238 hypostomatic slightly slightly Pleiocarpa cyclocytic curved curved pycnantha

24. 21-25-31 10-15-18 100-130-160 hypostomatic undulate undulate Rauvolfia paracytic caffra

25- 19-20.9-32 11-16-18 63-81-125 hypostomatic undulate undulate Rauvolfia paracytic mannii

26 20.8-27-29 11.5-14-17 125-138-175 hypostomatic straight undulate Rauvolfia paracytic mombasiana

27. 13-15-18 11-16 850-856 hypostomatic slightly slightly Saba anomocytic curved curved comorensis

28. 27-28.5-31 16-18-20.3 63-88-106 amphistomatic straight slightly Schizozygia paracytic angular curved coffaeiodes

29. 15-18-21 9-10-13 132-220-264 hypostomatic slightly undulate Stephanostma paracytic undulate stenocarpum

30. 24-27-33 17-19-29 63-100-144 amphistomatic slightly undulate Strophanthus paracytic curved courmontii anomocytic- angular actinocytic

3i- 30.6-34.8 25-27.8 31-56-81 hypostomatic straight slightly Tabernaemon- paracytic angular curved tana angular elegans

32. 29-32-33.4 21-22-23.6 31-56-69 hypostomatic straight ± straight Tabemaemon- paracytic angular angular tana pachysiphon

33- 26-28-36.5 19.5-21-25 99-109-123 hypostomatic straight ± straight Tabernaemon- paracytic angular angular tana stapfiana

72 WageningenAgricultural UniversityPapers 96-1 (1996) Species Stomatal Stomatal Densitype r Subsidiary Anticlinal Anticlinal lengthi n widthi n um mm cells, surface walls walls urn adaxial abaxial

34- 26-29-32 20-23-25 30-60-88 hypostomatic ±straigh t ±straigh t Tabernaemon- paracytic angular angular tana ventricosa

35- 26-28 18-20 60-80-120 amphistomatic ± ± Voacanga paracytic straight straight africana angular angular

36. 25-28-30 20-23-25 80-90-140 amphistomatic ± ± Voacanga paracytic straight straight thouarsii angular angular

WageningenAgricultural UniversityPapers 96-1 (1996) 73 Table3 .A summar yo fApocynacea especie si nEas tAfrica , includingthei rsub-divisio nint otribe s and subtribes.Tribe sar ei nbol dface , subtribesi nitalics ,* = specie sexamined ,Afr = Africa, As= Asia.

Subfamily Plumeriodeae l.Carisseae 2. Tabernaemontaneae 1.1 Carissinae 11. Voacanga 12Af r + As 1. Carissa 20 Afr + As 25. V. africana* i.C. bispinosa 26. V.thouarsii * 2.C . edulis* 12. Tabernaemontana 99 tro 3. C. tetramera* 27. T.elegans * 2. Acokanthera 5 Afr 28. T.odoratissim a 4. A. laevigata 29. T. pachysiphon* 5. A. oppositifolia 30. T. stapfiana* 6.A . schimperi 31. T. ventricosa* 1.2 Landolphiinae 13. Carvalhoa 1 Afr 3. Landolphia 60 Afr 32. C. campanulata* 7.L . buchananii* 14. Schizozygia 1Af r 8.L . eminiana 33. S. coffaeiodes* 9.L . kirkii 3. Plumerieae 10.L . landolphiodes 3.1 Aspidospermatinae 11. L. owariensis 15. Diplorhynchus 1Af r 12.L . parvifolia 34. D. condylocarpon* 13. L. watsoniana* 3.2 Alstoniinae 4. Clitandra 1 Afr 16. Alstonia 40 trop 14.C . cymulosa* 35. A. boonei 5. Dictyophleba 5 Afr 4. Alyxieae 15. D. lucida* 4.1 Rauvolfiinae 6. Ancylobotrys 7 Afr 17. Rauvolfia 60 trop 16.A . amoena 36. R. caffra* 17. A. petersiana* 37. R. mannii* 18. A. tayloris* 38. R. mombasiana* 7. Saba 3 Afr 39. R. volkensii 19. Saba comorensis* 40. R. vormitoria 1.3 Pleiocarpinae 5. Cerbereae 8. Picralima 1Af r 18. Cerbera 7 Seych-Oc 20. P.nitida * 41.C. manghas 9. Hunteria 12Af r + As 21. H. congolana* Subfamily Apocynoideae 22. H. zeylanica* 6. Wrighteae 10. Pleiocarpa 5 Afr 6.1 Neriinae 23. P. bicarpellata* 19. Adenium 5 Afr 24. P.pyenantha * 42. A. obesum*

74 WageningenAgricultural UniversityPapers 96-1 (1996) 6.2 Wrightiinae 27. Alafia 23 Afr 20. Wrightia 23 Afr + As 61. A. erythrophthalma 43. W. demartiniana 62. A. caudata 21. Pleioceras 5 Afr 63. A. lucida 44. P.oriental e 64. A. microstylis* 22. Stephanostema 1 Afr 65. A. orientalis 45. S. stenocarpum* 66. A. schumanii 23. Strophanthus 38 Afr + As 67. A. zambesiaca 46. S. courmontii* 7. Apocyneae 47. S. eminii 7.1 Ichnocarpinae 48. S. hispidus 28. Motandra 3 Afr 49. S. hypoleucus 68. M. guinensis* 50. S. kombe 29. Baissea 18 Afr 51. S. mirabilis 69. B. leonensis 52. S. petersianus 70. B. leontonori 53. S. preusii 71. B. major 54. S. sarmentosus 72. B. myrtifolia* 55. S. welwitschü 73. B. viridifolia 56. S. zimmermannianus 30. Oncinotis 7 Afr 6.3 Malouetiinae 74. O. glabrata 24. Funtumia 2 Afr 75. O. hirta 57.F . africana* 76. O. pontyi 58. F. elastica* 77. O. tenuiloba* 25. Mascarenhasia 8 Afr 59.M . arborescens* 6.4 Alafiinae 26. Holarrhena 4 Afr + As 60. H. pubescens*

WageningenAgricultural University Papers 96-1 (1996) 75 1.9 ANATOMY REFERENCES

Araujo, R.R. de, da Silva, A.M., & Gill, F.S., 1984. Plumeria rubra L. var. alba - Apocynaceae. Anatomia foliar Rodriguésia 36 (5a):67-72 . Anheisser, R., 1900.Übe r die aruncoide Blattspreite. Flora 87:64-94 . Areschoug, F.W.C., 1897. Über die physiologischen Leistungen und die Entwickelung des Grundgewebes des Blattes.Lund s Univ.Arsskr . 33,2 : 1-46. Baas, R, 1975. Vegetative anatomy and the affinities of Aquifoliaceae, Sphenoste- mon, Phelline, and Oncotheca. Blumea 22, 311-407. Ballard, C.W., 1926.Taxonom y andpharmacognos y ofth egenu sApocynum. Proc. Internat. Cong. Plant. Sei. 2: 1406-12. Chandra, V., Kapoor, S.L., Sharma, P.C., & Kapoor, L.D., 1969. Epidermal and venation studies in Apocynaceae I.Bull .Bot . Surv. India 11: 286-9. Chandra, V., Kapoor, S.L., Sharma, P.C., & Kapoor, L.D., 1972. Epidermal and venation studies in Apocynaceae. Bull.Bot . Surv. India. De Kruif, A.P.M., 1985. A revision of Oncinotis Benth. In: Leeuwenberg, A.J.M. (ed.), Series of revisions of Apocynaceae XVI-XVIII. Wageningen Agricultural University Papers 85,2 : 5-45. Fjell, I., 1983. Anatomy of the xeromorphic leaves of Allamandra neriifolia, Thevetiaperuviana and Vincaminor (Apocynaceae). Nord. J. Bot. 3: 383-392, struct. 027. Esau, K., 1965. Plant anatomy, ed. 2.John , Wiley, New York. Haberlandt, G., 1934. Über die Sonnen und Schattenblätter der Crataegomesplili und ihrerEltern . Sitz. Preuss. Akad. Wiss.,Phys-Math . KL (20):363-76 . Hartog van ter Tholen, R.M., & Baas, P. (1979). Epidermal characters of the Celastraceae (sensu-lato). Act. Bot. Neerl. 27:355-8 . Hickey, L.J., 1979. A revised classification of the architecture of dicotyledonous leaves. In: Metcalfe, CR., & Chalk, L., (eds), Anatomy of the dicotyledons ed. 2, Vol. 1:25-39 . Oxford, Clarendon Press. Holm, T., 1910.Apocynum cannabinum L. Merck's Pep. 19, 277-80. Jansen, W.T., & Baas, P., 1973. Comparative leaf anatomy of Kokoona and Lophopetalum (Celastraceae). Blumea 21: 153-78. Kapoor, S.L., Sharma, P.C., Chandra, V., & Kapoor, L.D., 1969. Epidermal and venation studies in Apocynaceae H.Bull .Bot . Surv. India 11: 372-6. Kapoor, S.L., &Mitra , R., 1979.Epiderma l and venation studies of Apocynaceae - VI. Bull. Bot. Surv. Ind. 21: 68-80. Karatela, Y.Y., & Gill, L.S., 1983. Leaf epidermal features of two species of Rauvolfia (Apocynaceae) from Nigeria. Philip.J . Biol. 12: 186-189. Karatela,Y.Y. ,& Gill,L.S. , 1988.Observation s onlea f epidermal features of some medicinal plants from Nigeria. Herba Hungarica 27: 7-18. Leeuwenberg,A.J.M. ,& Berndsen ,M. , 1988.Clitandra Benth .an d Chamaeclitan- dra(Stapf ) Pichon.Serie so fRevisio no fApocynacea eXXV .Bull .Jard .Bot .Nat . Belg. 58: 159-168. Metcalfe, CR., & Chalk, L., 1950.Anatom y of the dicotyledons. Vol.H :904-917 . Oxford, Clarendon Press.

76 WageningenAgricultural UniversityPapers 96-1 (1996) Newcombe, F., & Patel, B.A., 1966. Comparative anatomy of the leaves of V. schweinfurthii andV. africana. J. Pharmacy Pharmacol. 18, supp. 133-141. Ngan,P.T. , 1965.A revisio n ofth e Wrightia(Apocynaceae) .Ann .Miss .Not . Gard. 52: II4-I75- Nyawuame, H.G.K., & Gill, L.S., 1991. Cuticular studies of some West African specieso fth eApocynacea eo fmedicina lvalue .Fedde sRepertoriu m 102,1-2:87- 104. Paganelli Cappeletti, E.M., 1975. Studio morfologico al microscopio elletronico a scansione di foglie diAtropa belladonna L. Bot. Ital . 7:24-25 . Poulsen, V.A., 1917. Plantenanatomiske Bidrag, Anatomiske Bemaerk-ninger om Bladbygningen hos nogle Apocynaceer. Vidensk Medd. naturh. Foren. Kbh. 68: 299-307. Sayeedud-Din,M. , 1941.Som ecommo nIndia nherb swit hnote si nthei r anatomical characters. J. Bombay Nat. Hist. Soc.42 : 280-282. Sharma, G.K, &Dunn , D.B., 1968.Effec t of environment onth e cuticular features in Kelanchoëfedschenkoi. Bull.Torre y Bot. Cl. 95: 464-73. Sharma, P.C., Chandra, V., Kapoor, S.L., & Kapoor, L.D., 1970. Epidermal and venation studies in Apocynaceae - HI.Bull .Bot . Surv. India 12,1-4: 92-96. Solereder, H., 1908. Systematic anatomy of the dicotyledons, Vol. 1,2 (Translated by L.A. Boodle & E.Frisch , revised by D.H. Scott).Oxford , Clarendon press. Stace,C.A. , 1965.Cuticula r studiesa sa nai dt oplan ttaxonomy .Bull .Br .Mus .Nat . Hist. Bot. 4 (1): 1-78. Van Staveren, M.G.C., &Baas ,P. , 1973.Epiderma l leaf characters inth e Malesian Icacinaceae.Act a bot. neerl. 22:329-59 . Watson, R.W., 1942. Effect of cuticular hardening on the form of epidermal cells. New Phytol. 41: 223-9. Wilkinson, H.P., 1971. Leaf anatomy of the various Anacardiaceae with special reference to the epidermis and some contribution to the taxonomy of the genus Dracontomelon Blume. Thesis,Universit y of London. Wilkinson,H.P. ,1979 .Th eplan tsurfac e (mainlyleaf) .In :Metcalfe ,CR. , & Chalk, L. (eds),Anatom y of theDicotyledons , ed. 2,vol . 1:97-165. Oxford, Clarendon Press. Wilkinson, H.P., 1989. Leaf anatomy of the Menispermaceae, tribe Tiliacoreae Miers. Botanical Journal of the Linnaean Society, 99: 125-174.

WageningenAgricultural University Papers 96-1 (1996) 77 APPENDIX1

List ofMateria l examined:

i. Acokanthera oppositifolia (Lam.) Codd, Kenya, Sagalla Mts., E.A. Omino 133, (EA, WAG). 2. Adenium obesum (Forssk.) Roem. & Schult., Kenya, Silaloi, E.A. Omino 123, (EA,WAG) . 3.Alafia microstylis K.Schum,Kenya ,Luke &Robertson 22ç8, (K). 4.Ancylobotryspetersiana (Kl.)Pierre ,Kenya ,Arabuk oSokok eForest ,E.A. Omino 213 (EA,WAG) . 5.A. tayloris (Stapf) Pichon, Kenya,Bud a mafisini, E.A. Omino 199, (EA, WAG). 6. Baissea myrtifolia A.DC, Kenya, Makadara Forest, E.A. Omino 174, (EA, WAG). 7. Carissa edulis (Forssk.) Vahl,Kenya, Sagalla mts,E.A. Omino 131, (EA,WAG) . 8.C. tetramera (Sacl.)Stapf ,Kenya, Arabuk oSokok eForest ,E.A. Omino212, (EA , WAG);Kenya, Galana Ranch,Bally 16743(K) . 9. Carvalhoa campanulata K. Schum., Kenya, Makadara Forest, E.A. Omino 175 (EA,WAG) . 10. Clitandra cymulosa Benth., Uganda, Sese Islands,Maitland 339(K) . 11.Dictyophleba lucida(K .Schum )Pierre ,Kenya ,Shimb aLodge ,E.A. Omino 166, (EA, WAG); Kenya,Makadar a Forest, E.A. Omino 181(EA , WAG). 12. Diplorhynchus condylocarpon (Muell. Arg.) Pichon, Tanzania, Lovett & Congdon 1933, (EA,WAG) . 13.Funtumia elastica (Preuss) Stapf, Uganda, Mengo,B. styles 228 (K). 14.Holarrhenapubescens (Buch.-Ham)Wall ,e xG.Don ,Kenya ,Musau-Rong eRd. , E.A. Omino 124,(EA ,WAG) . 15.Hunteria congolana Kenya,Mt .Kulal ,Bally 3960 (K). 16.H. zeylanica Kenya, Shimba Lodge, E.A. Omino 163, (EA, WAG). 17. Landolphia buchananii (Hall.f.) Stapf, Kenya, Ngangao Forest, Beentje 2192 (K). 18.L. watsoniana Romburgh, Kenya, E.A. Omino 173B, E.A. Omino 211 B,(EA , WAG). 19. Mascarenhasia arborescens A.DC., Kenya, Marere Waterworks, Perdue & Kibuwa 10220 (K). 20.Motandra guineensis (Schum. &Thonn. ) A.DC,Uganda , Synnot 725, (EA). 21. Oncinotis tenuilobaStapf ,Kenya, Mkomb aRiver ,E.A. Omino 192,(EA ,WAG) . 22.Picralima nitida Cultivated, Wageningen Bot.Garden , TheNetherlands . 23.Pleiocarpa bicarpellata Stapf,Kenya ,Chawi aforest ,E.A. Omino157,158, (EA, WAG). 24.P. pycnantha (K.Schum.) Stapf, Kenya,Mwel eMdog oForest ,E.A. Omino 187, (EA, WAG);Kenya ,Mer u Nat. Park,Ament &Magogo 169 (EA).

78 WageningenAgricultural UniversityPapers 96-1 (1996) 25. Rauvolfia coffra Sond., Kenya,Mer u National Park, Gillet 20801(K) . 26.R. mannii Stapf, Kenya Chawia Forest, E.A. Omino 154, (EA, WAG). 27.R. mombasiana Stapf, Kenya,E.A. Omino 162B, (EA,WAG) ;Kenya , Marere, E.A. Omino 170, (EA,WAG) . 28. Saba comorensis (Bojer) Pichon, Kenya, Ukunda, E.A. Omino 161 B (EA, WAG). 29. Schizozygia coffaeoides Baill., Kenya, Marere, E.A. Omino 168, (EA, WAG). 30. Stephanostema steonocarpum Tanzania,J &J Lovett 500, (K);Tanzani a Luke etal 3754 (EA). 31. Strophanthus courmontii Kenya, Ramisi Mrima Rd, E.A. Omino 211 (EA, WAG). 32.Tabernaemontana elegans Kenya,Shimb aLodge ,E.A. Omino 167(EA ,WAG) . 33. T.pachysiphon Kenya,Bud a Mafisini E.A. Omino 107 (EA, WAG),. 34.T. stapfiana Kenya,E.A. Omino 127.(EA , WAG). 35.T. ventricosa Tanzania, Lumi Forest, Gardner 2956 (K). 36. Voacanga africana Tanzania, P.Cribb &G. Wilson10693 (K);Tanzania ,Poes 6277/B (EA). 37.V. thouarsii, Tanzania, TG, Ruffo 1533.

WageningenAgricultural UniversityPapers 96-1 (1996) 79 2 AMONOGRAP HO FTH E SUBTRIBEPLEIOCARPINA E

(APOCYNACEAE-PLUMERIOIDEAE-CARISSEAE)

Serieso frevision so fApocynacea eXL I

2.1 SUMMARY

The subtribe Pleiocarpinae, belonging in the tribe Carisseae of the subfamily Plumerioideaeha sbee nrevise d andcomprise sthre egenera :Pleiocarpa, Hunteria andPicralima, with 5, 12an d 1 speciesrespectively . Allspecie s arerestricte dt o continental Africa, butfo rHunteria zeylanica. Pleiocarpa hasfiv e species,includin g anewl y described one:P. brevistyla. A new combination for a probable sixth species, P. picralimoides, was made for Carpodinopsis (?)picralimoides, butdu et oa scarcit y ofmateria lthi s species re­ mains doubtful. Threene wHunteria specie swer edescribed :H. macrosiphon,H. myrianthaan d H.oxyantha, whilea ne wcombinatio nH. hexaloba wasmade ,a sth egenu snam e Tetradoafall s intosynonym y withHunteria. Themonotypi cgenu sPicralima remaine d assuch . Of all Pleiocarpinae typification, full synonymy andliteratur e references are given,an da ke yprovide sth emean st oidentif yth especies .Detaile ddescription sar e accompanied with illustrations, andth edistribution s aremapped . Notes present further information whererelevant .

2.2 INTRODUCTION

Thepresen tchapte ri sa monographi crevision o fth esubtrib ePleiocarpina eo fth e Plumeriodeae-Carisseae (Apocynaceae),represente db y3 gener aan d 18 speciesi n continental Africa.Thi s monographi smainl ybase d onstud y of herbariumspeci ­ mens,i nadditio n theautho r hasha dth e opportunity of studying 6specie s in the fieldan di ncultivatio na tWageningen .Th esubtrib ePleiocarpina ecomprise s3 gen ­ era,Pleiocarpa Benth (5 spp.);Hunteria Roxb.(1 2 spp.) andPicralima Pierre (1 sp.).On eo fth especies ,Hunteria zeylanica, extendst o Asia. Mosto fth etyp emateria lcoul db etraced ,a fe wlectotype swer edesignate dhere , whileonl yon eneotyp ewa schosen . Allmeasurement sar efro m driedmaterial .Th euse so fth eplant swer ecompile d from herbariumsheet san dth eavailabl eliterature .Loca lname swer eno trecorded , asmos to fthe mprove dt ob eunreliable .Th esourc eo ninformatio n fromherbariu m sheets isarrange d systematically, e.g (Scott Elliot5690 ,K , 1892, SierraLeone) , where Scott Elliot isth ecollector , 5690th ecollectio n number, K,th eherbariu m Kewlodgin gth especimen ,189 2th eyea ro fcollectio nan dSierr aLeon eth ecountr y of collection.

WageningenAgricultural University Papers 96-1 (1996) 81 2.3 GENERAL PART

23.1 History of the subtribe

The subtribe Pleiocarpinae was first described by K.Schuman n in 1895 as a tribe based on three genera Notonerium, Lepinia and Pleiocarpa. Lepinia currently be­ longs to the subtribeAlyxiinae although it has multicarpellate fruits, but these are drupaceous and not baccate. Pichon (1948)first describe d it as a subtribe basing it on 5 genera, Polyadoa, reduced to synonymy by him in 1953 under Hunteria; Tetradoa, reduced to synonymy by Huber (1963) under Hunteria; Pleiocarpa, Picralima andHunteria. Inth elas tcomplet erevisio n ofth e subtribe Pleiocarpinae in 1953 Pichon recognised 18 species in 7 genera, Pleuranthemum (1 sp.), Comularia (2 sp.) and Tetradoa (2 sp.) (all currently synonyms of Hunteria); Hunteria (6 sp.), Carpodinopsis (3 sp.),currentl y a synonym of Pleiocarpa (3 sp.) and Picralima (1sp.) .

2.3.2 History of the genera Hunteria Roxb.,Pleiocarpa Benth. andPicralima Pierre

The genus Hunteria was first proposed as Apocyno-Nerium by Linnaeus in 1747 basedo non especie sH. zeylanica.Th enam ewa slate rrejecte db yRoxburg hi n 1814 when he merely listed it asHunteria. Helate r describedth egenu s in 1824basin g it on Hunteria corymbosa, a synonym of H. zeylanica, which had already been de­ scribed by Retzius in Cameraria in 1786.Th e genus is named after Dr.Willia m Hunter of the Bengal Medical Establishment, an eminent botanist and author of variousAsiati cResearche s ofth ehistor y of Pegu.G.Do n (1837)enumerate d 5spe ­ cies, 4 later attributed to the genera Wrightia, Alyxia and Chilocarpus and A.D e Candolle (1844) included 7 species in the genus, 3 later attributed to the genus Alyxia and2 t oMelodinus and Chilocarpus. F.Mique l (1856)enumerate d 4 species, 2 later attributed to the genera Rauvolfia and Alyxia. Pichon (1948) enumerated 7 species which he placed under two sections, section Euhunteria with five species, 3 later attributed to the genera Alyxia, Wrightia and Rauvolfia and section Pleuranthemum with two species from Africa. In the most recent revision of Hunteria inWes t Africa by Huber (1963), 5 species were recognised.

The genus Pleiocarpa was first described by Bentham in 1876,base d on P.mutica (thelectotyp e of thegenus ) andP. rostrata.Th enam e comes from theGree k words Pleios,mor ean dcarpos ,fruits , referring toth efruit s withmor etha n twocarpels .P. bicarpellata wasfirst describe db y Stapf in 1894.K .Schuman n (1895) enumerated 3 species, and Stapf (1902) distinguished 9 species, 4 were laterreduce d to synon­ ymyunde rP. pycnantha and 1 reducedt osynonym y underP. mutica. Pichon (1953) recognised 3 species,P. pycnantha with twovarietie s andHube r (1963) recognised 4 species,P. pycnantha with two varieties.

82 WeigerungenAgricultural UniversityPapers 96-1 (1996) Pierre (1896) first described the genus Picralima basing it on P.klaineana, a syn­ onym of P. nitida, the only species of the genus which had already been described by Stapf (1895) inTabernaemontana. The name Picralima comes from the Greek wordpikro swhic hmean sbitter ,probabl y referring toth ebitte rseeds .Th egenu sha s since remained monotypic.

2.3.3 Geographical distribution

Hunteria zeylanica is the most widely distributed species which extends to Asia, from Sri-Lanka, India to S.Chin a (Hainan) including Malaysia and Indonesia. In Africa, its distribution is restricted to Somalia, Kenya,Tanzani a (including Pemba and Zanzibar) and Mozambique, mainly along the Zanzibar-Inhambane regional mosaic (sensu White 1979). It also extends to Zambezian and Somalia-Masai re­ gional centres of endemismi n Kenya and Somalia. Four species are found West to East of the Dahomey gap and these include Pleiocarpa pycnantha, the most widely distributed of the African species, which covers 8 phytochoria sensuWhit e (1979). H. umbellata is also widely distributed from Senegalt oZair ean dAngol abu trestricte dt oonl ytw ophytochoria . Pleiocarpa mutica extends from SierraLeon et o Gabon andPicralima nitida from Ivory Coast to Congo and Zaire, including Uganda. Two species ofHunteria, H. ghanensis andH. simii arerestricte d toth eWester n centreo fEndemism ,Wes to fth eDahome y gap.H. ghanensis isfoun d onlyi n Ghana and Ivory Coast, whereas the later extends from Guinea to Ivory Coast. Theres to f thespecie s isrestricte d toth eEaster n centreo fendemism ,Eas t of the Dahomeygap .H. camerunensis inCamerou n andGabon ;H. congolanai nZair ean d Kenya, in two phytochoria; H. macrosiphon in Gabon and Congo;H. oxyantha in Gabon, Congo and Zaire; Pleiocarpa bicarpellata extends further East to Kenya from Cameroun. P.rostrata is restricted toNigeria , Cameroun and Gabon. The remaining four species are endemic. Hunteria hexaloba and Pleiocarpa brevistylaar elocall yendemi ct oGabo nan dH. myriantha andH. densiflora i s found only in Zaire. Twospecies ,Pleiocarpa bicarpellata andHunteria congolana, showdisjunction s in their current distribution areas.

2.3.4 Morphology

Habit Themember s of the subtribe Pleiocarpinae areusuall y smalltree so rshrubs ,rarel y lianescent.The y are completely glabrous in all parts except inside thecoroll a tube, below orabov eth e stamens.Predominantl y shrubby species upt o 5m hig h include Pleiocarpa rostrata and P. brevistyla; Hunteria ballayi, H. camerunensis, H. hexaloba andH. macrosiphon.Th eres to fth especie srarel y exceeda heigh to f 20m. Pleiocarpa pycnantha may attain aheigh t of 30m an dPicralima nitida, aheigh t of 35 m.

WageningenAgricultural UniversityPapers 96-1 (1996) 83 Bark The bark of most species is thin, smooth and palet o dark grey or brown. In some species like Picralima nitida and Hunteria oxyantha the bark may be rough, hard and brittle or it may be longitudinally fissured with straight horizontal cracks in Pleiocarpa pycnantha. The inner bark is usually cream, pale yellowish-brown or orange.

Wood Thewoo do fth especie sattainin gexploitabl eheigh ti sver yhard ,dens ean ddurable , shinydar kbrow no rpal eyello wt oorange ,an di suse dt omak esmal lsundr y objects or sometimes used for local construction.

Leaves The leaves are opposite and decussate, rarely in whorls of 3-5 as in Pleiocarpa pycnantha andsometime sternat ea si nP. mutica.Th eblad ei spaper y to coriaceous when fresh, elliptict ooblon g with anacuminat e apexexcep t inHunteria zeylanica where the apex is variable especially in specimens from Asia.Th e smallest leaves arefoun d inHunteria oxyantha andH. ghanensis (upt o 13.5x 4.6cm )an dth elarg ­ esti nPleiocarpa mutica (upt o 29.5x 11.4c mlong) .Accordin gt oth e terminology proposed by Hickey (1979) for the architecture of dicotyledonous leaves, all the leaves inPleiocarpina e arecamptodromou sa sth esecondar y veinsd ono t terminate at the margin.Th e venation is either brachidodromous with secondariesjoine d to­ gether in a series of prominent arches or eucamptodromous with secondaries con­ nected to superadjacent veinsb y aserie s of cross-veins without forming prominent marginal loops.Al l the species have thebrachidodromou s type of venation except Hunteria myriantha which is eucamptodromous. H. simii, H. hexaloba, H. camerunensis and Pleiocarpa wstrata have leaves which are entirely brachido­ dromous or brachidodromous in the upper 0.5-0.75 and eucamptodromous in the lower part.

Inflorescence Theinflorescenc e inPleiocarpa ispredominantl yaxillar yan dsometime sals otermi ­ nal, sessile, ramiflorous, and when many-flowered, fasciculate. In Hunteria and Picralima theinflorescence sar epredominantl yterminal ,sometime sa tth esam etim e axillary,few -t omany-flowere d cymesexcep ti nH. ballayi whereinflorescence s are axillary, sessile and ramiflorous.Th e bracts are usually sepal-to scale-like.Th e pe­ duncle sizei shighl y variable inmos t species andrange s from ot o6 m m long in H. camerunensis to 15t o4 0m mlon gi nHunteria densiflora.Th epeduncl ean dpedice l base and inflorescence axes in Hunteria umbellata may appear swollen due to exudation of resin from the colleters and young buds.Th e inflorescences in H. zeylanica, H. ghanensis and H. myriantha arela x and manyflowered. Th e number of flowers is also very variable and may be as low as 1(-4 ) in Pleiocarpa wstrata and P.brevistyla to as high as 160i nHunteria myriantha.

84 WageningenAgricultural UniversityPapers 96-1 (1996) Flowers The subtribe Pleiocarpinae is distinguished from the rest of the members of the Carisseae by the ovary which is completely apocarpous with 2-5 carpels and the flowers which are completely glabrous except inside below or above the insertion of the stamens.

Calyx The calyx consists of five (sub)equal sepals which are almost free, completely gla­ brous on both sides inPleiocarpa, with colleters insideu pt o0.7 5 of theirlengt h in Hunteria (excepti nH. myriantha) andi n1- 4row s atth eextrem ebas ei n Picralima. The sepals are tightly clasped to the base of the corolla tube in Hunteria and Picralima due to the exudation of resin from the colleters and is persistent in fruit in all genera.Th e size varies from 0.6-4.5x 0.3-3 n™ m Hunteria and Pleiocarpa to 5-5-7-5x 3-5-5m mlon g in Picralima.Th e texture is variable and may be mem­ branaceous asi n Hunteria myriantha, to thick and coriaceous as inPicralima. The calyx lobes arealway s imbricate in bud,bu t are strongly imbricate even at anthesis in Picralima and rarely also inPleiocarpa rostrata.

Corolla Salverform, sympetalous,wit hfive lobes ,contorte d inbud .Th ecolou ri spur e white in Pleiocarpa except in P. pycnantha where it may vary from white via yellow to orange; whitet o lime-yellow inPicralima and white, cream, to yellow or even red inHunteria. The corolla head is usually oblong with an obtuse apex in mature bud inPleiocarpa andPicralima andmostl yovoi dwit ha nacut eape xi nHunteria except H. ghanensis where iti s globose andwit h arounde d apex.Th etub e ismor e or less cylindrical, widened at the base and at the insertion of the stamens.Th e tube is al­ ways thick and fleshy in Picralima, sometimes thick in Pleiocarpa rostrata and in some species of Hunteria asin H. zeylanica; it is very constricted at the orifice in H. camerunensis. Thelengt h varies from 3-4 mm inH. camerunensis to 12-32 mm long inPleiocarpa rostrata.Ther ei s abel to fpubescenc ebelo wth einsertio n of the stamens which may continue up to the mouth of the tube as in H. congolana, or may be reduced to minute tufts of hairs at the base of each filament as in H. camerunensis. Thecoroll alobe soverla pt oth elef t inbud ,ar etwiste di nbu di n some species of Hunteria and are always shorter than the tube in Pleiocarpa and shorter orlonge rtha nth etub ei nHunteria andPicralima. The shape isovat e in Picralima, ovate to oblong in Hunteria and variable in Pleiocarpa where it is obovate in P. rostrata.Th e apex is acute to obtuse in all species except in H. ghanensis andP. rostrata where it is rounded and is spreading to recurved at anthesis.

Fruits The fruits are composed of two to five separate, indéhiscent mericarps. The mericarps aregenerall y smooth except inH. ballayi (slightly rough and warty) and in some specimens of P. rostrata (partly or wholly rugose). They are mostly subgloboset oobovoi dexcep ti nHunteria simiiwher ethe yar eoblong .Th efruit s are divergent at an angle of 1800 except in H. ghanensis, 900 and usually turn yellow

WageningenAgricultural UniversityPapers 96-1 (1996) 85 andfinally orang ewhe nripe. Th esmalles t fruits arethos eo fH. ghanensis, 10-15 x 9-12 mm followed by those of H. macrosiphon, H. zeylanica, H. densiflora, H. bailayi, H. congolana, and H. hexaloba which are 15-28 x 20-21 mm, while those of H. oxyantha measure 28-35 x 20-26 mm. The largest fruits are those of H. umbellata: 31-60 x 40-50 mm.Th e fruits of H. simii measure 37-60.5 x 10-18 mm but are usually clearly longer than wide. The seeds of Hunteria bailayi are swallowed together with the sweet mesocarp to act as a fertility drug in Central African Republic, and the pulp and seeds of Picralima nitida are used as an arrow and fish poison.

2.3.5 General taxonomie references

Bentham, G. &J.D.Hooker , 1876. Genera Plantarum 2: 699. DeCandolle ,A. , 1844.Prodromu s systematisnaturali sregn ivegetabili s 8:349-351. Don, G., 1837.A general system of gardening and botany 4: 105. Rivington et al., London. Hickey, L.J. 1979. A revised classification of the architecture of dicotyledonous leaves. In:Metcalfe , CR., L.Chalk, C.Fagette & R.A.Howar d (eds). Anatomy of Dicotyledons, ed. 2,1: 25-39. Clarendon Press. Oxford. Huber,H. , 1963.Apocynaceae .In :Hepper ,F.N .(ed) .Flor ao f WestTropica l Africa ed. 2,2 : 62-64. Linnaeus, C, 1747.Flor a zeylanica, 404. Miquel, F., 1856. Apocyneae. Flora Indae Batavae 2 / Flora van Nederlandsen Indië 2:384-459 . Pichon, M., 1948. Classification des Apocynacées I, Carissées et Ambelaniées. Mémoires du Muséum National d'Histoire Naturelle, série 2, 24: 111-181. Pichon, M., 1953. Classification des Apocynacées: XXXVI, Révision des Pléïocarpinées. Boletim da Sociedade Broteriana 2a série, 27:73-153 . Pierre, L., 1896. Plantes du Gabon. Bulletin Mensuel de la Société Linnéenne de Paris 2: 127-1279 Retzius,A .J. , 1786. In Fasciculus Observationum Botanicarum 4: 24. Roxburgh,W. , 1814.Hortu s Bengalensis. Serampore: 84. Roxburgh,W. , 1824.Flor a Indica, ed. Carey. Serampore vol.2 : 531-533. Schumann, K., 1895. Apocynaceae. In: Engler and Prantl (eds), Die Natürlichen Pflanzenfamilien 4, 2: 133-135. Stapf, O., 1894.Apocynaceae , Diagnoses Africanae 1.Ke w Bulletin 1894: 19-25. Stapf, O., 1902.Apocynaceae . In:W.T.Thiselton-Dye r (ed.),Flor a of Tropical Af­ rica 4, i: 96-105 . Reeve & Co., London. White, F., 1979.Th evegetatio n of Africa. A descriptive memoir to accompany the UNESCO/AETFAT/UNSO vegetation map of Africa. UNESCO.

86 WageningenAgricultural UniversityPapers 96-1 (1996) 2.4 TAXONOMIC PART

2.4.1 Thesubtrib ePleiocarpina e

Subtribe Pleiocarpinae (K.Schum. )Picho ni nMém .Mus .Natn .Hist .Nat .sér . 2, 24: 158 (1948), Leeuwenberg inWageninge n Agr. Univ.Pap .94 , 3:5 3(1994) . - Typegenus : PleiocarpaBenth .

Basionym:Pleiocarpeae K .Schum . inEngle r& Prantl ,Nat .Pflanzenf . 4,2:133 (1895),partl y asfo rPleiocarpa.

Unarmedtree so rshrubs ,sometime slianescent ;branchlet sglabrous .Leaves oppo ­ site,o rles softe n inwhorl so f3-5 ,petiolate ;petiol eglabrous ;blad emostl yelliptic , entirean dglabrou so nbot hsides .Inflorescence i ndens efascicle s ori nla xt odens e cymes, terminal and/or axillary, sometimes ramiflorous, glabrous. Bracts small. Flowersactinomorphic ,5-merous .Sepals subequal ,fre e orconnat ea tth eextrem e base,persisten ti nfruit ,glabrou so rwit hcolleter sinsid eo nth einne rside ,no tciliate . Corollasalverform ,whit et oyellow ,rarel yals ored ;tub enearl ycylindrical ,widene d at theinsertio n of the stamens,no ttwisted ; lobes overlapping toth elef t in bud, spreadingan dofte n recurvedlater .Stamens fre e fromeac hother ,included ,inserte d ato rabov eth emiddl eo fth ecoroll atube ;anther sintrorse ,ovat eo roblong ,cordat e or saggitate atth ebase ,o ftw oparalle l cells,entirel y fertile orwit ha smal lsteril e acumen,glabrous .Pistil glabrous ;ovar ysuperior ,o f2- 5 separatecarpels ,unite da t theextrem ebas eb ya disk-lik ethickening ;pisti lhea dsubglobos et ooblon gan dwit h apronounce dstigmoi dape xi nHunteria an dPleiocarpa; ovule s1 -3 0(-150 )i neac h carpel.Frui to f2- 5fleshy, indéhiscen tmericarps ;mericarp s globoset ooblong , 1- 100seeded ;wal lthi nt othick fleshy, o rfibrous .Seeds variousl yshaped ,hilu mbasal , endosperm rather hard, starchy, leaving acavit y around the radicle base;embry o straight, spathulate;test athi nt othick ,membranaceou s tocoriaceous .

Threegener ai ntropica lAfrica .On especies ,Hunteria zeylanica, extend sint oAsia .

2.4.2 Keyt oth egener ao fth esubtrib ePleiocarpina e

1.Sepal scompletel yglabrou so nbot hsides ;pisti lhea dsubglobos et ooblong ;inflo ­ rescence sessile,predominantl y axillary andramiflorous ; bractsles stha n0. 4 mmlon g 3.Pleiocarp a Sepalsglabrou soutside ,wit hcolleter sinside ;pisti lhea dwit ha stigmati coblon g tosubglobos ebasa lpar tan da stigmoi dape xu pt o1. 5m mlong ;inflorescenc e predominantly terminal andsometime s axillary (excepti nH. ballayi); bracts atleas t0. 5m mlon g 2 2.Sepal simbricat eeve na tanthesis ,(3.5-)5.5-7- 5m mlong ,wit hcolleter su pt oo .1 8 of theirlength ;test acoriaceous ;cotyledon s withsecondar y nerves;fruit s 11- 20x 8-1 5 cm,wal lver y hardan dver yfibrou s 2.Picralim a

WageningenAgricultural UniversityPapers 96-1 (1996) 87 Sepals imbricate in bud, 0.7-2.5 mm long, with colleters covering atleas t 0.4 of the sepal length; testa membranaceous; cotyledons without visible secondary nerves;fruit s 2-5 x 1-5 cm, wall mostly fleshy 1. Hunteria

2.5 HUNTERIA ROXB.

Hunteria Roxb.i nFl .Ind .Seramp .ed .Care y &Wall .2 :53 1(1824 )& ed . 2,1:695 (1832);G.Do n inGen .Syst .4 : i05(i837);ThwaitesinEnum.Pl.Zeyl. 191(i860) ; Benthami nBentha m& Hooke r f., Gen.PI . 2:69 8(1876) ;Hooker , J.D. inFl . Brit. Ind. 3: 637 (1882); K.Schuman n in Engler Pflanzenw. Ost.-Afr. C: 317 (1895); Stapf inFI .Trop .Afr . 4,1:104 (1902) ;O .Kuntz ei nPos t& O .Kuntze ,Lexico n 288 (1904); Pichon inMém . Mus.Natn . Hist. Nat. 24: 160(1948 ) & in Bol. Soc. Brot, sér. 2,27: 88 (1953);H .Hube ri nFl .W.Trop .Afr . ed. 2,2:62 (1963);Hube r inRev . Handb. Fl. Ceylon 1: 10 (1973);Tsian gYin g in Fl. Rep. Pop. Sin. 63: 15 (1977); Markgraf inBlume a 30: 169(1984) ;Kupich a inFl .Zamb .7,2:43 0 (1985). -Type species:Hunteria corymbosa Roxb. (=H. zeylanica (Retz) Gardn. exThw.) .

Heterotypic synonyms:

Polyadoa Stapf in op. cit: 103; Pichon in Mem. Mus. Natn. Hist. Nat. 24: 159 (1948). Lectotype species: P.umbellata (K.Schum. ) Stapf (= H. umbellata (K. Schum.) Hall, f.), designated here. Tetradoa Pichon in Bull. Soc. Bot. France 93, 7-8: 251 (1946) & in Mém. Mus. Natn. Hist. Nat. 24: 160 (1948) & in Bull. Soc. Brot. sér. 2, 27: 119 (1953). - Type species:Tetradoa hexaloba Pichon, designated by Pichon in 1953 (= H. hexaloba (Pichon) Omino. Hunteria sect. Euhunteria Pichon in Mém. Mus. Nat. Hist. Nat. 24: 161 (1948), partly.-Typ e species:Hunteria zeylanica (Retz.)Gard ,e xThw. ,designate dhere . Hunteria sect.Pleuranthemum Pichon, I.e. - Lectotype species:H. ballayi, desig­ nated here. Pleuranthemum Pichon in Bol. Soc.Brot . sér. 2,27 : 112 (1953).-Typ e species: P. ballayi (Hua) Pichon (=H. ballayi Hua), designated by Pichon in 1953. Comularia Pichon op. cit. 114(1953) .-Typ e species:C. camerunensis (K.Schum . ex Hall, f.) Pichon (= Hunteria camerunensis K.Schum . ex Hall. f.).

Shrubs ortrees ,rarel y lianescent withwhit e latex or occasionally colourless (in H. zeylanica) inal lparts ;branchlet sglabrous .Leaves opposite,petiolate ,thos eo f apair equal or subequal;blad emostl y elliptic,entir ean dglabrou s onbot h sides.Inflores­ cencemostl yterminal ,sometime sals oaxillary ,few-t o many-flowered, denset olax , pedunculate; peduncle 2-45 mm long, glabrous. Flowers fragrant, actinomorphic except for the subequal sepals.Sepals lightgreen ,thi nt othic k andcoriaceous , free or connate at the extreme base, membranaceous atth e edge, glabrous outside, and with colleters inside up to 0.9 of their length. Corolla white to yellow to red, tube greenish in bud, with a belt of pubescence up to 3 mm wide insidejus t below the

88 WageningenAgricultural UniversityPapers 96-1 (1996) insertion ofth e stamens,sometime s extending toth emout h orreduce dt ominut e tufts ofhair sbelo weac hfilament; tub emuc hlonge rtha nth ecalyx ,almos tcylindri ­ cal, narrowed below the insertion of the stamens;lobe s overlapping toth elef ti n bud, sometimes twisted, shorter orlonge r than thetube , spreading andofte n re­ curvedlater .Stamens included ,inserte da to rabov eth emiddl eo fth ecoroll atube ; anthersovate ,obtus et oacuminat ea tth eapex ,cordat ea tth ebase ,glabrous ,o ftw o parallelcell slongitudinall ydehiscent .Pistil glabrous ;ovar ysubglobos et ooblong , composedo f2 separat ecarpels ,unite da tth eextrem ebas eb ya disk-lik ethickening ; pistilhea dcompose do fa stigmati csubglobos ebasa lpar tan da stigmoi dape xu pt o 1 mmlong ;ovule si-6(-30 )i neac hcarpel .Fruits yello wt oorange ,smoot ho rwarty , composedo ftw oseparat emericarp sdivergen ta t90-180° ;mericarp ssubglobos et o obovoid, with a rounded tobeake d apex, 1-9(-26)-seeded ; wall 2-12 mm thick, fleshy tofibrous . Seedsbrown ,variousl y shaped, somewhat angular, smooth.Em ­ bryo straight, spathulate, surrounded by thickrathe r starchy andhar d endosperm, leavinga hol earoun dth eradicl ebase ;cotyledon sthin ,leafy ;radicl ealmos tcylindri ­ cal,flattene d andnarrowe d atth eapex ,widene dtoward sth ebase .

DISTRIBUTION.Al l 12 speciesoccu ri nAfrica . Onespecies ,Hunteria zeylanica, isals owidel ysprea di n San dS EAsia .

Keyt oth especie so fHunteria

1. Corolla lobes wider than long, 1.3-2x 1.7-2.8 mm, broadly ovate with a rounded apex; corolla head globose in mature bud,0.22-0.2 4o f the bud length 5.H .ghanensi s Corollalobe slonge rtha nwide ,2.6-1 2x 0.5-3. 9mm ,ovate ,ellipti ct ooblon g witha nacut et oobtus eapex ;coroll ahea dovoi di nmatur ebud ,0.25-0.6 8 of thebu dlengt h 2 2. Corollatub e1.3-2. 1x a slon ga sth ecalyx ,3- 4m mlong ,insid ewit ho rwithou t minutetuft s ofhair sbelo weac hfilament ; stamensinserte d 1.7-2 mmfro m thebas eo fth ecoroll atub e 2.H .cameruneni s Corollatub e2.9-1 8x a slon ga sth ecalyx ,4-18. 5m mlong ,insid ewit ha bel t ofpubescenc e0.8-2. 0(-3.5 )m mbelo wth estamen san dsometime su pt oth e moutho fth etube ;stamen sinserte d 2.6-9.8m mfro m thebas e 3 3. Stamenswit hape x2.5-3. 7mm belowth emout ho fth ecoroll atub e 4 Stamenswit hape x0.2-1. 2m mbelo wth emout ho fth ecoroll atub e 5 4. Corolla tube 13-18x aslon g asth ecalyx , 12.5-18.5m mlong ; stamens with apex3.2-3. 7m mbelo wth emout ho fth ecoroll atube ,inserte d6- 8m mfro m thebas eo fth etube ;mericarp soblon gwit ha lon gslende rbeak ;Sierr aLeon e toIvor yCoas t 10.H . simii Corollatub e4. 6x a slon ga sth ecalyx ,6.5- 7 mmlong ;stamen swit hape x2. 5 mmbelo wth emout ho fth ecoroll atube ,inserte d2.6-2. 8m mfro mth ebas e of thetube ;mericarp s ellipsoid witha nacuminat ebeak ; Gabon 6.H .hexalob a

WageningenAgricultural UniversityPapers 96-1 (1996) 89 5. Inflorescencepredominantl yaxillary ,sessile ,ramiflorus , 1-1.2x1-1.5cm ;frui t warted,beake d 1. H.ballay i Inflorescence terminal andsometime s atth e same time axillary, 1.5-7.5x I_ 8.5cm ,fruit s smooth,rounde dt oobtus ea tth eape x 6 6. Secondaryvein soblique ,i n7-1 2pairs ,reachin gth emargi nwithou tloopin gt o jointhei rneighbours ,formin g anangl eo f50-60 0wit hth emidrib ;inflores ­ cence,lax ,i ndense ,umbellat ecyme su pt o 160-flowered 8.H .myrianth a Secondaryvein smor eo rles sstraight ,i n9-3 0pairs ,formin ga submargina lvei n anda n angleo f (55-)65-80 0 withth emidrib ;inflorescence , contractedo r lax,u pt o 150-flowered 7 7. Leaf acumen indistinct; inflorescence lax,u pt o96-flowered , Eastern Coast from Somaliat oMozambiqu e 12.H .zeylanic a Leaf acumendistinct ;Inflorescenc efew-flowere d orcontracted ,Wester nCoas t from Senegalt oZair ean dMt .Kula li nNorther n Kenya 8 8. Corollatub e9-1 3m mlong ;stamen sinserte da t7-9. 8m mfro mth ecoroll abas e 9 Corollatub e4-8. 2 mmlong ;stamen sinserte d at2.5-6. 2m mfro m thecoroll a base 10 9. Secondaryvein si n9-1 4pairs ,8-1 3 mmapar ta tth ecenta lpar to fth eleaf ; se­ pals membranaceous, colleters sparsely distributed within; ovary with 1 ovulepe rcarpe l 7.H .macrosipho n Secondary veins in 20-25pairs , 3-6m m apart atth ecentra l part of the leaf; sepalsthic ktoward sth emiddle ,coriaceou swit hcolleter su pt o0.7 5o fthei r length;ovar ywit h6- 7ovule spe rcarpe l 9.H .oxyanth a 10. Corollalobe s0.4-0. 5x a slon ga sth etube ,2.6- 3m mlong ;ovule s2 pe rcarpe l 4.H .densiflor a Corollalobe s0.8-2. 7x a slon ga sth etube ,4-1 2m mlong ;ovule s6-3 0pe rcar ­ pel 11 11. Ovarygraduall ynarrowe dint ostyle ,1.5- 2m mlong ,wit h5- 6ovule spe rcarpel ; fruit 20-26x 11-18cm ;Zair ean dKeny a 3.H .congolan a Ovary abruptly narrowed into style,0.6-1. 1m mlong , with 14-30ovule s per carpel;frui t 31-60x 40-5 0cm ;Senega lt oZair e 11.H .umbellat a

2.5.1 Hunteriaballay iHu a in Bull.Mus .Hist . Paris 7: 281 (1902).-Type : Cultivated ingreenhous e atPari s from seedscollecte di nGabo nb yBallay inMa y 1906(holotyp eP) . Fig.23 ,p . 91; map1 ,p .9 2

Homotypic synonym:

Pleuranthemumballayi (Hua )Picho ni nBull .Soc . Brot,sé r2,27:113 ,plat eE ,fi g 7-10,ma pA ,p . 85(1953).

90 WageningenAgricultural UniversityPapers 96-1 (1996) tav

Figure 23.Hunteria ballayi. 1,habi t( x 2/3); 2flower, bu d( x4) ;3 ,opene dcoroll a (x4) ;4 ,pisti l( x4) ; 5,fruitin g branch( x2/3) ; 6,frui t (x2) ;7 ,embry o( x4) .1 fromLeTest u8558 ;2- 4fro mva nde rMaese n etal .5579 ;5 fromWilk s 1453;6- 7 from Schlott 11.

WageningenAgricultural UniversityPapers 96-1 (1996) CI 20 10

10 10

20 Map i. Hunteriaballayi. Shrub i- 3 mhigh ,sometime slianescen tan du pt o5 m hig hwit hwhit elatex .Trun k slender; inner bark yellowish; wood very hard. Branchlets drying greyish green. Leaves opposite;petiol e 3-7 mm;blad epaper y when dried,ellipti c toslightl yob - ovate,2.2-3. 1x a slon ga swide ,6.7-20. 3x 2.2-8. 1cm ,acuminat ea tth eapex ,acu ­ men obtuse, 7-14 x 2 mm, acute at the base; midrib prominent beneath in dried leaves,dryin gbrownis horange ;secondar yvein sstraight ,i n20-3 0pairs ,formin ga submarginalvei nan da nangl eo f65-70 0wit hth ecosta ,5- 6m mapar ta tth eapproxi ­ matecentr eo fth eleaf .Inflorescence predominantl y axillaryan dterminal , fascicu­ late,ramiflorous , 1-1.2 x 1-1.5 cm,abou t io-flowered;peduncl e0- 2mm .Pedicel s 0.5-2 mm. Bracts 0.5-0.7x 0.4-0.5mm , acuminate. Flowers: Sepals light green, free, membranaceous atth eedge ,thicke rtoward sth emiddl ean da tth ebase ,erect , ovate,1.3- 3 xa slon ga swide ,1 -1. 5x 0.5-0. 8mm ,obtus et oacut ea tth eapex ,wit h rowso fcolleter sinsid eu pt o0. 4o fthei rlength ,colleter sver y small.Corolla dirt y white,pal eyello wi nbud ,7. 1-11. 2m mlon gi nmatur ebu dan dformin g acompara ­ tively small ovoid head 0.36-0.39o f thebu dlength , 2.8-4x 0.9-1.6mm , witha n obtuset orounde dapex ,wit ha bel to fpubescenc e 1 mmwid einsid ejus tbelo wth e insertion ofth estamens ,wit ha fe w hairsaroun dth estamens ,tub e3.3- 6 xa slon g asth ecalyx , 1.7-2.4 xa slon ga sth elobes ,5-7. 2 mmlong ,almos tcylindrical ,0.9 - 2m mwid eabov eth ebase ,narrowe dbelo wth einsertio n ofth estamen st o0.8-1. 6 mmwide ,agai nwidene daroun dth eanther st o 1.3-1.8 mmwide ;lobe sno ttwiste d inbud ,ovate ,0.4-0. 6x a slon ga sth etube ,1.9-2. 3 xa slon ga swide ,2.7-3. 5x 1.3 - 1.5mm ,obtus ea tth eapex ,entire ,spreadin go rerect .Stamens wit hape x0.2-0. 3 mm belowth emout ho fth ecoroll atube ,inserte d0.69-0.7 9o fth elengt ho fth ecoroll a tubea t3.5-5. 5m mfro mth ebase ;filament sshort ,filiform , 0.3-0.6m mlong ;anther s narrowly ovate,3.5-4. 5x a slon g aswide ,0.9-1. 4x 0.3-0. 4mm ,acuminat e atth e apex,acume n0.2-0. 3mm ,cordat e atth ebase .Pistil 4.6-5. 6m mlong ;ovar ynar ­ rowlyoblong ,1-1. 2x 0.4-0. 8x 0.3-0. 6mm ,abruptl ynarrowe dint oth estyle , with

92 WageningenAgricultural UniversityPapers 96-1 (1996) adisk-lik ethickenin gunitin gth e carpelsa tth ebase ,abou t0. 2o fth elength ,0. 2m m high,o ftw oseparat ecarpels ;styl e2.8-3. 7n ™long ;pisti lhea dcompose do f aglo ­ bosestigmati cbasa lpar t0.4-0. 5x o .15-0. 3m man d afilifor m stigmoidape x0.2-0. 6 x0. 1-o .1 5 mm. Ovules 2 ineac hcarpel .Fruits pal eyellow ,rough ,wart yt o smooth, oftw oseparat emericarps ,mericarp sobovoi do rellipsoid ,15-2 8X9-1 5x6- n mm, with a short beak at the apex,bea k 2-6mm , 2-seeded,wal l about 1-2mm ;pul p smooth or slightlyfibrous. Seeds ellipsoi d or subglobose, 8-13 x 5-9 x 3-7mm . Embryo4.3- 9m mlong ;cotyledon sovate ,1.2- 2x a s longa s wide 2.3-5 x I-8- 4mm , obtuse atth eapex ,rounde d atth ebase ;radicl e0.7- 1x aslon g ascotyledons , 2-4 x0.5- 2mm .

DISTRIBUTION. Cameroun,Centra lAfrica n Republic,Gabo nan dCongo . ECOLOGY.Ol dsecondar yforest ,floo dplai nfores t and drierfores twit hscattere d treesan dnumerou sgap sdominate db y Marantaceae. Alt.350-50 0m . Floweringi n April in Central African Republic, April and November in Gabon, and fruiting aroundJul yt oSeptembe ri nal lcountries .

USES. Used as afertilit y drug,wome nswallo wth esee dtogethe rwit hth eswee t mesocarp;chimp ssuc kth e sweetmesocar p (Fay 8127,MO , 1987,RCA) ; useda s afertilit y drugan dals ofoun d in gorilladroppings ,bu tneve ri n highdensit y(Harris & Fay 63, MO, 1988, RCA).

Specimens examined: CAMEROUN. Est: 65 km NNE of Moloundou,Letouzey & Villiers 10568 (P); 85K mN ofMolundou , Letouzey & Villiers 10383 (P); Ndankan, D. W. Thomas720 2 (WAG);Moloundou , Mildbraed 4038 (HBG).Ngongondj e Hill,Koufani 143 (P); Nyabessan,Mezili 120(P) . CENTRAL AFRICAN REPUBLIC. Sangha: NdankanGorill aStud yarea ,Harri s & Fay 63 (MO), Harris & Fay 547 (WAG),fa y 8127 (MO), Fay &Harris 878 2 (MO), 8835 (MO); 45 km S ofLidjombo , Schlott 11 (WAG). GABON. Haut Ogoué: Ngounie, Le Testu 8558 (BM, BR, P). Ogooué-Ivindo: 40 km N of Koumameyong,Wilks 1453(WAG) .Ogooué-Lolo : 10 kmfrom Lastoursville , Vander Maesen et ai. 5579 (WAG).Estuaire :Libreville , Klaine307 7 (P); BallayMa y 1906 (P, type). CONGO: Ngoko,Bouquet 162 9 (P); SembeForest ,Sita 337 3(WAG) ;nea rN'gbal ,Sita 345 1(BR) .

2.5.2 Hunteriacamerunensi s K. Schum.e xHal l f. inJahrb .Hamb .Wiss .Anst , 17Beih . 3: 187(1899) ;Picho n in Mém.Mus .Natn . Hist.Nat .24:161,1791 . 3fig. 2 0 (1948).-Type :Cameroun ,Bipindi ,Zenker 1620 (lectotype P, designatedb y Pichoni n 1953,isolectotype s B, BP, BR, E, G, HBG, L, M,N Y S,W ,WAG ,WU , Z). Fig.24 , p. 94; map 2, p.9 5

Homotypicsynonyms :

Pleiocarpa camerunensis (K. Schum.e xHall , f.) Stapf inFI .Trop .Afr . 4, 1:10 2 (1902 ). Polyadoa camerunensis (K. Schum. ex Hall, f.) Brenan in KewBull .

WageningenAgricultural University Papers96-1 (1996) 93 Figure 24. Hunteriacamerunensis. 1, habit (x 2/3); 2,flower bu d (x 1.4); 3, flower (x 10); 4, opened corolla( x io);5 ,sepa linsid e( x 1.4); 6,pisti l (x 1.4). 1- 6fro mZenke r1211.7 . Fruiti sTabernaemontana letestui.

94 WageningenAgricultural UniversityPapers 96-1 (1996) 5 15 10 n n 10 -> \

m 6 5. mt 0 %'

IS Map 2.Hunteria camenmensis.

!952-"45 1 (1952).Comularia camenmensis (K.Schum.e xHall ,f. ) Pichoni nBol . Soc.Brot . sér.2 ,27 : 116, 117 t. 3,fig 1-3 ,ma pA ,p . 85 (1953).

Shrub 1-3m high .Trun k slender, green up to 1.5 cm in diameter. Branches dark green,dryin ggre yt oblack ;branchlet squadrangula rt oslightl y four-ribbed, green, even when dried. Leaves opposite; petiole 4-10 mm;blad e thinly leathery when fresh, elliptic,oblon gt oobovate ,2-3.3(-5 )x a slon ga swide ,6-i8.8(-24.7 )x 1.4- 8 cm,acuminat ea tth eapex ,acume nobtuse ,5-2 0x 2- 4mm ,acute ,obtus et ocuneat e at thebase , smooth,gloss y above;midri bprominen t beneath and drying lightor ­ ange;secondar yvein smor eo rles sstraigh tt oslightl ycurved ,i n 11-13pairs ,form ­ inga nangl eo f60-75 °wit hm e midrib,6-1 8 mmapar ta tth eapproximat ecentr eo f theleaf .Inflorescence terminal,rarel y axillary,i ncompac tclusters ,0.5-1. 5x 0.5 - 2.3cm ,abou t5-12-flowered , sessileo rwit ha peduncl e1- 6m mlong .Pedicel s0.5 - 2mm .Bracteoles ,thre ea tth ebas eo feac hpedicel ,0.5-1. 5x 0.2- 1 mm,acuminat e atth eapex .Flowers: Sepalspal egreen ,free , erect, coveredb y alittl eresi no nth e innerside ,membranaceou sfo r about0. 2m mo fthei redge ,ovate ,1.9-3. 6x a slon g aswide , 1.5-2.5 x0.6-1. 2mm ,acuminat et oacut ea tth eapex ,acume n0.5- 1 mm, withrow so fcolleter su pt oon e0. 4o fthei rlengt hinside ,colleter sver ysmal l0. 2x 0.05mm .Corolla cream yt owhite ,ligh tgree ni nbud ,4.3- 8m mlon gi nmatur ebu d and forming a comparatively large ovoid or triangular head 0.44-0.63o f thebu d length,2.3-3. 7x 0.5-0.9mm ,wit h anacut eapex ,wit hminut etuft s of hairs atth e baseo feac hfilament inside ,tub e 1.3-2.1 x aslon ga sth ecalyx ,0.6-0. 9( _I-2)x a s longa sth elobes ,3- 4m mlong ,almos tcylindrical ,0.7-1. 8m mwid eabov eth ebase , narrowed below theinsertio n of the stamens to0.6-1. 2m mwide ,agai n widened aroundth eanther st o0.8-1. 5m mwide ,thickene dan dver yconstricte da tth ethroat ; lobespal epink ,twiste di nbud ,narrowl yoblon gt oovate ,(0.9- ) 1.1-1. 6x a slon ga s thetube , 2.4-5.9(" I0) x as l°n8 aswide , 2.8-5.5x °-5"2mm .obtus e at theapex , entire, spreading toerect .Stamens with apex0.2-0. 8m mbelo w themout h of the corollatube ,inserte d0.43-0.6 7o fth elengt ho fth ecoroll atube ,a t1.7- 2m mfro m thebase ;filaments 0.2-0. 5m mlong ;anther snarrowl yovat et ooblong ,4- 6x a slon g aswide ,0.8-1. 2x 0.2-0. 3mm ,obtus ea tth eapex ,subcordat ea tth ebase .Pistil 1.9-

WageningenAgricultural UniversityPapers 96-1 (1996) 95 3.i m mlong ,ovar ysubglobose ,0.6- 1. ox 0.4-0. 7x 0.2-0. 5mm .distinctl ynarrowe d into the style,wit h adisk-lik ethickenin gunitin gth e carpelsa t the base,abou t 0.25- 0.33 of the length, 0.2m mhigh , oftw o separate carpels; style 0.5-0.8m mlong ; pistilhea dcompose do fa subglobos et ooblon gstigmati cbasa lpar t0.3-0. 8xo.1 5x 0.2m man da cylindrica lstigmoi dape x0.3-0. 9x o .1 mm. Ovules 6-8 ineac hcarpel . Fruits orange, smooth,o ftw oseparat emericarps ,mericarp s divergenta ta n angle of 1800,ovoi dt osubglobose ,28-4 7 * I5"25x 10-18mm , bluntlyacuminat ea tth e apex, 6-8-seeded; wall about 2mm ; pulp smooth or slightly fibrous. Seeds subglobose toellipsoid ,7-1 4x 5.5-1 0x 3- 4 mm. Embryo 5.5-9 mmlong ;cotyle ­ donsovate ,1.2-2. 5x as long as wide3-4. 2 x2-2. 5mm .obtus ea t the apex,rounde d atth ebase ;radicl eo.75-o.83(-i.2 )x a slon ga scotyledons ,2.5-4. 8x 0.5-0. 9 mm.

DISTRIBUTION. Cameroun andGabon . ECOLOGY. Primary orsecondar y forest alongsmal lcreeks ,o rstrea mgullie so n forest border ori nrai n forest. Alt.50-65 0 m. Flowering and fruiting allth eyea r round.

Geographical selectiono fth e approximately 40specimen sexamined : CAMEROUN. Sud-Ouest: Limbe, Mildbraed 10716 (A); S Bakundu Forest Reserve, Beentje 1450 (WAG),Brenan 940 1 (FHO, K, P), Binuyo& Daramola FHI3509 0(K , P). Centre-Sud:8 k m No fKribi , De Wilde &De Wûde-Duyfies 2901 (WAG);6 0 km NNWo f Eseka,De Wilde &De Wilde-Duyjjes 2195 (WAG);Bipindi , Zenker121 1 (E,G ,HBG , L,M , P,S,W , Z), 1620 (B,BP ,BR , E,G ,HBG , L,M ,NY , P,W ,WAG ,WU ,Z ,typ eof« , camerunensis),231 1 (BR,E ,HBG ,L ,M , P, S,W,WU, Z), 3163 (BP, BR, E,G ,HBG , K,L ,M ,P , S,W,WAG,WU), 3611 (BP, BR,E , G,HBG ,K , L,M ,MA , P, S,US,W ,Z) ,411 3 (BP, BR,E ,G , HBG,K ,L ,M ,MA , P,S ,W , Z); Mimfia, ZenkerU 163(B ,C , FT, G,VC, US,U , WAG); 25 kmN Wo f Eseka, DeWilde &De Wüde-Duyjjes 144 2 (BR, K, P, WAG); Nkoemvone,Raynal 960 6 (P); 15k mS o f Ebolowa,De Wilde &De Wlde-Duyfjes 1974(BR , K,P , WAG). GABON.Woleu-Ntem : 29k mES Eo f Medouneu,Reitsma 185 8(NY , WAG). Ogooué-Ivindo:ca .20-4 0 kmNN Eo fKoumémayong ,Breteler e t al.865 4(WAG) ;1 0 km S ofMakokou ,Florence 15 4 (P), 566 (P, WAG). Ogooué-Lolo: 16k mo nLastoursville-Moand a Rd., Breteler & deWilde 748/78 (WAG);2 8 km NE ofLastoursville , Wieringa & Van derPoll 1479 (WAG).

2.53 Hunteria congolana Pichon inBol . Soc.Brot , sér2 ,27 : 101,t. i fig.9-13 ;ma pA p.8 5 (1953).-Type : Zaire, Haut-Zaire, 10k mEas to fYangambi , J.Louis 1083 (holotypeBR ; isotypesK ,P ,S) .

Fig. 25, p. 97; map 3, p. 98

Shrubo rsmal ltre e1.7-2 0 m highwit hwhit e to yellowlate x in all parts.Trun k5-3 0 cmi n diameter; woodyello wan dhard .Bar ko fbranche s thin, smooth, finelyfis­ sured.Leaves opposite ;petiol e8-1 8m mlong ;blad ethin ,paper ywhe ndried ,ellipti c tooblong , 2.4-4 x as long aswide ,6.3-18. 4x 1.5-5.9cm > acuminate atth eapex , acumen obtuse, 10-18x 2-4mm , acutet oobtus e atth ebase ;midri b paleyellow , prominentbeneath ;secondar yvein sslightl yprominen tbeneath ,i n 11-1 6pairs ,mor e orles s straight,formin g anangl eo f70-80 0wit hth emidrib ,6-1 4m mapar ta tth e approximatecentr eo fth eleaf .Inflorescence termina lo raxillary ,3-3. 5x 3- 5cm ,1 -

96 WageningenAgricultural UniversityPapers 96-1 (1996) Figure 25.Hunteria congolana. 1, habit( x 2/3);2 , calyx( x6) ;3 , corolla( x6) ;4 ,opene dcoroll a( x6) ; S,sepa linsid e( x 14);6 ,pisti l( x 10);7 ,drie dimmatur efruit ( x2/3) ;8 ,drie dfruit ( x2/3) ;9 ,drie dpar t offruit wal l( x 1);10 ,opene ddrie dfrui t (x 1);11 ,see d( x2) ;12 ,longitudina lsectio no fsee d( x2) ; seed (X2). 1 fromJ.Loui s6489 ;2- 6from J.Léonar d 1445; 7fro mNsol a 1261:8-13 fromJ.Loui s6328 .

WageningenAgricultural UniversityPapers 96-1 (1996) 97 Map3 .Hunteria congolana.

2x branched ,abou t7-1 8(-45)-flowered ;peduncl e2-1 4m mlong ,primar ybranche s 3-10m m long, secondary branches 2-4m m long; pedicels 2-5m m long. Bracts sepal-like, 1-1.5 x 0.5-0.7mm , with colleters inside.Flowers: Sepals palegreen , coriaceous, almost free, erect, membranaceous at the edge and the apex, thicker towardsth emiddl ean da tth ebase ,wit hmuc hwhit et oclea rresi nwithin ,broadl y to narrowly ovate, 1.1-2.7x aslon g as wide,0.8-1. 5* 0.3-0.8mm , acuminate to acute atth e apex, with rows of colleters inside upt o 0.9 of their length, colleters very small0. 2 x0.0 5 mm.Corolla whiteo r yellow,creamy-pin k inbud ,6.6-11. 6 mmlon gi nmatur ebu dan dformin g acomparativel y wideovoi dhea d0.38-0.5 5o f thebu dlengt h2.2- 6x 0.9-1. 7mm ,wit ha nobtus eapex ;wit ha bel to fpubescenc e 1-1.5 mmwid einsid ejus t belowth einsertio n of the stamens andu pt oth emout h of thetube ,tub e3.7-6. 3x aslon ga sth ecalyx ,0.7-1. 3x aslon ga sth elobes , 4.8- 5.5 mmlong ,almos tcylindrical ,1 -1. 8m mwid eabov eth ebase ,narrowe dbelo wth e insertion of thestamen st o0.8-1. 3nu nwide ,agai n widened aroundth eanther st o 1-2 mmwide ;lobe snarrowl yovat et ooblong ,0.8-1. 5x a slon ga sth etube ,1.7- 4 x aslon ga swide ,4- 8x 1.5-2.8 mm,obtus ea tth eapex ,spreading .Stamens wit hape x 0.2-1m mbelo wth emout ho f thecoroll atube ,inserte d 0.63-0.73 of thelengt ho f the corolla tube, at 3-4.7 mmfrom th e base; filaments 0.2-0.5 mml° ng; anthers narrowlyovat e3.7-7. 5x a slon ga swide ,1.1-1. 5 x0.2-0. 3mm ,obtusel yacuminat e atth eapex ,acume nu pt o0. 4mm ,cordat ea tth ebase .Pistil 3.4-4. 8m mlong ;ovar y oblong, 1.5-2 x0.6- 1x 0.3-0. 8mm ,graduall ynarrowe dint oth estyle ,wit ha disk ­ likethickenin g unitingth ecarpel s atth ebase ,abou t0.1-0. 2o f thelength ,0. 2m m high,o f twoseparat ecarpels ;styl e0.8-2. 1x 0.2-0. 4mm ;pisti l headcompose do f asubglobos estigmati cbasa lpar t0.2-0. 8x 0.2-0. 5n ™an da nconica lstigmoi dape x 0.4-1.0x 1-0.2 mm.Ovule s5- 6i neac hcarpe la tth eextrem ebase .Fruits yello wt o brightorange ,smooth ,dull ;mericarp sellipsoi dt osubglobose ,somewha t flattened,

98 WageningenAgricultural UniversityPapers 96-1 (1996) 20-26 XII-I8XII-I6 mm, roundedt oobtus e atth eapex ; 1-3-seeded; wallabou t 1-3 mmthick , slightlyfibrous within . Seeds ovoid to ellipsoid, 7-13x 5-7.5 x 3- 4mm .Embry o6.3-8. 7m mlong ;cotyledon sovat et oelliptic ,1.5-2. 6 (-5.7)xa slon g as wide,2.3-4. 2x 0.7- 2 mm,obtus e atth e apex andth ebase ;radicl e0.7-1. 7x a s longa scotyledons ,3-4. 7 x0.5-0. 8 mm.

DISTRIBUTION. Zairean dKenya . ECOLOGY.Dens eunderstore yo fprimary ,rarel ysecondar yfores to rgaller yfores t onplateau .Alt .470-170 0m . Flowering andfruitin g allth eyea rround .

Geographical selection of the approximately 48specimen s examined: ZAIRE. Equateur:Boende , L.Dubois 651 (BR); Yalifake, Evrard627 0 (BR);betwee nYokol o and Mondombe,Evrard 435 4 (BR). Haut-Zaire:Mobwasa ,Reygaert 88 4 (BR); EsaboIsland ,Germain 495 7 (BR,M) ; nearYangambi , J.Léonard 144 5 (BR); ibid., J.Louis 108 3(BR ,K ,P , S,type) , 3809(B ,BR , FT, K, NY), 6328(BR) , 6489(BR , C, NY); ibid.,Menavanza 14 3(BR ,K ,WAG) . Kasai-Oriental: Mukumari, Germain799 8 (BR, K, P). Kivu: Kampala, Pierlot 950 (BR);Kabunga ,A.Léonard 1831 (BR, MO, P, WAG); nearIrangi ,Troupin 12151 (BR); Masanga, A. Léonard587 9(BR ,MO , WAG); km8 9 Shabunda- KaseseRoad ,Michelson 914 (BR). KENYA. KI: NorthernFrontie rDistr. ,M L Kulal,Adamson 11 5(EA ,K) ; Northo f Gatab, sin. coll. 178 (EA.K).

Note.Th e corollao fthi sspecie salway sfall s off leavingonl yth e pistil,therefor e itprobabl y flowers duringth enight .I tca nb eclearl y distinguished from theothe r Hunteria species by the ovary which gradually narrows into the style. Collected twicei nKenya , in Mt. Kulalwher elobe s are muchlonge ran dleave smuc hsmaller . Fruits notknow nfro m Kenya.

2.5.4 Hunteriadensiflor aPicho n inBull .Jard .Bot .Brux . 23: 392fig, 3 8(1953) ,partl y excludingo fMichelson 85- Type:Zaire ,Maniema , 22k m onElila-Kail oroad ,Michelson 921 (holotype BR; isotypeP) . Fig. 26, p. 100; map 4, p.10 1

Treeu pt o 18 mhigh ;trun ku pt o5 5c mi ndiameter ;bar k smooth,dar kgreenish - brown;inne rbar kbrown ;woo dno tdifferentiated , hard,yellowish-orange . Leaves opposite,occasionall y ternate,petiolate ;petiol e 12-20m mlong ;blad ecoriaceous , elliptic, 2.8-2.9x as l°ng as wide, 6.5-15.3x 3.5-6.5cm , acuminate at the apex, acumen acute, 10-22x 3-4mm ,acut et oobtus e atth ebase ;midri bprominen tbe ­ neath; secondary veins prominent above andbeneath , in 22-30pairs , forming an angle of 70-800wit hth emidri b anda nea t submarginal vein,4- 6m mapar t atth e approximate centre of the leaf, tertiary veins more or less parallel. Inflorescence terminal,dense ,2 x branched ,i numbellat ecymes ,60-120-flowered ; peduncle 15- 40 mm, branches 2-8 mmlong ;pedicel s 2-3 mmlong .Flowers ver yfragrant .Sepals palegreen ,free ,erect ,membranaceou sa tth e edge,thic ktoward s the middlean dth e base,ovat e 1.4-1.7x aslon ga swide , 1-1.2x0.7mm ,acuminat ea tth eapex ,wit h

WageningenAgricultural University Papers 96-1 (1996) 99 Figure 26. Hunteria densiflora. 1, habit (x 2/3); 2, section of corolla (x 5); 3, pistil (x 10). 1-3 from Michelson 941. adapted from Pichon, 1953.

IOO WageningenAgricultural UniversityPapers 96-1 (1996) Map4 .Hunteria densiflora.

resinwithi nan dcolleter sinsid eu pt o0. 5o fth elength .Corolla white ,1 0m mlon g inmatur ebu dan dformin ga comparativel ysmal lovoi dhea d0.3 5o fth ebu dlength , 3.2x 1 mm,wit ha nobtus eapex ,wit habel to fpubescenc e1-1, 3 m wideinside ,jus t belowth einsertio no fth estamen san du pt oth emout ho fth ecoroll atube ;tub e 5- 5.5x aslon g asth ecalyx , 1.8-2.3 x as long asth elobes , 5.5-6m m long, almost cylindrical,0.8-1. 5m ™ wideabov eth ebase ,narrowe dbelo wth einsertio n ofth e stamenst o0.6-0. 8m mwide ,agai nwidene daroun dth eanther st o1.2-1. 3 mmwide ; lobesovate ,0.4-0. 5x a slon ga sth etube ,1.3-1. 4x a slon ga swide ;2.6- 3mm l°ng. obtusea tth eapex ,entire , spreading.Stamens with apex0. 4 mmbelo wth emout h ofth ecoroll atube ,inserte d0.8 7o fth elengt ho fth ecoroll atub ea t4. 8m fro mth e base;filament s0. 5m mlong ;anther sovate ,3.5- 4x a slon ga swide ,0.6-0. 7x 0.15- 0.2 mm, obtuse atth e apexan d sterilefo r0. 2 mm,cordat e atth ebase .Pistil 5.5 - 6.2m mlong ;ovar y subglobose,0.7- 1x 0.6-0. 7x 0.4-0.5mm ,abruptl y narrowed intoth e style,wit ha disk-lik ethickenin g unitingth ecarpel s atth ebase ,abou t0. 3 ofth elength ,0.2-0. 3m mhigh ,o ftw oseparat ecarpels ;styl e3.8-4. 3 mmlong ;pisti l headwit ha stigmati cellipsoi dbasa lpar t0.3-0. 5x 0. 3an da conica lstigmoi dape x 0.5 xo .15-0. 2mm .Ovule stw oi neac hcarpel .Fruits immature ,green ,o ftw osepa ­ ratemericarps ;mericarp s subglobosewit ha nobtus eapex ,2-seeded .

DISTRIBUTION.Zaire . ECOLOGY.Semi-deciduou sforest .Alt .50 0m .Flowerin gi nFebruar yan dfruitin g inMarch-April .

Specimensexamined : ZAIRE. Equateur:Bokata ,Territor y Bokungu Evrard,567 5 (BR,WAG) ;Maniema , 22k mo n Elila- KailoRoad ,Michelson 921 (BR, P, type).

WageningenAgricultural University Papers 96-1 (1996) IOI Notes. Mature fruits unknown.Th e immature fruits of Michelson 85 cited by Pichon (1953),wer eanalyze dan dfoun d tohav e7 seeds ,meanin gtha tth eovarie s havea tleas t 7ovules .H. densiflora hasonl ytw oovule spe rcarpe l andimmatur e fruits ofMichelson 92 1hav e2 seed sonly .Michelson 8 5 matchesth ene wspecies , Hunteria oxyantha Omino,fro m Gabon.

2.5.5 Hunteria ghanensisHal l& Leeuwenber g

inBull .Jard .Bot .Nat .Belg .49:422,423 fig., 1 (1979).-Type :Ghana ,Apa mjunc ­ tion,J. B. Hall G C4704 4(holotyp eWAG ; isotypesB ,BR ,K , P). Fig.27 ,p . 103;ma p5 ,p .10 4

Tree 1-12 mhigh .Trun k 20-25c m m diameter;bar kpal egrey ,shallowl y fissured, darkbrow no nsection ;woo dochraceous . Branchespal egrey ,wit hsmal llenticels ; branchletsthin ,smooth .Leaves opposite ;petiol e5-1 0mmlong ;blad esubcoriaceou s whenfresh , narrowlyellipti c(2- )2.7-5. 1x a slon ga swide ,3.5-13. 5x 0.8-3. 4 cm, abruptlyacuminat ea tth eapex ,acume n 12-15x2-4mm ,acute ,cuneat ea tth ebase ; midribprominen tbeneath ;secondar y veinsstraight ,inconspicuou s above,slightl y prominentbeneath ,i n 15-20pairs ,formin g asubmargina lvei nan da nangl eo f60 - 720 with the costa, 6-10m m apart at the approximate centre of the leaf;tertiar y venationparalle lt oth esecondar y veins.Inflorescence terminal ,almos tcompoun d umbellate,2 xbranched ,2- 4x 2.5-4cm ,30-150-flowered ;peduncl e3-1 8mmlong , branches5-1 8 mmlong ;pedicel s2- 4m mlong .Bract sver y small.Flowers: Sepals almost free, erect, with alittl eresin within , ovatet onarrowl y triangular, acutet o acuminatea tth eapex ,1.7-3. 4 xa slon ga swide ,1-1. 2x 0.35-0. 6mm ,wit hrow so f colleters inside upt o 0.5 of their length, colleters very small, scale-like. Corolla creamt opal eyellow ,5.5-8. 5m mlon gi nmatur ebu dan dformin g acomparativel y small,globos ehea d0.22-0.2 4o fth ebu dlength ,1.2- 2x 1 mm,wit ha rounde dapex , with abel t of pubescence 2.5-3.5nu n wide insidejus t below the insertion of the stamens,an daroun dth estamens ;tub e3.6-6. 2x a slon ga sth ecalyx ,2.7-4. 8x as longa sth elobes ,4.3-6. 2m mlong ,almos tcylindrical ,0.6-1. 5 nunwid eabov eth e base,narrowe dbelo wth einsertio n ofth estamen st o0.5-0. 8m mwide ,agai nwid ­ enedaroun dth eanther st o0.7-1. 5m mwide ;lobe sbroadl yovate ,0.2-0.3 5x a sl° ng as the tube,0.7-0. 9 x as long as wide, 1.3-2x t.7-2.8mm , rounded at the apex, entire, spreading to slightly recurved. Stamens with apex 0.3-0.5 mm below the moutho fth ecoroll atube ,inserte d0.77-0.9 3o fth elengt ho fth ecoroll atube ,a t4 - 5.2m mfro mth ebase ;filament s0.3-0. 5m mlong ;anther sovat et ooblong ,2.6-3. 5* aslon g aswide ,0.6-0. 8x 0.2-0. 3mm ,obtus e atth eapex ,subcordat e atth ebase . Pistil4.6-6. 0m mlong ,wit hape xapproximatel y halfway alongth eanthers ; ovary subglobose,0.6-0. 9x 0.5-0. 6x 0.3-0. 5mm ,abruptl y narrowedint oth estyle ,wit h adisk-lik ethickenin gunitin gth ecarpel sa tth ebase ,abou t0.22-0.2 5o fth elength , 0.1-0.2m mhigh , of two separate carpels; style 3-4.5 x 0.15-0.2 mm;pisti l head composedo fa subglobose ,stigmati cbasa lpart ,0.2-0. 4x 0. 2x 0.2 5m man da coni ­ cal stigmoid apex0.3-0. 5x 0. 1mm ,sometime s witha bilobe d apex.Ovule s i(-2) in eachcarpel .Fruits orange,smooth ;mericarp s diverging ata n angleo f 900,ob -

102 WageningenAgricultural UniversityPapers 96-1 (1996) Figure 27.Hunteria ghanensis. 1, habit( x 2/3);2 ,branc h( x 2/3); 3flower, bu d( x 8);4 flower, ( x 8);5 , openedcoroll a( x8) ;6, sepa linsid e( x16) ;7 ,pisti l( x8) ;8-9 ,fruit s (x2/3) ;10 ,see d( x2) ;11 ,transvers e sectiono f seed( x 2);12 ,longitudina l sectiono f seed( x 2);13 ,embry o( x6) . 1-3 from Leeuwenberge t al. 11208;4- 7fro mHal l& Swai n43741 ;8-1 3 from HallG C47044 .

WageningenAgricultural UniversityPapers 96-1 (1996) IO3 Map5 .Hunteria ghanensis. ovoid, 10-15 x 10-12 x 9-11 mm, rounded at the apex, stipitate; 1-2-seeded; wall about 1-2 mmthick , slightly fibrous within.Seeds bean-shaped orellipsoid , coated with astick y mucilage when fresh, olive green andtinge d withbrown ,pale-veined , smooth,7-1 0 x 5-8 x 3-5 mm,test asmoot horange .Embry o6-6. 4m ml° ng; cotyle­ donsovate ,2-2. 3x a slon ga swide ,2.8- 3 x I-2-I-5 mm,obtus ea tth e apex,rounde d at thebase ;radicl e 0.8-1 x as long as cotyledons, 2.5-3 x 0.5-0.6 mm.

DISTRIBUTION. Ivory Coast and Ghana. ECOLOGY. Dry forest or light secondary forest with an annual rainfall of 1000- 1200mm . Alt.50-10 0m . Floweringan dfruitin g betweenDecembe ran dJul yi nbot h countries.

USES. Bark together with that of Crvton zambesicus Muell. Arg. and Nauclea latifolia Sm. taken as a medicine for stomach ache and difficult urination (Leeuwenberg 1979).

Specimens examined: IVORY COAST. Abongoua Forest,Alee Assi 11222(WAG) . GHANA. Central:Apa mJunction ,Hall G C4704 4(BR ,K ,MO ,P ,WAG ,type) ;Hall G C4704 5(B ,BR , K,MO ,P ,WAG) ;2 k mE o f ApamJunction ,Leeuwenberg 1116 4(BR ,MO ,P ,WAG) ;1. 7k mE o f Apam Junction,Hall & Swain 4374 1 (K, MO, WAG).Eastern :nea rharbou ro f Akosombo,Leeuwenberg etal. , 11208 (B, BR, MO, P,WAG) . CULTIVATED. Ivory Coast: AdiopodoumeBo t Garden,AkéAssi 7478 (WAG), 13763(WAG) .

2.5.6 Hunteriahexalob a (Pichon) Omino,comb .nov .

Fig. 28, p. 105;ma p 6, p. 106 Basionym:

Tetradoahexaloba Pichon inBull . Soc.Bot .Franc e 93, 7-8:25 2 (1946) & in Bull. Soc. Brot. sér. 2, 27: 122, t. UI fig. 7, p.117; map C, p. 121 (1953). -Type: Gabon, Klaine 3158 (holotype P; isotype BR).

Shrub 1.6m high.Trun k slender. Branchlets drying dark brown to greyish-black. Leaves: petiole 7-10m m long;blad e elliptic, 1.8-2.5x as l°ng as wide, 10.3-21 x 4.5-11.3cm , acuminate atth e apex, acumen acuteo r obtuse, 6-13x 4-6 mm, acute to obtuse at the base; midrib prominent beneath; secondary veins very prominent

104 WageningenAgricultural UniversityPapers 96-1 (1996) Figure 28.Hunteria hexaloba. 1,habi t (x 2/3);2 flower, bu d (x 4);3 , sepal inside (x 30);4 ,opene d corolla( x4) ;5 ,pisti l( x4) ;6 ,pisti lwit hopene dovar y( x30) ;7 ,infructescenc e (x2/3) .1- 6fro mKlain e 3158;7 fro mTrille s 118.

WageningenAgricultural UniversityPapers 96-1 (1996) 105 10 15 Map6 . Hunteria hexaloba.

beneath in dried leaves, curved, in 7-10pairs ,th e lower veins reaching the margin without loopingt ojoi n theirneighbours ,formin g anangl eo f 55-650 withth ecosta , 13-30 mmapar t atth eapproximat e centreo fth eleaf .Inflorescence terminal,rarel y axillary, 1.5-2x 3-3.5 cm, about 10-20-flowered; peduncle 2-3 mm long; pedicels 3-6 mm long.Bract s very small.Flowers: Sepals pale green, free or connate at the base for about 0.2 of their length, membranaceous at the edge and the tip, thicker towards themiddl e andth ebase ,erec tan dwit hresi n within,ovate ,2. 3 x aslon g as wide, 1.3-1.5x 0.6-0.7nun , acute at the apex, with rows of colleters inside up to 0.75o fthei rlength ;colleter ssmall .Corolla reddish , 16-16.4m mlon gi nmatur ebu d and forming acomparativel y bigovoi dhea d0.56-0.5 7 of thebu d length,9-9. 4 x 1- 2 mm, with an obtuse apex, with a belt of pubescence 1-1.5m m wide inside just belowth einsertio n ofth estamen san du pt oth emout ho fth ecoroll atube ,tub e4. 6 x as long asth e calyx, 0.9 x as long asth e lobes, 6.5-7 nun l°ng» almost cylindrical, 1 mm wideabov eth ebase ,narrowe d below the insertion of the stamens to0. 8 mm wide, again widened around the anthers to 1m m wide; lobes narrowly oblong, twistedi nmatur ebud , 1.2x a slon ga sth etube ,4. 2x aslon ga swide ,7. 5 x 1.8mm , obtuse atth e apex, spreading and recurved later.Stamens with apex 2.5 mm below the mouth of the corolla tube,inserte d 0.45 of the length of the corolla tube at 2.6- 2.8m mfro mth ebase ;filament s 0.3-0.5m mlong ;anther snarrowl yovat et ooblong , 5.6 x aslon ga swide , 14-1.5 xo.25-0.5 mm,obtus ea tth eapex ,cordat ea tth ebase . Pistil 2.5-2.8 mm long; ovary subglobose, 0.3-0.5 x 0.3-0.5x 0.2 mm, abruptly narrowed into the style, with a disk-like thickening uniting the carpels at the base, abouto.2o fth elength ,0. 1 mmhigh ,o ftw oseparat ecarpels ;styl e 1.7-1.8m mlong ; pistil headcompose d of asubglobos e stigmaticbasa lpar t0. 4 x 0.2 mman d conical stigmoid apex 0.2 x 0.1 mm. Ovules 6 in each carpel. Fruits red, smooth, of two separatemericarps ;mericarp sellipsoid ,25-2 7 x 18-20x8-i o mm,acuminat ea tth e apex,acume n6x 4 mm,abou t4-seeded ; wallabou t4 m mthick ;pul pfibrous . Seeds obovatet ooblong , 16-17 x 11-12 x 6-7 mm,test asmooth ,orange .Embry o9. 5 mm long; cotyledons obovate, 1.4-1.7x aslon g as wide, 3.5-5 x 2.5-3 nun, rounded at the apex,rounde dt o obtusea tth ebase ;radicl e0.8-0. 9x aslon g ascotyledons , 3-4 x 0.8 mm.

DISTRIBUTION. Gabon. ECOLOGY. Forest. Alt.0-10 0 m. Flowering in November.

106 WageningenAgricultural UniversityPapers 96-1 (1996) Specimensexamined : GABON. Estuaire:nea rLibreville , Klaine315 8 (holotypeP ,isotyp eBR) ,Trille s 118 (P, paratype).

2.5.7 Hunteriamacrosipho n Omino,sp . nov.

Arbuscula gracilis foliis laminis angusteelliptici s papyraceis apicecaudati s venis submarginalis.Inflorescentiaterminali sve lrarite raxillaris .Flore sodorantes .Sepal a liberaerect aovat ave ltriangularia . Corollalaet elute aextu sglabr a intuspr opart e pubescens tubo angusto lobis patentibus. Stamina inclusa filamentis brevisbus filiformis glabrisantheri sovati sve loblongi sapic eobtusi sbas isubcordati sglabris . Ovarium ovoideum carpellis duabis separatis. Fructus luteus laevis mericarpiis duabusobovatu sdivaricatus .Seminaellipsoidea .-Type :Gabon ,nea rLastour sville , LeTestu 7085(holotyp eBR ,isotype sB ,M ,P , LISC). Fig.29 ,p . 108; map.7 p .10 9

Etymology: XJ.CIXQOÇ= long,of<|>(ov =siphon ,referrin g toth elon gcoroll atube .

Slendererec tshru b 1-4 mhigh ,wit ha littl ewhit elatex .Trun kslender .Branchlet s quadrangular, green. Leavesopposite ; petiole 5-8m m long; blade papery when fresh, elliptict ooblong ,2.4-3. 2x a slon ga swide ,5.6-19. 5x 1.8-6.4nun ,caudat e atth e apex, acumen obtuse, 13-25x 2-3mm , acutet o obtuse at thebase ;midri b prominent beneath; secondary veins prominent beneath in dried leaves, slightly curved,i n9-1 4pairs ,formin g asubmargina l vein anda nangl eo f 65-750 withth e midrib,8-1 3 mmapar t atth eapproximat ecentr eo f theleaf . Inflorescencei n i(-3) terminal,umbellat e cymes, 1.5-3 x I_3-5n™ .eac n 6-15-fiowered; sessileo rwit h apeduncl e2- 9 mmlong ;pedicel s 1- 4m mlong .Bract sver ysmall ,sepal-like .Flow­ ersfragrant . Sepalsfree ,erect ,membranaceous ,ovat et otriangular , 1-2.5 xa slon g aswide ,0.7-1. 1 x0.3-0. 7mm ,acut ea tth eapex ,wit hcolleter s sparsely distributed inside on the side; colleters small. Corolla bright yellow, 11.7-14.3 mm long in maturebu dan dformin g acomparativel ysmal lovói dhea d0.25-0.3 6m mo fth ebu d length, 2.9-6x 0.7-1. 8mm ,wit h anacut e apex,wit ha bel to f pubescence 1.2-3.5 mmwid einsid ejus tbelo wth einsertio n ofth estamen san du pt oth emout ho f the tube;tub e10-1 3x a slon ga sth ecalyx ,1.8-3. 9x a slon ga sth elobes ,9-1 3 mmlong , almostcylindrical ,0.7-1. 1m mwid eabov eth ebase ,narrowe dbelo wth einsertio n of the stamens to0.7-1. 5m mwide ,agai n widened around the antherst o 1-2 mm wide;lobe sovate ,no ttwiste di nbud ,0.3-0. 6x a slon ga sth etube ,1.6-2. 5x a slon g aswide ,2.8-6. 5x 1-8- 3nun ,obtus ea tth eapex ,entire ,spreadin gan drecurve dlater . Stamenswit hape x0.5-1. 2m mbelo wth emout ho fth ecoroll atube ,inserte d0.75 - 0.85m mo fth elengt ho fth ecoroll atube ,a t7-9. 8m mfro mth ebase ;filament s0.4 - 0.6 mmlong ;anther s ovate4-5. 7 x aslon g as wide, 1-1.9x 0.2-0.3mm ,obtuse , cordatea tth ebase .Pistil 8.8-10. 8m mlong ;ovar yovoid ,0.8-1. 4x 0.5-1x 0.3-0. 7 mm,abruptl ynarrowe dint oth estyle ,wit ha disk-lik ethickenin gunitin gth ecarpel s atth ebase ,abou t0.2-0. 3o fth elength ,0. 2m mhigh ,o ftw oseparat ecarpels ;styl e 5.5-8x 0. 2mm ;pisti lhea dcompose do f asubglobos estigmati cbasa lpart ,0.2-0. 5 x 0.2-0.3nu nan d aconica l stigmoid apex0.3-0. 7x 0.1-0. 2mm .Ovul e 1 ineac h

WageningenAgricultural University Papers 96-1 (1996) IO7 Figure29 .Hunteria macrosiphon. 1,habi t( x2/3) ;2 , sepalinsid e( x30) ; 3flower, abov e( x4) ;4 , flower bud( x4) ;5 ,opene dcoroll a( x4) ;6 ,pisti lwit hopene dovar y( x4) ;7 fruit, ( x2) ;8 ,longitudina lsectio n of seed( x4) ;8 ,embry o( x4) . 1-6from Bretele re tal .9934 ;7- 9from Breyn e4131 .

108 WageningenAgricultural UniversityPapers 96-1 (1996) 15

15 Map7 .Hunteria macrosiphon. carpel.Fruits yellow ,smooth ,o ftw odivaricat emericarps ,divergen ta ta nangl eo f 180°, mericarps obovate orellipsoid , slightly stipitate, 15-20x 9-12x 5-10 mm, witha nobtus eapex ,1 -seeded ;wal labou t2 m mthick ;pul psmooth .Seeds ellipsoid , 11x 5- 6x 4- 5mm . Embryo6.8- 7 r*1111long ;cotyledon sobovate ,1.9-2. 7x a slon g aswide ,3.5- 4 x 1.5-1.8 mm,rounde d atth eape xan da tth ebase ;radicl e0.8-0. 9 x aslon ga scotyledons ,3-3. 3x 0.5-0. 6 mm.

DISTRIBUTION. Gabon andCongo . ECOLOGY. Inrai n forest andsecondar y forest inCongo . Flowering inMarch - April andfruitin g betweenNovembe rt oJuly .

Specimensexamined : GABON. Ogooué-Lolo:3 0k mN Eo fLastoursville ,Breteler 992 0(WAG) ,993 4(WAG) ,986 7(WAG) ; 30k mE o fLastoursville ,Breleler et al. 11267 (WAG), 11351 (WAG);6 k mE o fLastoursville , Van der Maesene tal . 5534(WAG) ;Lastoursville ,Le Testa 7085 (BM,BR ,P , LISC,type) , 24-2-1930 (BR,BM , MO),4.3.193 1 (BM,BR ,LISC) .Wole uNtem : 35k mo nroa dfro mLebamb at oYeno ,J. deWilde Sc. Sosef 10454(WAG) . CONGO. Komono, Breyne413 1 (BR);2 5k mW o f Sibiti,Farron 4362 (P);Loundji , Artims9 8 (P); MbayaForest ,Sita 404 8 (P).

Note.Th esepal sar emembranaceou san dcolleter sar esparsel ydistribute do nth e innersid ea soppose dt oth eothe rspecie swher eth ecolleter s covera tleas thal fo f thelowe rpar to fth esepals .

2.5.8 Hunteriamyrianth aOmino ,sp . nov.

Frutexve larbo rparva .Foli aoppositapetiolat alammi selliptici sbrevite racuminatis . Inflorescentia terminalislaxa .Sepal apallid eviridi aliber atriangulari a apiceacut e usque ad acuminata. Corolla rubra tubo lobis ovatis longiore. Stamina inclusa antherisangust eovati sapic eobtusi sbas icordatis .Pistillu movari ocarpelli sduabu s separatis.Fructu signotus .-Type :Zaire ,Kasai-Occidental ,Kakenge ,Dechamps 5 (holotypeBR ;isotype sMO ,WAG) . Fig.30 ,p . 110; map8 ,p . Ill

WageningenAgricultural University Papers 96-1 (1996) IO9 Figure 30.Hunteria myriantha. 1,habi t (x 2/3); 2, leaf beneath (x 2/3); 3, flower bud (x 4); 4, opened corolla (x 12);S ,pisti l with opened ovary (x 12). 1 from Dechamps 5; 2fro m Dechamps 242;3- 5 from Declercq 32. no WageningenAgricultural UniversityPapers 96-1 (1996) 10 20 s / \ ' j *1 \( ; ~i i y

1 i \ V i ( 10 20 Map 8.Hunteria myriantha.

Etymology: nvploç,myrios ,ver ymany ;àv6oç ,anthos =flower, referrin g toth e manyflowered inflorescence .

Shrubo rsmal ltre e2-1 4m hig hwit hscanty ,white ,stick ylatex .Trun kstraight , 21- 25c mi ndiameter ; outerbar kgreyis h black,patche dpal ebrownis h yellow, 1 mm thick; inner bark orange brown, 4m m thick. Leaves opposite; petiole 10-13 mm long;blad eelliptic , 1.7-3.2x a slon g aswide ,6-2 0x 3.2-8.3shortl y acuminatea t theapex ,acume nobtuse ,5-14x2- 5mm ,acut ea tth ebase ;midri bprominen tbelo w indrie dleaves ;secondar yvein soblique ,obscur eabove ,i n7-1 2pairs ,reachin gth e marginwithou tloopin gt ojoi nthei rneighbour san dformin ga nangl eo f50-60 0wit h themidrib , 10-18m mapar ta tth eapproximat ecentr eo fth eleaf . Inflorescenceter ­ minal,lax ,i n umbellatecymes , 1 or2 x branched ,4.5-6. 5x 4- 7cm ,30-160-flow - ered; peduncle 13-32mm ;branche s 5-18mm ;pedicel s 2-3mm . Flowers: Sepals palegreen ,free , erect,triangular , membranaceous, 1.1-2.2x a slon ga swide , 0.7- 1.1 x0.4-0. 5 mm,acut et oacuminat ea tth eape xan dwithou tcolleter sinside . Co­ rollared ,7.7-8. 3m mlon gi nmatur ebu dan dformin g acomparativel y smallovoi d head0.3-0. 4o fth ebu d length, 2.5x 0. 8mm , with anobtus e apex, with abel t of pubescence1.2- 2 mmwid einsid ejus tbelo wth einsertio no fth estamen san dsome ­ timesu pt oth emout ho fth ecoroll atube ;tub e4.7-7. 9x a slon ga sth ecalyx ,1.4- 2 x aslon ga sth elobes ,5.2-5. 8m mlong ,almos tcylindrical ,0.8- 1m mwid eabov eth e base,narrowe dbelo wth einsertio no fth estamen st o0.7-0. 9m mwide ,agai nwid ­ enedaroun dth eanther st o1-1. 3 mmwide ;lobe sovate ,0.6-0. 7x aslon ga sth etube , 1.7-2.4x aslon g as wide, 2.8-3.9mm * obtuse atth eapex , entire andspreading . Stamens withape x0.2-0. 5 n™belo wth emout ho fth ecoroll atube ,inserte d0.77 - 0.82 of thelengt h of the corolla tube at4-4. 5 mm from thebase ;filaments 0.4 - 0.5m mlong ;anther s narrowly ovate,2- 4x a slon g aswide ,0.6- 1x 0.2-0. 3naa < obtusea tth eapex ,cordat ea tth ebase .Pistil 4.8-5. 5m mlong ;ovar ysubglobos e0.5 - 0.8 x0.4-0. 7x 0.25-0. 4mm ,abruptl ynarrowe dint oth estyle ,wit ha disk-lik ethick ­ eningunitin gth ecarpel sa tth ebase ,abou t0. 3o fth elength ,0. 4m mhigh ,o ftw o separatecarpels ;styl e3.2- 4m mlong ;pisti lhea dwit ha stigmati cellipsoi dbasa lpar t

WageningenAgricultural UniversityPapers 96-1 (1996) III 0.4-0.6x 0. 2 mman da stigmoi d apex0.4-0. 6x 0.1 5 mm.Ovul e 1 ineac hcarpel . Fruitsunknown .

DISTRIBUTION.Zaire . ECOLOGY.Secondar yforest .Alt . 520m .Flowerin gbetwee nJanuar yan dApril .

Specimens examined: ZAIRE. Kasai-Occidental: Benalongo,Declerq 3 2(BR) ;Kakenge ,Dechamps 5 (BR ,WAG) , 139 (BR), 242 (BR, MO,WAG) .

Note.Thi s species isver y close toH. densiflora andH. ghanensis, andca n be differentiated asfollows :

1. Corollalobe swide rtha nlon g H.ghanensi s Corollalobe slonge rtha nwid e 2 2.Secondar y veins oblique,i n7-1 2pairs ,reachin gth emargi n without loopingt o join theirneighbours ,formin g anangl eo f50-6o°wit hth ecosta ; sepals without colleters;inflorescenc e lax H.myrianth a Secondaryvein sstraight ,closel yspaced ,i n22-3 0pairs ,formin g anangl eo f70 - 80°wit hth ecosta ;sepal swit hcolleter s within;inflorescenc e dense H. densiflora

2.5.9 Hunteriaoxyanth aOmino ,sp . nov.

Arbuscula rarius scandens. Folia laminis papyraceis ellipticis veloblongi s apice acuminatis.Inflorescenti aterminali sve laxillari spauciflora . Sepalapallid eviridi a basi connata erecta ovata vel elliptica margine mebranacea apice rotundata vel obtusa.Coroll asordid ealb aextu sglabr aintu spr opart epubescen stub ocylindrace o lobiserectis .Stamin ainclus afilamentis brevibu sfiliformibus glabri santheri s fere ellipticis glabris. Ovarium subglobosum carpellis duabus. Fructus luteus vel aurantiacuslaevi smericarpii sduabu sdivaricati sellipsoideis .Semin aellipsoidea . - Type:Gabon ,Ogooué-Lolo ,40-5 0k mSE Eo fLastoursville ,Breteler e tal .1129 9 (holotypeWAG). Fig.31 ,p .113 ;ma p9 ,p . 114

Etymology:oë,vç, oxus ,acute ,ôvGoç ,anthos ,flower ,referrin g toth eacut eape x of theflowe rbud .

Treeo rshru b 1.5-18 mhigh ,rarel ya climber .Trun ku pt o3 0c mi ndiameter ;bar k rough,gree ngre ywit hwhit espot so flichen ,8 m mthick ,hard ,brittle ,brigh tyello w underneath; wood hard, dense, very brittle, with twisted fibres, lemon-yellow. Branchesgreyish-black finely, striate .Leaves opposite ;petiol e4- 6m mlong ;blad e papery tocoriaceou s when fresh, elliptict ooblong , 2.3-3 x aslon g aswide , 3.7- 12.8x 1.3-4.6 cm,acuminat ea tth eapex ,acume nobtuse ,4-1 9x 2- 3mm ,acut et o obtuse atth ebase ; midrib slightly prominent beneath; secondary veins straight,

112 WageningenAgricultural University Papers 96-1 (1996) Figure 31. Hunteriaoxyantha. 1, habit (x 2/3);2 , leaf beneath (x 2/3);3 , sepal inside (x 30);4 , opened corolla (x 4); 5, stamen outside (x 30); 6, pistil (x 4); fruiting branch (x 2/3). 1-7 from Breteler et al. ii299.

WageningenAgricultural UniversityPapers 96-1 (1996) 113 10 20 30

10 10

10 20 30 Map 9.Hunteria oxyantha. slightlyprominen tbeneath ,i n15-2 5 pairs,formin g asubmargina lvei nan da nangl e of75-8o°wit hth emidrib ,3- 6m mapar ta tth eapproximat ecentr eo fth eleaf .Inflo­ rescencetermina lo raxillary ,2- 3x 1-2 cm, 1 xbranched ,6-10-flowered ;peduncl e 1-11m m long; primary branches, 2-10m m long; pedicels 4-6m m long. Bracts sepal-like,0.5- 1x 0.4-0. 5mm . Flowers: Sepalspal egreen ,free , erect,membrana ­ ceous at the edge and the apex, thicker towards the middle and at the base,wit h much resin within, ovate, 1-2.1 x as long as wide, 1-1.3 x 0.6-1.1mm , obtuse to acutea tth eapex ,wit hrow so f colleters insideu pt o0.7 5o f theirlength ;colleter s verysmal l0. 2x 0. 1 mm.Corolla whit et odirt ywhite ,re di nbud , 15-15.5m mlon g inmatur ebu dan dformin g acomparativel y smallovoi dhead ,0.35-0.4 3o fth ebu d length,5.5-6. 5x 0.7-1.6mm ,wit h anacut e apex,wit h abel t of pubescence1.5- 2 mmwid einsid ejus tbelo wth einsertio no fth estamens ,tub e8.6-9. 5x asl° nI asth e calyx, 1.5 x aslon g asth elobes ,9.5-1 2m mlong , almostcylindrical ,0.8-1. 5 mm wideabov eth ebase ,narrowe d belowth e insertion of the stamenst o0.7-0. 9m m wide,agai n widened around theanther st o 1.5-2m mwide ;lobe soblong ,0. 7 xa s longa sth etube ,2.9-3. 8 xa slon ga swide ,7.5-7. 9x 2-2. 7vam > obtusea tth eapex , spreading andrecurve dlater .Stamens wit hape x 1 mmbelo wth emout ho f theco ­ rolla tube,inserte d 0.73m mo f thelengt h of thecoroll atube ,a t 8.8 mmfro m the base; filaments 0.4-0.5m mlong ; anthers almost elliptic,4-4. 5 x as long aswide , 1.8-2x 0.4-0. 5mm ,obtus ea tth eape xcordat ea tth ebase .Pistil 7.3-8. 5m mlong , with apex halfway along the anthers; ovary subglobose, 1-1.4X 0.6-0.7x 0.3- 0.7mm ,graduall y narrowed intoth e style,wit h a disk-likethickenin g unitingth e carpelsa tth ebase ,abou t0. 3o fth elength ,0.2-0. 4m mhigh ;o ftw oseparat ecarpels ; style5.3- 6 mm; pistilhea dcompose do f aglobos et oellipsoi d stigmaticbasa lpar t 0.5-0.7x 0.3-0.4an d abifi d stigmoid apex0.4-0. 5x 0.15-0.2mm .Ovule s 6-7i n eachcarpel .Fruits yello wt oorange ,smooth ,mericarp sellipsoid ,28-3 5x 20-26x 15-20mm ,bluntl y acuminatet orounde da tth eapex ,acume n4- 6mm ,3-7-seeded ; wallabou t 1 mmthick ;pul pslightl yfibrous. Seeds oblon go rellipsoid , 12-14x 8-

114 WageningenAgricultural University Papers96-1 (1996) io x 4-6 mm,test a smooth,orange .Embry o 7.5 mmlong ;cotyledon s ovate,1.5 - 1.6 x as long as wide,3.8- 4 x2. 5 mm, obtuse at the apex,rounde da tth e base;radi ­ cle0.9- 1x a slon ga scotyledons ,3.5-3. 7x 0.8- 1 mm.

DISTRIBUTION. Gabon,Cong oan dZaire . ECOLOGY. Oldsecondar yfores t orfores tborderin gsavanna h on humidrock so r riverbanks .Flowerin gan dfruitin g betweenMa y andSeptember .Alt .48 0m .

Specimensexamined : GABON.Woleu-Ntem :Chantie r Oveng,A.M.Louis 2101 (WAG).Ogooué-Ivindo :Mpass aFiel dSta ­ tion, Florence 584(P) ,61 4 (P),Caballé 365 (WAG),nearMakokou , Gentry3302 7(MO) ,3317 7(MO) , 33264 (MO);3350 6 (MO, WAG), Gentry& Emmons 33736 (MO). Ogooué-Lolo: 9 km Eo f Ndambi, Bretelere t al. 13233(WAG) ;40-5 0k m SEE ofLastoursville ,Breteler e tal . 11229 (WAG, type). Nyanga: 25 kmSS Wo f Doussala,Reitsma e tal . 976(WAG) . CONGO. Kouilou:Le sSaras ,Sita 1361(P) . ZAIRE. Kivu:Maniema ,Pangi , Michelson 85(BR) .

2.5.10 Hunteriasimi i (Stapf) H. Huber

inKe w Bull.15:43 7(1962 ) & inFl .W.Trop .Afr . ed. 2,2:62(i9Ó3).Type :Liberia , SinoDistrict ,Sim 1 6(holotype , K). Fig. 32, p. 116; map 10, p.11 7

Basionyman dhomotypi csynonyms :

Polyadoa (?) simiiStap f in H.H.Johnstone,Liberia ,2:62 4(1906) .Pleiocarpa simii (Stapf) Hutch. &Dalz . in Fl.W.Trop . Afr. ed. 1, 2: 38 (\ysi);Tetradoasimii (Stapf) Pichon in Bol. Soc.Brot . sér. 2: 120;p . 117t . 3,fig 4-6 ;ma p C,p .2 1 (1953)- Treeo rshru b0.3- 8 mhigh ,rarel yclimbe r 1 mhig h withwhit estick ylatex .Trun k slender, slashcreamish-yellow ;branche s greenish grey,branchlet s greyish-brown to green, with a few lenticels.Leaves opposite; petiole 4-9 mmlong ; bladecori ­ aceous orpaper y when fresh, smooth, ovatet oelliptic , 2.2-2.8 x aslon g aswide , 5.5-19.2x (1.8- )3.8-7. 4cm ,acuminat ea tth eapex ,acume nobtuse ,6-2 3x 2- 3mm , acutea tth ebase ;midri bprominen tbeneath , slightly channelled above,secondar y veinsoblique ,slightl yprominen tbeneath ,i n7- 9pairs ,formin g anangl eo f55-56 0 withth emidrib ,18-3 0m mapar ta tth e centralpar to fth e leaf.Inflorescence termina l oraxillary ,2-4. 5x °-5-3 cm> I"10flowered; peduncl e 2-10mm ,primar ybranche s 2-10mm ;pedicel s 5-6m mlong .Bract s 0.7-1.3mm .Flowers: Sepals palegreen , almostfree ,erect ,membranaceou sa tth e edge,thicke rtoward s the middle and atth e base,coate dwit hresi nwithin ,ovat e1.4-1. 6x a slon ga swide ,1 -1. 3x 0.7-0. 4mm , acuminate at the apex, with rows of colleters inside up to 0.8o f their length; colleters very small 0.2 x 0.1 mm. Corolla cream-coloured, 16-23.7 n™ l°ng m maturebu d andformin g a comparativelylarg eovoid ,sometime sfalcat ehead ,0.38 - 0.5 of thebu d length,7-11. 2x 1.3 mm, with anacut e apex;wit h abel t of pubes-

WageningenAgricultural University Papers96-1 (1996) 115 Figure 32.Hunteria simii. 1-2, habit (x 2/3);3 ,flower ( x4) ;4 ,opene dflower ( x4) ;5 ,sepa linsid e (x 20); 6-7,stame nin -an doutsid e (x 20);8 ,pisti l (x20) ;10 ,frui t (x 2/3); 11, seed (x 2);12 ,longitudina l section of seed (x 2); 13,embry o (x 4).1- 2 and 10-13ft 10111Bretele r 7388; 3-9 fromVa n Meer464 .

116 WageningenAgricultural University Papers96-1 (1996) Map 10.Hunteria simii. cence 1-1.2m m wide insidejus t below theinsertio n of the stamens,tub e greenish- white, 13-18 x aslon ga sth ecalyx , 1-2.3x a slon ga sth elobes , 12.5-18.5m mlong , almost cylindrical, 0.8-1.7 mm wide above the base, narrowed below the insertion of the stamens to0.5-1. 3m m wide, again widened around the anthers to 1-1.8m m wide;lobe s narrowly ovatet o oblong,0.4- 1x aslon g asth etube ,3.6-6. 7 x as long as wide, 7-12 x 1.8mm , obtuse at the apex, entire, spreading. Stamens with apex 3.2-3.7m mbelo w themout ho f thecoroll atube ,inserte d 0.48-0.62 of the length of the corolla tube, at 6-8 mm from the base;filament s 0.4-0.5m m long; anthers nar­ rowly ovatet ooblong ,4.8-5. 3x a slon ga swide , 1.6-1.9 x0.3-0. 4 mm,obtus ea tth e apex subcordate at the base. Pistil 6.5-7.8 mm long; ovary subglobose, 1.0x 0.5- 0.7 x0.3-0. 4mm ,abruptl y narrowed intoth estyle ,wit ha disk-lik ethickenin g unit­ ingth ecarpel s atth ebase ,abou t0. 2 ofth elength ,0.2-0. 3m mhigh ;o ftw o separate carpels; style 6-7 x 0.25 mm; pistil head composed of an ellipsoid stigmatic basal part 0.5-8 x 0.2-0.3m m and abifi d stigmoid apex 0.4-0.6 x 0.1-0.2 mm. Ovules 6 in each carpel.Fruits yellow to orange, smooth, of two divaricate follicles, oblong, 37-60.5 x 10-18 x 10-13mm , produced into a long slender beak atth e apex, beak 6-15 x 2-3 mm, 5-6-seeded; wall about 1m m thick; pulp smooth. Seeds oblong or ellipsoid, 11.5-15 x 6.5-8.5 x 5.5-6.5mm ,test asmooth , orange.Embry o 6-8.5 mm long; cotyledons ovate, thin and leafy, 2.2-4 * as long as wide, 3-4 x 1-1.8mm , obtuse at the apex, rounded at the base;radicl e 0.8-1.1 x as long as cotyledons, 3- 4.5 x 0.5-0.6 mm.

DISTRIBUTION. Guinea, Sierra Leone, Liberia and Ivory Coast. ECOLOGY. Densefores t orol dsecondar yforest .Flowerin gfro m Februaryt oJul y and fruiting all the year round in all four countries. Alt. 0-450 m.

Geographical selection of the approximately 40 specimens examined: GUINEA. Nzérékoré:Macenta ,Koyama ,Adam 59(MO) . SIERRA LEONE.Yonibana , N.W. Thomas 4764 (K); Gola Forest, block m, Small53 9 (K), 649 (K); KambuiFores t Reserve,Jordan 203 7 (K). LIBERIA. GennaTanyehun,Baldwin Jr. 10736 (K);Mecca ,Baldwin Jr. 10443 (K); 8k mN of Bomi Hills,Jansen 228 2 (WAG);Dukwi a R., Cooper 169(K) ;Fireston e Plantations, S of Kakata,Stoop-van de Kasteele 38 (WAG); HarbelForest ,Bos 202 8 (WAG); Belafani,Daniel 238 (MO); Ganta,Baldwin Jr. 12561 (K, MO); 10 km N of Tappeta, Van Meer 464 (WAG); Yeképa, Mt. Nimba,Adam 21395 (MO,

WageningenAgricultural UniversityPapers 06-1 (1996) II7 UPS); Sanikwele (= Sanniquellie), BaldwinJr. 9460 (K);Tappeta ,Adam 30299 (MO);sin . loc., Sim 16 (K,holotype) . IVORY COAST. 13 km NW of Tabou, Breteler 7388 (WAG); 14 km NW of Tabou, Breteler 7398 (WAG); Fort Binger (= Djiroubou), Chevalier 19535 (P);Tai ' Forest, De Namur 1423 (WAG); Kéeta, Chevalier1931 1 (P,K) ,Chevalier 19367 (P); ZozoroR. ,Chevalier 19036 (P); TékéForest ,Bamps 230 4 (BR,P) , 2388 (BR,P) . CULTIVATED. Nigeria: AkampaRubbe rEstate ,Latilo 196(K ) (= FHI40918 (A)).

2.5.11 Hunteria umbellate (K.Schum. ) Hall. f. inJahrb .Hamb .Wiss . Anst. 17, 3 Beih., 190(1900) ; Pichon inBull . Soc.Brot . sér. 2, 27: 99, t. fig. map A, p. 85 (1953). -Type: Cameroun, Lolodorf, Staudt 130, (holotype B t; lectotype K,designate d here;isotypes ,A , COI,E , P,PRE ,Z) . Fig. 33, p. 119;ma p 11, p. 120 Basionym and homotypic synonyms:

Carpodinus umbellata K.Schum . in Engler, Bot. Jahrb. 23: 221 (1896); Pierre in Bull. Soc. Linn. Paris 38, (1898). Polyadoa umbellata (K.Schum. ) Stapf in FI. Trop.Afr .4,1:10 3 (1902).Picralima umbellata (K.Schum. ) Stapf in KewBull . 1908: 302 (1908).

Heterotypic synonyms:

Polyadoa elliotii Stapf in op.cit . 104;Picho n inMem . Mus.Natn .Hist .Nat . sér.2 , 24: 174 (1948). Picralima elliotii (Stapf) Stapf in Kew Bull. 1908: 302 (1908). Hunteria elliotii (Stapf) Pichon in Bol.Soc .Brot . sér. 2,27:97,1.1fig. 5-8 ;ma p A, p. 85 (1953), syn. nov. - Type: Sierra Leone, near Makunde, Limba, Scott Elliot 5690 (lectotype K, designated by Pichon in 1953). Hunteria eburnea Pichon in Bol. Soc.Brot . sér. 2, 27: 91, t.i fig 1;ma p A (1953), syn. nov. - Type: Ivory Coast, Danir, between Yaou and Ayiame, Chevalier 17766, (holotype P; isotypes K, P, MO, WAG). Hunteria mayumbensis Pichon in Bol. Soc.Brot . sér. 2, 27:95 ,t. i fig. 2-4; map A (1953), syn. nov. - Type: Gabon, Mayumbe Bayaka, Inganga, Le Testu 1744 (holotype P;isotype s BR, K, NY,L) .

Tree or shrub 1.5-22 m high, with a dense leafy crown.Trun k sinuous or straight, fluted, (3-)10-4 0c m in diameter; outer bark grey to dirty brown, rough or smooth, 1m mthick ;inne rbar k 3-4m mthick , whitewit horang e stripeso r spots;woo d very hard, creamy, brown to yellow or orange. Branches greyish to dark brown; branchlets palegreen , smooth.Leaves opposite;petiol e 8-25 mm long;blade , thin, soft, leathery, subcoriaceous when dried, elliptic to oblong, 1.9-3.4 x as long as wide,5.8-22. 5x 2-11cm ,obtus et o acuminate atth eapex , acumen obtuseo racute , 5-18 x 3-7 mm, acute to obtuse at the base or decurrent into the petiole, midrib deeply channelled above, prominent beneath; secondary veins prominent beneath, moreo rles sstraight ,i n 13-25pairs ,formin g asubmargina l veinan da nangl eo f 65-

I18 WageningenAgricultural UniversityPapers 96-1 (1996) Figure33 .Hunteria umbellata. 1, habit( x 2/3);2 , leaf beneath( x2/3) ;3 flower, bu d( x4) ;4 flowers, ( x 4);5 ,opene dcoroll a( x 6);6 ,sepa linsid e( x 14); 7,pisti l( x 10);8 ,ope nfrui t (x 2/3); 9,see dwit hpul p (x 2);10 ,see d (x 2); 11, longitudinal section of seed (x 2);12 ,embry o( x 6).1- 7 from EspiritoSant o 3001;8-1 2 fromJanse n2147 .

WageningenAgricultural UniversityPapers 96-1 (1996) lic Map 11.Hunteria umbellata.

75°wit hth emidrib ,4-2 3m mapar ta tth ecentra lpar to fth eleaf .Inflorescence te r minai,rarel y axillary, 1.5-5x ï-3-5cm , denset o lax, about io-20(-8o)-flowered; pedunclesometime scovere dwit hresi na tth ebase ,2-1 3m mlong ,primar ybranche s 3-10m m long;pedicel s sometimes covered with resin at thebase , 3-7 mmlong ; bracts very small, 0.8 x 0.5mm , without colleters.Flowers very fragrant.Sepals palegreen ,almos tfree , thickertoward sth emiddl ean da tth ebase ,erec tan dstuc k toth ecoroll atub ewit hthic kresinou s substance,membranaceou sfo rabou t0.2-0. 3 mmo f theiredge ,broadl y ovatet otriangular ,0.7-2. 6x aslon g aswide ,0.6-1. 8x 0.5-1.6mm ,acut et oobtus ea tth eapex ,wit hrow so fcolleter sinsid eu pt o0.7 5o f theirlength ;colleter s verysmal l0. 2 x0. 1mm .Corolla white,cream yo rpal eyel ­ low, 8.5-17.5m m long in mature bud andformin g acomparativel y narrow ovoid head (o.45-)o.5-o.68 of thebu dlength , 5.5-9.5x 0.5-2.5mm ,wit ha nacut eapex ; with abel t of pubescence 0.8-1.7mm wideinsid ejus t below the insertion of the stamens,sometime s with afe w hairsaroun d the stamens,tub e2.9-1 2x aslon ga s thecalyx ,0.4- 1(-1.2 )x a slon ga sth elobes ,4-8. 2m mlong ,almos tcylindrical ,0.9 - 1.8 mmwid e aboveth ebase ,narrowe dbelo w theinsertio n of thestamen st o0.5 - 2.2m mwide ,agai nwidene daroun dth eanther st o0.7-2. 0m mwide ;lobe screamy , slightlypinkish ,narrowl yovat et ooblon gan dfalcate ,twiste di nbud ,(0.8- )1.2-2. 7 x aslon g asth etube , 2.5-7.5 x as long as wide,6.2-1 2x 1.3-3.5T^^ obtuse atth e apex,entire ,spreadin gan drecurve dlater .Stamens wit hape x0.3-1. 0m mbelo wth e

120 WageningenAgricultural UniversityPapers 96-1 (1996) moutho f the corollatube ,inserte d0.54-0.8 6m mo f the lengtho f the corollatube , at 2.5-6.2m mfro mth e base;filament s0.2-0. 8m mlong ;anther snarrowl yovat e 2.7- 5.5x a slon ga swide ,0.8-1. 4x 0.2-0.3 mm,obtus ea tth eape xcordat ea tth ebase . Pistil2.5-6. 4m m long;ovar ysubglobose ,0.6-1. 1x 0.5-1. 0x 0.3-0. 8mm ,abruptl y narrowedint oth estyle ,wit ha disk-lik ethickenin g unitingth ecarpel s at the base, about 0.25-0.4o f thelength ,0.2-0. 3mm high, oftw o separate carpels; style 0.7- 4.8m mlong ;pisti lhea dcompose do f astigmati cellipsoi dbasa lpar t0.3-1. 0x 0.2 - 0.4 mm and abilobe dstigmoi dape x0.2-0. 7XOA mm- Ovules 12-30i n eachcarpel . Fruitsyellow ,smoot ho ftw o separatemericarps ,mericarp ssubglobose ,3 1-6 0 x 40- 50x 30-40mm , 8-26-seeded; wall about 5-12m m thick, slightlyfibrous. Seeds oblongt oellipsoid flattened, a tth esid eadjacen t toon eanother , 11-17x5-11x4- 7mm .Embry o6-1 0m mlong ;cotyledon s elliptict oobovate , 1.3-2.4x a slon ga s wide,3-5. 5x 1.7-3. 5 mm,obtus e to rounded at the apexan da tth e base,radicl e0.7 - 1.4x a slon ga scotyledons ,3-4. 5 x0.5-0. 8 mm.

DISTRIBUTION. Senegalt oZair ean dAngola . ECOLOGY.Secondar yforest ,rai nfores t and galleryforest . Alt. 0-600 m. Flower­ ingan dfruitin g allth eyea rround .

USES.Th e wood is cream-coloured or deepbrow n to yellow-orangefine, graine d and very hard (Punch 138,E ,K , 1900,Nigeria ; Onochie FHI4936 ,FHO , 1945, Nigeria;Bakare e tal .FH I22857 ,K ,1948 ,Nigeria ;Letouzey 5477 , BR, K, P, 1963, Cameroun;Bur t& Hoyle1937) ,use dlocall yfo rcarvin gcombs ,spoons ,carpenter s planesan dweaver sshuttle s(Dalzie l 1937;Causdale 166, FHO, 1944,Ghana) ; for makingshuttle sfo rweavin gNativ eclot h (Mc Ainsh 890, Nigeria)an dfo rmakin g combs(Scott Elliot 5690 ,K , 1892,Sierr aLeone ;Mc Ainsh 890, Nigeria; Chevalier 19706, P, 1907,Ivor yCoast) .Th eforke d stemsar euse da s housepost san dar e con­ sideredver ydurabl e and immunet o termites(Dalzie l 1937).Fruit sar e consumedb y ants (Oldeman 642,BR ,K ,P ,WAG , 1963,Ivor y Coast).Roo t areuse da sa bitte r tonic (Akpabla no, K,P , 1948,Nigeria) .Bar k andfrui t aretake n locally against bellyach e (Bos 3065,WAG ,P , Cameroun).

Geographical selectiono f the approximately 200specimen sexamined : SENEGAL. Catió, Espirito Santo 3001 (BR, MO,P) ; Catanhez, Pereira 3163 (K, MA); Chitole Cusselinta,Espirito Santo 318 7 (MO,P , WAG). Oriental:Gouloumbo , Berhaut 2082(BR , P);Beig e du Nieriko, Berhaut166 9 (BR, P);Ouassadou ,Trochain $4&j (P).Casamance :Kayang aR. ,Monsier 259 4 (P)- GUINEA BISSAU. Canchungo,Esptrito Santo 1940 (K,MO ,P , WAG). GUINEA. Bóke, Pobéguin 2015 (P); Mt.Nimba ,Adam 28738 (MO). SIERRA LEONE.. Mafinta, Pyne 115 (K, P);nea rMakundi , ScottElliot 569 0(K ,typ eo fH. elliotii); Batkanu, N. W.Thomas 19 (K); Kasewe,King 23 1 (K), Lane-Poole12 9 (K); Mano, N. W.Thomas 10102 (K); Kambui Forest Reserve, Jordan 2120 (K), Samai 522 (K);Gol a Forest, Small 402 (K); Njala, Deighton501 0 (B, P,K) . LIBERIA. Mt. Huelliton,Jaeger 10019 (G); BomiHill-Man o Rd., VanMeer 38 0 (WAG);Gol aNationa l Forest, Voorhoeve3 6(BR ,WAG) , Voorhoeve 853(WAG) ;Saniquellie , Georges2781 4 (M); Mt.Tokadeh, Adam2141 4 (MO, P, UPS); 29k mS o fTchien ,Jansen 2147 (WAG). IVORY COAST. Sassandra R.basin ,Chevalier 1637 3(P) ;Cavall yR .basin ,Chevalier 1970 6 (P); near

WageningenAgricultural University Papers 96-1 (1996) 121 Tabou& Béréby, Chevalier1997 5 (P);Béréby , Oldeman 642 (BR, K,P ,WAG) ;Soubré ,AkéAssi 11578 (B); Monogaga, Geerling& Bokdam 2419 (BR, MO, WAG); 21 kmo n Rd.t oMonogaga , Leeuwenberg 12111 (WAG);Mafia , Garnier653 E(P) ;Agnieby , Chevalier 17043 (P);Bouroukrou , Chevalier 16519 (P.WAG); 36k mN E of Sassandra,Breteler 6102 (BR, K,P ,US ,WAG) ; Banco Forest,Ate 'Assi 15904 (BR, WAG); Sofalco Plantations,Leeuwenberg 10722 (WAG);Aboubo u Forest, Leeuwenberg238 6 (B, BR,FHO ,K ,LISC ,P ,WAG , Z); 20k mS Wo f Abengourou(BR ,WAG) ; Boiobo, Chevalier 17659(MO , P,WAG); between Yaou & Ayiatne, Chevalier 17766 (K, P, MO, WAG, type of H. eburnea); Assinie, Chevalier 17865 (P);Aboisso , Chevalier 16310A (K, P, WAG). GHANA. Western: Ankasa Forest Reserve, EntiR 97 3 (MO);nea r Simpa, Enti 6920 (B, BR, K, P); Benoo,Andoh FHI547 1 (BR, K, US, WAG);Subr iR. ,Hall & Abbiw 45134 (MO, WAG). Eastern: Abua- Kwa, Causdale16 6 (FHO);Essubon i (=Asuboni) Forest Reserve, Juma1 (FHO);Kumasi , Vigne 2701 (FHO,US) ;Brong-Ahafo :Bi aNationa l Park,Short G C4700 8 (MO). NIGERIA. Ogun:Olokomej i Forest Reserve, Gentry &Pilz 32674 (US, WAG); Omo ForestReserve , Gentry & Pilz 32810 (BR, MO, WAG). Erin, Ross 126 (K). Lagos: Lagos, Foster 5 (K, P,W). Oyo: Gambari, Van Eijnatten 1266(WAG) ;Ibada nSout hFores tReserve ,Keay FHI 25705 (B,K , P); Ago-Owu Forest Reserve, Wit & Geerling 1260 (K, MO);Shash a Forest Reserve, Ross 8 (MO, S);Bende l State: OkomuFores t Reserve,Akpabla 110 (K, P).Cros s River:Nort h Forest Reserve, Latilo FHI 31840(K) ; Ogqja,Keay FHI2825 0(K) . CAMEROUN. Sud-Ouest: KorupNationa l Park,D. W.Thomas 3207 (K, MO, P, WAG); Masore, D. W. Thomas 4178 B (K);Kumba ,D. W.Thomas 4195 (BR, K, MO, P, WAG); Abonando, Rudatis 59 (G,K , P, W, Z); Wernevillage, Nemba e tal .58 2 (MO, WAG).Centr e Sud: Bafia-Ndikimeki Rd.,Benoit 38 2(P) ; 50k mN Wo f Eseka,J. deWilde 1272 (P, WAG); Banko f NyongR .nea rth e newbridge ,/ . deWilde 2863 (WAG);Lolodorf , Staudt13 0(A , COI, E, K,P ,PRE , S, Z, type);9 kmN of Kribi,Bos 4978 (K, M,P , WAG); 2 km S of Kribi on Grand Batanga Road, Bos 3065 (WAG, P); 12 km on Grand Batanga to Campo, Wieringa &Haegens 215 6 (WAG);6 0 kmo n Kribi-Edea Rd.,Bos 7106 (P, WAG); nearKribi , Mildbraed609 8 (B,HBG) ;1 0k mN o f Kribi,J. deWilde 2922 (P, WAG); Bipindi,Zenker 170 7(BP , BR, COI, E, F, G, HBG, K, L, NY, P, S,W,WAG , Z), 1729 (BP, BR, E, G, HBG, K, L, MO, P, S,W,WAG) , 3705 (BR,E ,G ,HBG , K,L ,MO ,P ,S , US,W , Z),448 8 (B,BR , E,G , HBG,K ,L ,MO ,P ,S,W,WU ,Z) , H 287 (B, BR, C, G, GH, LD, P,UC , US, WAG); Akoakas rock, J. de Wilde 8371 (MO, P, WAG). Est: nearKolembong ,Letouzey 547 7 (BR, K,P) . GABON. Estuaire: 100k mS of Libreville, J.de Wilde et al. 812/83 (MO, WAG). Ogooué-Maritime: Gamba, Wieringa &. Haegens 2524 (WAG);Mossomala , Wieringa 1285 (WAG);Doudo u Mts.,Reitsma 1952 (NY,WAG). Nyanga:Tcbibanga ,Le Testu 1744(BR ,K ,NY ,P ,typ e of H. mayumbensis); Moyen- Ogooué:7 kmo n Ndjole-Bijoun Rd.,Leeuwenberg 12464 (MO, WAG). Ngounié: Maghounga,Le Testu 6408 (B, BR, K, MO, P). Nyanga: Mayombe, Le Testu802 5 (BR, P). Ogooué-Lolo: 30 km NE of Lastoursville,Breteler 999 3(WAG) . CONGO. Kouilou: Kakamoéka,Sita 134 5(P) ; Dimonika,Groulez i2.io.i954(P) ;Mayombe ,Dowsett- Lemaire 1447 (WAG);Kouilou ,Sargos 75 (P). ZAÏRE. Equateur:Bosawa , Evrard 1614 (BR). Haut-Zaire: Yangambi, Germain27 9 (BR, C, K,NY , WAG);HeTutuku , Germain 357 (BR, K). ANGOLA. Cabinda:Chiluango , Gossweiler 7764 (BM,K ,MO) . CULTIVATED. Netherlands: Wageningen Bot. Gardens, Van Setten 179 (WAG). Ivory Coast: AdiopodouméBo t Garden, Gamier 653 A (P);Adiopodoumé , Leeuwenberg 12026(WAG) .

Notes.Hardl yope n andope nflower s arepresen t onth etre eitself ,bu tabundan t on the forest floor. Probably night flowering ( Wieringa &Haegens 2524, WAG, 1994,Gabon) .

2.5.12 Hunteriazeylanic a(Retz. )Gard ,e x Thw. inEnum .PI .Zeyl .19 1 (i860);Tsian gYin gi nSunyatseni a3:13 5(1936) ;Picho ni n Bull. Soc. Brot, sér 2,27:104(1953) ;Wyatt-Smit h& Kochummeni nMai . For. Rec.

122 WageningenAgricultural UniversityPapers 96-1 (1996) 17: 72 (1965); Huber in Handb. Fl. Ceylon 1: 10 (1973);Tsian g Ying in Fl. Reip. Pop.Sin .63:15-17 ,fig. 4 (1977) ;Markgra f inBlume a30:16 9 (1984)).-Type:Sri - Lanka, Koenig s.n. (holotype C, isotypes BM, K-W 1608C , MO). Fig.34 ,p . 124;ma p 12,p . 125

Basionym and homotypic synonym:

Cameraria zeylanica Retz.i n Obs.Bot . 4: 24 (1786); A.DC, Prod. 8: 389 (1844). Hunteria corymbosa var. zeylanica (Retz) Hall, f., Jahrb. Hamb.Wiss . Anst. XVII, Bein. 3: 195 (1900).

Heterotypic synonyms:

Hunteria corymbosa Roxb., Hort. Beng. 84 (1814) nomen & Fl. Ind. ed. Carey et Wall., 2: 531 (1824) & ed. 2: 695 (1832);Wigh t Ic. 2 t. 428 (1843); A.DC, op. cit. 350 (1844);Hooke r f. in Fl. Brit. Ind. 3:63 7 (1882);Boerlage , in Bull. Inst. Bot. Btzg 5: 13 (1900); Halber f. in op. cit. 195 (1900); Boerlage, Handl. 2: 93 (1907);Hunte r in J.Str .Br .R .As . Soc. 53:8 0 (1909);Rendl e inJ.Bot .6 3 suppl. 67 (1925);Burkil l & Hend. in Gard. Bull. Str. Settl. 3: 396 (1925); Pitard in Fl. Indo-Chine 1129 (1933); Sealy in Kew. Bull. 1956 348 (1956); Gamble in Fl. Madras 568 (1957);Desc h inMan .Mal.Timb . 39,t . 6 (1957);Whitmor e inTre e H. Mal.2:1 5(1973) .-Type :India ,Bengal ,sin .loc ,Hunter July (holotypeBM) . H. corymbosa var. genuina Hall f. op.cit . 195. Tabemaemontana salcifoliaWall. in Bot. Reg, 15: sub t. 1273 (1829); A. De Candolle, Prodr. 8: 376 (1844).?: parviflora Heyne ex Wall. Cat. 1580 & ex Hook, f., Fl. Brit. Ind. 3: 637 (1882), in syn., non Poiret (1817), nor Roxburg (1832), nor Decaisne (1834), nor Bojer (1837), nor Heyne exWallic h Cat. 4453 A. H. corymbosa var. salcifolia (Wall.) Hall f. in op. cit. 195; H. zeylanica var. salicifolia (Wall.)Picho ni nop .cit . 111.-Type :India , sin.loc ,Heyne inWallich 1580a (holotype K-W; isotype BM). Hunteria lanceolataWaü. ex A.DC. op. cit. 350.-Type : Burma,Tavoy, Gomez in Wallich161 1(holotyp e G-DC, isotypeK-W) .Gynopogon lanceolatum (Wall,e x A.DC.) Kurz, For. Fl. Brit. Burma TL: 177 (1877). Hunteria roxburghianaWight, Ic. PI. Ind. Or. 4, 2: 2, t. 1294 (1850); Thwaites, Enum.PI .Zeyl . 192(i860) ,H. corymbosa var. roxburghiana (Wight)Trime n ex J.Gamble ,Fl .Pres .Madra sH :80 8(1923) .-Type :India ,Tami lNadu : Courtallum (=Kuttalam) , Wight 1874(lectotyp eG ,designate d here;isolectotype s GH,L ,M , P,S,W). Hunteria qfricanaK.Schum. inEnglerPflanzenw. Ost-Afr.C: 317(1895).//imteri a zeylanica var.africana (K.Schum. ) Pichon inop .cit . 112,syn .nov .- Syntypes: Tanzania,T3 , Pangani, Stuhlmann 77 (B f); Bagamoyo Stuhlmann 206 (B f). Neotype:Tanzania ,T3,TangaDistr. , Sawa,Faulkner 1626(neotyp eK ,designate d here; isoneotypes B,BR , FT,P) . Hunteria legocii Livera,Ann . Roy. Bot. Gard. Peradeniya 10: 140 (1926). -Type: Sri-Lanka,Paradeniya ,Thwaite s C.P.251 8(holotyp ePDA ,no tseen ;isotype s G, K,P,W).

WageningenAgricultural UniversityPapers 96-1 (1996) 123 Figure34 .Hunteria zeylanica. 1,habi t( x2/3) ; 2,sepa linsid e( x30) ;3 ,opene dcoroll a( x4) ;4 ,pisti l( x 6);5 ,opene d ovary (x 30);6 ,fruitin g branch (x 2/3). 1 from Maxwell 85-675(L) ,Thailand :2- 5 from MycologistBo t GardenPeradeniya ,Sr iLank a2 1 Nov.192 1 (A); 6fro mMatuo lF.D .22004 ,Selangor ,Malaysi a(FHO) .

124 WageningenAgricultural UniversityPapers 96-1 (1996) 30 40 50

10 10

20 20

30 40 50 Map 12.Hunteria zeylanica, African part of area of distribution. Tree or shrub 1-15(-40? ) m high, with colourless to milky latex in all parts.Trun k 2-30 cm in diameter; bark pale to dark grey to dark brownish-yellow, smooth or rough, shallowly fissured; inner bark cream, yellow to bright orange; wood very hard,dense ,whitis ht oyellow .Branche sbrownish-grey ,lenticellate ;branchlet sthin . Leaves: petiole 6-17 mm long;blad e coriaceous, elliptic or oblong to obovate,1.4 - 5.7 x as long as wide, 2-20.7 x 0.9-6.7 cm,rounde d to acuminate at the apex, acu­ men 4-17 x 2-3 mm, cuneate to acute to rounded at the base, glossy and very dark green above, dull and pale green beneath; midrib prominent beneath; secondary veinsstraigh tt over y slightlycurved ,i n 12-30pairs ,formin g asubmargina lvei nan d an angle of 50-770 with the midrib, 3-15m m apart at the central part of the leaf. Inflorescence a compound dichasial cyme, lax, terminal, rarely also axillary and sometimesovertoppe db ya late raxillar yvegetativ ebranch ,3.2-7. 5 x 2.5-8.5 cm,3 - 4 xbranched, IO-65(-Q6)flowered;- peduncle ,(5-)io-4 0 mmlong ;primar ybranche s 6-20 mm long; secondary branches 4-10 mm long, tertiary branches 5-7 mm; pedicels 3.8-10m m long. Bracts ovate to triangular, 0.7-1.3 x 0.5-1mm . Flowers fragrant. Sepals green,thicke rtoward sth ebas ean d atth emiddle ,erect ,wit ha littl e resinwithin ,membranaceou s atth eedg ean dth eapex ,ovat et otriangula ro relliptic , obtuse to shortly acuminate at the apex, 1.2-3x as long as wide, 1.0-2.5x 0.5-

WageningenAgricultural UniversityPapers 96-1 (1996) 125 i .4 mm, withrow so fcolleter sinsid e up to0. 7o fthei rlength ;colleter sver ysmal l 0.2-0.4x 0. 1 mm. Corollawhit e to paleyellow ,10-17. 3 mmlon gi n the mature bud andformin ga comparativel ywid eovoi dhea d0.35-0.5 3m mo fth ebu dlength ,4- 7 x 1-2.5 nun,wit ha nobtus eapex ,wit ha bel to fpubescenc e 1-3 mmwid einsid ejus t belowth einsertio no fth estamens ,sometime swit ha fe whair saroun dth estamen s andu pt oth ethroat ;tub ecreamy-green ,3.7-7. 7x a slon ga sth ecalyx ,1.1-2. 2x a s long asth elobes , 6-10m m long, almost cylindrical, 0.8-1.8m m wide above the base,narrowe d belowth einsertio no f the stamenst o0.7-1. 5m mwide ,agai nwid ­ enedaroun dth eanther st o 1.2-2.0m mwide ;lobe spur ewhite ,ovate ,0.5-0. 9x a s long as the tube,1.4-3. 2x a s long as wide,3.7-8. 8x 1.8-3. 5 mm,obtus e at the apex, spreading or recurvedlater . Stamenswit hape x0.2-0. 8m m belowth emout ho f the corollatube ,inserte d0.67-0.8 3 ofth e lengtho fth e corollatube ,a t4.5- 8m mfro m thebase ;filaments 0.3-0. 9m mlong ;anther s narrowly ovatet ooblong ,3.0- 8x a s longa swide , (I-)I.3-I.5 x0.2-0. 5 mra,obtus ea tth eapex ,subcordat ea tth ebase . Pistil5.5-9. 5m m long;ovar yoblong ,0.9-1. 8x 0.6- 1x 0.4-0. 6mm , graduallynar ­ rowedint oth e style,wit ha disk-lik ethickenin gunitin gth e carpelsa tth ebase ,abou t 0.3-0.4o f the length,0.2-0. 4m mhigh ,o ftw oseparat ecarpels ;styl e3.5-7. 2x 0.1 - 0.3mm ;pisti l headcompose do fa subglobos e stigmaticbasa lpart ,0.3-0. 7x 0.25 x 0.5 mman da conica l stigmoid apex0.3-0. 8x 0. 2mm .Ovule s 2a tth eextrem e base in each carpel.Fruits yellow toorange , smooth, of two separate mericarps; mericarpsobovoi dt o globose,13-3 0x 10-1 5x 7_9ram, rounde d at the apex,some ­ times stipitatea tth ebase ;stip e4- 8m mlong ,2-seeded ;wal l about 2-4m mthick ; pulpsmooth .Seeds oblon go rellipsoid ,8.5-1 5x 6-1 0x 4- 6mm ,test asmooth ,or ­ ange.Embry o 3.8 mm long;cotyledon sovate , 2 xa s longa s wide,2x 1 mm,obtus e atth eapex ,rounde da tth ebase ;radicl e0. 9x a slon ga scotyledons ,1. 8 x0. 4mm .

DISTRIBUTION. Somali, Kenya,Tanzania , and Mozambique, India, Sri-Lanka, Burma,Thailand ,Cambodia ,Laos ,Vietnam ,S .Chin a(Hainan) ,Malaysia ,Indonesia . ECOLOGY. Drycoasta l forest, mixed evergreen forest gallery forest, or thick woodedbushlan do nsand ysoi lo rcora lroc kalon gth ecoas tan do nsand ygravell y soilso n rockyriver banks ,inland .Alt . 0-335ra. Flowerin gi nJanuar y and February inMozambique ,i nSeptembe rt o Januaryi nKeny aan dfruitin g inMarch-Apri lan d flowering allth eyea rroun d in Tanzania.

USES.Woo duse dt omak ecomb s (Kimberly-Medley 231, K7,Kenya , 1987), as boxwood(Bally 16718, K7, Kenya) and to makebow sfo rhuntin g(Burstyn 75/83A , K7, Kenya, 1975).Woo dyellow ,dens ean dno tattacke db y termites(Poilane 6774 , P, 1923,Vietnam) ,use dt omak eax ehandle s (Poilane 6721,P , 1923,Vietnam) .

Geographical selectiono fth eapproximatel y 200specimen sexamined : SOMALIA. Bibahal,Paoli 850 (FT); ShootoForest , Syrmot 1949 (EA,K) ;Uabi ,Scassellati 5 6 (FT); Jess,Hemming & Deskmukhs.a., (EA); Giuba,Paoli 47 8 (FT); 73k mN of Bardera,Tardelli 43 4 (WAG); Bardera,Paoli 81 5 (FT); 25mile sN o fGelib ,Mooney 766 6(EA ,FT ,K , S). KENYA. K I: Nothem FrontierDistr. , Chebele, Bally 6068 (K).Mer u Distr., MeruNationa lPark , MasandukuCamp ,Hamilton 93 (EA); RojweruR. , Gillett 18905(BR ,EA ,P) ; Banko fUr aR. ,Ament & Magogo 388 (EA, K).K1/7 : Nothem Frontier/TanaTiver :Garissa ,Greenway 886 0 (EA,K ,NY) ;2 km

126 WageningenAgricultural University Papers 96-1 (1996) ESEof Mararani,Kuchar 1347 8(EA) .K4 : Kitui Distr.,Ukassi ,Edwards 101 (EA); EndauHill ,Gatheri etal .79/6 7 (EA,K) .K 7 : LamuDistr. ,Mangai-Kiung a Rd.,Gilbert & Kuchar586 0(EA ,K ,UPS) ; Witu, RawlinsJan . 1957 (EA, K). TanaRive rDistr. ,Tan aR. , Battiscombe K 226 (EA);Mchelel o Forest, Luke etal .TP R3 1 (EA, K). Kilifi Distr.,nea rAd u (= Hadu),Dale 1077 (EA, FHO);Marafa ,Polhill & Paulo 818 (B,BR , EA,FT ,K , P,UPS) ;6. 4k mN of Malindi,Polhill & Paulo75 5 (B,BR , EA, FT,K ,P , UPS); ArabukoSokok e Forest, Graham197 6(EA ,FHO ,K ,NY) ;9 kmE o f Bamba,Reitsma 14 4 (EA, WAG); Vipingo,Bally 471 2 (EA, K).TaitaDistr. ,Bura ,Bally 2042(EA , K). KwaleDistr. , KayaWaa,Robertson 3506 (E,K ,MO , WAG); ShimbaLodge , Omino 64 (EA, WAG), 89 (EA, WAG); Diani Forest,Robertson & Coverdale 3668 (EA, K,MO , WAG); Likoni,Jex-Blake 4641 (EA, K);Dian i Beach,Hiepko 266 3 (B, EA);Mwel eMdog oForest ,Spjut 274 5(BR ,EA ,K , UC); 35k mS ofMombasa ,Leeuwenberg 10800(EA , WAG);Marenj e Forest, Faden& Faden 74/321 (EA, K, MO, WAG); Vanga, Graham220 5 (EA, FHO, K,P ,NY) ;Galan aRanch ,Bally 16718 (EA, K). K4/7 : Kitui/Kilifi Distr., LaU Hüls,Adamson 34(EA) . TANZANIA. T 3: TangaDistr. , 13k mS o f Moa,Drummond & Hemsley 3655 (B,BR , K,S) ;8 k mS E of Ngomeni, Drummond& Hemsley 356 3 (B, BR, EA, FT,K , S);Sawa ,Faulkner 1626 (B, BR, FT, K, P); Kumbatoni,Tanne r246 3(BR , K).Lushot oDistr. ,Sangaraw e (= Sangerawe),Zimmermann 23.3.1917 (K). PanganiDistr. ,Msumbugw eFores tReserve ,Hawthorne 81 2(K) ; MkaramoParish ,Tanner 336 6(BR , K, NY, UC,WAG) ; Mkwapa (= Mkwaja), Tanner372 2 (BR, K, NY, UC).T 6: Bagamoyo Distr., Bagamoyo, Sacleux48 2 (P);Kikok a Forest Reserve, Semsei 3723 (BR, EA, K);nea rKaw e (= Kawi). UzaramoDistr. ,Harris 2828 (EA, K, WAG); Dar-es-Salaam, Kirk March 1968 (K); Vikindu ForestRe ­ serve,Paulo 13 3 (EA, K). Rufiji Distr., Uranzi, Greenway507 7 (EA, K). P:Msit u Mkuu,Beentje 438 1 (WAG);Pemba , Vaughan 2078 (EA, FHO), 2729 (EA).Z :Zanzibar , Vaughan 885 (EA,FHO) . MOZAMBIQUE. Zambezia, Mopeia, Bowbrick JC10 (LISC, SRGH). Sofala: Maringuè, Bond 9837 (LISC, SRGH);Vill aFonte s (= Caia), Greenwood 12(LISC ,SRGH) ;2 5 kmfro mLacerdónia ,Muller & Pope 1926 (K, LISC, PRE, SRGH); 5 km E of Inhamitanga, Muller & Pope 1865 (K, LISC, SRGH); Gorongasa National Park, Macedo 1258 (WAG); Sangarassa Forest,Tmley 236 9 (HBG, K, M, MO,P , SRGH,WAG) .Manica :Chimoi , Simâo 273/48 (LISC).

Notes.Picho n (1953)divided// , zeylanicaint o 3 varieties asfollows : var.zeylanica, leaves over2. 5c mwid ean dwit ha wel ldefine d acumen, var.salicifolia (Wall exA .DC. ) Pichon,leave s rarely moretha n 2.3c mwid ean d without adistinc t acumen. var. africana (K. Schum.)Pichon ,leave smostl yove r2. 5cm ,bu twithou ta distinc t acumen.

In myopinion , this species cannot be divided into varieties on the basis of the width of the leaves which proved tob e very variable over the whole distribution range, e.g. TardelliAßA, Somalia(0.9- 2c mwide) ;Reitsma 144,Keny a(2-4. 2cm) ; Drummond& Hemsley 3563,Tanzania(i.4-4. i cm);Forbes 2722 ,Sumatr a(0.9-3. 2 cm); Poilane5562 ,N.Vietna m(2.1-3. 7cm ) and Van Steenis 870, AnambasIsland s (5.4-6.7 cm). In Africa the leaves are mostly rounded (Gardner 2205, Kenya; Greenway 8860,Kenya ;Tanner 3366,Tanzania ) toobtus e atth e apex (Robertson 3506Kenya) ,althoug hthe y may sometimes bebluntl y acutet oacuminat e witha blunt acumen (Beentje 4381).Th e leaf isver y variable in size, shape and apexi n Asia.Th elarges tleave sar efoun d inMalaysi a(u pt o20. 7x 5 cm)an dth esmalles t inSr iLank a(5. 5x 1.6 cm). Ina majorit y of theAsia n specimens,th elea f apexi s verydistinc tbu t therear eals oman yexample so f specimenswithou t adistinc tacu ­ men,especiall y from Hainan and Vietnam (How 71618,Hainan ;Poilane 3002an d 6774,Vietnam) .Th e leaf acumen of all the specimens formerly placed undervar . salicifoliai sals over yvariabl ean dma yb ebluntl yacuminat e(Wight 543 , India)t o

WageningenAgricultural UniversityPapers 96-1 (1996) 127 distinctly acuminate (Alston 262, SriLanka) .I havetherefor e notmaintaine d the varietiespropose db yPicho n(1953 )i nth ecurren ttaxonomi etreatment . Inherb .Simäo 273/48(LISC )th eovar yexist so f 3separat ecarpels .

2.6 PlCRALIMA PIERRE

PicralimaPierr ei nBull .Soc .Linn .Pari s2,127 8 (1896);Stap f inH .Trop . Afr.4 , 1:9 6(1902) ;Picho n inMém .Mus .Na tHist . Nat. 24: 158(1948 ) &i n Bol.Soc . Brot,sé r2 ,27 :8 1(1953) ;H.Hube ri nFl .W.Trop .Afr . ed.2 ,2 :6 2(1963) .-Typ e species:Picralima klaineana (=P. nitida (Stapf)Th .& H.Dur.) .

Tree or shrub with white latex in all parts;branchlet s glabrous.Leave s opposite, petiolate,thos eo fa pai requa lo rsubequal ;petiol eglabrous ;blad eellipti ct ooblong , entirean dglabrou so nbot hsides .Inflorescence a compound ,umbellat ecyme ,termi ­ nalo rsometime saxillary ,pedunculate ;peduncl eglabrous .Flowers fragran t ornot , actinomorphic exceptfo r thesubequa lsepals .Sepals pal egreen ,imbricat eeve na t anthesis,connat ea tth eextrem ebase ,glabrou soutside ,no tciliat ean dwit h2- 4row s ofcolleter sa tth eextrem ebase .Corolla whitet oyellow ,tub eofte n greenish,wit h abel t of pubescenceu pt o7. 3 mmwid einsid ejus t belowth einsertio n of thesta ­ mens;tub ealmos tcylindrical ,thic kan dfleshy ,narrowe dbelo wth einsertio no fth e stamens,glabrou soutside ;lobe soverlappin gt oth elef ti nbud ,no tciliate ,glabrous , entire,spreadin gan drecurve dlater .Stamens included ,inserte dabov eth emiddl eo f the corolla tube; filaments short, filiform, glabrous; anthers ovate, cordate at the base,glabrous ,o ftw oparalle lcell slongitudinall ydehiscent .Pistil glabrous ;ovar y composedo ftw oseparat ecarpels ,unite da tth eextrem ebas eb ya disk-lik ethicken ­ ing;pisti l headcompose d of a stigmaticoblon gbasa lpar t anda filiform stigmoi d apexu pt o1. 5mm .long . Ovulesnumerou si neac hcarpel .Fruits yello wt oorange , smooth, composed of two separate divaricate mericarps; mericarps obovoid to ellipsoid,wit ha rounde dapex ,u pt o80-seeded ; wallver ythic kan dfibrous. Seeds brown, variously shaped, somewhat angular smooth; testa hard brown. Embryo straight,spathulate ,surrounde db ythic krathe rhar dan dstarch yendosper mleavin g ahol earoun dth eradicl eapex ;cotyledon sthick ;radicl ealmos tcylindrical , flattened andnarro w atth eapex ,widene dtoward sth ebase .

2.6.1 Picralimanitid a(Stapf )Th .& H .Dur . inBull .Jard .Bot .Etat .Brux .2:33 8(1910) ;Picho ni nBol .Soc .Brot . sér. 2,27: 82, mapA p . 85(1953 )& i nMém . Mus.Natn .Hist .Nat .24 : 158(1948) ;H .Hube ri n Fl.W.Trop . Afr. ed. 2, 2: 62 (1963). -Type: Cameroun, Ambas Bay,Mann 71 0 (holotypeK ;isotype s GH, P). Fig. 35,p . 129;ma p13 ,p .13 0

128 WageningenAgricultural UniversityPapers 96-1 (1996) Figure35 .Picralima nitida. 1,habi t( x2/3) ; 2,branchle t( x2/3) ;3 flower, bu d( x 1);4 flower, ( x 1);5-6 , flower above( x1) ;7 ,coroll abeneat h( x 1);8 ,sepa linsid e( x2) ;9 ,opene dcoroll a( x2) ;10 ,pisti l( x4.6) ; 11-13,stamen s( x6.4) .1- 2 fromW.d eWild ee tal . 1952;3-1 3 fromReitsm a899 .

WageningenAgricultural UniversityPapers 96-1 (1996) 129 30

10 3D Map 13.Picralima nitida.

Basionym:

Tabernaemontana nitida Stapf in Kew Bull. 1894: 22 (1894); De Wildeman & Durand in Ann. Mus. Congo Bot. séï. 2, vol. 1, 1: 39 (1899) & sér. 3, vol. 1,2 : 153 (1901).

Heterotypic synonym:

Picralima klaineana Pierre in Bull. Soc. Linn. Paris, 2: 1279 (1896); Stapf in Hooker,Ic .PI .28 :t .2745-274 6 (1902).-Type : Gabon,Libreville ,Klaine 299o f 11.9.1896 (lectotype P,designate d here).

Treeo rshru b4-3 5m hig h withver ydens ecrown .Trun k 5-60 cmi ndiameter , cylin­ drical; bark hard, brittle, pale to dark greyish black to brown, smooth to slightly rought ofinely striate ,sometime slenticellate ,ver ybitter ;inne rbar k6-1 2m mthick , brownt oyello w with orange-brown spots,granular , fibrous; wood very hard,resis ­ tant, whitish-yellow to pale brown or orange. Branches dark green, smooth; branchlets,smooth ,dar kgree nt odar kbrownis hblack .Leaves opposite;petiol e 10- 22m m long; blade thickly papery to thinly leathery, papery to coriaceous when dried, ovate, broadly to narrowly elliptic or oblong, 2.1-4.3x as long as wide, (5- )io.8-26.6 x 1.9-13cm ,abruptly acuminat ea tth eapex ,acume nacute ,6-1 4 x2 mm, cuneate, acuteo rrarel y rounded to obtuse atth ebase ,gloss y anddar kgree n above, dull and paler beneath; midrib impressed above, vividly shiny and pale green be­ neath; secondary veins more or less straight, curved towards the margin, in 14-23

130 WageningenAgricultural UniversityPapers 96-1 (1996) pairs,formin g asubmargina lvei nan da nangl eo f70-72 0wit hth emidrib ,8-1 6m m aparta tth ecentra lpar to fth eleaf ;tertiar yvein sobscur ebelow .Inflorescence 6-1 0 x7.5- 9cm ,12-36-flowered , 1 xbranched ;peduncl emediu mt opal egreen ,2-3 7m m long, with 3 primary branches each bearing 6-12flowers; branche s 3-15mm ; pedicelsmediu mt opal egreen ,2-2 0m mlong .Bract sver ysmall .Sepals vividl ypal e tomediu mgreen ,fleshy ,coriaceou swhe ndried ,fre eo rconnat ea tth ebas efo r0.08 - o.1 8o fthei rlength ,0.5- 1 mmthic ka tth ebase ,erect ,membranaceou sfo rabou t0. 3 mmo fth eextrem eedge , broadly ovatet osuborbicular , (O.9-)I.I-I.6 xa slon ga s wide,(3.5- )5-5-7- 5x 3.5-5mm ,obtus et orounde d atth eapex ,an dwit h2- 4row s ofcolleter sinsid ea tth eextrem ebase ,colleter s small0.7- 1x 0.1-0. 2mm . Corolla whitet olime-yellow ,coriaceou s when dried, 25.3-47m ml° ng m maturebu dan d forming acomparativel ylarg eovoi dhea d0.47-0. 7o fth ebu dlength , 14.3-33 x 3-7 mm,wit ha nobtus eapex ,glabrou soutside ,no tciliat ea tlobes ,wit ha bel to fpubes ­ cence3-7. 3m mwid einside ,jus tbelo wth einsertio no fth estamens ,sometime swit h a few hairsaroun dth estamens ;tub epal egreen fleshy,, coriaceous ,2-3.5(-6 )x a s longa sth ecalyx ,0.6-1. 2x a slon ga sth elobes , 14.5-21 mmlong ,almos tcylindri ­ cal,3- 6m mwid eabov eth ebase ,narrowe dbelo wth einsertio no fth estamen st o2 - 4.5m mwide ,agai nwidene daroun dth eanther st o3- 5m mwide ;lobe spal egree n toyello wi nbud ,cream ya tanthesis ,ovate ,no ttwiste di nbud ,0.8-1.5 7x a slon ga s thetube ,1.9- 3 xa slon ga swide , 13.5-29.8 x5.5-1 0mm ,obtus ea tth eapex ,entire , spreadingo rerect .Stamens wit hape x1.2- 4m mbelo wth emout ho fth ecoroll atube , inserted 0.58-0.71o f the length of the corolla tube, at9.5-1 5m m from thebase ; filaments 0.7-1.4x 0.1-0. 2m mlong ;anther sovate ,3.4-6. 4x a slon ga swide , 2.8- 3.7 xo.5-1mm ,obtus eo racuminat ea tth eapex ,acume n0. 3mm ,sterile .Pistil 9.7 - 12.5m mlong ;ovar y ovoid, 2-3x 1-2.5 x 0.5-1.8mm ,wit ha disk-lik ethickenin g 0.2-0.3o fth elength ,0.5-0. 6m mhigh ; style5-7. 3x 0.3-0. 4mm ;pisti l headwit h apex 2m mbelo w stamens, 1.6-3.1 x0.5- 1mm ,stigmoi d apex0.6-1. 5x 0.1-0.15 mm.Ovule s 120-150i neac hcarpel .Fruits: mericarps irregularlypyrifor m or ob- ovoidt oellipsoid ,laterall ycompressed ,wit ha nadaxia llin eo fdehiscence ,11-2 0 x 8-15x13-14cm ,rounde da tth eapex ;many-seeded ;wal lver yhard ,whit et ocream y on section, 4-8m m thick, inside with a layer of large fibres like inth ecoconut . Seedsembedde d in a soft white or orange pulp orange or brown, smooth,flat, obliquelyovate ,obovat et ooblong ,flattene d andmor eo rles sangular ,24-4 4x 15- 27x 5- 8mm ,test a smooth,coriaceous .Embry ostraight , 16.5-22m mlong ,endo ­ spermexcessivel ybitter ;cotyledon sovat et oobovat et ooblong ,thic kan dleafy ,1.4 - 3.3x a slon ga swid e 10.5-13x 4- 8mm ,obtus et orounde da tth eapex ,subcordat e torounde da tth ebase ;radicl e0.6-0. 8x a slon ga scotyledons ,6.5-9. 5x 1.5- 2 mm.

DISTRIBUTION.Ivor yCoast ,Ghana ,Nigeria ,Cameroun ,Gabon ,Cong oan dZair e andUganda . ECOLOGY. Understore ytre ei nrai nfores to rmatur eol dsecondar yforest ,remnan t forest onplateau xi nZaire ,an dsemi-deciduou s forests byriver bank so rroadsid e growing on sandy clay soil. Alt. 20-900 m.Flowerin g andfruitin g allth e year round.

WageningenAgricultural University Papers 96-1 (1996) 131 USES.Al lpart so fth eplan tar ebitte r(Burkil l 1985).Th ewoo di spal eyellow , fine grained, hard and elasticfinishes, an d polishes well (Dalziel 1937; Irvine 1961); usedt omak epaddle s (Flamigni7116 ,BR , 1944,Zaire ;Dewèvre 847,BR ,Zaire) ; bows(De Wilde 393,BR ,MO ,WAG , Zaire;Bates 1421 ,MO ,Z , 1919,Cameroun ; Harris& Fay504 , MO, 1988, Central African Republic); shuttles for weaving clothes,doll san dcomb s(Vigne 864 ,K ,1922 ,Ghana ;Dalzie l 1937);walkin gstick s inGhan a(Dalzie l 1937;Irvin e 1961);pestl ean dMorta r(Gille 100,BR ,C ,MO , P, WAG,1938 ,Zaire) ;i nGabo nth ewoo di suse dfo rcarvings ,paddles ,incens ehold ­ ers (Walker & Sillans 1965); spade handles (N. WTnomas2061 , 2101, K, 1912, Nigeria;Dalzie l 1937);arrow san dspoon s (Benoit235 ,P , 1956, Cameroun). Immaturefruit s poundedan dthrow nint owate rt oac ta sa fish poiso n(Michelson 231,BR ,Zaire ;Dewèvre 945 ,BR ,Zaire ;Irvin e 1961,Ghana) .Pul pwhic hseparate s thegrai n is used in the composition of arrow poison (Dewèvre 1113,BR ,Zaire) . Seedsar eintensel ybitte r(Dalzie l 1937)an dar eextensivel y usedb yth enative si n place of Quinine (Oppenheimer, son& Co.s.n. , K);crushe d and eaten for chest complaints,pneumoni aan dacut estomac hach ewithou tbein gregarde da sa purga ­ tive(Dalzie l 1937;Irvin e 1961);decoctio n of seedsi suse d asa nenem ai nGhan a (Dalziel 1937; Irvine 1961). Fruit used to cure cough (Benoit 235, P, 1956, Cameroun);eate nb yelephant s(Harris & Fay 504 ,MO , 1988,Centra lAfrica nRe ­ public);use dt opoiso narrow sfo rhuntin gMonkey s(Tisseront 509 ,P ,1952 ,Centra l African Republic). Smalldipper s or spoons made from the hard shell of the fruit (Dalziel 1937);th efrui t contentsar eremove d andth eshel filledl wit h palmwin e whichi sdrun kwhe ni tha sabsorbe dth ebitte rprincipl e(Irvin e 1961).I nGabo nth e Pahouinche wa littl eo fth efrui t andbar kt oalla yhunge rwhil eo nlon gmarche si n thebus h (Raymond-Hamet 1951).I n Gabon theseed s arerecognise d astoxi can d useappear st ob erestricte dfo r externaltreatmen tfo r ancesses(Aubrevill e 1959). Thebitte rbar ki sboile d ina po twit htw oteaspoon s of sugaran dth edecoctio n isdrun kagains tfood/drin kpoisonin g(A. M. Louis 1844,WAG ,1985 ,Gabon) ;use d against stomach ache (J.M. Reitsma et al. 1866,NY ,WAG , 1986, Gabon; Benoit 232, P, 1956,Cameroun) ;chewe d by theBunyor o as avermifug e (Dawe 707,K , Uganda; Eggeling & Dale 1952); used as a febrifuge (Howes 1157,FHO , 1926, Ghana; Dalziel 1937, Aubreville 1959); used in Gabon against venereal disease (Walker 1953;Walke r& Sillan s 1961);bar kdecoctio ntake ni nCong oa sa nanthel ­ mintican dpurgative ,t otrea thernia ,wit hothe rdru gplant st oreliev eblennorrhoe a (Bouquet 1969),i nIvor yCoas tagains tjaundic ean dyello wfeve r(Kerhar o& Bou ­ quet 1950)an dals oi nNigeri a (Ainslie 1937). Theroot san dbar kar euse dt oeas eches tcomplaint san dcomba tfeve rwit hmil d jaundice,popularl ycalle d 'yellowfever ' (Dalziel 1937);th eroot sar euse da sa ver ­ mifuge in Gabon (Walker 1953;Walke r & Sillans 1961) and against fevers and pneumonia(Ainsli e 1937,Aubrevill e 1959). Leaf-sapi sdrippe dint oear sfo rotiti si nCong o(Bouque t 1969).A coole ddecoc ­ tion of leaves and bark is used against ascaris (Floret et al. 1633,WAG , 1983, Gabon). Seedshav ebee nfoun d tocontai n about 10alkaloids ,seve no f whichhav ebee n crystallized;o ntria lalkaloid sar eineffectiv e onbir dmalaria ;bar kan dleave syiel d

132 WageningenAgricultural University Papers 96-1 (1996) onlyamorphou salkaloid s(Henr y& Sharp1927,1932) .Akuammin ei s theprincipa l alkaloidfro m the seeds,i sa powerfu l sympathicostenic andha s aloca lanaestheti c actionequa lt o thato fcocaine .Akuammadin ei s hypotensive,an d weakerbu tlonge r lastingi neffec t thanYohimbin e(Burkil l 1985).

Geographical selection of the approximately 200specimen sexamined : IVORY COAST. Near Bingerville, Chevalier 15600 (P); Anoumabo, Chevalier 22391B (B, LG, P, WAG),2244 0 (P). GHANA.Western :nearTarkwa , Vigne20 8 (FHO).Ashanti :Obuasi ,Seward 65 1 (K);Adiembra , Vigne 864 (K);Kumasi , Vigne 3956 (K,P) . NIGERIA. Ogun: Agege, E.W.Forster 22 0 (K). Oyo: Gambari, Van Eijnatten 1526 (WAG);Ibadan , Onyeachusim FHI48005 (K,MO) ;Shash aF.R. , Osseyemeh 3341 (BR,MO , S);ibid. ,Ros s 7 (BR,MO , S). Ondo:Akw eF .R. , Van Meer 792 (WAG); Okeluse F.R. ,Onochie FH I3337 0 (K,MO) .Bendel :Oshi , Kennedy205 6 (FHO, K); 15k mS W of Ekenwan,Leeuwenberg 11348 (WAG);Okom u F.R , Brenan & Richards 8910 (K); Sapoba, Kennedy 212 (FHO); Ogwasbi, N.W. Thomas 206 1 (K); Onicha Olona, N. W.Thomas 194 1 (K). Benue:Jones 98 2 (FHO). Cross River: sin. loc.,Talbot 21 9 (BM, K);Eket ,Talbot 3146 (BM, K,Z) ;Eke tDistrict ,Talbot 325 6 (BM, K,Z) , s.n. (BM, K); Oban,Talbot 169 0 (B, BM,Z) ; ObanDistrict ,Talbot s.n . (BM, K);Iyamoyong ,Binuyo FH I4129 7(BR ,FHO ,K ,WAG) ; km11 4Calabar - Mamfe Road,Latilo FHI5399 7 (K, P, WAG). CAMEROUN. Sud-Ouest:Victori a (=Limbe) ,Maitland 28 4(B , K); AmbasBay ,Mann 710(GH ,K , P, type); Ikilliwindi, 20 km N of Kumba, Etuge & D.W.Thomas 499 (MO, NY, WAG); near Kumba, D. W.Thomas 4684 (BR, MO, P, WAG); NE corner of Korup Nat Park,D. W.Thomas 3313 (MO, WAG). Ouest:Wu mDistrict , Ujor FHI 29271 (K). Littoral: near Maleke, Benoist 326 (P);;nea rNkongsamba , Benoist 256 (P). Centre-Sud: Nkolbewa, nearKribi ,Benoist 232 (P);Bipindi , Zenker302 8 (B, BM,BP , BR, E, G, HBG, K, L, MO, P, S,W,WAG,WU), n 283 (B, BR, C, F,G , GH, MO, NY, P, U, UC, US, WAG);Melen , km 27 on Kribi-Ebolowa Road,Bos 6373 (WAG);Mvimi , 35 km Eo f Campo, Kaji 50 (P); Ebolowa, W. deWilde 195 2 (B,BR , K,MO ,P ,WAG ,Z) ;Koumo u Forest,Benoist 149 (P); between Zoetelean dNkolebang ,Asonganyi 12 8(P ,WAG) ; nearYaoundé ,Letouzey m o (P); Bitye,c a6 0k mEN E of Sangmélima, Bates 1421 (BM, MO, Z); near Nanga Eboko, Mildbraed 8285 (HBG, K). Est: near Ndjangani,4 6 kmN Wo f Bertoua,Breteler 1358(BR , P, WAG); 21 kmS o f Mboy I,Letouzey 506 9(K , P); 19 kmW of Yokadouma, 5 kmW of Mbol, Leeuwenberg617 6 (BR, K,MO , P, WAG); 70 kmE of Djoum,Letouzey 1180 4(BR ,HBG ,K ,P ,WAG) ; 5k mW o f BatekaMalemba ,Vtlliers 65 2(K ,P) ; Nginda, 21 kmN o f Moloundou, Mildbraed 4138 (HBG). CENTRAL AFRICAN REPUBLIC. Sangha: Harris & Fay 504 (MO). Ombella-Mpoko: Boukoko, Tisseront50 9 (K,P) . GABON. Estuaire: near Libreville, Klaine 106 (P), 299 of 11 Sept. 1896 (P, lectotype; possible isolectotypesBR ,K) ,29 9o f9 Aug . 1896(K ,P , paratype).Woleu-Ntem :Oveng ,A. M. Louis211 4(WAG) ; nearAbanga ,Reitsma e tal . 1866(NY ,WAG) ; Otuoma,Dibata 956 (BR); Ovengi,Reitsma e tal .89 9(NY , WAG);Mindoumo , A. M. Louis 1884 (WAG).Moyen-Ogooué : Onangé Lake,Pobéguin 4 5 (P).Nyanga : Dibilila,Le Testu 1223(BM ,BR ,G ,P) ;Tchibanga ,Le Testu 1841(BM ,BR ,E ,K ,P , US, WAG). Ogooué- Ivindo: 10-15km W of Akouba,Floret et al . 1633(P.WAG) ;M'pass aFiel dStation ,Gentry 3351 2(MO) ; Makokou, Hladik 1314, 2098, (P); near Ekobakoba, Reitsma 3132 (NY,WAG) ; Belinga, Halle & Le Thomas64 1 (P); Booué, Vadons.n . (P).Ngounié : SE of Sindara,Leeuwenberg & Persoon1369 5 (WAG); Gnyoungou, Le Testu603 5 (B,BR ,K , P); Mimongo,Le Testu589 8 (B,BR ,HBG , MO, P). Ogooué-Lolo: Poungui,Le Testu 7429 (P). CONGO. Kouilou: Pointe Noire, Farron 4828 (P); Kouilou, Sargos 180 (BR, K, P, US); Region du Kouilou,Moutsamboté, 4-9-1990, (BR).Bouenza : Loubofo, Bouquet67 2 (P); Massia, Bouquet81 9(P) . Le Koumou:Wes t of Sibiti, Farron424 7 (P). Pool: Mayombe, les Saras, Bouquet 1917 (P). Cuvette: Odzala,Dowsett-Lemaire 1833 (B). Sangha: Ouesso,Hanie 7 4(P) . ZAIRE. Bas-Zaire: Borna, Luki, Hombert556 (BR, K);Luki , Maudoux17 1 (BR, MO, P);Kimbuya , Donis 165(BR) ; Kinshasha,Claessens 6 6(BR) . Bandundu: Lac LeopoldI I( =LacTumba) ,Claessens 756 (BR); BoloboWaterfall ,Flamigni 711 6(BR) ; Ganya,Flamigni 1038 3(BR ,WAG) ; Ulongonga,Sapi nDec . 1907 (BR);Ipamu , Vanderyst 11997 (BR). Equateur:Lukolela ,Tofa i5 7 (BR);Eala , Corbisier 103 (BR,

WfcgenmgenAgricultural UniversityPapers 96-1 (1996) 133 K); Eala, Corbisier 720 (BR, C,FHO ,K ,MO ,NY) ;Befale , Evrard317 8 (BR, K); Tokaya,Evrard 785 (BR); Yambata,De Giorgi 1672(BR ,Z) .Kasai-Occidental :Sankuru , Van denBossche & Luja 3 1 (BR); Sankuru, Demeuse 105 (BR, HBG); Bena Longo, Dechamps 171 (B, BR, L, UPS,W , WAG, Z). Haut- Zaire:Mobwasa , Lemaire38 8 (BR);Mobwasa , Vermoesen 287 (BR);Zobia , &>«>* 873 (BR);Itukpwo , G/tóert 425 (BR); Bambesa, G

2.7 PLEIOCARPA BENTH.

Pleiocarpa Benth. in Benth. & Hook. f. Gen. PI. 2: 699, (1876); in Hooker, Ic. PI. 12:71, (1876); K. Schumann inEngle r& Prant l Nat. Pflanzenf. 4,2:134 (1895);D e Wildeman & Durand In Ann. Mus. Congo Bot. sér. 2 vol. 1, 1: 37 (1899) & sér. 3 vol. 1, 2: 150 (1901); Stapf in FI.Trop .Afr . 4, 1:9 7 (1902); Pichon in Mém. Mus. Natn. Hist Nat. 24: 161 (1948) & in Bol. Soc. Brot. sér. 2, 27: 123 (1953); Hutchinson & Dalziel in H. W.Trop . Afr. éd. 1, 2: 37 (1931); H.Huber in Fl.W . Trop. Afr. éd. 2, 2: 63 (1963); Kupicha in FI. Zamb. 7, 2:43 2 (1985). - Lectotype species: P. mutica Benth. (designated by Pichon in Bol. Soc. Brot. sér. 2, 27: 123 (1953)-

Heterotypic synonym:

Carpodinopsis Pichon in Bol. Soc. Brot. sér. 2, 27: 142 (1953). -Type species: C. rostrata (Benth.) Pichon (= P. rostrataBenth.) , designated by Pichon 1953.

Shrubs orsmal ltrees ,rarel y lianescent, withwhit elate x inal lparts ;branchlet s gla­ brous. Leaves opposite or in whorls of 3-5, petiolate, those of a pair equal or subequal; blade mostly elliptic, entire and glabrous on both sides. Inflorescence predominantly axillary, sometimes at the same time terminal, ramiflorous, usually fasciculate; sessile.Flowers usually fragrant, rarely malodorous (in P. pycnantha), actinomorphic exceptfo r thesubequa l sepals.Sepals palegreen ,mostl ythin , rarely coriaceous, free or connate atth e extreme base, membranaceous atth e edge, com­ pletely glabrous onbot h sides,withou t colleters. Corollapur ewhit et oyellow , tube often greenish in bud, with a belt of pubescence up to 10.5 mm wide inside just below the insertion of the stamens; lobes overlapping to the left in bud, always shortertha nth etube ,spreadin g andofte n recurved later.Stamens included, inserted ato r aboveth e middleo f the corolla tube; anthers ovate,obtus et o acuminate at the

134 WageningenAgricultural UniversityPapers 96-1 (1996) apex,cordat ea tth ebase ,glabrous ,o ftw oparalle l cells,longitudinall y dehiscent. Pistilglabrous ;ovar ycompose do f2- 5separat ecarpels ,unite da tth eextrem ebas e by a disk-like thickening; pistil head oblong orellipsoid ; ovules i-4(-6) in each carpel.Fruit yello wt obrigh t orange,smoot ho rrugose ,compose do f2- 5separat e mericarps; mericarps variously shaped, witha rounde d tohook-shape d apex,1-6 - seeded; wall 2-4m mthick , fleshy andsometime s slightlyfibrous. Seed s brown, variouslyshaped , somewhatangular . Embryostraight , spathulate,surrounde db ya thick, rather starchy and hard endosperm, leaving ahol e around theradicl ebase ; cotyledonselliptic ,thi nan dleafy ;radicl ealmos tcylindrical flattened, an dnarrowe r atth eapex ,widene dtoward sth ebase .

Keyt oth especie so fPleiocarpa

1. Leavesternat e oro nth esam ebranc hbot hopposit ean dternate ,occasionall y inwhorl so f4- 5 2 Leavesopposit e 3 2. Mature bud 8-13m mlon g (rarely upt o 15m mlon g inZaire) ;coroll a lobes 1.3-4.8 mmlong ;ovar y2-carpellate ;secondar yvein susuall ymor etha n1 2 pairs;leave si nwhorl so f3- 5i nth ewhol edistributio nrang e 4.P . pycnantha Maturebu d14-26. 6m mlong ;coroll alobe s(5-)6-i 3m mlong ;secondar yvein s in9-1 2pairs ;ovar y(3-) 5carpellate ;leave sternat eonl yi nspecimen s from Liberiaan dIvor yCoas t 3.P . mutica 3. Stamensinserte dc .0. 5o fth elengt ho fth ecoroll atub ea t4.3-5. 5m mfro mth e base;styl e 1.2-1.5m mlon g 2.P . brevistyla Stamensinserte d0.58-0.9 2o fth elengt ho fth ecoroll atub ea t6.7-1 8m mfro m thebase ;styl e5.5-1 6m mlon g 4 4. Stamens with apex 3-8m mbelo w themout h of the corolla tube; secondary veinsoblique ,ver yprominen tbeneath ,i n6-9(-i2 )pairs ;inflorescenc e 1-4- flowered, fruits smootht orugos ewit ha rounde dt obeake d apex 5.P . rostrata Stamenswit hape x0.2-1. 3m mbelo wth emout ho fth ecoroll atube ;secondar y veinsi n9-3 0pairs ;inflorescenc e io-i5(-35)flowered; fruit s smoothwit ha rounded apex 5 5. Secondaryvein si n9-1 2pairs ,8-2 0m ma tth ecentra lpar to fth eleaf ;ovar y 3- 5-carpellat e with 1 ovulepe rcarpe l 3.P . mutica Secondary veins numerous,i n 19-25pairs ,4- 9m ma tth ecentra lpar to fth e leaf;ovar y 2-carpellate with 2 ovulespe rcarpe l 1.P .bicarpellat a

2.7.1 Pleiocarpabicarpellat aStap f inKe wBull . 1894:21(1894) ,partl ya sfo rth electotyp e& i nFI .Trop .Afr . 4,1:9 9 (1902); Pichon in Bol.Soc .Brot . sér.2 , 27: 125, p. 117 1.3, fig. 8-10, (1953); Hutchinson &Dalzie l inFl .W .Trop .Afr .ed . 1,2 :3 8(1931) ;H .Hube r inFl .W . Trop.Afr .ed . 2,2 :6 3(1963) .-Type : Cameroun Mountain, 1200m ,Mann 121 3

WageningenAgricultural UniversityPapers 96-1 (1996) 135 (lectotypeK ,designate db yPicho ni n 1953;isolectotype , P). Fig.36 ,p .137 : map14 ,p . 138 Heterotypic synonym:

Hunteria ambiens K.Schum . in Engler, Bot.Jahrb . 23:22 3(1896) ; Hallier f. in Jahrb. Hamb.Wiss . Anst. 17 Bein 3: 191 (1899).Type : Cameroun, Lolodorf, Staudt9 3(lectotyp eK ,designate d here;isolectotype sA ,E ,G , Z).

Shrubo rsmal ltre e 1.5-8 mhig h with whitelate xi nal lparts .Trun k very slender, smooth;bar k andbranche s very palebrownish-gre y topal e silvery grey, smooth; branchlets thin, quadrangular when young, later terete, very palet o dark brown. Leavesopposite ;petiol e4- 1i(-i4 )m mlong ,glaucou sgreen ;blad epaper yt othinl y leatheryi nfres h leaves,narrowl yellipti ct ooblong ,(i.6-)2.o-3. 8x a slon ga swide , 5.1-16.8x 1.8-7.2cm , acuminatea tth e apex,acume n obtuset oacute , 10-22x 2- 4mm ,acut eo rcuneat ea tth ebas ean ddecurren tint oth epetiole ,midri bprominen t infres h leaves;secondar y veins straight,i n 19-30pairs ,obscur ebeneath , forming anea tsubmargina lvei nan da nangl eo f65-80 0wit hth emidrib ,4- 9m mapar ta tth e approximatecentr eo fth eleaf ,tertiar yvenatio ninconspicuou sbeneat han dmor eo r lessparalle lt oth esecondar y veins. Inflorescence axillary,ramiflorous , sometimes atth esam etim eterminal ,fasciculate , 1.5-2.5X2.5- 3cm ,io(-i5 )flowered, sessile . Pedicels 1-2 mmlong .Bract sver ysmall ,scale-like .Flowers ver yfragrant . Sepals very pale green, free, erect, membranaceous for about 0.2-0.4m mo f their edge, ovate,0.6-2. 1(-3. 1) x a slon ga swide ,1. 1-2. 8x 0.6-1. 7mm ,acut ea tth eape xwher e iti susuall ydamage dan dappearin gdentate .Corolla pur ewhite , 15-18.5m mlon g inmatur ebu dan dformin g acomparativel y largeoblon ghea d0.31-0. 4o fth ebu d length,(5- )6.5- 8 x 1.5-3 mm,wit ha rounde dapex ,wit ha bel to fpubescenc e2.8 - 4.5m mwid einsid ejus tbelo wth einsertio no fstamens ,wit hlonges thair sjus tbelo w theanthers ;tub e5.1-14. 3 xa slon ga sth ecalyx ,1.3-2. 4x a slon ga sth elobes , (9.2- )io-i5 mmlong , almost cylindrical, 0.8-1.8m m wide above thebase , narrowed belowth einsertio n ofth e stamenst o0.4-0. 8m mwide ,agai n widened aroundth e antherst o0.7- 2m mwide ;lobe snarrowl yoblon gt obroadl yovate ,0.4-0. 8x a slon g asth etube ,1.3-3. 4x as longa swide ,5-10. 2x (i.5-)2.i-5. 5mm ,obtuse ,spreadin g and recurved later. Stamens with apex 0.2-1 mmbelo w themout h of thecoroll a tube, inserted 0.68-0.86 of the length of the corolla tube, at7.5-1 2m m fromth e base;filaments 0.3-0. 6m mlong ;anther snarrowl yovat et ooblong ,1.8-5. 5x a slon g aswide , 1-1. 4x 0.2-0.3 5mm .acuminat ea tth eapex ,acume nu pt o0. 3mm ,cordat e atth ebase .Pistil 7.5-12.2m mlong ;ovar yovoi do rsubglobose ,0.7- 1x 0.5-0. 7x 0.3-0.7mm ,abruptl ynarrowe dint oth estyle ,wit ha disk-lik ethickenin gunitin gth e carpelsa tth ebase ,abou t0.25-0. 4o fth elength ,0.2-0. 4nu nhigh ,o ftw oseparat e carpels; style 5.5-10.7 mmlong ; pistil head oblong to ellipsoid, 0.3-1x 0.15- 0.3mm . Ovules two in each carpel. Fruits orange, smooth, of two separate mericarps;mericarp sobovoi do rellipsoi dt oobliquel yellipsoi di nimmatur efruits , 11-18x7-12x8-9 mm,rounde d atth eapex ;2-seeded , wallabou t 1-2 mm. Seeds ellipsoid,oblon go rsubglobose ,9-1 1 X5-7x 3- 4mm .Embry o5-7. 8 mmlong ,coty ­ ledonsellipti ct oovate ,1.75- 2x a slon ga swid e2- 4x 1-2 mm,obtus eo rrounde d

136 WageningenAgricultural UniversityPapers 96-1 (1996) Figure36 .Pleiocarpa bicarpellata. 1,habi t( x2/3) ; 2,branc h( x2/3) ; 3,sepa l( x20) ;4 ,coroll a( x4) ;5 , openedcoroll a( x4) ;6 ,caly xwit hovar y( x 6);7 ,pisti l( x 6).1- 2 from ZenkerII242 ;3- 7fro mZenke r 1660.

WageningenAgricultural UniversityPapers 96-1 (1996) 137 O 10 Map 14.Pleiocarpa bicarpellata.

atth eapex ,rounde d atth ebase ; radicle 0.87-1.5x aslon ga scotyledons , 3-4 x 0.5- 0.8 mm.

DISTRIBUTION. Cameroun, Gabon, Zaire,Angol a and Kenya. ECOLOGY. LOWbush , secondary or primary forest, sometimes in stream ravine. Alt. 300-1200 m. Flowering and fruiting all the year round.

Geographical selection of the approximately 70 specimens examined: CAMEROUN. Sud-Ouest:Mamf eRd. ,aroun dEkon aMombo ,Etuge &D.W. Thomas 41 9 (MO,WAG) ; BarombiLake ,Brenan & Jones 945 8(FHO ,K ,P) ;Baromb iLake ,Preuss 4 4 (K);Kumba ,Staxidt 57 3(G , K,P , S, US,WU ,Z) ,68 3(EA ,G ,K ,P ,Z) ,79 1 (FHO);Bambuk oF .R ,Keay FHI3745 7(K ,P) ;Camerou n ML, Mann 1213 (K, P,type) ; NE slope of MT.Etinde , Wieringa207 0 (WAG). Littoral: near Douala, Fleuryi n Chevalier3334 8 (P);Ede aReserve ,nea rL .Tissongo ,D. W. Thomas 294(K) .Centre-Sud :N of MLElephan t (Rouillon), Bos 6502 (BR, K,M , P,WAG) ;Calvar y ML,Bido u I,Bos 6215 (BR, C, K, P, UPS,WAG) ;Bipindi ,Zenker 165 8(B ,BP ,BR ,E ,G ,K ,L ,M ,NY ,S ,W,WU,WAG ,Z) , 1660(B ,BP ,BR , E, G, HBG, K, L, M, P, MO, S,W ,WAG ,WU , Z), 1728 (B, BP, BR, E, G, HBG, K, L, M, NY, S,W , WAG,WU ,Z) , 2501 B,BP ,BR , E, G, HBG, K,L ,M ,MO , P,W,WAG,WU , Z), 2858 (B, BR, BP,E , G, HBG, K, L, M, P, S,W,WAG,WU , Z), 2893 (BR, BP, E, G, HBG, K, L, M, P, S,WAG ,WU , Z),423 1 (BP, BR, E, G, HBG, K,L ,M , MO, P, S,W, Z),447 0 (B,BR , E, G, K, L, M, P, S,W, Z),463 5 (B,BP , BR,G , HBG, K,L ,M ,MO ,P ,S ,W ,Z) ,488 2 (BR,G ,HBG , K,L ,M ,MO ,P ,S,W , Z);488 9 (B,BP ,BR , G, HBG, K,L ,M , MO,P ,S,W , Z),H 11 7 (B, C,NY ,US , U.WAG), n 242 (B, BR, C, FT,G , GH,MO , NY,P ,US ,WAG) ; Efoulen,Bates 368 (G, K,Z) ;Lolodorf , Staudtg^ (A,E , G, K,P,W ,Z) ;Ndikinimeki , Asonganyi41 4 (P,WAG) ;2 5 kmES Eo f Ebolowa,Letouzey 985 0 A (P);nea rOveng , Breteler 2650 (K, P,WAG) ; c. 5k mS of M'balmayo,De Wilde &De Wtlde-Duyßes 1798B (BR, P,WAG) ; OtotomoFores t Reserve,Farron 7261 (P);Yaoundé ,Zenker & Staudt 61 1 (B,M ,US ,WU) , 753 (K);Meyan g Hill,Dang 614 (P). GABON. Estuaire: Monts de Cristal, Halle & Villiers508 0 (P). Ogooué-Ivindo: near Ekobakoba, Reitsma 2717 (WAG), 3051 (NY.WAG); 10k mSS Wo f Makokou,Leeuwenberg 11451 (WAG);Boou é

138 WageningenAgricultural UniversityPapers 96-1 (1996) Forest,Florence 179 8 (P). WoleuNtem : Eo f Médouneu, Halle& Villiers 5008(P) . ZAIRE. Kivu: Kasongo, Herman 2-7-1955 (Z). Shaba: Galerie Luba, Herman 2179, 2248 (BR); Kaniama, Gathy 1908(BR) . ANGOLA. Cabinda:Buc o Zau,Gossweiler 6549 (K), 6687 (BM),664 8 (COI), 8024 (BM). KENYA. K7,TaitaTavetaDistr . Chawia forest, Omino 82 (EA), 157,158 (EA, WAG).

Notes.Whe n Stapf (1894)describe d P. bicarpellata, he distinguished it onth e basiso f its oppositeleave san dbicarpellar yovar ywit h2 ovule seach .Al lth e speci­ mensanalyze dfro mCameroun ,Gabo nan dTait ahill smaintai nthes echaracteristics . In Equatorial Guinea, the only specimen analyzed has 3 carpels and 2 ovules (Tessmann 549). Ason eapproache sCongo ,specimen stha tloo kquit esimila rt oP. bicarpellataar efoun dtha thav e5 carpel sinstea do fth e usual2 (Sita2977 ,Bouquet 514). Unfortunately, none of these specimens have flowers, therefore making it difficult toplac ethe ma tth e moment. In Zairea fe wspecimen sfro mKaniam amain ­ tainth eorigina lP. bicarpellata characteristics ,bu thav ecomparativel ymuc hsmalle r flowers(Herman 2248,217 9an dGathy 1908).Th e resto fth e specimensfro mZair e have an extremelyvariabl enumbe ro fcarpel s(3-5) , eveni n the sameinflorescence , one or two ovules per carpel and very short corolla lobes (less than 3.4 mm), (Toussaint2411 ,Donis 1967,Germain 7622 ,EdLuja, Jun e1910 ,Vanderyst 10270 , 10271,an d 15900). I, howeverhesitat et o placethes especimen sunde rbicarpellata untilthoroug hfieldwork ha sbee ncarrie d out. Inm yopinio nth egenu sPleiocarpa hasprove dt o be extremelyvariabl ea s awhole ,an don ecanno tmerel yunrave lthi s mysterybase do n morphologyalone .Othe rdiscipline snee dt ob e employedt o study thevariou spopulation st obette runderstan d thevariation .

2.7.2 Pleiocarpabrevistyl a Ominosp . nov.

Frutex foliis oppositis petiolatis lamina coriacea elliptica apice acuminata. Inflorescentia plerumqueaxillari srante rterminali sve l ramiflora.Flore sodorantes . Sepalapallid eviridi aovat aapic eobtus ausqu e ad rotundata.Coroll aalb atub olobi s ovatislongiore .Stamin ainclus acircite rmedi otub icorolla einsert aantheri sovati s apice obtusis basi cordatis. Pistillum ovario carpellis quinque separatis. Fructus mericarpis quinque separitis. - Type: Gabon, Ogooué-Maritime, 5 km East of GambaAirport , Wieringa 1267(holotyp eWAG , photo. WAG). Fig. 37, p. 140; map 15, p. 141

Shrub 1-1.2m high .Trun k very slender. Branchlets drying greyish-black. Leaves opposite; petiole 5-7 mm long; blade leathery when fresh, broadly to narrowly elliptic, 2.2-4.2x as long as wide, 4.5-16.8 x 2.1-5.9cm > acuminate at the apex, acumen7-1 7x 2-3mm , acutet o cuneate atth ebas eo r decurrent intoth epetiole ; midrib slightly prominent beneath in dried leaves; secondary veins more or less straight and slightlycurve dtoward sth emargin ,i n 12-18pairs ,formin g asubmar ­ ginalvei nan da nangl eo f60-70 0wit hth e costa,4-1 0m mapar t at the approximate centre of the leaf. Inflorescence predominantly axillary, rarely terminal or ramiflorous, i(-2)flowered, sessile .Flowers fragrant . Sepalspal egreenis hyellow ,

WageningenAgricultural UniversityPapers 96-1 (1996) 139 Figure 37.Pleiocarpa brevistyla. 1,habi t( x 2/3);2 ,sepa l( x 12);3 , openedcoroll aan dabov ei tpisti lwit h part of calyx (x 4);4 ,pisti l with opened ovary and parto f calyx (x 12);5 ,branchle t with young fruits (x 2/3). 1fro mWiering a 1267;2- 4fro mVa nde r Bürgt48 ; 5 fromWiering a& Haegens 2502.

140 WageningenAgricultural University Papers96-1 (1996) 15

15 Map 15. Pleiocarpabrevistyla. erect,ovate ,0.8-1. 2x a slon ga swide , 1.4-1.5 mm,obtus et orounde da tth eapex . Corollawhite ,wit ha bel to fpubescenc e1.5- 2m mwid einsid ejus tbelo wth einser ­ tiono fth estamens ;tub e5.8- 6 xa slon ga sth ecalyx ,1.9-2. 2x a slon ga slobes ,8.5 - 10.5m mlong ,almos tcylindrical ,1. 9 mmwid eabov eth ebase ,narrowe dbelo wth e insertion of the stamens to 1.5 mm wide, again widened around the anthers to 1.9 mm;lobe sovate ,0. 5x a slon ga sth etube ,1.6-1. 8x a slon ga swide ,4.5-4. 8 mm long,obtus ea tth eapex ,recurved .Stamens wit hth eape x3-3. 2m mbelo wth emout h ofth ecoroll atube ,inserte d0.5-0.5 2o fth elengt ho fth ecoroll atub ea t4.3-5. 5m m from thebase ; filaments short, filiform, 0.5m mlong ; anthers ovate, 3.7-4.8x a s longa swide ,1.1-1. 2x 0.25-0. 3 mm,obtus ea tth eapex ,cordat e atth ebase , Pistil 2.5-3n ™long ;ovar ybroadl y subglobose,0.8-1. 2x 1-1.5 x0.6-1. 3 mm,abruptl y narrowed intoth estyle ,o f 5 separate carpels; style 1.2-1.5m m long; pistil head narrowly ovoid to ellipsoid, 0.5-0.6x 0.2-0.3mm . Ovules (1-) 2i n each carpel. Fruits offive separat e mericarps, mericarps immature, rounded, triangular from above,c .9 x 6 mm .

DISTRIBUTION.Gabon . ECOLOGY.Dens eforest . Alt.3 0m .Flowerin gbetwee nJul yan dMarch ,fruitin g inMarc ht oMay .

Specimensexamined : GABON. Ogooué-Maritime: Gamba,J. De Wilde 1137s (WAG); Gambanea rNamb aLake , Van der Bürgt 48 (WAG); 5 km E of Gamba Airport, Wieringa 1267 (WAG); Wieringa& Haegens 2502 (WAG).

Notes.Th eleave so fPleiocarpa brevistyla resemblethos eo fP. mutica, thetw o speciesca nb edistinguishe d asfollows :

Stamenswit hape x3-3. 2m mbelo wth emout ho fth ecoroll atube ,inserte d0.5-0.5 2 of thelengt ho fth ecoroll atub ea t4.3-5. 5m mfro m thebase ; style 1.2-1.5 mm long P. brevistyla

WageningenAgricultural UniversityPapers 96-1 (1996) 141 Stamenswit hape x0.8-1. 3 mmbelo wth emout ho fth ecoroll atube ,inserte d0.73 - 0.92o fth elengt ho fth ecoroll atub ea t8-1 8m mfro mth ebase ;styl e7.9-1 6m m long P. mutica

2.7.3 Pleiocarpamutic aBenth . in Hooker. Ic. PI. 12: 71, PI. 1181(1876) ;K .Schuman n in Engler &Prantl ,Nat . Pflanzenf. 4, 2: 135,fig. 52 ,C & D ,p . 134(1895) ;Stap f inFI .Trop .Afr . 4, 1: 98 (1902);Picho ni nBol .Soc .Brot .sér .2 ,27 : 138,ma pc p . 121 (1953);Hutchinso n &Dalzie li nH .W .Trop .Afr . éd. 1,2 :3 8(1931) ;H.Hube ri nFI .W .Trop .Afr . éd. 2, 2: 63,fig. 214 , (1963).-Type : Nigeria, Cross River State, Old Calabar River, Mann 2277 (holotypeK) . Fig. 38,p . 143; map16 ,p .14 4

Homotypicsynonym :

Hunteriapleiocarpa Hall .f . in Jahrb.Hamb .Wiss .Anst . 17,Beih . 3: 193(1899) , partly asfo rth etype .

Heterotypic synonyms:

Pleiocarpa salcifolia Stapf op. cit. 99 (1902); Hutchinson & Dalziel op. cit. 38 (1931). -Type: SierraLeone ,sin .loc ,Barter 8 May 1851,(holotyp eK) . P. tricarpellata Stapf inKe wBull . 1915:4 7(1915) ;Hutchinso n &Dalzie lop .cit . 38(1938) .-Type :Sierr aLeone ,Falaba ,Alymer 3 5(holotyp eK) .

Shrubo rsmal ltre e0.5-7. 6m high ,rarel ya climbin gshrub ,4- 9m high .Trun k1.5 - 5 cmi ndiameter ;bar ksmooth ,dar kbrow nt opal egrey ;woo dpal ebrow nt oyellow , hard,ver yclose-grained .Branche sdar kgree nwit hbrownis hpatches ,brow nt odar k brown,o rdar kgrey ,lenticellate ;branchlet sbrownish ,pal et omediu mgreen .Leaves opposite,sometime sternat e(i nplant sfro m Liberiaan dIvor yCoast) ;petiole ,5-1 3 mmlong ;blad e papery tothinl y leathery when fresh, rather stiff, broadly to nar­ x rowlyelliptic ,oblon gt oovat e 1.6-5 aslon ga swide , 5.i-i9-5(-29.5)x I.9-8.I(- 11.4) cm, acuminate at the apex, acumen acute,4-2 5 x 2-5mm , acute, obtuse or cuneatea tth ebas ean ddecurren t intoth epetiole ,smoot han dshin yo nbot hsides , dryingreddis hbeneath ;midri bslighd ychannelle d above,prominen tbeneath ;sec ­ ondaryvein si n9-1 2pairs ,formin g asubmargina lvei nan da nangl eo f50-70 0wit h themidrib ,8-20(-3o )m mapar t atth ecentr eo f theleaf ;tertiar y veinsobscur ebe ­ neath.Inflorescence a naxillar ycluste ro rramiflorous ,rarel yterminal ,1.5-2. 5x 2- 3 cm,abou t 10(-35)-flowered ;sessile .Pedicel sbrown ,0.5- 2 mm.Bract sver ysmall , 0.8 x0. 5mm .Flowers veryfragrant . Sepals palet odar k green,fre e orconnat ea t the base up to 0.2-0.4m m of their length, erect, membranaceous for about0.3 - 0.4m mo fthei redge ,ovat eo relliptic ,0.7-2. 6x a slon ga swide ,1.3- 2(-2.4 )x 0.7 - 1.9mm ,obtus ea tth eapex . Corollapur ewhite ,14-26. 6m mlon gi nmatur ebu dan d

142 WageningenAgricultural UniversityPapers 96-1 (1996) Figure 38. Pleiocarpa mutica. 1,habi t (x 2/3);2 , flower (x 3.4);3 ,opene d corolla (x4) ;4 ,opene d calyx withpisti l (6);S ,infructescenc e (x 2/3);7 ,longitudina l section of seedi nfrui t (x 1);8 ,pul p( x 1);9 ,see d (x 1); 10,embry o( x 2). 1 from J.J .d eWild e 7548;2- 4 from Leeuwenberg 12219;5-1 0fro m De Koning 5790.

WageningenAgricultural University Papers96-1 (1996) 143 Map 16.Pleiocarpa mutica. forming acomparativel y largeoblon ghea d0.29-0.3 7o fth ebu dlength ,5-1 1 x2.5 - 3.5mm ,wit ha rounde dapex ,wit ha bel to fpubescenc e2.5- 6m mwid einsid ejus t belowth einsertio no fstamens ,longes thair sjus tbelo wth eanther sbecomin gshorte r andscarc efurthe r downan dsometime swit ha fe w hairsaroun dth estamen su pt o themouth ;tub e5.3-1 3(-18.4 )x a slon ga sth ecalyx , i.4-2.4(-2.8)x a slon ga sth e lobes, 10.8-22m mlong ,almos tcylindrical ,1- 3m mwid eabov eth ebase ,narrowe d belowth einsertio n of the stamenst o0.4-1. 8m mwide ,agai n widened aroundth e antherst o0.9-1.8(-2.5 )m mwide ;lobe sovate ,ellipti ct onarrowl yoblong ,0.4-0. 7x aslon g asth etube , 1.4-5.2x aslon g aswide ,(5-)6-i 3x 2.1-4.3mm > rounded or obtuse at the apex, entire, spreading and recurved later. Stamens with apex 0.8- 1.3 mmbelo wmout ho fth ecoroll atub einserte d0.73-0.9 2o fth elengt ho fth eco ­ rollatube ,a t8-1 8 mmfro mth ebase ;filament s 0.2-0.6m mlong ,glabrous ;anther s narrowlyovat et ooblong ,yello wwit hre dstripes ,2.8- 6x a slon ga swide ,1-1. 9 x 0.25-0.6mm ,obtus e atth eape xan dsteril efo r 0.2 mm,cordat e atth ebase . Pistil 9.2-i8mmlong;ovar ysubglobose ,0.7-1. 2xo.6-1. 5xo.5- 1mm ,abruptl ynarrowe d intoth estyle ,wit ha disk-lik ethickenin gunitin gth ecarpel sa tth ebase ,abou t0.4 - 0.5o fth elength ,0.3-0. 4m mhigh ,o f 5separat ecarpels ,(occasionall y 3-4carpel s found inth eflowers o f thesam einflorescenc e orbranch) ;styl e white,7.9-1 6 mm long;pisti lhea dwhite ,oblon gt oellipsoid ,0.6- 1x 0.2-0. 4m mrarel ywit ha filiform stigmoidape xu pt o0. 3m mlong .Ovul eon ei neac hcarpel .Fruits yello wt obrigh t orange,smooth ,dull ,slightl ywrinkle dt ofinely lenticellate ,5 (rarel y3-4 )separat e mericarps;mericarp sobovoid ,subglobose ,ovoid ,o rellipsoid ,sometime ssomewha t curved and with apointe d apex, often in clusters, 13.5-20x 8.5-20x 7-11.6mm , rounded atth eapex ,alway s 1-seeded ;wal labou t 2-3m mthick .Seeds subglobose tooblong ,7.5-1 2x 4.5-9. 5x 4- 8mm .Embry o7.1- 9m mlong ;cotyledon sovat et o

144 WageningenAgricultural UniversityPapers 96-1 {1996) elliptic,thi nan dleafy , i.2-2. 3x aslon ga s wide,3- 4x 1.5-3mm ,obtus e at the apex androunde d atth ebase ;radicl e0.9-2. 3 xa slon ga sth ecotyledons , 3.4-5 x0.6-0. 8 mm.

DISTRIBUTION. Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroun and Gabon. ECOLOGY. Denseprimar y orsecondar y forest inswamp y areasric hi n epiphytes oralon gth erive rbank s onsand y soilo rlimeston e out-crops.Alt .0-60 0m . Flower­ ingan dfruitin g allth eyea rroun d inIvor y Coast, flowering inJanuar y toApri l and fruiting between Julyt oSeptembe r inth eothe rcountries .

USES.Th e wood is yellow, hard, heavy, close-grained and tough (Vigne 269, FHO,K , P, Ghana; Burt-Davy 1937;Irvin e 1969);use d tomak e combs (Deighton 3270, K, P, 1936, Sierra Leone); for combs and planes (Vigne 269, FHO, K, P, Ghana);fo r sundry small objects (Aubreville 1959);use d in Ivory Coast for canoe paddlesan dpestle s(Bouque t & Debray 1974).A nornamenta l plantwit hcluster s of narrow, tubular, white sweet-scented flowers known as the stove plant in Europe (Dalziel 1937;Irvin e i96i).InIvoryCoas tth eplan ti sconsidere d agenera lpanace a for all ills by the coastal people, the Anyi use a decoction of the grated bark for stomachpains ,an da similarpreparatio n is usedb yth eKyam aagains toedem ao fth e legs,probabl ydu et okidne ymalfunctio n (Dalziel 1937);a decoctio n ofth eroo tbar k used against kidney diseases and malaria (Leeuwenberg 12302,WAG , 1981,Ivor y Coast); bark ground and rubbed on the body against fever (Deighton 3270, K,P , 1936,Sierr a Leone). Anumbe r of alkaloids havebee n reported from the bark and roots (Willaman &Li 1970).

Aselectio n of the approximately 130specimen s examined: SIERRA LEONE.Yor kPeninsula ,Morton 23.11.196 3(K) ,SL126 8 (WAG);Falaba ,Alymer 35 (K ,typ e of P. tricarpellata);nea rFalama ,Deighton 386 9 (K),Deighton 387 1 (K);Giehun ,Marmo 6 5 (K);Gol a Forest,Small 70 1 (B,K , P); Yonibana,N. W.Thomas 423 5 (K);Kambu iFores tReserve ,Lane-Poole 33 3 (K); Mabonto, Deighton 3270 (K, P); Sugar Loaf Mt., Hepper 2539 (K);Nea r Sugar Loaf,Richards 6.2.1971 (K), Roberty 17318 (G); Gbinti, Deighton 2501 (K); Kafa, Alymer 216 (K); sin. loc. Barter 8.5.1851 (K,typ eoff . salicifolia). LIBERIA. Bonge Range, Voorhoeve 1140 (WAG),Leeuwenberg & Voorhoeve494 5 (B, BR, HBG,K , MO, P, WAG). Eastern Prov.:Tchie n Distr., Baldwin 7013 (K); 16 km NW of Tchien, Jansen 1255 (WAG); AlongDukwa iR. , Cooper 248(A , FHO, GH,K ,NY ,US) ; Monrovia,Dinklage 297 6 (Z); Grand CapeMt. ,Baldwin 1081 4 A (K); Greenville,Baldwin Jr. 11552 (K,MO) ;Ganta ,Harley 317 (K);Roa d toTappita,Jansen 2112 (P); ZeahTown ,Baldwin 699 1 (K);Fireston ePlantation ,Linder 2.8.192 6(A) . IVORY COAST. MontCope , Chevalier 19722 (P); Yokobo, Leeuwenberg1230 2 (WAG);For tBinge r (= Djiroutou), Chevalier 19432 (P);Béréby , Chevalier 20011 (K, P);Sa nPedro ,Thoiré 796 (P);2 2k m NE of Monogaga, Geerling &Bokdam 237 7 (BR, MO, WAG); Sassandra,Thijssen 11 8 (WAG);6 0k m N of Sassandra, left bank of Davo River,Leeuwenberg 283 5 (BR, FHO,GENT , K,L , MO, NY, P);5 0 kmNN Eo f Sassandra,Breteler 612 0(K , L,P ,U , US, WAG, Z); Westo f Fresco,Breteler 535 4 (BR, K, P); YapoForest ,Bamps 2559 (BR);Banc oForest ,De Koning 5790 (WAG); ibid., Oldeman 961 (B,BR , K, M, WAG); Between Port Bouet and Grand Bassam, J. de Wilde320 4 (A, B, BR, K, NY, P, WAG); Aboisso, Sanvi, Chevalier 17862 (P); Assinie, Chevalier 17869(P) . GHANA.Western :Enchi ,Andoh 5639 (BR); TanoNimri ,Enti F H754 4(K , UPS);Ankas aGam eRe ­ serve,Jongkind & Abbiw 2073 (WAG);Asanwinso , Vigne 269 (FHO, K,P) ;Axim , Irvine 2173 (E, K,

WageningenAgricultural University Papers 96-1 (1996) 145 MO); AsenanyoFores tReserve ,Andoh 430 5(BR , K);Ndumfr iR .Fores tReserve ,Enti & Hall GC 38392 (FHO, K);Prince sTown ,Akpabla 792 (K);Neun g Forest Reserve, Enti2296 A (MO);Daboase ,Hall & Abbiw GC 45117 (K).Central :Eguafo ,Hall 294 4 (K). Eastern:Atew aRang eFores tReserve , Vigne434 5 (K). NIGERIA. CrossRive r Odukpani,Ariwaodo inFHI8871 5 (MO);Abangara-Ndiakor oRd. , Ujor Fffl 30169 (K);Ol dCalaba rR. , Mann227 7 (K,type) ;Stubb s CreekFores tReserve , Van Meer 1178(WAG) ; EkintaR.Fores tReserve , Van Meer 1491 (WAG);Oba nGrou pFores t Reserve, Van Meer 1266(WAG) ; Oban,Talbot 1565 (K), 1654(K) , 1911-1912 (K). CAMEROUN. Sud-Ouest: Korup National Park, D. W. Thomas 4575 (P, MO,WAG) ; Kondotiti- MundembaRd. , Gentry&D.W. Thomas 52681 (MO, WAG); Near Ekondotiti, Lowe 3807 (K);4 kmW of OngeRiver , Watts 760 (K); Idenao,Tchouto 740 (K); Weke, Kumba,Smith 75/36 (FHO).Centre-Sud : Lolodorf, Staudt 322 (G, K, P, S); 5 km NE of Lolodorf, Letouzey 12784 (P, YA); NearMeyo , Ntem, Koufani 107 (P, WAG), Letouzey 15209 (P); Ebolowa, De Wilde & de Wilde-Duyßes 1964 (BR, K,P , WAG); Station du Cacaoyer de N'koemvone, J. de Wilde 7548 (BR, P, MO,WAG) ; 10 km SW of Ambam,De Wilde &De Wilde-Duyßes203 2 (WAG);2 5 kmES Eo f Ebolwa,Letouzey 985 0 B (P). GABON. Nyanga:Gamba ,Breteler & Van Raalte563 5 (BR,WAG) . Ogooué-Lolo:nea rLastoursville , Breteler 6558 (BR, MO, WAG); 14k mN Eo f Bambidie,Breteler et al., 13338(WAG) . CULTIVATED. Netherlands:Wageninge n Bot. Garden, Leeuwenberg 14054 (WAG), Van Setten 415 (WAG), seedling of De Koning 576; Ivory Coast: Adiopodoume, Breteler 6207 (WAG); Université d'Abidjan, Fredoux58 9 (P). Singapore:Botani c Garden,Leeuwenberg 13340(WAG) .

Notes.Th eonl yconstan tcharacte ri nP. muticai sth enumbe ro fovule swhic hi s always onepe rcarpe l in all the specimens examined, even when thecarpel s vary from 3-5.Th e leaves,ar eopposit e and sometimes ternate especially in specimens from IvoryCoast .

2.7.4 Pleiocarpapycnanth a (K. Schum.)Stap f in FI.Trop . Afr.4,1:9 9(1902) ;Eggelin g & Dalei nIndig .Tree sUgand aProt . ed. 2: 29 (I95I);Picho ni n Bol. Soc.Brot .sér .2 ,27 : 128,t .4 fig . 1 (1953);F.Whit ei n F. F.N.R.35 0(1962) ;H . Huberi nFl .Trop .Afr . ed.2 ,2 :6 3(1963) ;R.B.Drummon d in Kirkia 10:26 9 (1975);Kupich ai nFl .Zamb .7 ,2 :432 ,tab.99 ,p .43 3(1985) . - Type:Uganda ,Sess eIslands ,Stuhlmann 1216,(holotyp eBf , isotypeK) . Fig.39 , p. 147; map 17, p.14 8

Basionym:

Hunteria pycnantha K.Schum . in Engler, Bot. Jahrb. 23:22 2 (1896); Hall. f. in Jahrb.Hamb .Wiss .Anst . 17,Beih . 3: 191 (1899).

Heterotypicsynonyms :

Pleiocarpatubicina Stap f in Kew Bull.1898 : 304(1898) ;D eWildema n& Duran d in Ann. Mus. Congo Bot. Ser. 2 vol. 1, 1: 37 (1899) & 1, 2: 150 (1901). P. pycnantha var. tubicina (Stapf) Pichon op. cit. 132,ma p C, p. 121.;H .Hube rI.e . -Type:Zaire ,Lowe rCong oR. ,Dewèvre 94 5(holotyp eBR ;isotyp eK) .

146 WageningenAgricultural UniversityPapers 96-1 (1996) Figure 39.Pleiocarpa pycnantha. 1,habi t (x 2/3);2 ,flower bu d (x 4);3 ,flower ( x 4);4 ,flower abov e (x 4); S, sepal inside (x 8);6 ,opene d corolla (x 6);7 ,pisti l (x 8). 1fro m J.Loui s 12898; 2-7 from Breteler & DeWild e 636778.

WageningenAgricultural UniversityPapers 96-1 (1996) 147 10 o Map 17.Pleiocarpa pycrumtha.

P. welwitschii Stapfex Hiern inCat .Welw .Afr . PI. 1,3:665 (1898);Stap f inop .cit . 100. -Type: Angola, Golongo Alto, left bank of R.Cuango, Welwitsch 5981 (holotype K; isotypes B,BM , C, G, P). Hunteria breviloba Hall. f. in Jahrb. Hamb.Wiss. Anst. 17, Beih 3: 189 (1899); Schlechteri nWestafr . Kautschuk.Exped .30 6(1900) .Pleiocarpa breviloba (Hall. f.) Stapf op. cit 102. -Type: Congo, near Sanga R., betweenWos o and Bonga, Schlechter 12706(holotyp e B t)- Pleiocarpa micrantha Stapf inop .cit . 100;Hutchinso n &Dalzie li nF lW .Trop . Afr. ed. i, 2:3 8 (1931);Irvin e inWood y pi. Ghana, 630 (1961).-Type : Ghana, base of Aburi hills,Johnson 623 (holotype K). P.flavescens Stapf in op.cit . 101;Hutchinso n & Dalziel I.e.-Type : Ghana, Aburi hills,Johnson 616 (lectotype K, designated here). P. microcarpa Stapf in op. cit. 102. - Type: Zaire, Haute-Zaire Niamniamland, Mbrowole R., Schweinfurth 3073 (holotype K). P.bagshawei S.Moore, in J.Bot. 14: 49 (1907). -Type: Uganda, U2,Toro Distr., Durro (= Dura) Forest, Bagshawe 1086 (holotype BM). P. swynnertonii S.Moore, in J.Linn. Soc.Bot . 40: 138 (1911).-Type : Zimbabwe, Chirinda Forest, Swynnerton 14 (lectotype BM, designated by Kupicha 1985; isolectotypes K, SRGH, US,Z) .

148 WageningenAgricultural UniversityPapers 96-1 (1996) Shrubo rsmal ltre e i.5-2 0(-30 )m high ,rarel ylianescent .Trun k2-5 0c mi ndiame ­ ter;bar k3- 6m mthick ,smoot ho rslightl yrough fissured, t oreticulat ewit hirregula r longitudinal and straight horizontal cracks 1-2 cmdeep ,pal et odar k grey,pal et o reddish-brown,yellowis ht obrow no nsection ,sometime slenticellate ,mor eo rles s flaky andpeelin goff ; innerbar kcreamy-whit et opal ebrow nwit hdar kyello wstria - tions;woo dver yhar dan ddurable ,smooth ,dar kbrown .Branchlet sterete ,ver ypal e todar kbrow ni n dried specimens,sometime s lenticellate, often withbar kpeelin g off. Leavesopposit e and in whorls of 3-5, always whorled in one node of each branchlet of aplant ; petiole 5-20m mlong ;blad e coriaceous when fresh, papery when dried,narrowl y elliptict ooblong ,rarel yobovate , 1.9-5.4x aslon ga swide , 4-22.3x 1.1-8 cm,acute ,obtus et oacuminat ea tth eapex ,acume n4-1 2x 2- 5mm , acute or obtuse to rounded, or cuneate at thebas e and decurrent into thepetiole , sometimes undulate, glossy, with 15-25pair s of secondary veins,formin g a neat submarginalvei nan dangl eo f50-80 0wit hth emidrib ,tertiar yvenatio ninconspicu ­ ous.Inflorescence axillary ,rarel yals oterminal ,fasciculate ,sometime sramiflorous , 1-2 x1- 3 cm,io-3o(-4o)-flowered ;sessil eo rwit ha peduncl e1- 2 mmlong .Pedicel s 1-3 mmlong .Bract s very small, scale-like.Flowers very fragrant tomalodorous . Sepalsgreen ,fre e orconnat e atth ebas efo r upt o0. 2 of theirlength ,erect , often spreading in driedflowers, membranaceou s for about 0.2-0.4 mm of their edge, broadlyovat eo rovat et oelliptic ,0.9-2. 5x a slon ga swide ,1 .i-2.5(-3 ) x0.6- 2 mm, acutet orounde da tth eapex .Corolla pur ewhit et oorange-yellow ,8-i2.8(-i5 )m m longi nth ematur ebu dan dformin g acomparativel ynarro wovoi dhea d0.14-0. 4o f thebu dlength ,2-4. 5x 0-5-2-5 mm,wit ha rounde dapex ,wit ha bel to fpubescenc e 1-3m mwid einsid ejus tbelo wth einsertio no fstamens ,hair slonges tjus tbelo wth e stamens;tub egreenish-white ,3.2-7. 5(-8.8 )x a slon ga sth ecalyx , 1.7-4.8(-5.4 )x aslon ga sth elobes ,6.2-1 0(-10.5 )m mlong ,almos tcylindrical ,0.6- 2(-3 )m mwid e aboveth ebase ,narrowe dbelo wth einsertio n of thestamen st o0.6-1. 5m mwide , again widened around the anthers to 0.7-2 mm wide; lobes ovate or elliptic to suborbicular,o.2-o.5(-o.6 )xa slon ga sth etube ,0.9-2 .i(-2.5 )xa slon ga swide , 1.3- 4.5(-4.8)x o.7-3.2(-3.5)mm , rounded to acute at the apex, entire, spreading and recurvedlate ran dwit ha nupcurve dapex .Stamens wit hape x0.2- 1 mmbelo wth e moutho fth ecoroll atube ,rarel yvisibl ea tth emouth ,inserte d0.64-0.8 7(-0.97 ) of thelengt ho fth ecoroll atube ,a t4.6- 8 (-9.5)m mfro mth ebase ;filament swhite ,0.3 - 0.7 mm long; anthers yellow, ovate, 1.75-5.3 x as l°ng as wide, 0.6-1.3x 0.15- 0.5mm ,acuminat e toobtus e atth eapex ,cordat e atth ebase .Pistil glabrous,4.6 - 8.6 (-9.7)m mlong ,ovar yovoi do rsubglobose , O.5-I.4(-I.7) x0.3-1.o(-1.2 )x 0.2 - o.8(-i)mm ,abruptl ynarrowe dint oth estyle ,wit ha disk-lik ethickenin gunitin gth e carpels atth ebase ,abou t0.25-0. 4o f thelength ,0.2-0. 5m mhigh ,o f twoseparat e carpels;styl e3.8-7. 2(-8 ) mmlong ;pisti lhea dellipsoi dt oovoi d0.2-0. 8(-1.0 )x 0.2 - 0.5mm .Ovule stw oi neac hcarpel ,( 4i nTroupin 4024 ,WA Gan dGoldsmith 38/65 , SRGH).Fruits yello wt oorange ;mericarp ssmooth ,sometime srugose ,subglobos e orpyriformt oellipsoid ,sometime sstipitate , 13-23(-30 ) x6.5-2 4x 5-1 5 mm,acu ­ minatet orounde d atth eapex , 2seeded , wall about 2-5m mthick , sometimesfi­ brous.Seeds ellipsoid tooblong ,6.5-13. 5x 4.5-9. 5x 2-4mm .Embry o4.7- 9m m long,cotyledon sovat et oelliptic ,thi nan dleafy , 1.1-3. 7x a slon ga swide ,2- 4x 0.8 -

WageningenAgricultural University Papers 96-1 (1996) 149 2. o mm, rounded at theape xan d at thebase ;radicle ,0.4-1. 9x a slon g as thecotyle ­ dons, 1.4-5 x 0.4-0.9mm .

DISTRIBUTION. Tropical Africa from Senegal in the West to Kenya in the East andAngol aan dMozambiqu e inth e South. ECOLOGY. Montane forest, secondary forest or rain forest understorey rich in lianas,mosse san depiphytes ,o rgaller yforest . Alt. 0-2300m . Floweringan d fruiting allth eyea r round.

USES.Th e wood is very hard (Dalziel 1937) and is used for local construction (Flamigny 6123,BR) ;fo r making combs (Irvine 1961). Roots ground, mixed with guinea grains and palm wine and used to promote freedom of bowels, or used in palm wine to make it strong (Irvine 1759, E, K, 1932, Ghana); leaves are bitter (Irvine 1961).

A selection of theapproximatel y 200 specimens examined: SENEGAL.Sin eSaloum : Deltad u SaloumNationa lPark ,Lykke 35 6(WAG) .Casamance : Djibelor, Van denBerghen 5661 (BR);Oussouy eAdam 13686 (MO); Boukitimgo,Berhaut 692 2 (BR, M,P) . GUINEA BISSAU. Canchungo, Emapaca, EspiritoSanto 1941 (BR, MO, P, WAG). Bolama,Entacha , EspiritoSanto 191 9 (MO, WAG); Fulacunda,Espirito Santo204 2(WAG) . GHANA. Central:Winneb ajunction ,Hall & Abbiw 4709 6(BR ,MO , WAG); Awutu-WinnebaRd. , Enti, SP 393 (F, MO, NY). Eastern: Akropong, Akwapim, Irvine 1759 (E); 1k m S of Aburi, Leeuwenberg 11042(WAG) . TOGO, nearLome , Wamecke48 1 (EA,P) . BENIN. Ouidah,Le Testu 70 (P); nearSakété , Chevalier2287 3( p)- NIGERIA. Oyo: Ilesha, Patel 51694 (WAG); Igbajo, Latilo FHI 31754 (WAG); Iressa, Latilo FHI 58433(EA) .Ondo :Akure-Ow oRd. , Gbile et al. FHI 20580 (P, WAG); Iyin,Ekiti ,Olurunfemi &Fagbemi FHI7075 7 (WAG).Kwara :Kabba ,Latilo FH I6055 6 (MO).Anambra :Abakaliki-Obubr a Rd., Okafor& EmwiogbonFH I6604 0 (WAG).Benue : Ankpa, OkaforFH I3688 6 (FHO). Cross River:Ogoja , Binuyo FHI4124 0 (BR, WAG). CAMEROUN. Sud-Ouest:Mbilishe ,D. W.Thomas 7432 (MO).Ouest :Bangwa ,De Wilde & DeWilde- Duyßes 2420 (P, WAG). Centre-Sud:M t Eloumdren, Lejoly 88/601 (BR);Lob o R., Letouzey 4589 (P). Est:3 0k mN of Bertoua,Breteler 2203 (BR, P, WAG); 53k mS Eo f Ngoila, Letouzey1196 3 (BR, K, P, WAG); Moloundou,MUdbraed 458 8 (HBG);Nea rNdongo ,Letouzey 12087 (BR, G, HBG, P,WAG) ; 65 kmNN Eo f Moloundou,Letouzey & Villiers1049 2(K ,P) . CENTRAL AFRICAN REPUBLIC. Nana-Mambéré: Along Ndongue Forest, Haxaire 1345 (WAG). Lobaye: Ebure Forest,Tisseront 2411 (BR, K, P); Torogurade,Tisseront 2412 (K, P). Ombella-Mpoko: Bangui, Chevalier 10930 (P). Kemo-Gribingui:nea rSibut , Chevalier 10793 (P). Ouaka: KuduR. , near Moroubas,lisseront 1461 (P).Haute-Kotto : Yalinga,Le Testu 4524 B (P,US) ,456 6 (BR,P) . GABON. Ogooué-Ivindo:Beiinga , Halle 3773 (P). Woleu-Ntem: Oyem,Le Testu 9461 (EA, MO,P) ; Babiel North,Breteler & De Wilde 636/78(WAG) . CONGO. Pool:Kak aForest ,Koechlin 5065 (P).Plateaux :Ng o Forest,Sita 3075 (P).Cuvette :Sangh a R., Pobéguin15 7 (P); Kassa, Chevalier508 4 (P).Likouala :Ibeng aForest ,Sita 3797(BR) . ZAIRE. Bas-Zaire:Matadi ,Dacremont 32 9 (BR, EA, MO, P, UPS, WAG); Mbuami,Dubois 63 (BR, EA, K, WAG); between Dembo and Krango, Butaye Oct 1900 (BR); Nzuma, Nsimundele93 7 (BR); Kinshasha: Maluku, Breyne 790 (BR), 2318 (BR). Bandundu: Bokoro, Jans 4.4.1948 (BR); Buna, Flamigny 6123 (BR); Kamtsha R., Vanderyst 11305 (BR), Kikwit, Renier 24.5.1944 (BR); Ipamu, Vanderyst931 4 (BR), 10432 (BR);Libue ,Luja 292 (BR). EquateurWangata ,Lebrun 915 (BR, C,EA , MO, P,WAG) ; Hœnge,Huyghe Ma y 1907 (BR); Ingende,Evrard 611 9 (BR); Djoa,Evrard 408 2 (BR, K); Makanza,D e Gi'o/gi'133 9(BR ,G , US); Bomputu,Ghesquière 2782(B ,BR ,K , P); betweenKaraw aan d Businga, Lebrun 1950 (BR, WAG); Bobutu, Evrad 481 (BR); Bodangabo, Evrard 327 (BR);Businga ,

150 WageningenAgricultural UniversityPapers 96-1 (1996) Lebrun198 7 (BR, MO, WAG); Likimi,Malchair 107(BR) ; Molegbue, Gutzwiller58 1 (BR,MO ,WAG) ; Lisala, Leontovitch22 8 (BR, MO);Yambata , £>e Giorgi 1737 (BR); Dundusana, Reygaert 125 (BR); Ubangi,£<*n m 2167 (BR, EA, MO, WAG); flcela, Hulstaert 1306 (BR);Bokungu ,Jespersen 86 (BR). Haut-Zaire:Yobahonde ,Germain 815 0(BR ,K ,M ,P) ; Barumbu,Laurent 137 6(BR) ; Mobwasa,Reygaert 1326 (BR); Kulu, V&n den Brande 380 (BR);Buta , Leen« 2479 (BR, MO, P, WAG); Yangambi,Louis 4275(BR ,C ,K ,NY) , 12898(BR ,C) ; Isangi,Louis 634 0(BR ,B ,C ,EA , NY, US); Kisangani,Lc/o/ y193 3 (BR); Kisangani, Lisowski4786 5 (BR); Wanie-Rukula, Lisowski 17126 (BR, K), 43425 (BR, K);nea r Bangoka,Lisowski 4550 8 (BR, K,WAG) , 45511 (BR, K); KongoloIsland ,Mosango 53 9 (BR); Kabondo, Mosango43 1 (BR); ZobiaForest ,5e/» *89 3 (BR); Tukpwo,Gérard 274 6 (BR,WAG) , 3732(BR) ; Batite, Gérard554 2(BR ,WAG) ; nearPanga ,Bequaert 1623(BR) ; Bafwaboli, Claessens25 8(BR) ; Doruma,£> e Graer23 7 (BR); Nala,Boon e1 1(BR) ; Mabwasa,/far t 508 (BR,MO) ; 4k m SSWo f Epulu,Ge/ea u538 5 (WAG);Penghe ,Bequaert 229 8(BR) ,222 9(BR , K); Mniamnian,Mbrowole ,Schweinfitrth 307 3(K ,typ e of P. macrocarpa);Gombari , Seret48 2 (BR);Garamb aNationa l Park,Troupin 7 (BR, EA, K, P, WAG), 547(BR ,WAG) ; Kurukwata,Gérard 358 0(BR ,WAG) ; Mokatji, Vander Ben 1433(BR ,K , M); Mahagi, Afeyers29 7 (BR, C,K , M); Lekwa,Lebrun 989 8 (BR,MO ,P ,WAG) ; DjuguForest ,Bamps 26 1 (BR,K) ; DjuguReserve ,Meyers 195(BR ,C ,EA , MO,WAG) ; KawaForest , Van der Ben 1262(BR , K,M) .Kivu : Rumangabo,Germain 3796 (BR,EA ,MO , WAG);Kahe ,Gutzwiller 1983(BR ) & 1995(BR) ; Rutshuru, Deru 344 (BR); Burungu, A.Léonard 5230 (BR, EA, MO, WAG); Irangi,Troupin 3515 (BR, EA, K, WAG); Kilongo, Gutzwiller 1117 (BR, MO, WAG); Kavumu-Walikale Rd„ Pierlot 2341 (BR,WAG) ; Walikale,Lebrun 512 1 (BR, MO, P.WAG);Bukavu-Shabund a Rd., Pierlot 2523(BR ,C ,EA ,MO ,WAG) ; Bukavu-WalikaleRd. ,Stauffer 102 1 (BR, K, UPS, WAG); Mt. Kaluli,Pierlot 2390 ((BR,WAG) ; Bitale, Pierlot 1045(BR) ; Lusheni,A. Léonard470 4(BR ,EA , MO, P,WAG) ; Kitole,A .Léonard 394 8 (BR,EA , MO, WAG); Nyakigunda,A.A. Léonard 387 1(EA ,MO ,P) ; Urega,Lebrun 558 6(BR ,WAG) ; nearKindu , Lebrun608 8(BR ,M)) ; Elundu,Henrad^ji (BR); Maniema,Michelson 13 3(BR) ; Lumuna,Lebrun 587 7 (BR, MO, WAG); Kapanga,Michelson 73 6(BR ,P) ; Mutambo,A. Léonard569 0(BR ,EA ,MO ,P.WAG) ; Kasongo, Dewèvre 945 (BR, K, type of P. tubicina); Kahusi, A.Léonard 3981 (BR, EA, MO, WAG). Kasai-Occidental:Lutshuad i Forest,Deffet 10(BR , P).Kasai-Oriental : between Lodjaan dKole ,Lebrun 6220 (BR,WAG);Bena-Kabindi , Liben 1835 (BR). Shaba: Monono,Thiebaud 203 (BR);nea r Kilwesi National Park,De Witte 419 8(BR , EA, WAG); UpembaNationa l Park,De Witte 694 2(BR ,WAG) ; N'zilo nearKolwezi , Schmitz 3704 (BR);nea rLuamb o& Bunkeya ,Devlaux 40 9 (BR);Lukaf u Forest, Schmitz 7334 (BR, MO), Schaijes 3119 (BR);Boa , Malaisse 6763 (BR, WAG); Kafubu Forest, Malaisse 8599 (BR). RWANDA. Rangiro,Troupin 14693(BR ,EA) . BURUNDI. Bugarama,Niyongere 1/84(BR) ;Siguvyay e valley,Lewalle 3990(BR) . ANGOLA. Cabinda:Sumba ,Peco ,Gossweiler 729(K) ,872 9 (US).Malanje :Golungo-Alto ,Wfe/wittc n 5981 (B,BM ,BR ,C ,G , K,MO ,P , typeo f P. welwitschii).Congo :Uige ,Gossweile r 1139(K ,P) .Lunda : Dundo,Gossweile r 13862(K) , 14051 (B, K, P,US) . SUDAN. Sourceo f YubuR. ,Hoyle 725 (FHO);Azz aForest ,Turner 160(K) . UGANDA. U 2:Kigez i Distr., Kigezi Forest,Forest Dept. EAH 12437 (EA, K).Toro Distr., Mungilo, BwambaFores tReserve ,Paulo 60 7(EA , K); HimaForest ,Osmaston 68 4(EA , K); Durro( =Dura )Forest , Bagshawe 1086(BM ,typ eof P. bagshawei). AnkoleDistr. ,Kalinz uFores tReserve ,Osmaston 2767(K) , Paulo61 9 (BR, EA, K);Ruampara , Kikagati near Kagera River (EA, K). Masaka Distr., W of Katera, Katende 1429 (EA);Malabigamb o Forest,Drummond & Hemsley 4546 (K). Mengo Distr., nearBudo , Eggeling 517 (BR,EA ,K) ; 16k mo nBorb oRoad , Chandler212 5 (B,BR , K,P) ;Kyewag a (=Kyiwaga ) Forest,Maitland 606 (K);Sess e Islands,Stuhlmann 1216 (K,type) . KENYA. K4 : MemDistr. ,Ngai aForest ,Wachiori 4 1(EA , K); MeruNationa lPark ,Rojwer oR. ,Ament & Magogo 169(BR , EA, K). K7 : Kwale Distr.,Mrim aHill ,Faden & Faden 77/751 (K, US), Kokwaro 3959(EA) ; ShimbaHill sLodge ,MkombaR. ,Robertson & Luke515 3 (EA); ShimbaHills ,Mwel eMdog o Forest,Omino 3 2(EA ,WAG) .Kilif i Distr.,Kay aRibe ,Robertson & Luke483 0(EA , K); ArabukoSokok e Forest, Omino6 9 (EA, WAG). LamuDistr. ,Wit uFores t Reserve,Luke & Robertson 1405 (EA,K) . TANZANIA. T 1: Bukoba Distr., Kaige, Gillman 352 (BR, EA, K).T 3: Lushoto Distr., Kwamkoro Forest Reserve,Ruffo & Mmari 2129 (K);Lushoto/Tang aDistr. , Longuza,Ruffo &Mmari 1873 (K).T 4: Kigoma Distr., Kasakati, Suzuki B 23 (EA);Mpand a Distr., near Selimwenguru below Kungwe Mt., Harley958 7 A(B ,BR , EA, K);Kungw eMt. ,Harley 9596 (BR, K). T 6: UlangaDistr. ,Ifakara ,Vikund u

WageningenAgricultural UniversityPapers 96-1 (1996) 151 ForestReserv e( =Vikindu) ,Haerdi 033-196 3(Z) .Morogor oDistr. ,Ulugur uMts . nearRuv uBridge ,Luke 747 (EA); Morogoro,Wallace 47 5(EA ,FHO ,K). T 8: LindiDistr. ,Mchinjiri ,Rond oPlateau ,Semsei 67 9 (BR, EA, K). Z: 27 kmfro m Chwaka, Faulkner 3022 (BR, P,UPS) ;Ufufuma , Vaughan 1777 (EA, K); Kizi Kazi Cave, Vaughan 1820(EA , K);Zanzibar , sin. loc., Sacleux145 7(P) . ZAMBIA. ChembeFerry ,Angus 21 8(BR , FHO, K, MO). Nothern:Brenan & Greenway 801 1(BR ,EA , FHO, K). ZIMBAMBWE. Melsetter (= Chimanimani), Swynnerton 21 (PRE);Chirind aForest , Goldsmith38/6 5 (BR,COI ,K ,MO ,PRE ,SRGH) ,Swynnerton 14, (BM, K,SRGH ,US ,Z , typeo f P. swynnertonii). MOZAMBIQUE.Zambezia :Goben eForest,7bni e& Correia 1454 1(COI ,LISC) , 16199(LISC) , 17015 (LISC).Manica :Serr aMacuta ,Muller & Gordon178 4(LISC ,SRGH) ;Chibabava ,Carvalho 1228(BR) . Sofala: Gorongosa National Park,alon g UremaRd. ,Tinley 2718 (K, MO, SRGH);Chiniziua , Gomes & Sousa440 7 (COI,K ,MO ,PRE) ;2 5k mfro mLacerdonia ,Müller & Pope 1916 (LISC,K ,PRE ,SRGH) ; Serracaod e Corboso,Torre 5904 (LISC). Gaza:Macia ,Balsinas 1453 (LISC, WAG).

Notes.P. pycnantha ca neasil yb edistinguishe dfro mth eothe rPleiocarpa specie s byth epresenc eo fa tleas ton e nodeo never ybranchle twit hleave si nwhorl so f3- 5 and from theternat e P. mutica specimens by themor enumerou s secondary veins and shortercoroll alob ean dtube .Th e carpels arealway s2 excep ti nLebrun 9898 where 3-carpelled ovules arereporte db y Pichon (1953).Th e numbero f ovulesi s almostalway s 2 except'rnTroupin 402 4WAG , andGoldsmith 38/65 ,SRGH ,wher e four ovuleswer efound ,bu tth e leavesoccu ri nwhorl so f 3-5. The habitan dsiz eo f the corolla tube and lobes is greatly affected by the habitat. Plants growing near rivers tend tob e small shrubs, 2-3 (-6) mhig h withbar k of thebranchlet s easily peelingoff .Th eflower s are verysmall ,wit hver yshor tcoroll alobe san dofte nhav e anunpleasan t odour (Breteler 2203,Letouzey 4589,Tisseront 1461& 2412).Al l thesespecimen s werepreviousl y placedunde r var. tubicina. All the materialfro mUsambar aMts. , citedbelow ,hav eopposit eleave sunlik e the resto f the materialan dal lflowers analyze dfro mthi sregio nhav e4-ovulat ecarpels . Moremateria li s neededt o verify whetherth e leaves are alwaysopposit eo r not, and toverif y thetaxonomi epositio no fthes especimens .

TANZANIA.T 3: Tanga Distr., Usambara Mts. Bulwa, nearAmani , Zimmermann G7712 (NY),Peter 58222 (B); Kwamkoro,Peter 58229 (B, WAG), 58267 (B), 58305 (B),Zimmermann 27.12.1916 (K),G 7724 (EA);Amani , East Usambara Mts.,Botany Students 25 2 (WAG),Peter 58199 (B, WAG); Mlinga Peak, EastUsambaras ,Drummond & Hemsley 1449(B ,BR , EA,K , S), Greenway490 5 (EA, FHO,K) .

2.7.5 Pleiocarparostrat aBenth . in Hook. Ic. PI. 12: 71, t. 1182 (1876); K.Schuman n in Engler & Prantl, Nat. Pflanzenf. 4, 2: 135, fig 52 E (1895); Stapf in Fl. Trop. Afr. 4, 1: 98 (1902); Hutchinson &Dalzie l in Fl.W.Trop .Afr . ed. 1,2 : 38(1931) .-Type : Cameroun, WouriR. ,Mann 72 0(holotyp eK ,photo . MO). Fig. 40, p. 153; map 18, p. 154 Homotypicsynonyms : Hunteria rostrata (Benth.) Hall. f. in Jahrb. Hamb.Wiss . Anst. 17, Beih 3: 194 (1899).Carpodinopsis rostrata (Benth. )Picho ni n Bol. Soc.Brot . sér. 2,27:147, t.5, 4 p. 145.ma pc , p. 121 (1953).

152 WageningenAgricultural University Papers 96-1 (1996) Figure40 .Pleiocarpa rostrata. 1 and5 habi t( x2/3) ;2 an d6 ,opene dcoroll a( xs) ;3, 4an d7 ,pisti lwit h openedovar y( x 10).1- 3 fromTalbo t i037;4fromLeTestu 8871;5- 7fro mZenke r4362 .

WageningenAgricultural UniversityPapers 96-1 (1996) 153 Map 18.Pleiocarpa rostrata.

Heterotypic synonyms:

Pleiocarpa fa/èoftï Wernham in CatTalb. Nig. pi. 62 (1913); H.Huber in F.W.T . Aed .2,2:6 4 (1963),syn .nov .Carpodinopsis talbotii (Wernh.)Picho ni nop .cit . 144,t .5,1-3 ,ma pC ,p . 121.-Type :Nigeria ,Oba ndistrict ,Talbot 103 7(holotyp e BM; isotype K). Carpodinopsis unifloraPicho n op. cit. 148,t . 5, 5-7, syn. nov.-Type : Cameroun, Bipindi,Zenker 1181-b(holotyp eP ;isotype sBM ,E ,G ,HBG ,K ,L ,M ,W ,WU) .

Shrub or small tree 0.6-5 m high with a semi-scrambling habit and white, sticky latex.Trun ku pt o4 c mi ndiameter ;bar k smooth,ligh tt odar kbrown , purple-brown ordar k grey,inne r bark palegree n or creamt o yellow, wood dark cream. Branches brownt odar kolive-brow n orgrey ;branchlet s thin,quadrangula r whenyoung , dark brown, with a few lenticels. Leaves opposite; petiole 5-14 mm long; blade cori­ aceouso rsubcoriaceou swhe nfresh , broadlyt onarrowl y elliptict oobovate ,2.1-3. 5 x aslon ga swide , 10.6-28 x 3.8-11.3cm ,acuminate ,rarel y obtuse atth eapex ,acu ­ men 4-12x3 mm, acutea tth ebas e ordecurren t intoth epetiole ;midri b prominent beneath; secondary veins oblique,ver y prominent beneath, in7-1 2pairs ,formin g a submarginal vein onth eentir eo ronl y theuppe r0.3-0.7 5o f thelea f and anangl eo f 40-580 withth emidrib , 12-37m m aparta tth ecentra lpar to f theleaf . Inflorescence axillary,rarel y terminal,ramiflorous , 1.5-3.5x 2"4 cm,abou t 1-4-flowered, sessile. Pedicels 0.5-5 mm long.Bracts very small, scale-like. Flowers: Sepals pale green, free orconnat e atth eextrem ebase ,erect , membranaceous atth eedg ean dth e apex, ovate, 1-2.7x aslon ga swide ,2-4. 5 x J-2-3 ^ coriaceous ornot ,acut et o obtuse at the apex. Corolla white,cream y to pale green in bud, 13-37nu n long in mature bud and forming acomparativel y wideoblon g head 0.25-0.43 of thebu d length, 6- 15x 1.5-4mm , with a rounded apex, with a belt of pubescence 3-10.5 mm wide insidejus tbelo w theinsertio n of stamens;tub eyello w atth enarro wthroat ,thic k or not, 4-16 x as long as the calyx, 1.4-2.9x as long asth e lobes, 12-32 mm long, al­ most cylindrical, 1-3 mm wideabov eth ebase ,narrowe d below the insertion of the

154 WageningenAgricultural UniversityPapers 96-1 (1996) stamenst o0.5-2. 0 mm wide,agai nwidene d aroundth eanther s to 0.5-3.5m mwide ; lobesobovat e to elliptic,0.3-0. 9x a slon ga s thetube ,1.6-3. 1x a s longa s wide,5.5 - 26x 4-1 5mm ,rounded , spreading torecurved . Stamens with apex 3-8 mmbelo w themout h ofth ecoroll atube ,inserte d 0.58-0.74o fth elengt ho fth ecoroll atube ,a t 7-18m mfro m thebase ;filaments 0.5- 1 mmlong ;anther s ovatet oelliptic , 3.3-7 x as long as wide, 1.8-4 x 0.4-0.9mm , acuminate at the apex, acumen 0.2-0.5m m long, sterile,cordat e atth ebase .Pistil 10-13. 8m mlong ,wit h apex 1-5 mmbelo w thestamens ;ovar y subglobose, 0.9-1.5x 0.6-1. 5x 0.4-0. 9mm ,abruptl y narrowed intoth estyle ,wit h adisk-lik e thickening uniting thecarpel s atth ebase ,abou t0.3 - 0.5 of the length, 0.3-0.8m mhigh , of 3-5 separate carpels; style6.5-1 1x 0.2 mm long;pisti l head ellipsoid to oblong 1.5-3.4* 0.2-0.4mm -Ovule s 3-4(-6)i n each carpel. Fruits bright orange, waxy, smooth, minutely tubercled to rugose, of 3-5 separatemericarps ;mericarp s narrowly ellipsoid, ovoid to slightly crescent-shaped, 20-50 x 15-25x 10-13mm » rounded orwit h astraigh t tohook-shape d beak at the apex, beak 4-15x 2-4mm ; 1-5-seeded;wal l about 2-4m m thick. Seeds orange, smooth, 14-17x 8-11x 4-5 mm.Embry o 9.5 mm.long ;cotyledon s elliptic, 1.8 x aslon ga swide ,5. 5 x 3 mm.obtus e at the apexan da t thebase ,radicl e0. 7 xa slon g asth ecotyledons , 4x 0.5 mm.

DISTRIBUTION. Nigeria, Cameroun and Gabon. ECOLOGY. Denserai nforest , primary orsecondar y forest alongrivers ,creek san d swampsi nclayis h andsand y soil.Alt .0-140 0 m. Flowering andfruitin g inNovem ­ bert o January.

USES. Coldinfusio n used against stomach ache (Watts 954,Cameroun , 1993).

Geographical selection of the approximately 60specimen s examined: NIGERIA. CrossRiver :Akamp aRubbe rEstate ,Latilo FHI40918 (K);Oban ,Talbot 1037 (K,P ,typ e of P. talbotii), 1313(K) ;Oba nFores tReserve ,Onochie FH I3629 3(FHO ,K) ;Nort hFores tReserve , Van Meer 1725 (WAG); Ayuk,Latilo FHI6775 5 (K);Britis h Obokum,Keay FH I 28277 (K). CAMEROUN. Sud-Ouest:Mbilishe ,D. W.Thomas 744 4 (MO, WAG); EjaghamFores tReserve ,Satabie 743 (YA); Bokossi Mts.,D. W.Thomas & Mcleod 5339(MO ,P ,WAG) ; SBakund uFores tReserve ,Keay FHI 28571 (K, P); near Fabe, Manning91 0 (MO); Korup National Park, D. W. Thomas 4136 (MO,P , WAG);bigEkombe Forest,Nemba &D.W.Thomas450(MO); Même, Wheately 691 (K);Idenau , Watts 954 (K); Etinde, Cable 289 (K);Limbe , Wieringa204 0 (WAG). Ouest: 40 km WNW of Ndikinimeki, Letouzey1122 9(P , WAG). Littoral: 15k m No f Yabassi,Letouze y 14421(P) ; 5k mSS Eo fNkongsamba , Letouzey 14483 (P.WAG); Likomba plantation, Mildbraed 10692 (A); Wouri R., Mann72 0 (K,Type), Mann218 9 (K);5 0k mN Wo f Eseka,De Wilde &De Wilde-Duyßes124 5 (K, P,WAG);3 0k mEN Eo f Edea, Letouzey 12378 (YA);Betwee n Dibamba andEbo , Letouzey 11028 (BR, K, P,YA,WAG); 7 km E oYingui , Leeuwenberg909 1 (WAG). Centre-Sud:Dibombe-Mabomb e ForestReserve ,Beentje 1501 (WAG);1 5k mN o f Eseka,J. deWilde etal . 3887 (MO, P, WAG); Eseka,Bamps 130 7(BR ,WAG) ; 2k m N of Longii Forest,Bos 491 6 (WAG); Bipindi,Annet 35 5 (P),Zenker 1181b (E ,G ,HBG ,K ,L ,M , P,W , WU, Z,typ eo f Carpodinopsisuniflora), 4362 (B,BR , E,G , K,L ,M ,MO ,P , S, W, Z); 8k mS o f Kribi, Bos 3968 (WAG);Njabilobé , Raynal 10403 (P);Mamb e Forest,Letouzey 12275 (P, YA); 29 kmN W of Eseka,De Wilde & De Wilde-Duyßes127 8 (B,K ,MO ,P , WAG). GABON. Boudinga, LeTestu 8871 (P).

WageningenAgricultural University Papers 96-1 (1996) 155 Notes.Th esecondar y veinsma yfor m asubmargina l vein alongth eentir eedg eo r only in the upper 0.3-0.75 of the leaf.Th e corolla tube and lobes are very variable in size and in Beentje 1501 both small and large flowers are found on the same branch.Th e carpels also vary from 3-5, D. W.Thomas & McLeod 5339 has 4-5- carpellateovarie so nth esam ebranch .Th efruit s areals over yvariabl ei nsize ,shape , texture and at the apex. Watt60 8 and Wheatly 691 have rugose fruits which are smoothtoward sth ecentre ,th efruit s mayals ob eentirel yrugos eo rsmoot han dma y have a rounded to rostrate apex.Th e inflorescence always has very few flowers, rarely more than four.

2.8 DOUBTFUL SPECIES

Pleiocarpa picralimoides (Pichon) Omino, comb. nov.

Basionym:

Carpodinopsis (?)picralimoides Pichon in Bol. Soc. Brot. sér. 2, 27: 143, t. 4 fig. 2(1953) .-Type : Angola, Cabinda, nearBelize , Gossweiler 7994(LISU ;isotyp e K).

Shrub (sarmentose) to 2 m high.Trun k very slender. Leaves opposite; petiole 5-10 mmlong ;blad eelliptic ,acuminat e atth eapex ,cuneat ea tth ebase ;secondar y veins in 11-18pairs , forming a submarginal vein and an angle of 60-700 with the costa. Flowers unknown. Fruits yellow to orange, smooth to rugose, of 4-5 separate mericarps,mericarp s 14-30x 9.5-1 6mm ,rounde d atth e apex, 3-4 seeded, wall 2-5 mm thick. Seeds variously shaped, 11-15 x 6.2-10 x 4-8 mm.

DISTRIBUTION. Angola. ECOLOGY. Forest, sometimes riverine. Alt. low.

Other specimens examined: GABON. Moyen-Ogooué. Lac Ezanga,N.Hallé 2087 (P). CONGO. Kouilou:Mayumbe ,Thollon 1139 (P); ibid.,Bitsindou 8 Jan. 1977 (P); BetweenPoung aan d Dimonika, Cusset54 6 (P);Dimonika , Dechamps 13016 (BR);ibid. , De Foresta88 3 (P);ibid. ,F.Hallé 1851 (P);ibid. ,Moutsambote 68 (WAG); ibid., Sita469 8 (WAG):Cofibois , De Foresta 1184(P) .

Notes.Picho n (1953) first described the above species under Carpodinopsis which is currently a synonym of Pleiocarpa. Since then more specimens have been col­ lected in Angola and Congo, but unfortunately all are either fruiting or with very immatureflower buds .I ti stherefor e notver y easy tocommen t onth e currenttaxo ­ nomie position of this species which is definitely aPleiocarpa. In my opinion it is not close to P. mutica which always has one ovule per carpel as opposed to its 4 ovules per carpel. It is, however, very close to P. rostrata which also has 3-4 (-6) ovulespe rcarpe lbu tha s9 (-12 )strongl ycurve d andwidel y spaced secondary veins

156 WageningenAgricultural UniversityPapers 96-1 (1996) asoppose dt oit s 11-1 8pair so f slightlycurve dsecondar y veins.Perhap s whenflow­ ering specimens become available it will be easier to tell how close it is to P. rostrata.

2.9 EXCLUDED SPECIES

Hunteria atrvvirensWall, ex G.Don, Gen. Syst. 4: 105 (1837) = Chilocarpus denudatus Bl. Hunteria coriacea Wall ex G.Don, Gen. Syst. 4: 105 (1837); A.DC.Prod. 8: 350 (1844) =Alyxia coriaceaWall , ex Roxb. Hunteria? cuspidataWaü. exA.DC .i n Prod. 8: 350 (1844)= Melodinus orientalis Bl. Hunteria eugenifoliaWaü ex G.Don, Gen. Syst 4: 105 (1837) = Wrightia arborea (Dennst.) Mabberley. Hunteria fascicularis G.Don., Gen. syst. 4: 105 (1837); A.DC in Prod. 8: 350 (1844) =Alyxia fascicularis Benth. Hunteria gracilisWail, ex A.DC, Prod. 8: 350 (1844) =Alyxia gracilis (Wall, ex A.DC. ) Benth. Hunteria sundana Miq. in Fl. Ind. Bat. 2:40 9 (1856) = Rauvolfia serpentina (L.) Benth ex. Kurz.

WageningenAgricultural UniversityPapers 96-1 (1996) 157 2.io INDEX OF EXSICCATAE

Hunteria ballayi (ba),H. camerunensis (ca),H. congolana (co),H. densiflora (d), H. ghanensis (g), H. hexaloba (h), H. macrosiphon (ma), H. myriantha (my), H. oxyantha (o),H. simii (s),H . umbellata (u),// . zeylanica (z);Picralima nitida (n); Pleiocarpa bicarpellata (bi),/ ? brevistyla (br),P .mutica (mu),P .pycnantha (p), £ rostrata (r).

Adam,J.G.,59(s),6549(mu), 13606(p),i3686(p), 16636(5), i6708(s), 16759(5), 17334(u) . 17361(u) , 17937(u) ,2139 5(s) ,2141 4(r) ,2142 2(s) ,2160 6(s) ,2414 0 (s), 25768 (s), 26048 (s), 27452 (s), 27599 (s).2850 0 (s), 28608 (s), 28733 (u), 30466 (p),3029 4 (s), 30299 (s). Adams, CD., 3789 (p);433 1 (p). Adamson, J., 22 (z), 34 (z), 37 (z), 224 (z). Adebusuyi, J.K., FHI43383 (mu). AkéAssi ,L. ,747 8 (g),973 8 (u), 11578(u) , 11222(g) , 12676(s) , 13665(u) , 13763 (g), 15143 (n), 15904 (u), 17243 (mu). Akpabla, G.K 110(u) ,79 2 (mu). Alymer, G., 35 (mu),21 6 (mu). Ament, J., 169(p) , 388 (z). Andrada, E.C . da, 1062(z) . Andoh, J.E., 3335 (n),422 2 (n),430 5 (mu),FHI547 1 (u),563 9 (mu). Angus, A., 218 (p). Annet, E., 355 (r). Ariwaodo, J.O., FHI 88715 (mu),FH I 89126 (r). Asonganyi, J.N., 128 (n),41 4 (bi). Attims.Y., 98 (ma). Aubréville, M.A. , 23 (u),9 5 (s), 350 (mu), 1342 (mu), 1343(u) , 1383(n) . Bagshawe, A.G. , 1086(p) . Bakare, FHI 22857 (")• Baldwin J.T.Jr., 6991 (mu), 7013 (mu), 9460 (s), 10443 (s). 10736 (s), 10814 A (mu), 11552 (mu), 12561 (s). Ballay, M., May 1906(ba) . Bally, P.R.O., 2042 (z), 2157 (z),471 2 (z),591 9 (z),606 8 (z), 16718 (z). Balsinhas, A., 1453(p) . Bamps, P., 225 (p), 261 (p), 1307 (r), 1351 (r), 2035 (mu), 2047 (mu), 2304 (s), 2388 (s), 2559 (mu). Barker, A.J.D. , 1139(s) . Barret B.,BJ H 6542 (z). Bartin, D., 1858(p) . Bates, G.L., 368 (bi), 1421(n) . Battiscombe, E., K 226 (z). Beentje, H., 1450(ca) , 1501 (r),438 1 (z). Begné, 3120 (mu). Benoit,M. ,14 9(n) ,23 2(n) ,23 5(n) ,25 6(n) ,32 6(n) ,38 2(u) .

158 WageningenAgricultural UniversityPapers 96-1 (1996) Bequaert,J.C. ,14 8(mu) ,162 3(p) ,213 4(p) ,222 9(p) ,229 8(p) . Berhaut,H .J. ,166 9(u) ,208 1(u) ,208 2(u) ,407 9(u) ,412 1(u) ,412 4(u) ,429 2(u) , 4801(u) ,480 2(u) ,575 0(p) ,576 6(p) ,597 4(p) ,676 2(p) ,692 2(p) ,714 3(p) . Bernadi,L. ,813 8(mu) ,852 2(mu) ,855 0(mu) . Binuyo,A. ,FH I3509 0(ca) ,FHI4124 0(p) ,FHI4129 7(n) . Bolema,D. ,32 8(co) ,70 9(p) ,108 2(co) . Bond,W. ,983 7(z) ,988 3(z) ,1085 0(p) . Boone,A. ,1 1(p) , Bos,J .J. ,202 8(s) ,306 5(u) ,396 8(r) ,450 0(r) ,491 6(r) ,497 7(u) ,497 8(u) ,501 8 (r),610 2(u) ,621 5(bi) ,630 1(bi) ,637 3(n) ,650 2(bi) ,672 5(bi) ,686 8(bi) ,710 4 (r),710 6(u) ,742 1(u) . Bouquet,A .67 2(n) ,81 9(n) ,94 6(u) ,162 9(ba) ,174 8(n) ,191 7(n) ,197 7(u) . Bowbrick,J.M. ,J C1 0(z) . Brenan,J.P.M. ,801 1(p) ,891 0(n) ,940 1(ca) ,940 1A (ca) ,940 1B (ca) ,945 8(bi) . Breteler,F.J. ,636/7 8(p) ,748/7 8(ca) ,130 8(n) ,220 3(p) ,265 0(bi) ,521 5(mu) , 5354(mu) ,563 5(mu) ,588 6(mu )600 5(mu) ,612 0(mu) ,620 7(mu) ,655 8(mu) , 7388(s) ,739 8(s) ,759 2(n) ,865 4(ca) ,987 6(ma) ,992 0(ma) ,993 4(ma) ,999 3 (u),1126 7(ma) ,1129 9(o) ,1135 1(ma) ,1135 8(mu) ,1323 3(o) ,1333 8(mu) . Breyne,H. ,79 0(p) ,231 8(p) ,413 1(ma) . Buch,W.R. ,62/19 3(z) . Burstyn, P.,75/8 3 A (z). Butaye, R.P., oct. 1900(p) . Buetner, 621 (p). Caballé, G., 364 (ca),36 5 (o),40 7 (ca),41 4 (bi). Cable, S., 289 (r), 514 (r). Casier, P.,41 2 (p). Causdale, G.S., 166(u) . Chandler, P.212 5 (p). Chapman, J.D., 4581 (p). Cheek, M., 5504 (r). Chevalier,A. ,(p) ,59 7(p) ,508 4(p) ,1079 3(p) >1093 0(p) ,1380 4(mu) ,1384 5(p) , 13928(u) ,1396 6(u) ,1518 1(u) ,1519 8(u) ,1519 9(u) ,1520 0(n) ,1546 2(u) , 15600(n) ,1560 4(u) ,1631 0A (u) ,1631 0B (mu) ,1637 3(u) .1651 9(u) ,1658 1 (u),1664 0(u) ,1667 1(u) ,1687 1(u) ,1704 3(u) ,1745 1(mu) ,1765 9(u) ,1776 6 (u),1786 2(mu) ,1786 5(u) ,1786 9(mu) ,1795 2(mu) ,1903 6(s) ,1931 1(s) ,1936 7 (s),1936 8(s) ,1943 2(mu) ,1943 4(mu) ,1951 9(mu) ,1953 5(s) ,1957 2(mu) , 19706(u) ,1972 2(mu) ,1993 4(mu) ,1994 8(u) ,1997 5(u) ,1999 4(mu) ,2001 1 (mu),2001 2(mu) ,2001 3( u)>2004 4(mu) ,2239 1B (n) ,2244 0(n) ,2284 4(p) . 22873(P) .3416 4(mu) ,3424 9(mu) . Christiaensen,A .R. ,40 9(n) . Claessens,J. ,6 6(n) ,20 9(p) ,25 8(p) ,75 6(n) . Corbisier,A. ,10 3(n) ,72 0(n) ,72 6(n) . Cremers,G. ,51 8(n) ,78 3(u) ,86 8(u) . Cook,A.C. ,81 5(mu) . Cooper,G.P. ,16 9(s) ,24 8(mu) .

WageningenAgricultural UniversityPapers 96-1 (1996) 159 Dacremont, A., 329 (p). Dale, LR., 487 (p), 856 (p), 1077 (z).Dalziel, J.M., 8240 (r). Dang, D.,61 4 (bi). Daniel, P.M., 238 (s). Dawe, A., 707 (n),71 9 (n). De Briey, C, 257 (n). De Bruijn, 922 (n). Dechamps, R., 5 (my), 139(my) , 171 (n), 242 (my). Declercq, A., 32 (my). Decloix, G., 168(p) , D'Elzius, C, 499 (p). Deffet, 10(p) . De Carvalho, M.F., 1228 (p). De Giorgi, S., 241 (p), 1339 (P). 1672 (n), 1731 (p). De Graer, RA.M., 237 (p). Deighton, F.C., 2501 (mu), 3270 (mu), 3869 (mu), 3871 (mu)4348 (s),501 0 (u). Dekker, A. J.F.M., 84 (mu). De Kruif, A.P .M. , 4 (n), 5 (u). Demeuse, J., 105 (n),42 7 (mu). De Namur, C, 1423(s) . D'Orey, J., 376 (u). Deru, 344 (p). De Saeger, H., 122G (p), 1611 (p). De Schlippe, 267 (p),29 9 (p). Deville, A., 22 (p),24 6 (p),29 0 (p),48 8 (p). Devlaux, J., 409 (p). Dewèvre, A., 847 (n),94 5 (p), 1113 (n). Devred, R., 1779 (p),402 7 (co),419 3 (co). DeWilde ,J. , 393 (n). DeWilde ,J .J.F.E. ,81 2(u) ,99 3(mu) ,320 4(mu) ,754 8(mu) ,815 0(mu) ,837 1(u) , U375(br). DeWilde ,W.J.J.O. ,30 7(mu) ,124 5(r) ,127 2(u) ,127 8(r) ,144 2(ca) ,149 8(ca) , 1498B (ca) ,179 8(bi) ,179 8B (bi) ,195 2(n) ,196 4(mu) ,196 4B (mu) ,197 4(ca) , 1974B (ca) ,203 2(mu) ,206 2(bi) ,210 6(r) ,219 5(ca) ,242 0(p) ,286 3(u) ,290 1 (ca),292 2(u) ,388 7(r) ,1045 4(ma) . DeWit ,H .CD. ,A 285 0(p) ,943 5(mu) . DeWitte ,CF. ,419 8(p) ,694 2(p) . Dibata,J. ,95 6(n) . Dinklage,M. ,108 8(r) ,143 2(r) ,297 6(mu) . Donis,C ,16 5(n) ,213 4(n) ,273 8(co) ,283 7(p) ,300 2(p) ,306 3(p) ,312 2(co) , 3129(p) ,340 5(p) ,356 1(p) ,357 9(p) . Doumenge,54 6(r) . Dowsett-Lemaire,F. ,144 7(u) ,183 3(n) . Drummond,J.H. ,356 3(z) ,365 5(z) ,454 6(p) . Dubois,J. ,6 3(p) .

160 WageningenAgricultural UniversityPapers 96-1 (1996) Dubois,L. , 401 (p),65 1 (co), 851 (co). Earle,P.G., 5PID(z). Edwards, L.C., E 101 (z), 178 (z). Edwardson.T.E., 41 (u). Eggeling.W.J. , 517 (p), 3115 (p), 3854 (n),397 2 (n),404 2 (n). Ejiofor, M.C., FHI14010 (n),FHI2689 7 (")• Else, 10(z) , 16(z) . Emwiogbon,J .A. , FHI 33321 (p),FH I 45328 (n),FH I 61669. Enti,A.A. , R 973 (u),R 1168(mu) ,F E 1203(u) ,F E 2175 (p),229 6 A (mu), 2296 B (u) SP39 3 (p),F E 1875(n) ,F H 6920 (u),F H 7544(mu) ,G C 38392 (mu),G C 38426 (u). Espïrito Santo,J .V .d a Graca do, 1940(u) , 1941 (p), 1919 (p), 2042 (p), 2257 (u), 3001 (u), 3016 (u), Santo (u). Etuge,M. , 207 (r),41 9 (bi),49 9 (n). Evrard,C , 327(p) ,48 1(p) ,57 1(p) ,78 5(n) ,161 4(u) ,317 8(n) ,356 2(p) ,408 2(p) , 4306 (n),435 4 (co), 5482 (co),567 5 (d) 6119 (p),627 0 (co). Faden, R.B., 70/200 (z),74/32 1 (z),77/64 8 (z),77/75 1 (p) 77/760 (z). Fanshawe, D.B., 3904 (p),467 6 (p). Farmar, L., 540 (n). Farron C, 4247 (n),436 2 (ma),482 8 (n),726 1 (bi),755 4 (ca). Faulkner, H.G. , 1280 (z), 1626(z) , 3022 (p). Fay,J.M. , 8127 (ba), 8782 (ba), 8835 (ba). Fischer, 202 (z). Flamigni, A., 6014 (p), 6123 (p),711 6 (n), 10383(n) . Fleury in Chevalier 33348 (bi), 33099 (u). Florence, J., 154, 514, 516, 566 (ca),58 4 (o),61 4 (o), 1026(ca) , 1798 (bi). Floret, J.J., 1633 (n). Forest Dept., EAH 12437(P) - Foster, E.W. , 5 (u), 182 (p), 220 (n). Fox, 10(u) , 11(u) . Frédoux, A., 46 (n), 182 (K),41 3 (mu), 584 (mu), 589 (mu). Gardner, H.M. , 2205 (z). Gamier, 652 A (u),65 3 B (u), 653 C (u),65 3 D (u), 653 E (u). Gatheri, G.W. ,79/6 7 (z). Gathy 1908 (bi). Gautier-Béquin, D.,73 7 (mu),79 2 (mu), 800 (mu),96 7 (n). Gbile, Z.O. ,FH I 20580 (p) Geerling, G., 1516 (mu), 2361 (mu), 2377 (mu), 2379 (mu), 2419 (u). Gentry, A.L., 32674 (u), 32703 (u), 32758 (u), 32793 (u), 32810 (u), 33027 (o), 33177A (o) ,3317 7B (ca) ,3326 4(o) ,3350 3(n) ,3350 6(o) ,3351 2(n) ,3373 6(o) , 52681 (mu). Georges,J. , 27814 (u). Gérard, P.H. , 1701 (p), 2746 (p),358 0 (p), 3689 (p), 3732 (p), 3796 (p), 3839 (n), 5542 (p). Gereau, R.E. 5253 (p),538 5 (p).

WageningenAgricultural UniversityPapers 96-1 (1996) 161 Germain, R., 279 (u),35 7 (u),36 2 (u),316 2 (p),524 4(n) ,776 4(u) ,815 0(p) ,818 9 (p), 7998 (co). Ghesquière, J., 2782 (p),495 7 (co), 8593 (co). Gilbert, G.H., 425 (n), 1013(p) , 1058(p) , 1066(co) , 1180(p) , 1279(p) , 1296(co) , 2050 (co), 2169 (p), 2347 (p), 5860 (z), 8366 (p), 9043 (p),923 4 (u),947 3 (P). 10147 (p). Gille, P., 100(n) . Gillett, J.B., 18905 (z). Gillman, H., 352 (p). Goldsmith, B.,38/6 5 (p), 144/67 (p). Gomes, A., 4407 (p). Goosens,V. ,251 6 (p). Gossweiler,J .72 9(p) ,113 9(p) ,625 8(n) ,625 8B (n) ,654 9(bi) ,664 8 (bi),723 2(n) , 8729 (p), 13682 (p), 14051 (p), 14153(p) . Graham, R.M., 1976, 2205 (z). Greenway, RJ. , 5077 (z), 8860 (z),923 7 (z). Greenwood, K., 12(z) . Guile, D.P .M. , ii20(P). Gutzwiller, R., 201 (p) 581 (p), n 17(p) , 1983 (p), 1995 (p), 2008 (p). Hack,R.B., i(p). Hall,J.B. ,294 4(mu) ,G C4208 1(mu) ,G C4374 1(g) ,G C4513 4(u) ,G C4704 4(g) , GC 47045 (g), GC 47096 (p),G C4511 7 (mu). Halle,N. ,64 1 (n),67 8 (p),377 3 (p),500 8 (bi),508 0 (bi). Hamilton, F.H., 93 (z). Hamilton, PH., 683 (p),73 8 (p). Hanie,A. , 74 (n). Hansen, O.J., 73 (z). Harley, R.M., 9596 (p),958 7 A (p). Harley.W.J. , 317 (mu). Harris, D.J. , 63 (ba),50 4 (n),54 7 (ba),379 7 (r),387 3W - Harris, B.J. , BJH 2828 (z),BJ H 3636 (z),555 1 (z). Hart,T.9i(n),374(p),508(p). Hawthorne,W. ,81 2 (z). Haxaire, C, 1345(p) . Hedin, L., 1332 (n), 1628 (n). Hendrickx, F.L., 3627 (p). Henrard, A., 331 (p). Hepper., EN., 2539 (mu). Herman, G., 2179 (bi), 2248 (bi). Hiepko, A., 2663 (z). Hladik, A., 1314(n) ,209 8 (n). Holland, J.H., 106(mu) . Hombert, J., 87 (n), 556 (n). Howes, H57(n). Hoyle, A.C., 725 (p),78 0 (p).

162 WageningenAgricultural UniversityPapers 96-1 (1996) Hulstaert,K.N. , 1306 (p). Ichakawa,6 0 (n). Irvine,RR. , 1759(p) ,217 3(mu) ,281 7(n) . Jacques-Felix,H. ,289 8(r) . Jaeger,P. ,675 0(n) , 10019 (u). Jansen,J.W.A. , 1255(mu) , 1780(s) ,211 2(mu) ,214 7(u) ,228 2(s) ,256 4(mu) . Jawse,D. ,D 6(z) . Jasi,A. , 10 (p). Jeanty,CE. , I34(p). Jespersen,J. ,8 6(p) . Jex-Blake,M. ,464 1(z) . Jones,A.P.D. ,98 2(n) . Jongkind,C.C.H. ,207 3(mu) . Johnson,W .H. ,62 3(p) ,61 6(p) ,80 3(p) . Jordan,H.D. ,203 7(s) ,204 6(s) ,212 0(u) . Jumah,J.A. ,1 (u) ,2 (u) ,3 (u) . Jungner,J.R. ,14 7(r) . Kaji,M. ,5 0(n) . Katende,A.B. ,K 142 9(p) ,212 7(n) . Kayasha,E. ,14 7(z) . Keay,R .W.J. ,FH I2570 5(u) ,FH I2825 0(u) ,FH I2827 7(r) ,FH I2857 1(r) ,FH I 36605(n),FH I3745 7(bi) . Kennedy,J.D. ,21 2(n) ,205 6(n) ,218 2(n) ,234 8(n) . Key,Mc ,5 7(bi) . Kibuwa,S.P. ,248 1(z) . King,E.L. ,13 2B (mu) . King,H .C ,23 1(u) . Klaine.T.J.,10 6(n) ,29 9(n) ,307 7(ba) ,315 8(h) . Kodambo.T.J.,18 5(u) . Koechlin,J. ,506 5(p) . Kokwaro,J .O. ,395 9(p) . Koufani,A. ,10 7(mu) ,14 3(ba) . Kouakou,42 5(u) . Kuchar,P. ,1343 8a(z) ,1347 8(z) ,1349 1(z) ,1350 5(z) ,1353 0(z) . Lane-Poole,CE. ,12 9(u) ,33 3(mu) . Latilo,M .G. ,19 6(s) ,FH I3175 4(p) ,FH I3175 8(p) ,FH I3184 0(u) ,FH I4091 8(r) , FHI5399 7(n) ,FH I5843 3(P) ,6055 6(p) ,FH I6775 5(r) . Laurent,M. ,137 6(p) . Lebrun,J. ,91 5(p) ,195 0(p) ,198 7(p) ,215 7(p) ,236 3(mu) ,247 9(p) ,268 1(mu) , 2835(mu) ,284 0(mu) ,437 8(n) ,494 5(mu) ,558 6(p) ,512 1(p) ,587 7(p) ,608 8 (p),626 0(p) ,989 8(p) . Leeuwenberg,A .J.M. ,238 6(u) ,617 6(n) ,909 1(r) ,916 6(r) ,1072 2(u) ,1074 0(u) , 10778(n) ,1077 9(n) ,1080 0(z) ,1104 2(p) ,1116 4(g) ,1120 8(g) ,1134 8(n) , 11451(bi) ,H9i4(p) ,1202 5(n) ,1202 6(u) ,1208 2(mu) ,1211 1(u) ,I2i45(mu) , 12219(mu) ,1230 2(mu) ,1246 4(u) ,1334 0(mu) ,1369 5(n) ,1405 4(mu) .

WageningenAgricultural University Papers 96-1 (1996) 163 Lejoly,J. ,347 9(P) ,508 5(p) ,86/60 1(p) ,193 3(p) . Lemaire,H. ,9 7(p) ,38 8(n) . Léonard,A. ,9 8(p) ,75 4(co) ,183 1(co) ,331 2(p) ,387 1(p) ,394 8(p) ,398 1(p) , 4704(p) ,523 0(p) ,569 0(p) ,587 1(co) . Léonard,J. ,145 5(co) . Leontovitch,22 8(p) . Lescrauwaet,E. ,190 4(n) . LeTestu,G. ,174 4(u) ,184 1(n) ,122 3(n) ,452 4A (bi) ,452 4B (p) ,456 6(p) ,473 8 (p),589 8(n) ,603 5(n) ,640 8(u) ,708 5(ma) ,742 9(n) ,802 5(u) ,855 8(ba) ,887 1 (r),946 1(p) . Letouzey,R. ,111 0(n) ,458 9(p) ,506 9(n) ,547 7(u) ,900 2(bi) ,985 0A (bi) ,985 0 B (mu),1038 3(ba) ,1049 2(p) ,1056 8(ba) ,1102 8(r) ,1122 9(r) ,1180 4(n) , 11963(p) ,1208 7(P) .1227 5(r) ,1228 3(r) ,1237 8(r) ,1277 7(bi) ,1278 4(mu) , 14421(r) ,1448 3(r) ,1512 4(r) ,1520 9(mu) ,1527 4(mu) . Lewalle,J. ,318 9(p) ,399 0(p) . Liben,L. ,402 0(p) ,183 5(p) . Linder,D.H. ,350 8(p) . Lisowski,S. ,1584 6(co) ,1712 6(p) ,4342 5(p) ,455o 8(p) ,455 H (p).4786 5(p) , 52344(p) . Louis,A.M. ,188 4(n) ,210 1(o) ,211 4(n) . Louis,J. ,30 5(co) ,44 0(p) ,66 7(p) ,72 4(co) ,100 8(p) ,108 3(co) ,145 5(P) .154 3 (p),158 8(p) ,231 4(co) ,247 6(co) ,251 5(p) ,262 2(co) ,315 6(p) ,380 9(co) ,386 1 (p),396 6(co) ,427 5(p) ,575 1(co) ,632 8(co) ,622 8(co) ,633 7(co) ,633 8A (p) , 6338B (co) ,634 0(p) ,648 9(co) ,782 4(p) ,829 3(p) ,842 9(p) ,967 7(p) ,972 2 (co),993 8(co) ,1024 3(p) ,1261 2(p) ,1289 8(p) ,1318 0(p) ,1344 5(p) ,1514 3(P) , 15669(co) ,1630 9(co) ,1638 5(p) . Lowe,J. ,380 7(mu) . Luja,E. ,29 2(p) . Luke,Q..TP R3 1(z) ,74 7(p) ,89 3A (p),140 5(p) . Lykke,A.M. ,35 6(p) . Macedo,A. ,125 8(z) . MacGregor,W.D.,52 2(u) . Maitland,T.D.,28 4(n) ,30 6(p) ,76 4(n) ,78 7(n) . Malaisse,E ,675 3(p) ,859 9(p) . Malchair,10 7(p) . Mandango,196 9(p) . Mann,G. ,71 0(n) ,72 0(r) ,121 3(bi) ,218 9(r) ,227 7(mu) . Manning,S.D. ,91 0(r) . Marmo,V. ,6 5(mu) . Martineau,23 5(mu) ,26 0(u) . Maudoux,E. ,5 (n) ,17 1(n) ,59 0(p) ,117 6(p) ,118 2(p) . Mbambi,5 4(n) ,7 0(n) . Mbatchou,G .T. , 209(n) ,21 6(bi) , 449(r) . MeAinsh ,89 0 (u). Medley,K. , 231 (z).

164 WageningenAgricultural University Papers 96-1 (1996) Menavanza,F. , 143(co) . Mendonga, F.A. , 4447 (p). Meyers, S., 195 (p), 262 (p), 290 (p), 297 (p). Mezili, P., 120(ba) . Michelson, A., 85 (0), 133 (p),73 6 (p), 231 (n),91 4 (co),92 1 (d). Mildbraed, J., 4038 (ba),413 8 (n),458 8 (p), 6098 (u), 8285 (n), 10692 (r), 10716 (ca). MMer.J.S. ,7893(11) . Mooney, H.F. , 7666 (z). Moonmaw, J.C., 1295 (z). Moor, A.W., 1151(u) ,224 1 (u). Mortehan, M.G. , 520 (p). Morton, J.K., S.L. 1268 (mu). Mosango, 431 (p),53 9 (p). Mosnier, M., 2537 (u),259 4 (u). Muchiri, J., 424 (z). Müller,T. , 1320 (p), 1784 (p), 1865 (z), 1916(p) , 1926(z) . Muwin, A.H. , 26 (u). Mwasumbi, L.B., 13861 (p). Nchami,V.C. , FHI14341 (u). Ndele, 759 (co),76 7 (p),77 1 (p). Nemba, J., 281 (r),45 0 (r),58 2 (u). Niyongere, L., 1/84(p) . Nolde, B.von , 493 (p), 646 (p),72 7 (p). Nsimundele, 937 (p), 1077 (p). Nsola, 1261(co) . Obermeyer, A.A., 37485 (p). Ohnson, M., 903 (p). Okafor, J.C.FHI 36886 (p). Oldeman, R.A.A., 642 (u),96 1 (mu). Olorunfemi, J., FHI 30579 (u),FH I 70757 (p). Omino,E .A. , 32 (p),6 3 (z),6 9 (p), 82 (bi), 89 (z), 157 (bi), 158(bi) . Onochie,C , FHI 4936 (u),FH I 33370 (n),FH I 36293 (r). Onyeachusim, FHI 48005 (n). Orokoro, 228 (p). Osmaston, H.A., 684 (p),264 4 (n),276 7 (p). Osseyemeh, J., 3341(n) . Patel,M .B. , 51694(p) . Paoli, G., 815 (z),47 8 (z), 850 (z). Pardy, A.A., 2 (p). Paulo, S., 133(z) ,60 7 (p),61 9(p) . Perdue, R.E., 10012 (z). Pereira,J .A. , 1889 (u), 2704 (u), 3163(u) . Pierlot,R . 947 (n), 1045(p) , 1844(p) > 1936(p) ,202 5 (p),218 1 (p),234 1(p) ,243 7 (p), 2523 (p),259 6 (co), 2811 (n), 3210 (p),318 4 (p),95 0 (co).

WageningenAgricultural UniversityPapers 96-1 (1996) 165 Pobéguin, H.H., 45 (n), 157 (p), 2015 (u). Polhill, R.M., 755 (z), 818 (z),491 6 (p). Preuss,R. , 44 (bi). Punch, C, 138 (u). Putman, P.T.L., 126 (p),45 7 (n). Pyne,C.T., ii5(u). Raynal, J., 9606 (ca), 10403(r) . Reitsma,J.M. , 144(z) , 159(z) ,97 6(o) ,89 9(n) , 1858(ca) , 1866(n) , 1952(u) ,271 7 (bi), 3051 (bi), 3132 (n). Reygaert, F., 125 (p), 884 (co), 1290(p) , 1326(p) . Richards RW. 3211 (u). Robertson,S.A. , 3506(z) ,366 8 (z),413 8(z) ,414 5(z) ,483 0(p) ,515 3(p) ,557 3(z) , 5669 (z). Roberty, G., 15718 (mu), 17318 (mu), 15721 (u). Ross R., 7 (n), (u),7 9 (u), 126 (u), 142 (u). Rudatis, H., 59 (u). Ruffo, CK., 1873 (p).212 9 (p). Sacleux, C.H., 482 (z),67 8 (z), 1457(p) . Samai, S.K., 522 (u). Sangai, G.W., 15731(z) . Sargos, 53 (u),7 5 (u), 85 (n), 180 (n), 244 (n). Satabié, B.,74 3 (r). Scassellati, 56 (z). Schaijes, M., 3119 (p). Schlott,R.L., 11(ba) . Schmitz, A., 3704 (p),733 4 (p). Schnell, R., 6067 (s). Schweinfurth, G., 3073 (p). Scott Elliot, G.F., 4288 (p), 5690 (u). Semsei, S.R., 679 (p),372 3 (z). Seret, F, 482 (p), 873 (n),89 3 (p). Sharland, R.E., 329 (p),s n 25.1.81 (p). Short, J., GC 47008 (u). Sim, D., 16(s) . Simâo, J., 1201 (z),273/4 8 (z). Simpson, B.L., 138 (z), 235 (z), 265 (z). Sita, P., 1345 (u), 1361 (o), 3075 (p),337 3 (ba), 3451 (ba),379 7 (p),404 8 (ma). Soward, 651 (n). Small, D.,40 2 (u),53 9 (s),64 9 (s),70 1 (mu). Smith, J., 75/36 (mu). Spire, 33 (p). Spjut, R.W. ,271 0 (z),274 5 (z),461 7 (z). Staudt, 93 (bi), 130(u) , 322 (mu), 573 (bi), 683 (bi),79 1 (bi). Stauffer, H., 1021(p) . Stoop-van de Kasteele, F.S.C., 38 (s), 171(s) .

166 WageningenAgricultural University Papers 96-1 (1996) Straub,F .C , 287 (s). Stuhlmann, 1216(p) . Suzuki, A., B 23 (p). Swynnerton, J.C., 14 (p), 21 (p),650 3 (p). Synnot,T.J., 410 (p), 1949(z) . Talbot, P.A., 219 (n), 1037(r) , 1313(r) , 1565 (mu), 1654(mu) , 1690(n) ,314 6(n) , 3256 (n). Tanner, R.E.S., 2463 (z), 3366 (z),372 2 (z). Tardelli, M., 434 (z). Taten, 174(p) . Tawse, D., D 6(z) . Tchouto, P.,71 2 (r),74 0 (mu), 879 (r). Thairu, N., 69 (z). Thiébaud, 203 (p). Thyssen, M.T. , 118 (mu). Thoiré, M., 796 (mu). Thomas,D.W. , 294(bi) ,61 7(u) ,232 0 (r),320 7 (u),331 3(n) ,413 6(r) ,417 8A (r), 4178 B (u), 4195 (u), 4575 (mu), 4588 (u), 4684 (n), 5205 (u), 5339 (r), 7202 (ba), 7432 (p), 7444 00,7694 (r). Thomas,N .W. , 19(u) ,95 4(mu) ,96 3(mu) , 1941(n) ,206 1(n) ,210 1(n) ,423 5(mu) , 4764 (s),500 8 (s),524 5 (u),524 5 B? (s) 10102 (u), 10283(u) . Thompson, B.,E 5245 (p). Thomson,W.C. , 38 (mu). Tinley, K.L., 2369 (z), 2718 (p). Tisserant, C, 509 (n), 1461 (p),241 1 (p),241 2 (p). Toilliez, J., 291 (mu), 384 (n). Toka, L., 57 (n), 163(n) . Tondeur, 35 (n). Toree,A.R. , 5904 (p), I454I (p). 16199(p) , 17015(p) . Tou, 70 (p). Trilles, 118(h). Trochan, J., 3467 (u). Troupin, G., 7 (p), 547 (p), 759 (p), 3515 (p), 3769 (p), 3897 (p), 3961, 3962 (p), 3973 (P)> 3975 (P).402 4 (p), 4282 (p),448 7 (n), 4587 (n), 6402 (n), 7277 (n), 7748 (p), 10025 (P). 10296 (p), 10944 (p), 10637 (P). 12151 (co), 12496 (p), 14693(P) - Turner, L., 160(p) . Ujor, E., FHI29271 (n),3016 9(mu) . Umwin, A.H., 26 (u). Van den Berghen, 5116 (p),566 1 (p). Van den Bossche, 31 (n). Van den Brande, J., 380 (p). Van der Ben, D., 1262 (p), 1433(p) . Van der Bürgt,X.M. , 48 (br). Van derMaesen , L.J .G , 5534 (ma), 5579 (ba).

WageningenAgricultural UniversityPapers 96-1 (1996) 167 VanMeer ,P.P.C. ,38 0(u) ,46 4(s) ,101 7(u) ,117 8(mu) ,145 6(mu) ,149 1(mu) , 1266(mu) ,187 5(u) . Vanderyst,H. ,931 4(p) ,982 7(p) ,10215 ,1043 2(p) ,1091 0(p) ,1100 8(p) ,1105 3 (p),11121 ,11174 ,1130 5(p) ,1199 7(n) ,1204 2(p) . VanEijnatten ,CL.M. ,126 6(u) ,151 5(u) ,151 9(n) ,152 6(n) ,169 8(u) . VanMeer ,P.P.C. ,79 2(n) ,172 5(r) . VanSetten ,K. ,17 9(u) ,41 5(mu) ,42 3(n) ,47 1(n) ,58 6(u) ,79 3(mu) ,79 8(u) . Vaughan,J.H. ,88 5(z) ,177 7(p) ,181 9(p) ,182 0(p) ,196 6(p) ;207 8(z) ,272 9(z) , 2783(z) . Verdick,E. ,59 4(p) . Vermoesen,F. ,8 2(p) ,28 7(n) . Versteegh,C ,58 6(u) ,68 8(n) . Vigne,C ,20 8(n) ,26 9(mu) ,86 4(n) ,166 0(u) ,195 7(u) ,197 5(mu) ,270 1(u) ,316 6 (mu),333 5(n) ,395 6(n) ,434 5(mu) . Villiers,J.F.,652(n). Voorhoeve,A .G. ,3 6(u) ,85 3(u) ,93 3(mu) ,114 0(mu) . Wachiori,P. ,4 1(p) . Wagemans,J. ,49 4(n) ,55 9(n) ,114 2(n) . Wallace,G.B ,5 3(s) ,45 0(p) ,47 5(p) . Wanters,M. ,168/8 0(z) . Watts,J. ,59 9(r) ,60 2(mu) ,60 8(r) ,66 7(mu) ,76 0(mu) ,80 6(mu) ,81 3(r) ,95 4(r) . Warnecke,0,48 1(p) . Welwitsch,455 1(p) ,598 1(p) . Wheatly.J.1,69 1(r) . Wieringa,J .J. ,85 6(mu) ,126 7(br) ,128 5(u) ,147 9(ca) ,204 0(r) ,207 0(bi) ,209 0 (u),215 6(u) ,250 2(br) ,252 4(u) . Wilks,CM. ,145 3(ba) . Winkler,H. ,66 0(n) ,103 5(u) . Wit,P. ,125 7(u) ,126 0(u) . Yafunga,F ,8 1(p) ,12 9(p) . Zenker,G. ,11 7(bi) ,16 3(ca) ,24 2(bi) ,28 3(n) ,II28 7A (n) ,II28 7B (u) ,61 1(bi) , 753(bi) ,118 1b (r) ,121 1(ca) ,162 0(ca) ,165 8(bi) ,166 0(bi) ,170 7(u) ,172 8 (bi),172 9(u) ,231 1(ca) ,250 1(bi) ,285 8(bi) ,289 3(bi) ,302 8(n) ,316 3(ca) ,361 1 (ca),367 6(bi) ,370 5(u) ,411 3(ca) ,423 1(bi) ,436 2(r) ,448 8(u) ,447 0(bi) ,456 4 (u),468 5(bi) ,488 2(bi) ,488 9(bi) ,Nov.189 6(bi) .

168 WageningenAgricultural UniversityPapers 96-1 (1996) INDEX OFSCIENTIFI CNAME S

New names are in bold face, synonyms in italics. Page number of principal entries are in bold face, those of figures and maps in italics.

Acokanthera G.Don 3,4, 8,9,10, 34, 38-40, 74 laevigata Kupicha 74 oblongifolia (Höchst.) Codd 38 oppositifolia (Lam.) Codd 5-7, 9,10, 35,36,40 , 74, 78 schimperi (A.DC. ) Benth 74 spectabilis (Sond.) Hook.f. 38 Adenium Roem. & Schult 9,11, 74 obesum (Forsk.) Roem. & Schult 4, 6-9,11,12, 36,37 , 74,78 AlafiaThouars 12, 75 barteri Oliv 13 caudata Stapf 75 erythrophthalma (K. Schum.) Leeuwenberg 75 landolphioides (A.DC. ) K. Schum 13 lucida Stapf. 75 microstylis K.Schu m 6,12,75, 78 multiflora (Stapf) Stapf 13 orientalis K. Schum. ex De Wild 75 scandens De Wild 13 zambesiaca Kupicha 75 Alafiinae Pichon ex Leeuwenberg 75 Allamanda L. blanchetii A.D C 35.3 6 cathartica L 36 nereifolia Hook 3 schottii Pohl 3 violacea Gardn. &Fiel d 35.3 6 Alstonia R.Br. 4, 74 boonei De Wild 74 Alstoniinae (G.Don ) K. Schum 74 Alyxia R.Br. 82 coriaceaWall , ex Roxb 157 fascicularis Benth 157 gracilis (Wall, ex A.DC. ) Benth 157 Alyxieae G. Don 74 Alyxiinae Pichon exLeeuwenber g 82 Amsonia Walt, orientalis Dcne 9 Anacardiaceae Lindl 37

WageningenAgricultural UniversityPapers 96-1 (1996) 169 Ancylobotrys Pierre 4, 8, 9,13, 34, 35, 39,41 amoena Hua 74 petersiana (Klotzsch) Pierre 7, 8,13,14, 36,40,74, 78 tayloris (Stapf) Pichon 5-7,13, 14, 38,74,78 Apocyneae Allorge ex Leeuwenberg 75 Apocynoideae 3,74 Apocyno-Nerium L 82 ApocynumL 3 cannabinum L 3 Aspidosperma Mart. & Zucc. quebracho-blanco Schldl 34 AspidispermatinaePicho n ex Van der Ploeg & Leeuwenberg 74 Atropa L. belladonna L 35 Baissea A.DC 14,75 leonensis Benth 75 leontonori Van Dilst 75 major (Stapf) Hiern 75 myrtifolia (Benth.) Pichon 4-7,9,14, 36, 37 ,39,40,75,78 viridifolia (K. Schum.) De Kruif 75 Cameraria L 82 zeylanica Retz 123 CarissaL 3,4 , 8,9,15, 34,35 , 38,40,74 bispinosa (L.) Desf. ex Brenan 74 edulis (Forsk.)Vah l 5,7,9,15 , 16, 74, 78 grandiflora (E.Mey.) A.DC 36 hirsuta Roth 35 macrocarpa (Eckl.) A.DC 36 spinarum L 35 tetramera (Sacl.) Stapf 4, 5,7-9,15, 16, 35, 36,40,74,78 Carisseae Endl 74, 81 Carissinae A. DC 74 Carpodinopsis Pichon 82, 134, 156 picralimoides Pichon 81,156 rostrata (Benth.) Pichon 134, 152 talbotii (Wernham) Pichon 154 unifloraPicho n 154 Carpodinus R.Br .e x G.Do n umbellata K.Schu m 118 Carvalhoa K.Schu m 3,16,74 campanulata K.Schui n 4-7,16,17, 36,74,78 Catharanthus G.Do n pusillus (Murr.) G.Don 3,35 ,36 , 38 roseus (L.) G.Do n 36

170 WageningenAgricultural University Papers ç6-i (1996) Cerbera L 74 manghas L 74 Cerbereae (Benth. & Hook, f.) Leeuwenberg 74 Chilocarpus BI 82 denudatus BI 157 Clitandra Benth 8,17, 39, 41, 74 cymulosaBent h 4, 5,7-9,17, 35,39 ,74 , 78 Combretaceae R. Br. 37 Combretum Loefl 37 Comularia Pichon 82, 88 camerunensis (K.Schum .e x Hall.f.) Pichon 88,95 Croton L. zambesiacus Muell. Arg 104 Dictyophleba Pierre 4, 9,18, 35,39 , 41,74 lucida (K.Schum. ) Pierre 5, 8, 9,18,40, 74, 78 DiplorhynchusWelw .e x Fic. & Hiern 18, 39, 74 condylocarpon (Muell. Arg.) Pichon 5-7,9,18, 74,78 Echites Jacq. peltataVéü 35 Ervatamia (A.DC. ) Stapf coronaria (Jacq.) Stapf 35 Funtumia Stapf 3,19, 41, 75 africana (Benth.) Stapf. 75 elastica (Preuss) Stapf 6-9,19, 75,7 8 Gynopogon Forst. lanceolatum (Wall.e x A.DC.) Kurz 123 Holarrhena R.Br . 20, 39, 75 antidysenterica Wall, ex A. DC 21 pubescensWall , ex G.Don 4-8,20, 21, 35,37 , 75,78 Hunteria Roxb 3,4, 8, 9, 21, 39,41, 74, 81-87, 88. 89, 99 africana K.Schu m 123 ambiens K.Schu m 136 afrovire/wWall. ex G.Don 157 ballayi Hua 83-86,90,91,92, 158 breviloba Hall.f. 148 camerunensis K.Schum . ex HaLLf. 83-85, 88, 89,93, 94,95, 158 congolana Pichon 5,7, 21, 22,74, 78, 83-86, 90, 96,97,98, 158 coriaceaWall .e x G.Do n 157 corymbosa Roxb 82, 88, 123 var.genuina Hall.f. 123 var. roxburghiana (Wight)Trime n ex J. Gamble 123 var. salicifolia (Wall.) Hall.f. 123 var.zeylanica (Retz.) HalLf. 123 cuspidataWaü. ex A.DC 157 densiflora Pichon 83,84 , 86,90, 99,100,101,112, 158

WageningenAgricultural UniversityPapers 96-1 (1996) IJl eburnea Pichon 118 elliotii (Stapf) Pichon 118 eugenifolia'Waü. ex G.Don 157 sect.Euhunteria Pichon 82, 88 fascicularis Benth 157 ghanensis Hall &Leeuwenber g 83-86, 89,102,103,104, 112, 158 gracilisWall ,e x A.D C 157 hexaloba (Pichon) Omino 81-89,I04> I05< IO^, 158 lanceolataWall, ex A.DC 123 legocii Livera 123 macrosiphon Omino 81, 83,86 ,90, 107,70S,109, 158 mayumbensis Pichon 118 myriantha Omino 81-85, 90,109, no, in, 112, 158 oxyantha Omino 81-86,90,112,113,114, 158 pleiocarpa Hall.f. 143 sect.Pleuranthemum Pichon 82, 88 pycnantha K.Schu m 83, 146 rostrata(Benth. ) Hall.f. 152 roxburghianaWigh t 123 simii (Stapf) H.Huber 83-86, 89,115,116,117, 158 sundana Miq 157 umbellata (K.Schum. ) Hall.f 22, 83-88,90,118,119,120, 158 zeylanica(Retz.)Gardn.exThw. . 8,21,35,74,78,81-90,122,724,725,127,158 var.africana (K.Schum. ) Pichon 123, 127 var.salicifolia (Wall.) Pichon 123, 127 Icacinaceae Miers 37 Ichnocarpinae (Benth. & Hook, f.) Allorge 75 Ichnocarpus R.Br, frutescens (L.)W .T .Aito n 35 IsonemaR.Br. buchholzii Engl 35 Kalanchoë Adans. fedschenkoi Hamet & Perrier 36 Landolphia P.Beauv 8, 9, 22, 39,41,74 buchananii (Hall.f.) Stapf 7, 22,23 , 74,78 calabrica (Stapf) E.A .Bruc e 23 dulcis (R.Br. ex Sabine) Pichon 23,3 5 var.barteri (Stapf) Pichon 23,3 5 eminianaHall . f. 74 kirkii Dyer 74 landolphioides (Hall, f.) A. Chev 74 owariensis P.Beau v 23,35 , 74 parvifolia K. Schum 74 watsoniana Romburgh 5, 7, 22, 23, 74,78 Landolphiinae K. Schum 74

172 WageningenAgricultural UniversityPapers 96-1 (1996) Lepinia Decne 82 Lochnera Rchb. ex Endl. pusilla (Murr.) K.Schu m 3 Macropteranthes F.Muel l 37 Malouetiinae (Muell.Arg. ) Pichon 75 Marantaceae Petersen 93 Mascarenhasia A.D C 3,23 ,35 ,41 , 75 arborescens A.DC 5, 6, 8,9, 23,75 ,7 8 Melodinus Forst 82 orientalis BI 157 Motandra A. DC 3,4, 9, 24, 25,41,75 guineensis (Schum. & Thonn.) A. DC 3-9,24. 35»75 ,7 8 Nauclea L. latifolia Sm 104 Neriinae Boiteau 74 Nerium L. oleander L 34. 37 Notonerium Benth 82 Oncinotis Benth 3,4, 25, 41, 75 glabrata (Baill.) Stapf ex Hiern 26, 75 gracilis Stapf 26, 35 hirta Oliv 35,40 , 75 nitida Benth 26 pontyi Dubard 75 tenuiloba Stapf 5, 6, 8,25, 37,75 ,7 8 tomentella Radlk 35 PeltastesWoods , peltatus (Veil.)Wood s 35 Picralima Pierre 3,9, 26, 39,41, 74, 81-87,128 elliotii (Stapf) Stapf 118 klaineana Pierre 83, 130 nitida (Stapf) Th. & H.Dur. 6, 26,36,74,78 , 83,84, 128,129,130, 158 umbellata (K.Schum. ) Stapf 118 Pistacia L. terebinthus L 37 Pleiocarpa Benth 3,4, 5, 9, 26, 27, 38,41, 74, 81, 82, 85,87, 134, 135, 156 bagshawei S.Moore 148 bicarpellata Stapf 7, 8, 26, 27,74,78 , 82, 83,135,137,138, 158 breviloba (Hall.f.) Stapf 148 brevistyla Omino 81-84,135,139,140,141,158 camerunensis (K.Schum . ex Hall.f.) Stapf 93 flavescens Stapf 148 micrantha Stapf 148 microcarpa Stapf 148 mutica Benth 28, 82-84, 135» 141. *42,143,144,156,158

WageningenAgricultural UniversityPapers 96-1 (1996) 173 picralimoides (Pichon) Omino 81,156 pycnantha(K.Schum. )Stap f . 7,26,27,28,74,78,82-85, !35.Mó .*47> 148,158 var. tubicina (Stapf) Pichon 28,146 rostrata Benth 28, 82-85, 134, 135,152,153,154, 156,158 salicifolia Stapf 142 simii (Stapf) Hutch. & Dalz 115 swynnertonii S.Moore 148 talbotiiWemh 28,154 tricarpellata Stapf 142 tubicina Stapf 146 welwitschii Stapf ex Hiern 148 Pleiocarpeae K.Schu m 87 Pleioceras Baill 75 orientale Vollesen 75 Pleiocarpinae (K.Schum. ) Pichon 3,74. 81-83, 85,87 Pleuranthemum Pichon 82,8 8 ballayi (Hua) Pichon 88,9 0 Plumeria L. rubra L 3,3 5 Plumerieae Endl 74 Plumerioideae K.Schu m 3,74 ,8 1 Polyadoa Stapf 82, 88 camerunensis (K.Schum . ex Hall.f.) Brenan 93 elliotii (Stapf) Pichon 118 (?) simii Stapf 115 umbellata (K.Schum. ) Stapf 118 Rauvolfia L 4, 28,38 ,74 ,8 2 caffra Sond 4, 7, 8, 28,29 , 36,74,79 mannii Stapf 5, 6, 28,29 , 36,74,79 mombasiana Stapf 6, 8,28, 29, 74,79 serpentina (L.) Benth. ex Kurz 157 volkensii (K. Schum.) Stapf 74 vomitoria Afzel 74 Rauvolfiinae Benth. &Hook . f. 74 Rhazya Dcne. orientalis A. DC 9 Saba (Pichon) Pichon 29, 39,74 comorensis (Boj.) Pichon 6-9, 29, 34,74,79 Schizozygia Baill 3, 30,74 coffaeoides Baill 5, 8, 9, 30,74, 79 Stephanostema K.Schu m 30, 75 stenocarpum K.Schu m 4, 5,30 ,39 , 75,79 Strophantus DC 31.75 barteri Franch 32 courmontii Sacl. ex Franch 5-8, 31, 35-38,75,79

174 WageningenAgricultural UniversityPapers ç6-i (içç6) eminii Asch. &Pa x 75 gracilis K.Schum . &Pa x 32 hispidus DC 32, 35,7 5 hypoleucos Stapf 4, 75 kombe Oliv 75 mirabilis Gilg 4, 75 petersianus Klotzsch 75 preussii Engl. &Pa x 32,75 sarmentosus DC 75 welwitschii (Baill.) K. Schum 75 zimmermannianus Monach 75 Tabernaemontana L 3, 8, 9,32, 38,74, 83 divaricata (L.) R.Br .e x Roem. & Schult 35 eglandulosa Stapf 33 elegans Stapf 7, 8, 32,33 ,38 ,39 , 74,79 nitida Stapf 130 odoratissima (Stapf) Leeuwenberg 74 pachysiphon Stapf 5-7, 32,33-36 , 40, 74, 79 parvifiora Heyne exWal l 123 penduliflora K.Schu m 33 salicifolia Wall 5-7, 32, 33,35 , 123 stapfiana Britten 5-7, 32,33 ,35 ,74 , 79 ventricosa Höchst, ex A.D C 32,33 ,74 , 79 Tabernaemontaneae G. Don 74 TetradoaPicho n 81, 82, 88 hexaloba Pichon 88 simii (Stapf) Pichon 115 Thevetia L. peruviana (Pers.) K.Schu m 3,3 6 Trachelospermum Lern, jasminioides (Lindl.)Ler n 36 VincaL. minorL 3 roseaL 36 VoacangaThouars 3.33 . 38,74 africana Stapf 3-5,34 ,74 , 79 schweinfurthii Stapf 3 thouarsii Roem. & Schult 5,6 , 34, 39,74 , 79 WrightiaR.Br. 3,75 ,8 2 arborea (Dennst.) Mabberley 35, 157 demartiniana Chiov 75 laevis Hook.f. 35 saligna (R.Br.)RMuell . ex Benth 34, 35,3 7 tinctoria R.Br . 35 tomentosa Roem. & Schult 35

WageningenAgricultural UniversityPapers 96-1 (1996) 175 WrighteaeG .Do n 74 WrightiinaePicho ne xLeeuwenber g 75

176 WageningenAgricultural University Papers 96-1 (1996) ACKNOWLEDGEMENTS

Theautho rwoul dlik et othan kth efollowin g persons orinstitutions : Dr.A .J .M .Leeuwenber gan dProf .dr .L .J .G .va nde rMaese no fth eDepartmen t of PlantTaxonomy ,Wageninge n Agricultural University, for initiating the Ph.D. programme. Dr.A .J .M .Leeuwenber gfo rhi sinvaluabl eprofessiona l guidancethroughou tth e courseo fth ePleiocarpina emonograp han dhi sthoroug hcheckin go fth efinal text . Dr.Davi dCutle ro fth eJodrel lLaborator y (RoyalBotani cGardens ,Kew )fo rhi s valuabledirectio nan dcritica lremark so nth eanatom ychapte ran dfo rwaverin gth e benchfee s forth eAnatom yLab. ,i nth eJodrel lLaboratory . Prof.dr .L .J .G .va nde rMaese nfo rtrimmin gan dcorrectin gbot hmanuscript san d proofs, andfo r hispatience ,stimulatin ginteres t andcritica lremarks . Dr.Jj.J .J .Bo sfo r hisusefu l commentso nth emanuscript . MissY .F .Ta n and Mr.J .M . deVrie s for their professional illustrations for the Pleiocarpinae chapter, and Mrs.Wi lWesse l for preparing the fine drawings and mountingo f allth efigures i nth eanatom y chapter. Prof.J .O .Kokwar o(Universit yo fNairobi ,Kenya )fo rintroducin gm et oth e field ofplan ttaxonomy ,fo r suggestingth etopi co nApocynaceae fo r myM .Sc .an dfo r hisinteres t andencouragemen tthroughou tthi swork . Mr.Ti mLawrenc e of theJodrel l Laboratory (Royal Botanic Gardens,Kew ) for allth evaluabl etechnica lassistanc ean dhi shelpfu l remarksi nconnectio nwit hth e anatomy chapter. Dr. HazelWilkinso n of theJodrel lLab .fo r theencouragemen t and informative tips. Allth eStaf f ofth eJodrel lLaborator yfo ran yassistanc ethe ygav ei nconnectio n withthi swork . Mr.Ti m Pierce of PCPU (Plant Conservation and Propagation Unit, National Museumso fKenya )fo rassistin gwit hal lth efield wor ki nKeny aan dth eres to fth e PCPUStaf f for allth eassistanc ethe ygav ei nconnectio nt othi s work. TheWageninge nAgricultura lUniversit yfo rprovidin gth e i yearsandwic hgran t andth egrant stha tenable dm et ovisi tth eHerbari ai nBelgiu man dParis . TheDepartmen t of PlantTaxonom y for arranging thegran t that enabled met o worki nth eKe wHerbariu man dJodrel lLab .fo r threemonths . TheBritis hcouncil ,Nairobi ,fo rprovidin ga retur n ourticke tt oLondo n(Roya l BotanicGarden sKew) . All the staff and collaborators of the Department of Plant Taxonomy of the Wageningen Agricultural University for allthei r helpi nconnectio n withthi sthe ­ sis. The Jomo Kenyatta University of Agriculture andTechnolog y for granting me studyleav efo r theentir eresearc hperiod . Thestaf f ofth eEas tAfrica n Herbariuman dth eNationa lMuseum so f Kenyafo r allth eassistanc ethe ygav ei nconnectio nwit hthi sresearch .

WageningenAgricultural UniversityPapers 96-1 (1996) 177 ToMrs .Judit hva nMedenbac hd eRoo yfo r allth emora lsuppor tdurin gm ysta y inth eNetherlands . ToMis s Marjo Buitelaar for all the help shegav e me during the last stageso f preparationo fthi spublicatio n andfo r being sucha grea t friend. To myparents ,Dotti ean dJoe lOmino ,fo r theirmora l supportan dinspiration . ToMargare tMuen iMusa ufo rtakin gcar eo f myfamil y duringm yabsence . Specialthank s aredu et om yhusband , Daniel S.Omino ,fo r his continuousen ­ couragement, utmostpatienc ean dsacrifices , andfo r hisassistanc ei nman yways . Thedirector s and/orcurator so fth efollowin g herbariafo rputtin gthei rmateria l atm ydisposal :A ,B ,BM ,BP ,BR ,C ,COI ,E ,EA ,F ,FHO ,FT ,G ,GH ,HBG ,K , L, LD, LG, LISC, LISU, M, MO, NY, P,PRE , S, SRGH, U, UC, UPS, US,W, WAG,WU,YA,Z.

178 WageningenAgricultural UniversityPapers 96-1 (1996)