a revision of holarrhena r. br.

582.937.2

MEDEDELINGEN LANDBOUWHOGESCHOOL W A G E N I N G E N • N E D E R L A N D »81-2(1981)

A REVISION OF H O L A R R H E N A R. BR. (APOCYNACEAE)

A. P. M. DE K R U I F Laboratory of Plant Taxonomy andPlant Geography, Agricultural University, Wageningen, The Netherlands

Received 23-X-1980 Published 23-11-1981

B I S T . t o iT.'.-"/.ie

H. VEENMAN & ZONEN B.V.-WAGENINGEN-1981

CONTENTS

Introduction

1

History of the genus

1

Geographical distribution

2

Habit, growth and flowering seasons

2

Vernacular names

2

Uses

3

Relationship to other genera

4

Cytology

4

Genus diagnosis

4

Key to the species

6

Species descriptions: 1.H. curtisii 2. H.floribunda 3. H. mitis 4. H.pubescens

6 10 15 17

'H. congolensis'

25

Nomina nuda

25

Excluded species

26

References-

26

Acknowledgements

28

Index of exsiccatae Phytochemistry of Holarrhena R. Br. byN. G. BISSET Register

29 37 38

INTRODUCTION The present publication is a monograph of the genus Holarrhena. The genus occurs in tropical Africa and tropical Asia. The present author had the opportunity to study H. floribunda in the field in West Africa and collected in Ivory Coast.Inaddition thispublication isbasedonastudyoftheavailable herbarium material.

HISTORY OF THE GENUS The genus Holarrhena was proposed by ROBERT BROWN in 1811, based on a singlespecies,CarissamitisVahl,describedin 1790.Asecondcombination inthe genus was proposed by G. DON in 1837. This species has a rather complicated history. In 1678 VAN RHEEDE TOT DRAAKENSTEIN depicted and described this plant as Codaga-palain his famous Hortus Indicus Malabaricus. Van Rheede's plateiscited intheprotologue oïNerium antidysentericum L.of 1753, otherwise based on a specimen in Herb. Hermann, which, at present, is accepted as Wrightia antidysenterica. This combination was effected by ROBERT BROWN in 1811,citingtheHermann specimen astype. BUCHANAN-HAMILTON consolidated

FIG. 1. The 'Codaga-pala' of VAN RHEEDE TOT DRAAKENSTEIN 1678. Hortus Indicus Malabaricus 1 : t.47. Meded. Landbouwhogeschool Wageningen81-2 (1981)

1

thisinterpretation byproposing thename Echitespubescens for theVan Rheede element, while G. DON transferred it to Holarrhena in 1837 after consulting WALLICH'S catalogue. The third combination in Holarrhena was effected by DURAND & SCHINZ in 1896 based on Rondeletia floribunda G. Don of 1834 (Rubiaceae).After thedescriptionofH. curtisiibyKING&GAMBLEin 1908,more than 50 combinations have been proposed on Holarrhena but in the present publication only four species have been upheld.

GEOGRAPHICAL DISTRIBUTION The genus Holarrhena is restricted to tropical Africa, Southern continental Asia and SriLanka. Themost widelydistributed species,H.pubescens, occursin open forests and savannas in Eastern and Southeastern Africa and in Southern Asia, with a markedly discontinuous area. H. curtisii occurs in tropical continentalSoutheasternAsia,completelywithinthedistribution ofH.pubescens. H. floribunda occurs in open forests and savannas from Senegal to Zaire, while H. mitis isendemic in Sri Lanka.

HABIT, G R O W T H AND F L O W E R I N G SEASONS All Holarrhena species are deciduous (under)shrubs or trees.They shed their leaves at the end of the dry season. New leaves appear at the beginning of the rainy season.Theflowers appear shortlyafter theleaves.Theflowering period is rathershort;thefruits maturesomemonthsafter theflowering season,usuallyat the end of the dry season. H.floribunda,H. mitis and H.pubescensshowthearchitectural modelof Troll (HALLE & OLDEMAN 1970: 125; - & TOMLINSON 1978: 242). As the present author observed, in savannas H. floribunda can survive bush fires by producing sucker shootsfrom burned-down stumps.When suchshootsarenotdamaged by fire for several years,they show oncemoreTroll's model and inthisway several treeletscan originate from a single stump.

VERNACULAR NAMES Only names occurring on herbarium labels and covering a single species are listed. Doubtful or obscure names have been omitted. H. curtisii: Cambodia: (daeum) tuk das. Thailand: kood kok; lunmok; lut tanong; num nom sio; sap kun.

Meded. Landbouwhogeschool Wageningen81-2 (1981)

H. floribunda: Cameroun: Ewond o : nom-ekouk. Central African Republic: C i s s o n g o : molo-mondimbo. Ghana: A s h a n t i : slae. Guinea-Bissau: Bijagó: ete-éxi. Guinea (Conakry): F u l a n i : indama; M a l i n k é : indama, kounajana; Susu: yaté, yeté. Ivory Coast: A t t i é : pri, sohoué; Baoulé: sébé, sèbé. M a l i : B a m b a r a : fou-fou, kouma jui. Nigeria: Hausa: bakim mayu, bakim mutum, ga(m)ma(n) sauwa; l b o : okemba. Senegal: D i o l a :fournatiaf; L e b o u ; saoulou; M a d i n k a : tiariko; M a l i n k é : dierin-dierine;Wolof: salahal,soonlow,soulali. Sierra Leone: L o k o : mats; M a d i n k a : nono kende; M e n d e : boboneh, bubuwi, nookwi, nukoi, nukui; T e m n e : a-mat(s), an-watia, ka-mat, kabore; Y a l u n k a : kenyekhigina. Upper Volta:Mossi:jabarawgo. Zaire: K i k o n g o : kimbuki, kinzenze, yakoko; Z a n d é : bagu. H. mitis: Sri Lanka: genu-dena-mal, kir(r)i-walla(-gass). H. pubescens: Africa: Kenya: K i b o n i : kepako; L a m u & K i s w a h i l i : mtimeupe. Malawi: N y a n j a : c(h)ipeta, kachamba, t(h)ombosi; Y a o : chiwimbe, mkwale, mvumbamvula, nfiriti. Moçambique: C h i n d a o : muashu, rutshene; P o r t u g u e s e : quina. South Africa: A f r i k a a n s : koorspeul; English -.jasminetree; L o n g a : kahlakahla; V e n d a : khàthàkhàthàné, muhatu. Zambia: K i l u n g u : chipipa. Asia: Burma: khleng-kong. India: A r a b i a n & Persian:lissan al asafeer; Bengali: kurchi; E n g l i s h : bitter oleander, dysentery rosebay, easier tree, oval leaved rosebay; H i n d i : kura, kurchi; L u s h a i : thlengpa; M a l a y : codaga-pala;P o r t u g u e s e : herba malabarica, herbas contra camaras; U r i y a : kuruni. Thailand:mok man, mok yai. USES There is an impressive amount of publications on this subject. All species contain numerous alkaloids. Only the most important applications are menMeded. Landbouwhogeschool Wageningen81-2 (1981)

3

tioned here. Dr. N. G. BISSET (School of Pharmacy, University of London) publishes more data on this subject with this paper. Of H. pubescens the bark and less often the seeds are widely used as antidysentericmedicine.Theother species have thesameproperty butareless often used.Additionally H.pubescenswasusedbythePortugueseinMoçambiqueasa tonicand febrifuge. The wood of allspecies(except H. curtisii) islocallyused as timber of low quality.

RELATIONSHIPTOOTHER GENERA Holarrhena shows aclavuncula which iscompletely free from theanthers and therefore itisplacedinthePlumerioideae,although itresemblesseveralgenerain Apocynoideae (Echitoideae) byits aestivation, which isturned to the right, and by its seedswith a coma, resembling Compositae fruits. However, there are two other generainthePlumerioideae, Carruthersiaand Spirolobium, that haveseeds with a coma, and the former shows an aestivation turned to the right as well. Most genera ofthePlumerioideaehaveseedslackingacoma,contained in sappy fruits inthe Carisseae,orincapsulesina.o.thePlumerieae.Inthelatter tribe the seedsare often winged, but exceptionally they may bear a coma. Thisisthecase inAlstonia, whichshowsseedswithacomaatbothextremities.After considering thedata stated above,thepresentauthor hasdecided tosupport PICHON'S (1950) view regarding the position of Holarrhena: it belongs to the subtribe Holarrheninae of the tribe Plumerieae (Alstonieae) of the subfamily Plumerioideae, while Alstonia is referred to the Alstoniinae of the same tribe. In PICHON'S publication the Holarrheninae are composed of Carruthersia, Holarrhena and Spirolobium.

CYTOLOGY Holarrhena pubescens: 2n= 22 (SEN & ROY TAPADAR 1957: 19; RAGHAVAN 1957:294-298; ROYTAPADAR 1964:103-138;DATTA&MATTI1972:9-19;inall these cases:no voucher specimens preserved). Holarrhena floribunda: 2n= 20 (MIÈGE & JOSSERAND 1972:283-292;no voucher specimen preserved).

GENUS DIAGNOSIS Holarrhena R. Br. 1811: 62; G. Don 1837: 78; De Candolle 1844: 413; Schumann 1895b: 137; Stapf 1902: 161-162; Ridley 1923: 349; Pichon 1950: 157-158; Codd 1963:263. Type species: H. mitis (Vahl) R. Br. Heterotypic synonym: Physetobasis Hassk. 1857: 104-105; 1859: 258. Type 4

Meded. Landbouwhogeschool Wageningen81-2 (1981)

species: Physetobasis macrocarpa Hassk. ( = H.pubescens (Buch.-Ham.) Wall. ex G. Don. Deciduous undershrubs, shrubsortreesupto25mhigh,with abundant white latex in bark, branches, leaves and fruits. Branches unarmed, terete, sulcate when dry. Branchlets terete, sulcate when dry. Stipules intrapetiolate and often obscure or none. Leaves opposite or subopposite, those of a pair equal, shortly petiolate, without colleters in the axils;petiole glandular near the base; blade simple,medium green, paler beneath, entire;on each side 5-25 secondary veins which arecurved towards the margin, anastomosing; tertiary venation more or less conspicuous, reticulate. Inflorescences terminal and/or seemingly axillary, cymose, many-flowered, rather lax or dense. Bracts entire, linear, acute at the apex, without colleters in the axils. Flowers 5-merous, sweet-scented, actinoT morphic or with only sepals unequal, fragrant. Calyx persistent under the fruit. Sepals green, free or connate at the extreme base only, subequal or sometimes unequal, valvate, much shorter than the corolla, narrowly elliptic to linear, outsidepubescent withglandular and/or ordinaryhairsorglabrous,ciliate,with 0-10colletersinsideatthebasewhicharealternatingwiththem,narrowlyelliptic totriangular. Corrollainthemature bud salver-shaped;tube whiteor less often greenish white,cylindrical, slender, wider near the base and apex; on both sides variously pubescent or almost glabrous, inside, however, at the base always glabrous and at the throat always pubescent; lobes white, in the bud contorted and overlapping to the right, narrowly triangular to narrowly elliptic. Stamens included,inserted atthebaseofthecorolla tube;filamentsveryshort, pubescent at the base only; anthers introrse, elliptic or narrowly elliptic, glabrous, basifixed, rounded at the base, mucronate at the apex;cells two, discrete, parallel, dehiscentthroughout byalongitudinalslit.Pistilglabrousorpartiallyorentirely pubescent;ovary superior, carpelstwo,connate attheextreme baseonly, ovoid, graduallynarrowed intothestyle;disknone;stylecylindrical, slightlysplitat the base,somewhat laterallycompressed towardstheapex,reaching theinsertion of thestamens,composed oftwoconnate strands;clavuncula ovoid, mostlywith2 longitudinal indented lines;stigma mostly bifid;placentas adaxial;ovules numerousin4-8 rows.Fruitcomposed oftwo scarcely spreading mericarps, which arefollicular, slender,moreorlessconstricted betweentheseeds,dehiscent along an adaxial line of dehiscence;wall thinly coriaceous, outside smooth or sulcate when dry, glabrous, sometimes with lenticels or white-dotted or -spotted. Seeds numerous,intwoormorerows,linear toverynarrowly oblong,striate,grooved, glabrous, with a dense tuft of hairs at the apex which is 3-6 x as long as the seeds;embryo large, surrounded by very scanty endosperm; cotyledons broad, complicate, larger than the short straight radicle. D i s t r i b u t i o n ^ species in Africa and Asia. N o t e : The sections cited by PICHON (1950) are not maintained here, as they are artificial. Meded. Landbouwhogeschool Wageningen81-2 (1981)

K E Y TO T H E S P E C I E S 1. Leaves papery when dry, mostly acuminate or acute at the apex; fruits pendulous; shrub or tree, 0.60-15(-25) m 2 Leaves chartaceous when dry, mostly rounded or emarginate at the apex; fruits erect at least when young; (under)shrub, 0.30-2.50 m. S o u t h e a s t e r n c o n t i n e n t a l t r o p i c a l Asia l.H.curtisii 2. Leavesovate toelliptic, 1-3.3 x aslongaswide;corolla lobes2-5 x as long as wide 3 Leavesnarrowly elliptic,3-5 x aslongaswide,corolla lobes6-10 x aslong as wide. Sri L a n k a 3.H. mitis 3. Corolla 12-19 mm long, tube 6.5-9 mm long, lobes 7-11.5 x 2.2-3.5 mm; inflorescence verydense;fruits 30-60cmlong;branchesand fruits with few lenticels.West and Central Africa 2.H. floribunda Corolla 20-38 mm long,'tube 9-19 mm long, lobes 10-24x3-8 mm; inflorescence rather lax; fruits 20-37.5 cm long; branches and fruits often with many lenticels. E a s t e r n and S o u t h e r n Africa, S o u t h e r n Asia 4. H. pubescens N o t e . See 'ƒ/.

congolensis\

SPECIES

DESCRIPTIONS

1. Holarrhena curtisii King & Gamble 1908: 446^47; Pitard 1933: 1171; Craib 1939: 450. Fig. 2,p.7; Map 1,p.9. Type:Thailand:Trang, Curtis 3392(K, lectotype). Heterotypic synonyms: H. pulcherrima Ridley 1911: 130-131. Type: Malay Peninsula: Perlis, Kanga, Ridley 14982 (K,lectotype;isotype: BM). H. latifolia Ridley 1911: 131.Type: Thailand: Bangtaphan, Keith 19/3/90 (type not seen, probably not preserved). H. densiflora Ridley 1911: 131-132; Craib 1939: 450. Syntypes: Thailand: Bangtaphan, Keith s.n.; Thailand: Between Singgora and Nongchik, St. V. B. Downs.n. (syntypes not seen, probably not preserved). H.similisCraib 1913:70;Pitard 1933:1172.Type:Thailand :Lampun,MeTa, A. F. G.Kerr 2548(BM, lectotype; isotypes: E, K). Undershrub or shrub,0.30-2.50 m high. Shoots tufted from a perennial rootstock. Branches pale grey to dark grey, often lenticellate or white-dotted or -spotted;branchlets pale grey to brown, pubescent. Leaves: petiole pubescent, 0.5-6 mm long; blade ovate to elliptic or narrowly so, 2-3.3 x as long aswide, 2.3-9.7 x 1.2-4.4cm,acuminate,rounded oremarginate attheapex,cuneate or rounded at the base, chartaceous when dry, pubescent above, more densely so beneath;that in the leaves of the first pair or the first two pairs of a branchlet often suborbicular, rounded, obtuse or emarginate at the apex and smaller; on 6

Meded. Landbouwhogeschool Wageningen81-2 (1981)

FIG.2.Holarrhenacurtisü: 1.floweringbranch,f x ; 2.apexofleaf,| x ; 3.partofuppersurfaceof leaf, 2x ; 4. flower in bud, 2x ; 5. flower, 2x ; 6. sepal inside with colleter, 6x ; 7. part of flowershowingstamensandpistil,2x ; 8.partofcorollawithstamens,8x ; 9-10.partofcalyxwith pistil,6x ; 11.fruit, f x ; 12.seed,fx . - ( 1 , 3. VanBeusekom &Phengkhlai I245;2. A. F.G. Kerr 5693;4-9.Garrett293; 10.d Orleanss.n. (received23 June 1892); 11-12.Coudercannis 1883-1885).

each side 9-15 secondary veins. Inflorescences terminal, less often seemingly axillary or leaf-opposed, lax. Peduncle, branches and pedicels pubescent; peduncle0.5-1.6cmlong;bractslinear,2.0-5.0mmlong,pubescent onboth"sides; pedicels0.6-1.6 cm long. Sepalsgreen,elliptictonarrowly so, 1.9-6.3 x aslong aswide,2.2-6.9 x 0.8-2.0 mm,free orupto0.5mmconnateat thebase, outside pubescent with ordinary and some glandular hairs, inside less pubescent with ordinary hairs only, mostly glabrous at the basefor 0.5-3 mm,ciliate,with 0-10 colleters inside at the base which are alternating with them. Colleters narrowly elliptictotriangular, 1-3 x aslongaswide,0.2-0.5 x 0.1-0.2mm. Corolla'va.the mature bud 4.4-11.8 x as long asthe calyx, 20-34 mm long; tube white or less often greenish-white, 2.3-6.9 x aslongasthecalyx, 12-22mmlong, 0.5-2.0mm in diameter, inside densely pubescent with ordinary and also some glandular hairs,except for thebasal 1.1-2.1mm; lobeswhite,narrowly triangular, 0.8—1.3 x as long as the tube, 2.4-AA x as long as wide, 10-23 x 3-6 mm, recurved, pubescent and ciliate or entirely glabrous. Stamens inserted at 1.8-2.0mm from the base of the tube, anthers 1.3-3.0 x as long as the filaments, narrowly triangular, 1.2-2.2mm long,with a0.2(-0.5)mm longmucro. Pistil2.3-2.6 mm long;ovary0.9-1.2x0.5-0.7mm,glabrousorshortlypubescentattheapex;style 0.5-1.0 x 0.2-0.3 mm, pubescent or less often glabrous at the base;clavuncula 0.5-1.0 x 0.2-0.3 mm, pubescent or glabrous; stigma 0.1-0.3 mm long, shortly pubescent or glabrous. Fruits pale to dark brown;carpels erect, at least when young, 11 -33 cm long, 3-8 mm in diameter, outside sometimes with a few lenticels or white-dotted or -spotted. Seeds 8-14 mm long with a dense, 25-45 mm long tuft of hairs at the apex. Cotyledons 1.8-2.8mm long. D i s t r i b u t i o n : Southeastern, continental, tropical Asia. E c o l o g y : Mangrove swamps, savannas and in open forest such as: deciduous mixed forest, bamboo forest, scrub woodland or open places in dense forest (a.o. Dipterocarpaceae-foresi). Altitude: 0^400m. A selection of the a b o u t 100 specimens e x a m i n e d : THAILAND: C h i a n g M a i : Ban Ngiu Thao (fl., fr. Mar.) Winit 238 (BM, K); along Ban Bo Luang-Hot road (fl. June) VanBeusekom &Phengkhlai 1245(AAU, E,K, L,P).C h i a n g R a i : Doi Pui(fl.June) Garrett293(A, BM.E,K, L).L a m p h u n :DoiKai Khia (fl.July) Winit397(K);MeLi ( = Li Nam Mae R.) (fr. Aug.) Winit 398 (K); MeTa (bud, fl. Apr.) A. F. G.Kerr 2548(BM, E, K, lectotype ofH.similis). P h r a e : near Rawng Kwang (= Rong Kwang) (fr. Feb.) A. F.G.Kerr2357 (BM,E,K, paratype ofH. similis) ;DenJaya (= Den Chai)(fl.Oct.)Franck 477(A,BR, C,COI, L, P). U t t a r a d i t : Huai Nam Pat (fl. Apr.) Phengkhlai 141(L, P ) . P h i t s a n u l o k : Muang Jom (fl., fr. Mar.)A. F.G.Kerr5693 (BM,K, P,UC).U d o n T h a n i : BanNa Lat (fl.) Smiless.n. (K, paratype). P h e t c h a b u n : Nong Phai (fl., fr. May) Smitinand 10887(E, K, L, P). K h o n K a e n : 45 km N. of Khon Kaen (fl. Mar.) S^irensenetal.2193(K).Sara Buri :Thap Kwang (fl. May) Smitinand 10893 (K, L, P).N a k h o n R a t c h a s i m a : Pak Chaung (fl. May) A. F. G.Kerr 20450 (BM, K); Korat (fl. May) Bourke, May 1924(BM,UC). B u r i r a m : Ban Chum Saeng(bud, fl. May) Noe 157(A, AAU, E, L).S i s a k e t : Kantharom (fl.June) Smitinand 1596(K).U b o n R a t c h a t h a n i : Warin (fl. June) Lakshnakara 882 (BM, K). P r a c h i n Buri: Aranyaprathet (fl., fr. May) A. F. G. Kerr 19322(A, BM, E, K, P). C h o n Buri: Si Racha (fl. May) A. F. G. Kerr 4250 (BM, K). P r a c h u a p K h i r i K h a n : Hua Hin(fl.Aug.) Marcan 375(BM,K);ibid,(fl.,fr. Jan.)A. F.G.Kerr8322(BM, E.K, P); ibid. (fl. Apr.) Udell205 (K).C h u m p h o n : Bang Son (fl. Mar) Put 1487(KW, BM, E, K); Sapli (= Saphli Sathani) (fl. May) Haniff& Nur 4272(H); 15km N. ofChumphon (fl.Aug.) Larsen etal. 8

Meded. Landbouwhogeschool Wageningen81-2 (1981)

MAP 1.Holarrhena curtisii. 1445(AAU, L,P); Lansuan (fl.,fr. June) Put 1681 (BM, K). P h a n g n g a : Nop Pring(fl. Mar.) A. F. G.Kerr 18531 (AAU, BM,K).S u r a t T h a n i : Naung Wai(fl. Mar.) A. F.G.Kerr12266(BM, K, P); Konchanadit(fl.Aug.)A.F.G. Kerr 13094(AAU, BM,E,K,P);BanNa(fl. Sep.) Yuang44(BM, K). T r a n g : Trang (fl., fr. Mar.) Curtis 3392 (K, lectotype); ibid. (fl., fr. Mar.) Kunstler 1402 (K, paratype); ibid. (fl. Mar.) Vanpruk 648 (E, K). S A T U N : Adang Islands (fl.,fr. Apr.) Ridley 15864 (BM,K).S o n g k h l a : Sadao (fl.July) A. F.G.Kerr 15889(AAU, BM,K, P); UTaphao Khlongffl. May)A.F.G. Kerr3688(BM,K);Rattàphum (fl.June)Smitinand11028 (K,L,P);PadangBesar (fl. Dec.) A.F.G. Kerr 13583(BM, E, L, P). MALAYSIA: M a l a y P e n i n s u l a : Perlis (fl. Nov.) Henderson 22935 (K); ibid. (fl. Mar.) Ridley 14981 (BM, K, paratype of H. pukherrima); ibid, (fl., fr. Mar.) Ridley 14982 (K, lectotype of H. pulcherrima) ; Kedah, Kuala Nerang (fl. July) Corner31519 (K). LAOS: Nuang Pakxan (fl. Mar.) A.F.G. Kerr20739(BM, K); Ban Tha Ngon Na ( = Ban Na) (fl. May) Talbot de Malahide 13(A, BM); ibid. (ft.Apr.) Talbot de Malahide 145(BM); km 85 route Savannakhet-Quangtri (fr. Jan.) Poilane 11602\f); Muang Phin (fr. Jan.) Poilane 11483 (P). Sin. loc.:(fl.) dOrleans s.n. (received 23June 1892) (P);(fr.) Spire, Dec. 1903(P). CAMBODIA: Between Angkor Wat and Sisophön (fl., fr. July) Sfirensen et al. 7306 (E, K); Kol Tötung (fl. July, fr. Oct.) Martin 10(L,P); Kompong-chhnang (fr. Mar.) Chevalier31949(P);ibid. (fl. June)Godefroy-Leboeufl89 (B, K,P,paratypé);Kampot (fl.Mar.)Geoffray444(P);ibid.(fl., fr.) Pierre 174,July 1874(L, P). VIETNAM: Baria (fl., fr.) Pierre 174,Aug. 1867(K, NY, P);ibid. (fl., fr.) Pierre 174,July 1866(A, NY,P);ibid.(fl.,fr.) Pierre4486(A,BR,L,MO,NY,P);ibid.(fl.July)Pierre5243(A,BR,NY,P); Sin. loc.:(fl. Mar.) Coudercannis 1883-1885(P,UC); (fl. Sept.-Nov.) Thorel 1409(B,K, P).

N o t e . Unfortunately the present author was unable to trace the types of H. latifolia and H. densiflora. However, the descriptions are clearly based on materialwhichmustbelongtoH. curtisiiaccordingtothepresentstateofaffairs and consequently H. latifola and H. densiflora are reduced to synonyms here. Meded. Landbouwhogeschool Wageningen81-2 (1981)

2. Holarrhenafloribunda(G. Don) Dur. & Schinz 1896: 190; Schumann 1898: 122; Durand & Schinz 1909: 346; Schmit 1950; Huber 1963: 68-69; Keay, Onochie & Stanfield 1964: 382,pi. 165;Berhaut 1967: 124; 1971: 385,pi. 384. Fig.3,p.11;Phot. 1,p. 10;Phot. 2,p. 12;Map2,p.13. Basionym: Rondeletia floribunda G. Don 1834:516. Type: Sierra Leone: Sin.loc, amongbushesinthelowlands, G.Dons.n.(BM, holotype;isotypes:CGE, K). Heterotypic synonyms: H. ovata A. DC. 1844: 414; Stapf 1902: 163-164; Chevalier 1920: 418. Type: Guinea (Conakry): Rio Nunez, Heudelot 795(De Candolle erroneously 1795) (G-DC, holotype;isotypes:G, K, OXF, P, TCD). Homotypic synonym:H. floribunda var. tomentella Huber 1962:437; 1963:69. H. africanaA. DC. 1844: 138; Bentham 1849: 450; Oliver 1877: 44; Wulfsberg 1880; Schumann 1895b: 138; Morris (?) 1896: 47^18; Stapf 1902: 164; Chevalier 1920:417;Aubréville1936:170,pi.308;Dalziel 1937:372;DelaMensbrugge 1966: 324,pi. 325.Type: Sierra Leone: Sin. loc, Smeathmann? (G-DC, holotype). H. wulfsbergiiStapf 1902: 164; De Wildeman 1903a: 68; Durand & Schinz 1909:346;Chevalier 1920:418;Aubréville 1936:170;Dalziel 1937:372;Aubréville1950:443-^144.Type:Togo:Badja, Schlechter 12969(L,lectotype;isotypes: BM, BR, G, K, P, Z). Shrub or tree, (0.50-)2-25 m high, often regrowing from burned-down stumps.Trunk 5-30 cmindiam.Bark inyoungtreesnearlysmooth andmore or lessdistinctlylenticellate,laterroughandcorky,longitudinallyfissured, palegrey todark brown; innerbark palebuff with green outerlayer; wood soft. Branches lenticellate or not, grey to dark brown; branchlets lenticellate or not, grey to brown.Leaves: petiolepubescent, 0.5-7(-10) mmlong;blade ovatetoellipticor narrowly so, 1.3-3 x aslongaswide,2-14 (-19) x 2.3—8(—11)cm,acuminate or acute at the apex, cuneate or rounded at the base, herbaceous when living, paperywhendry,abovepubescent toglabrous,beneath moredensely pubescent allover or only on the main veinsor also glabrous;that inthe leaves of the first

PHOT. 1. Holarrhena floribunda (De Krui/ 25). 10

Meded.Landbouwhogeschool Wageningen81-2 (1981)

3.leafbase withpetiole from FIG. 3.Holarrhenafloribunda: 1.flowering branch,f x ; 2.leaf,• above, 2x ; 4. part of branch, | x ; 5. flower, 4x ; 6. flower in bud, 4 x ; 7. part of corolla showing stamens and pistil,4x ; 8.part ofcalyxwith pistil,8x ; 9.part ofcorolla with stamens, 11. seed, | x ; 12-13. detail seed both sides, 2 x . - ( l , 5 . Sanford 6780; 2, 10. fruit, 11-13. De Koning 6555; 3 ^ , 6 - 9 . Leeuwenberg 9664; 10.Espirito Santo 208).

PHOT. 2. Holarrhenafloribunda (De Kruif444).

pair orthefirsttwopairsofabranchlet often suborbicular, rounded orobtuse at the apex and smaller; on each side 5-14secondary veins. Inflorescences usually seemingly axillary or leaf-opposed, sometimes terminal,rather dense. Peduncle, branches and pedicels pubescent, glabrescent but never entirely glabrous;peduncle 0.5-1.3 cm long, bracts linear, 1.0-2.1mm long, pubescent on both sides; pedicels 0.4-0.8 cm long. Sepals green, elliptic, subequal, 1-3.6 x as long as wide, 1.1-4.4 x 0.6-2.0 mm, free or up to 1mm connate at the base, outside pubescent with ordinary and some glandular hairs, inside pubescent with ordinary hairsonly,glabrousat thebasefor 0.5-1.0mm,ciliate,with0-10 colleters inside at the base which are alternating with them. Colleters narrowly elliptic to triangular, 1.5-3 x aslongaswide,0.2-0.3 x 0.1-0.2mm. Corollainthe mature bud 3-11 x as long as the calyx, 12-19 mm long; tube white or sometimes greenish-white,2-5 x aslongasthecalyx,6.5-9mmlong,0.5-1.5mmindiameter,outsidepubescent withordinary and someglandular hairs,exceptfor thebasal0.5-3.1mm,sometimesentirelyglabrous,insidepubescent from thethroat to the insertion of the stamens with an indumentum which is dense in the throat, whichgraduallydiminishesdownwardsandmaybeagainmoredenseneartheinsertion of the stamens, and which iscomposed of ordinary and glandular hairs; lobes white, narrowly triangular, 0.6-1.5 x as long as the tube, 2-5 x long as wide, 7.0-11.5 x 2.2-3.5 mm, recurved, pubescent and ciliate or entirely glabrous. Stamens inserted at 2.1-3.0mm from the base of the tube;anthers 1.5-3.1 x aslongasthefilaments, narrowly triangular, 1.1-1.7mm long,with a 0.1-0.4 mmlongmucro.Pistil 2.5-4mmlong; ovary0.9-1.5 x 0.5-1.0mm,glabrous or 12

Meded. Landbouwhogeschool Wageningen81-2 (1981)

sometimes shortly pubescent at the apex; style 1.0-1.8 x 0.2-0.3 mm, shortly pubescent at the apex or glabrous; clavuncula 0.4-1.1 x 0.3-0.5 mm, shortly pubescent or less often glabrous; stigma 0.1-0.4 mm long, shortly pubescent or lessoften glabrous.Fruitspalegreytodark brown;carpels pendulous, 30-60cm long,5-7mmindiameter, outsidesometimeswithafewlenticelsor white-dotted or-spotted, Seeds 11-16mmlongwithadense,40-60mmlongtuft ofhairsatthe apex. Cotyledons 2.5-3.1 mm long. D i s t r i b u t i o n : West and Central Africa. E c o l o g y : Deciduous forests, open places in more dense forests, woodlands, and savannas, on clay, sand or rocky outcrops; alt. 0-1000 m. A selection of t h e a b o u t 500 specimens e x a m i n e d : SENEGAL:Sangalkam (fl., fr. Aug.) Berhaut 718(BR, P,Z);Rufisque (fl., fr. May) Leprieuranno 1826 (G, P, TCD). GAMBIA : Sin. loc. (fr. Feb.) Dawe 40 (FHO, K).

GUINEA-BISSAU:UnaFormosa, Abu (fl.Apr.) Espirito Santo 1978 (COI,K,MO,P);Farim (fl., fr. June) Espirito Santo 2393 (COI, K, LISU, P); Bafata (fl., fr. July) Espirito Santo 208(COI, LISU). GUINEA (CONAKRY): Rio Nunez (fl.)Heudelot 795 (G,G-DC, K,OXF,P,TCD, typeofH. ovata): Boké(fl.Apr.) Paroisse47(FHO, P);Dalaba (fl. Mar.) Dalziel8402(E,K, P,PRE);Friguiagbé (fr. Aug.) Chillou606(BR, COI, K, P,PRE);Timbo(fl.,fr. Feb.) Pobéguin140(A,K, P); Macenta (fr, Oct.) Baldwin 9793(K, MO): Beyla (fr. Oct.) Adam 28104 (C, MO). MALI: Kita (fr, Mar.) Dubois 203 (P, S); between Kangaba and Kindolé (fl. June) Chevalier857 (BR, G, K, P); Bamako (fl. June) Waterlot 1122(P);between Ouassada and Dandéla (fl., fr. Mar.) Chevalier620(BR,K, P);ibid.(fl. Mar.) Chevalier621 (COI,G, L,W,Z);Kléla,40kmN.of Sikasso (fl. May) Démange 2087(P). SIERRA LEONE:PortoLoko(fl.Apr.)Morion 1296 (K,MO,WAG);Freetown,Mt.Aureole(fl. Jan.) Dalziel 971(K. Z); Kenema (fr. Aug.) Edwardson 123(BM, FHO, MO); Sin. loc. G.Dons.n. (BM. CGE, K, type); Sin. loc. Smeathmann ?(G-DC, type of H. africana). LIBERIA: Voinjama (fr. Oct.) Baldwin 9883(K);Gbarnga (fr. Sept.) Baldwin 13279 (K, MO); Mt.

MAP 2. Holarrhena floribunda. Meded. Landbouwhogeschool Wageningen81-2 (1981)

13

Nimba (fr. Dec.) Adam 16477(P). IVORYCOAST: Boyo,km 53 road Boundiali-Tingréla (fr. Oct.)DeKruif620 (UCI,WAG);Kakpin, Comoë National Park (fl. Apr.) De Kruif 27 (UCI, WAG); Djebonoua, km 19 road Bouaké-Yamassoukro (fl. Apr.) De Kruif 25 &26 (UCI, WAG); km 5 road Seguéla-Worofla (fr. Oct.) De Kruif 444(UCI, WAG); km 44road Bouaké-Yamassoukro (fr. Aug.) De Kruif 274 (UCI, WAG);5kmNW. ofDuékoué,alongroad to Man (fl.Apr.) Leeuwenberg3888(WAG);Issia(fl., fr. Feb.) De Koning 6555 (WAG); Orumboboka (fr. July) Aubréville 64(BR, P, K). UPPER VOLTA: 22 km S. of Ouahigouya (fl. June) Leeuwenberg 4393 (B, BR, K, MO, P, PRE, WAG);10km S. ofTougan, N.ofDiouroum(fl.June)Leeuwenberg4376(B, BR, MO,P,PRE, UC, WAG); Gampéla, 10km N. of Ouagadougou (fl., fr. Aug.) Chevalier 24610 (P); Diébougou (fl.) Aubréville 2732(BR, P); Sindou (fl. Apr.) Laferrere 102(K, MO). GHANA: N.of Busumkyekye, along Mampong-Ejura road (fl. Mar.) Leeuwenberg 11990(WAG); 3kmS.ofDzakePeki,7kmN.ofAsikuma(fl. Mar.) Leeuwenberg 11194(WAG);Ho,Togoland (fl. Mar.) Dalziel 156(C, E, K, PRE); Kumasi (fl. Mar.) Vigne 1648(A, BM, FHO, K); 4 km NE. of Dodowa, NW. of Aiyikuma (fl. Feb.) Leeuwenberg 11911(WAG); Asuboi (fl. Feb.) Leeuwenberg 11046(WAG); 30 km E. of Secondi (fl. Apr.) Morton A489 (K); Aburi (fl. Mar.) Brown 682 (K, paratype of H. wulfsbergii); Sin. loc. (fl. May) Brown 326 (K. paratype of H. wulfsbergii). TOGO: 5km S. of Kandé (fl., fr. May) Hakkiet al.980(B); Niamtougou (fl. Mar.) Sanford6780 (MO); Badja (fl. June) Schlechter 12969(BM, BR, G, K, L, P, Z, lectotype of H. wulfsbergii). BENIN REPUBLIC:Birni (fl. May) Aubréville 95D(P,WAG);Boguila, Abomey (fl. Feb.) Chevalier 23192 (P); Sin. loc, Adja Ouéré (?) (fl.) Le Testu 122 (A, BM, BR, FHO, FI, K, LISC, MO, NY, P, S, UC). NIGERIA: S o k o t o S t a t e : W. of Sokoto (fr. Nov.) Dalziel 370 (K, P). K w a r a S t a t e : Along Ajaokuta-Okene road (fl.Apr.) Daramola& OsanyinlusiFH190198(PRE). Oyo S t a t e : Ibadan (fl., fr. June) Hambier 10 (B, BR, K, P); km 1Ife-Ondo road (fl. Mar.) Leeuwenberg 11235(WAG). Ogun S t a t e : Abeokuta (fl.) Barter 3334(K, paratype of//, wulfsbergii) ; ibid. (fl.) Irving s.n. (K, paratype of//, wulfsbergii). L a g o s S t a t e : Lagos (fr.) Denton s.n. (K, paratype of//, wulfsbergii); ibid. (fl. Oct.) Moloney anno 1883(K, paratype of H. wulfsbergii) ;ibid. (fl. Mar.) Rowland, 12 Mar. 1814 (P,paratype of//, wulfsbergii) ;Ikorodu (buds,Mar.) Schlechter 12296(BR, K, Z,paratype of H. wulfsbergii). O n d o S t a t e : Idanre (fl., fr. Nov.) Brenan & Keay 8655 (FHO, K). K a d u n a S t a t e : Katsina (fl.Mar.) Meikle 1355 (BR,K, P);YashiForest Reservenear Yashi (fl.July)Keay & Okeke FH1 37775 (FHO. K). Bauchi S t a t e : Dukka (fr. Dec.) De Leeuw 54 (WAG). P l a t e a u S t a t e : Wamba (fl. Mar.) Thornewill42 (FHO, K). Bendel S t a t e : Illushi, E. of Ubiaja (fl. Mar.) Lowe 1280(K, P, UCI). A n a m b a S t a t e : Obosi (fl. Apr.) Chesters 131 (FHO). C r o s s River State: Nsan, Calabar (fl. May) Smith 51 (FHO). TCHAD:Niellim (fl.June) Chevalier8525(BR, K, P);Golé (fl., fr. July) Fotius 1594 (P);Ndélé (fl. Apr.) Chevalier8019 (BR, G, K, P). CAMEROUN: Djinglia, 12 km NNE. of Mekolo (fr. Sept.) Letouzey 6836 (BR, K, P); Chutes d'Ekom, Nkam R. (fl. Apr.) Leeuwenberg 9664(WAG); Nguélémendouka (fl. Nov.) Breteler 2103 (A, BR, K, LISC, M, P, U, UC, WAG, Z); between Letta and Viali(fl., fr. Feb.) Letouzey 2977 (K, P); Moloundou (fl. Mar.) Mildbraed4616 (HBG). CENTRALAFRICAN REPUBLIC: IroR.(fl.July)Chevalier8767(P);Bozoum(fl.Apr.) Tisseront3295 (BM,P); km 20Mbaïki-Boda road (fr. Nov.) Badré 301 (P); Boukoko (fl. Mar.) Le Testu 2501 (B). GABON: Haut-Ogooué, Lastoursville Region(fl.Dec.)Le Testu 7803(BM,BR,LISC,P);ibid. (fl., fr. Nov.) Le Testu 8540 (BM, BR, LISC, P); Mouila (fl. Nov.) Le Testu 5053(BM, P). CONGO (BRAZZAVILLE): Chutes de la Foulakari, Plateau des Catarades (fl. Oct.) Sita 1857(P); between Mpioka and Manyanga, Boko Region (fl. Nov.) Trochain80 bis(P). ZAÏRE: B a n d u n d u : Nioki(fl..fr.)Flamigni7084(BR,P).B a s - Z a ï r e : Parcdelà Nkula,INEACLuki (fr. Sept.) Wagemans856 (BR, W); INEAC-Luki, Vallée de la Minkudu (fr. Aug.) Toussaint 2444(BR, FHO,K, P).Ki n s h a s a : INEAC-M'Vuazi(buds,Oct.)Dubois 75 (BR,K);ibid.,Forêt de Nkolo (fl. Dec.) Devred 412 (BR, K); Dila (fl. Oct.) Compère 691 (BR, K). H a u t - Z a ï r e : Parc National de laGaramba (fl. Mar.) Troupin507(BR, K); ibid. (fr. Aug.) Troupin 1728 (BR, K. UC). SUDAN: Summit of Mt. Otogo (fl. Dec.) Meyers 13605(K). ANGOLA:C a b i n d a : Pungo Mongo (fr. Jan.) Gossweiler6155(LISJC); Nkanda, Mbaku (fr. Jan.) Gossweiler9062(LISJC); Buco Zau (fl. Nov.) Gossweiler6822 (BM. LISJC). 14

Meded. Landbouwhogeschool Wageningen81-2 (1981)

3.Holarrhenamitis(Vahl)R.Br. 1811:62;DeCandolle 1844:413;G.Don 1837: 78;Thwaitesl860: 194;Hooker f. 1882:645;Huber 1973:14. Fig. 4,p. 16;Map3,p.17. Basionym:Carissamitis Vahl 1790: 44,pi. 59. Type: Sri Lanka: König s.n. (BM, holotype; isotype: L). Shrubortree,4-15mhigh.Trunk 5-20cmindiameter. Bark smooth, whitish, without lenticels;wood soft. Branches slender, drooping, dark purplish-brown, with inconspicuous lenticels;branchlets glabrous. Leaves: petiole glabrous, 2-8 (-15) mm long; blade narrowly elliptic, 3-5 x as long as wide, (0.5—)1—3 x (1-) 2-13cm, acuminate, acute or sometimes obtuse at the apex, cuneate at the base, papery when dry, glabrous; on each side 5-15 secondary veins. Inflorescences usually seemingly axillary or leaf-opposed, sometimes terminal, rather lax. Peduncle, branches and pedicels pubescent; peduncle 0.5-0.9 cm long, bracts linear, 1.1-2.8 mm long, pubescent or less often glabrous on both sides; pedicels 0.3-1.5 cm long. Sepals palegreen, narrowly elliptic to linear, subequal orsometimesunequal, 1.5-15 x aslongaswide, 1-7 x 0.5-1.8mm,free orupto 1 mm connate at the base, outside pubescent with ordinary and also some glandular hairs,insidepubescent with ordinary hairsonly allover,or onlyat the apex,ciliate,with0-5colletersinsideatthebasewhicharealternatingwith them. Colleters 1-2 x as long as wide, narrowly elliptic to triangular, 0.2 x 0.1 mm. Corollainthemature bud (3-)5-20(-30) x aslongasthecalyx, 20-35mmlong; tube white, 2-10(-20) x as long as the calyx, 9-13 mm long, 0.5-1.5 mm in diameter, outside pubescent with ordinary and some glandular hairs except for the basal 1.0-1.5 mm, inside pubescent from the throat to the insertion of the stamens with an indumentum which is dense in the throat, which gradually diminishes downwards and may be again more dense near the insertion of the stamens, and which is composed of ordinary and some glandular hairs; lobes white, 1-2 x long as the tube, linear, 6-10 x as long as wide, 11-15 x 1.5-2.8 mm, pubescent and ciliate. Stamens inserted at 1.5-1.8mm from the base of the tube;anthers1.5-2.2 x aslongasthe filaments, narrowlytriangular, 1.1-1.3mm long, with a 0.1-0.2 mm long mucro. Pistil 2.2-2.5 mm long; ovary 0.8-1.2 x 0.5-0.8 mm, pubescent at the apex; style 0.5-0.7 x 0.2-0.3 mm, glabrous; clavuncula 0.3-0.7 x 0.2-0.3 mm,glabrous; stigma 0.1-0.2 mm long, glabrous. Fruits pale grey to dark brown;carpels pendulous, 3 0 ^ 5 cm long, 3-4 mm in diameter, outside sometimes with afew inconspicuous lenticels. Seeds 6-11 mm long, with adense, 25-40 mm long tuft of hairs at the apex. Cotyledons 1.5-2.5 mm long. D i s t r i b u t i o n : Endemic in Sri Lanka. E c o l o g y : Open placesin forests, both inthehumid and inthedry zone. Alt. 0-800 m. M o s t of the specimens e x a m i n e d : SRI LANKA: N o r t h C e n t r a l P r o v i n c e : Gunner's Quoin, Polonnaruwa Distr. (fl. Oct.) Herb. Peradeniya493(UPS). E a s t e r n P r o v i n c e : Batticaloa(fl. Aug.)Fetch, 16Aug. 1922(A). C e n t r a l Meded. Landbouwhogeschool Wageningen81-2 (1981)

15

FIG.4. Holarrhenamitis: 1.flowering branch,f x ; 2.leafbeneath,§ x ; 3.flower,2x; 4.partof flower showing stamens and pistil,4x ; 5.part ofcorolla with stamens,8x ; 6.part ofcalyxwith pistil,6x ; 7.transversesectionofovary,20x ; 8.fruit, 9.seed, -(1-2,8-9. Macraeanno 1851; 3-7. Bernardi 15507).

MAP 3.Holarrhena mitts. P r o v i n c e : Hillcrest, Kandy Distr. (fl. Feb.) Worthington 4625 (BM); Gampola, Tembillyalla, Kandy Distr. (fl. Feb.) Worthington 4539(BM). Uva P r o v i n c e : Kataragama Peak, Monaragala Distr. (fl. Oct.) Bernardi 15507(G. K, MO, Z);ibid. (fl. Oct.) Huber 34 (BM, K, MO). S o u t h e r n P r o v i n c e : Kalutara. Kalutara Distr. (fl.) Macrae anno 1851 (FI); E. of Hiniduma.Galle Distr. (fl. May )Jayasuriya etal.793(K,MO);Sin.loc.(fl.) Königs.n. (BM.L.type);Sin.loc.(fl., fr.) Thwaites CP756 (BM, BR, G, K, LE, P), CULT.: Ivory C o a s t : Sofalco plantations near Abobo (fl. Dec.) Leeuwenberg 10709(L, WAG, introduced from Sri Lanka); Sri L a n k a : Hort. Peradeniya (fl. May) Hallier, J May 1903(HBG).

4. Holarrhena pubescens (Buch.-Ham.) Wall, ex G. Don 1837: 78; De Candolle 1844:413^114;Codd 1963:263;Palgrave 1977:786,pi. 262;Killick 1977: 4, pi. 1758. Fig. 5,p. 19;Fig.6, p.20;Map4,p.22;Map 5,p. 23. Basionym: Echitespubescens Buch.-Ham. 1822: 521-525 (not: Hook.& Arnott 1841:34). Type:Burma:Ava, Canoungaklay, Buchanan-Hamilton s.n. (BM, lectotype). Homotypic synonyms: Chonemorphapubescens (Buch.-Ham.) Sweet 1839:458. H. antidysenterica var.pubescens (Buch.-Ham.) Steward & Brandis 1874:326. Heterotypic synonyms: H. antidysenterica (L.) Wall, ex A. DC. 1844:413, partly asto specimensinK-WALL; Hook. f. 1882:644; Schumann 1895b: 134, Meded. Landbouwhogeschool Wageningen81-2 (1981)

17

138; Cooke 1904: 133; King & Gamble 1908: 445; Craib 1912: 130; Ridley 1923: 349;Craib 1939:448;Pitard 1933: 1169;Cowen 1950: 75;Biswas 1956: 67-68. pi. 32.Type: 'Codaga-pala', Van Rheede tot Draakenstein 1678: t. 47. exceptfor thefruit. Thisplateerroneously hasbeencited by Linnaeus 1753:209 with Nerium antidysentericum, of which the other elements belong to Wrightia antidysenterica (L.)R. Br.according tothepresent dayopinion (seeNgan 1965: 166). Homotypic synonyms: Echues antidysenterica (L.) Roxb. 1810: 166, partly, as for the above cited type. Chonemorpha antidysenterica (L.) G. Don 1837: 76; 1840-1843:t. 439. H. codagaG. Don 1837:78;Wight 1848:t. 1297; DeCandollel844:414. H. malaccensisWight 1848:t. 1298.Type: Malacca, Wight s.n.(K,holotype). H.febrifuga Klotzsch 1862: 277; Schumann 1895a: 137; 1895b: 315; Stapf 1902:162; DeWildeman 1903b:101 ;Durand &Schinz 1909:345;Engler 1910: 392, pi. 329; Codd 1951: 154;Palgrave 1957: 26; Dale & Greenway 1961: 47. Types:Moçambique: Tete,Sena, Peterss.n.(holotypenot seen,destroyed in B; no isotypes seen); Moçambique: Tete, Nyampanda, Gillen 17510(WAG, neotype;iso-neotypes:K, L, SRGH). H. tettensis Klotzsch 1862:278.Types: Moçambique: Tete, Sena, Peters s.n. (holotype not seen,destroyed in B;no isotypes seen);Moçambique:Namagoa, Milange road, 3 0 ^ 0 km from Mocuba, Faulkner 488 (BR, neotype; isoneotypes: COI, K, S, SRGH). H. glabra Klotzsch 1862: 279. Types: Moçambique: Tete, Peters s.n. (holotype not seen, destroyed in B;no isotypes seen);Moçambique:Niassa, Monapo,km 11 ofNamialo-Meserepane road, Torre&Paiva9272(LISC,neotype). H. antidysenterica var. glabra Steward & Brandis 1874: 326. Type: not designated. H.febrifuga var. glabra Oliver 1875: 108.Type: Tanzania: Khutu, Mbwiga, Speke &Grant, Nov. 1860(K, holotype). H.macrocarpa(Hassk.)Villar 1880:130.Basionym:Physetobasis macrocarpa Hassk. 1857: 104. Type: not designated; India: Pondichery, Hortus Pondicheriensis, Perrottet 323 (L, neotype; iso-neotypes: B, G, NY, P,W). H. fischeri Schum. 1895a: 316; 1895b: 137. Type: Tanzania: Uniamwezi, Usuri, Fischer378(holotype not seen, destroyed in B;lectotype:K, fragment). H. febrifuga forma grandiflora Stapf ex De Wild. 1903b: 101; Durand & Schinz 1909:346.Type:Zaire:Lukafu, Verdick103(Stapfand Durand &Schinz erroneously 133) (BR, holotype). H. antidysenterica v&v.macrantha Kerr ex Craib 1939: 449. Type: Thailand: Kanburi (= Kanchanaburi), Marcan 951(K, holotype;isotype:BM). H. glaberrima Markgraf 1950: 28. Type: Tanzania, Morogoro, Uluguru MountainsWestside,MiomboForest, Schlieben3326(M,holotype;isotypes: B, BM, BR, G, LISC, P, PRE, Z). Shrubor tree,0.60-18mhigh.Trunk 12-25cmindiameter. Bark smooth and more or less distinctly lenticellate or rough and corky, longitudinally fissured, pale to dark grey;innerbark pale buff, with green outerlayer, with much white 18

Meded. Landbouwhogeschool Wageningen81-2 (1981)

FIG. 5. Holarrhenapubescens: 1.floweringbranch,§x ; 2.partofbranchwithlenticels,fx ; 3.bark, f x ; 4.flower,2x; 5.part of flower with stamens and pistil, 2x ; 6.sepal insidewith colleters, 6 x ; 7. sepal inside with colleter, 8x ; 8. part of calyx with sepals, 4 x ; 9.part of corolla with stamens, 8 x ; 10. fruit, f x ; 11. seed, f x. - (1. Pawek 7734; 2. Thulin & Mhoro 1256; 3-4. Leeuwenberg 10867; 5-6, 8-9. Leeuwenberg 10866; 1. Squires 778; 10.Lawton 1212; 11. Harmand 23).

FIG. 6. Holarrhenapubescens: seedling, 3months,

(Fanshawe9068).

latex; wood soft. Branches lenticellate; branchlets pubescent or lessoften glabrous. Leaves very variable in shape and size; petiole pubescent or less often glabrous,0.2-12mmlong;bladeovatetoellipticornarrowlyso, 1-3.3 x aslong as wide, 1.7-19.5 x 1.3-11.2 cm, acuminate or acute at the apex, cuneate or rounded at the base, papery when dry, pubescent to glabrous on both sides:if hairymostly more sobeneath; bladeintheleavesofthefirst pair orthefirst two pairs of a branchlet often suborbicular, rounded or obtuse at the apex and smaller; on each side 5-25 secondary veins. Inflorescences usually seemingly axillaryorleaf-opposed, sometimesterminal,rather lax.Peduncle,branches and pedicels pubescent or less often glabrous; peduncle 0.9-1.7 cm long; bracts linear, 1.5-4.5mm long,pubescent orlessoften glabrous on both sides;pedicels 0.3-3cm long. Sepals green, elliptic to linear, subequal or sometimes unequal, 1-20 x aslong aswide,(1—)2—12 x 0.5-1.7 mm, free or up to 1 mm connate at 20

Meded. Landbouwhogeschool Wageningen81-2 (1981)

thebase,outside pubescent with ordinary and alsosomeglandular hairsall over or only at the apex, less often entirely glabrous, inside pubescent with ordinary hairs only all over or at the apex and then the basal 0.5-3 mm glabrous, or less often entirely glabrous, ciliate, with 0-10 colleters inside at the base which are alternating with them. Colleters 1.3-3 x as long as wide, narrowly elliptic to triangular, 0.2-0.4 x 0.1-0.3mm. Corrollainthemature bud 2-13 x aslongas thecalyx,20-38mm long;tubewhite 1.2-5.5 x long asthecalyx,9-19mmlong, 0.5-20 mm in diameter, outside pubescent with ordinary and also some glandular hairs except for the basal 0.5-3 mm, sometimes entirely glabrous, inside pubescent from the throat to the insertion of the stamens with an indumentum whichisdenseinthethroat, which gradually diminishesdownwards and may be again more dense near the insertion of the stamens, and which is composed of ordinary and someglandular hairs;lobeswhite,narrowly triangular, 0.9-1.7 x as long as the tube, 2-5 x as long as wide, 10-24(-30) x 3-8 mm, recurved, pubescent andciliateorglabrous. Stamens inserted at 1.5-2.5mmfrom the base of the tube; anthers 0.8-5 x as long as the filaments, narrowly triangular, 0.8-1.8 mm long, with a0.1-0.3 mm longmucro. Pistil 1.8-3.1mm long; ovary 0.7-1.2 x 0.5-1.0mm,glabrous;style0.3-1.2 x 0.2-0.3 mm, shortly pubescent at the apex or entirely glabrous; clavuncula 0.3-1.2 x 0.2-0.5 mm, shortly pubescent or less often glabrous; stigma 0.1-0.3 mm long, shortly pubescent or glabrous. Fruits pale grey to dark brown; carpels pendulous, 20-37.5 cm long, 2-9 mm in diameter, outside sometimes with many lenticels or white-dotted or -spotted. Seeds 9-16mm long, with a dense, 2 5 ^ 5 mm long tuft of hairs at the apex. Cotyledons 2-3 mm long. D i s t r i b u t i o n : Eastern and Southern Africa, Southern Asia. Ecology: Deciduous mixed forests, bamboo forests, scrub woodlands, savannas, or open places in heavy forests, on clay, sand or rocky outcrops. Alt. 0-1500m. A selection of the a b o u t 750 African and 650 A s i a n specimens examined: ZAÏRE: B a s - Z a ï r e : between Tchoa (= Tshoa)and Borna (fl. Nov.) Cabra3(BR, K, type of'//. congolensis') . S h a b a : Pare National de l'Upemba, Mabwe (fl. Dec.) J. de Wilde 692 (BR, K); Musosa (fl. Oct.) Bredo 3521 (BR); Haut Lomami (fl. Sept.) Leistner 1471 (M, PRE); Lukafu (fl. Oct.) Verdick 103(BR ; type of H.febrifuga forma grandiflora). KENYA: C o a s t : Mambosasa (fl., fr. Feb.) Graham316 (A, BM, FHO, K);Taita Hills (fl. Nov.) Bally 13610(BR, G, K); Sokoke (fl. Jan.) Dale 1073(BR, FHO, P). TANZANIA: Lake P r o v i n c e : Mbarika, Mwanza Distr. (fl. Sept.) Tanner 991(BR, K, MO, NY, UC, WAG); ibid. (fr. May) Tanner 1475 (B, BR, K, LISC, S, UC). T a n g a P r o v i n c e : Segoma Forest Reserve, Korogwe Distr. (fl. Nov.) Faulkner 4304 (C, K, MO, P); Mashewa Usambara (fr. Aug.) Hoist 8837(COI, HBG, K, LE, M, P,W, Z);Kideliko, Handeni Distr. (fl., fr. Apr.) Faulkner 1426(B,BR, K, S).W e s t e r n P r o v i n c e : Ikuu Distr. (fl.Jan.) Bullock 2287(E, BR, K, S);ibid. (fl. Jan.) Bullock 2316(B,BR, K, S).C e n t r a l P r o v i n c e : Manyomi, near Kilimatinde (fl. Dec.) Burn 5399 (BM, BR, FHO, K); Usur (= Ussure) (fl.) Fischer378 (K, lectotype of H. fischeri) ; km 30 Manyoni-Dodoma road (fl. Nov.) Leeuwenberg 10866 (WAG); ibid, (fl., fr. Nov.) Leeuwenberg 10867 (WAG). E a s t e r n P r o v i n c e : Mtibwa Forest Reserve, Morogoro Distr. (fl. Nov.) Semsei 1503 (B. BM,BR,FHO,K, MO);Mbwiga,Khutu(= Ukutu)(fl.Nov.)Speke &Grantanno 1860 (K, type of H.febrifuga var. glabra); Morogoro (fl. Nov.) Milne-Redhead & Taylor 7368(B, BR, K, Meded. Landbouwhogeschool Wageningen81-2 (1981)

21

MAP 4. • Holarrhena pubescens, Western part of area. A Holarrhena floribunda x Holarrhena pubescens. LISC); Uluguru Mts. (fi. Jan.) Schlieben 3326 (B, BM, BR, G, K, LISC, M, MO, P, PRE, Z). S o u t h e r n H i g h l a n d s P r o v i n c e : SW. Ruaha National Park, Iringa Distr. (fl.) Greenway & Kanuri 14812(K, M, MO, PRE); Madibira (fl. Sept.) Thulin &Mhoro 1256(K, UPS). S o u t h e r n P r o v i n c e : Lutamba-lake, Lindi Distr. (fl. Oct.) Schlieben5440 (B,BM, BR, G, LISC, M, P, PRE, S,Z);W. of R. Mbangala, Masasi Distr. (fl. Dec.) Milne-Redhead & Taylor 7679(B,BR,K, LISC). Z a n z i b a r : Mahonda (fl. Jan.) Sacleux 2214 (P); ibid. (fl. Jan.) Sacleux 2215 (P). ZAMBIA: Chiengi (fl.Oct.)Bullock 1244 (BR, K);Mbesuma Ranch, Kasama Distr. (fl. Dec.) Astle 1061 (SRGH);Kitwe(seedling,Dec.)Fanshawe9068(K);ibid,(seedling,Sept.)Fanshawe9303(K); Jumbe (fl., fr. Oct.) Robson 81 (BR, BM, LISC, PRE, SRGH); Luangwe R. near Beit Bridge (fr, Apr.) White 2401 (BR, FHO, K); Nega-Nega near Kafue (fl. Nov.) West 3455 (K, LISC, SRGH, PRE); Mazabuka Research Station, Mazabuka (fr.Apr.) Lawton 1212(FHO, SRGH); Mapanza, Chôma (fl., fr. Oct.) Robinson 2474 (K, M, PRE, SRGH); Kariba North Access road, Mazabuka Distr. (fl. Dec.) Goodier437 (BR, K, LISC, P, PRE, SRGH). MALAWI:W.ofKaronga (fl.Jan.)Pawek 7734(MO);40km S. ofKaronga(fl.Jan.)Robinson3128 (K,M, MO,PRE, SRGH);near SaniHill(fl., fr. Nov.)Benson599(BM,BR,K, LD,PRE, SRGH); between Lake Nyasa Hotel and Senga Bay Hotel (fr. Feb.) Robson 1639(BM, BR, K, LISC, PRE, SRGH); Fort Johnston (fl., fr. Nov.) Jackson 1378(BM, BR, FHO, K). ZIMBABWE: Mzarabani, Darwin Distr. (fr. Apr.) Bingham 1449(BR, LISC, M, PRE, SRGH); Mazoe dam (fl. Jan.) Müller 215 (L, LISC, PRE, SRGH); M'tarazi Falls (fl., fr. Nov.) Chase1319 (BM,COI, K, LISC,SRGH);BangaraFalls(fl.Dec.) Wild4372 {BR, K, MO,PRE,S,SRGH);Beit Bridge (fl. Nov.) Davies 377 (MO, SRGH). MOÇAMBIQUE:Niassa: Metangula (fl.Oct.) Mendonça 738(LISC).C a b o De1gad o:Mouth of M'salo ( = Rio Messalo) (fl. Nov.) Allen 110(FHO, K, STE). N a m p u l a : Namialo, Monapo (fl.

22

Meded. Landbouwhogeschool Wageningen81-2 (1981)

Nov.) Torre & Paiva9272(LISC, neotypeof H.glabra). Z a m b e z i a : Namagoa, Mocuba Distr. (fl. Nov.-Dec.)Faulkner 328(BR, G, K, LD, PRE, SRGH); Milange road, 30-40km from Mocuba (fl. Nov.)Faulkner 488(BR,COI,K, S,SRGH, neotype ofH.tettensis). Tete: Tette (= Tete)(fr. Feb.) Kirk anno 1859 (K, LE);Nyampanda (fl.Oct.)Gillen 17510(K, LISC,SRGH, WAG, neotype of//. febrifuga); Sena (fr. Feb.) Kirk anno 1859(K, LE). M a n i c a e Sofala: Vila Paiva deAndrada (fl. Oct.)GomeseSousa 4733(COI,K, LISC,PRE);Chibabava (fl., fr. Dec.) Swynnerton 209a(BM, K, SRGH). G a z a : Massangena (fr. July) Smuts 358 (BM, K, NY, PRE). SOUTH AFRICA: T r a n s v a a l : Kruger National Park, near Punda Maria (-Ruskamp) (fl. Nov.) Lang 32126(B,K, NH, PRE). INDIA: H i m a c h a l P r a d e s h : Dalhousie (fl. June) Stewart 2117 (A, K, MO, NY). P u n j a b : Pathankot (fr. Feb.) Stewart 974(K, NY). H a y a n a : Malabar (fl., fr.) Stocks s.n. (BM, FI, G, LD, M, NY, P, S,W). D e l h i : Delhi (fr. Dec.) Kuntze 7267(NY). Gurajat:Gir forest, 5km N of Sasan (fl. May) Hodd 75(K). M a d h y a P r a d e s h : Gwalior (fl. Mar.) Maries anno 1890(BM); ibid. (fr. Aug.) Maries anno 1890(BM);Dorhora (= Duhara), Chhatapur Dt. (fl.) Wallich 1672Lpart a(KWALL); Rai Pur (fl., fr. Sept.)Sant Bam Chamla28(G). M a h a r a s h t r a : BorivliNational Park (fl. June) Hepper4400(K); Matheran (fl., fr. Apr.) Meebold2997 (G).A n d h r a P r a d e s h : Vicarabad, 85kmW.ofHyderabad(fr. Feb.) Van der Maesen3136(WAG);Secundera.bad(n.) Campbell 332(E). G o a : Bardez, Noronha 20 (LISC). M y s o r e : Belgaum (fl. Feb.) Ritchie 447 (K); Shiradi (fl. Apr.) Saldanha 13273(K, MO); Mangalore (fl.) Hohenackeranno 1849 (A,B,BM,BP,FI, G, HBG, K, L, M, MPU, P, S, UC, UPS, W, Z). K e r a l a : Taliparamba (fl. Apr.) Anstead 20101 (A). Tamil N a d u : Madras (fl. July) Gamble 16532(HBG, K);ibid. (fl. Oci.)Wight 1875(A, G, K, L, LE, S); Hosur, Salem Distr. (fl. June) Kanoth Yeshoda 146(A, B);Pondichery (fl. Aug.) Lacour anno1864 (P); Coimbatore(fl.) Miers 15428(BM).U t t a r P r a d e s h : Dehra Dun (fl. May) Singh337(NY, S); ibid, (fl.) Wallich 1673.La (K-WALL); Shrinagar, Garwhal Dt. (fr.) Wallich 1673.4 (K-WALL); Kumaon ( = Almora & Pithoragarh Dts.) (fl.) Wallich 1673.3(K-WALL); Moradabad (fl. July) Thomson 217 (K); Rampur (fr.) Wallich 1673.Lb (K-WALL). Bihar: Monghur (fl. Apr.) Wallich 1673H (K-WALL): Banka, Bhagalpur Dt. (fr. Nov.) Wallich 1673H (K-WALL); Rajmahal (fr. Aug.) Wallich 1672.3(K-WALL); Kuju, 38km S. ofHazaribagh (fl., fr. May) F. H. W. Kerr 2104B

80

90

100

MAP 5. Holarrhenapubescens, Eastern part of area. Meded. Landbouwhogeschool Wageningen81-2 (1981)

23

(BM,L).O r i s s a : AngulDistr. (fr. Dec.)Lace2580(E, WAG);betweenSalurand Koraput (fl. May) Rao 18407(L). West Bengal: Darjeeling (tl.) Cowans.n. (E); between Siliguri and Kurseong (fl. Apr.)Kanai & Murata, 29Apr. I960 (BM).Sikkim: Sukna(fr. Dec.)Rihu& Rhomoo3281 (L):Sin. loc. (fl., fr. May)Hookerf., 12 May 1850 (A,BM,FI,G, K, L,M,P,W).A s s a m : Goalpara, Wallich 1672L. a (K-WALL); Tura, Garo Hills (fl. Mar.) Koek 25013 (L, UC). M i z o r a n : Lushai Hills, Koladyne R., Parry 503 (K). N e p a l :Rapti Valley,near Ghorai (fr. Oct.)Poluninetal.5938(BM,UPS);ChuriaHills(fr. Aug.) Williams &Stainton 8247; Noakole ( = Nawakot) (bud, fl.) Wallich 1673.2 (BM, G, K-WALL); Rawa Khola (fl., fr.) Dobremez 276(BM,G); Dharan Bazar, Biratnagar (fl. Apr.) Stainton 44(BM, UPS). BANGLADESH: Sylhet(fl.) Pierre227(h, P);ibid,(fl.) Wallich1672.2(BR,K-WALL);Dacca(fl„ fr, Apr.) Clarke 6783(BM); Chittagong (fl.) Wallich1672.1 (K-WALL, W) Cox's Bayas (fl., fr. Apr.) Sinclair 2922(E). BURMA:Haka (fl.Apr.) Dickason 7590(A, E,L);Kyaukpyu (fl.May) Wallace84 (BM, K); Mount Popa (fr. Nov.) Dickason 6639(A, E,K, L);Ava, on the R. Irrawaddy (fl. Apr.) Wallich 1673.5(KWALL); Ava, Canoungaklay (fl.) Buchanan-Hamilton s.n. (BM, lectotype); Ava, Loungyee (fl.) Buchanan-Hamilton s.n. (BM, paratype); Pegu (fl. Apr.) Meebold2993 (G); Rangoon (fr.) Lelland s.n. (P.L,W);Martaban (fl.) Wallich1673.6(K-WALL); Tavoy,Tenasserim(fr.) Wallich 1673.7(G, K-WALL); Mergui (fl. Mar.) Meebold 14843(BP, Z). CHINA: Y u n n a n : Che-li (fr. Aug.) Wang 77682(A); ibid. (fr. Sept.) Wang 78076(A). Sin. loc, West-Tibet (fl., fr.) Falconers.n. (FI. K, L, LE, M, P, S, W). THAILAND: C h i a n g M a i : Doi Suthepffl. Apr.) A. F.G.Kerr 1129(BU, K, P).C h i a n g R a i : S. of Chiengkhan (fl., fr. Apr.) Garrett 146(BM, K, L). L a m p a n g : Ban Den (fr. Nov.) A. F. G.Kerr 4538(BM, K, P). P h e t c h a b u n : Phetchabun (fl., fr. Mar.) A. F. G. Kerr20344 (BM, E, K, L, P). N a k h o n P h a n o m : Dawn Tan (fl., fr. May) A. F.G.Kerr21523(BM, E, K, L, P, paratype of H. antidysenterica var. macrantha); Takli (fr. Nov.) Put 2094 (BM, E, K, L. P). K a n c h a n a b u r i : Kanburi ( = Kanchanaburi) (fl.) Marcan 951 (BM, K, lectotype of//, antidysenterica var. macrantha). Phet Buri: Cha Am (fl. June) Lakshnakara 400 (K. L. P); Chumphon (fl. Mar.) Put 1474 (K, L, P). LAOS: Attopeu (fl. Mar.) Harmand 1316 (B,HBG, P); Lao Bao (fl. Nov.) Poilane 1326(P); Cam Mon (= Ban Khammouan) (fr.) Spire 34(K, type of H.perotii) ;ibid. (fr.) Spire 36(LE,type of H. pierrei); Borikhane (fl., fr. Mar.) A. F.G. Kerr20753(BM, K, P); Sin. loc, borders of Mê-Khong (fr.) Harmand 23(FHO, HBG. K, P). CAMBODIA: Kamchay, Kampot (fl. May) Pierre 4424 (NY, P); Mt. Chereer, Samrong-tong (fl. May) Pierre4422(NY, P, UC); Kratié (ff Mar.) Chevalier31850(P). VIETNAM:Baria(fr. Nov.)Pierre1892B(HBG, K,NY,P);Dalat (fl., bark sample) Squires 778 (A, BM,BR,G, K, M,MO,P,S);Ca Na (fl.Mar.)Poilane5713(FHO,K, P); Cho-Ganh, Ninh Binh (fl. Sept.) Petelot 1444(P,UC);Thu Duc(fl., fr. Nov.) Pierre1829(h, MO,NY, P);Tam Dao (fl. Apr.) Petelot 4916(P); Lang-than (fl., fr.) Thorel 1041 (B,FHO, HBG, K, NY, P, UC). CULT.:Indi a:Bot.Garden Calcutta (fl.) Pierre3585(NY);ibid.(fl.) Pierre3989(NY);ibid. (fl.) Pierre4424 anno 1863 (P);ibid.(fl.,fr.) Wallich1672.4(BM,K, L,LE,M); Bot. Garden Pondichery (fl., fr.) Perrottet 323 (B, G, L, NY, P, W, neotype of H. macrocarpa). I n d o n e s i a : Bot. garden Bogor (fl. Mar.) Hallier C65a (G); ibid. (fl. Feb.) Hallier C65b (L). M a l a y s i a : Malay Peninsula: Sin. loc. (fl.) Wight anno 1848 (K, type of//, malaccensis) . M a u r i t i u s : Royal Bot. Gardens Pamplemousses (fl.Jan.) Gueho10543(MAU); ibid.(fl.Jan.) Vanghan11701(MAU);ibid.(fr. May) Vanghan10758(MAU). U.SA.: Plant Introduction Station, Miami, Florida (fl.June) Gillis10936; (S) ;ibid. (fl. July) Gillis 11405(BM, S).

N o t e s . BUCHANAN-HAMILTON citesin hisprotologue of this species a manuscript he wrote in 1795, which the present author has consulted in the British Museum. In thismanuscript hecitesthree specimenswith their localities,two of whicharebothmounted onasinglesheetpreserved intheBritishMuseum. With the manuscript a drawing is preserved as well which doubtlessly is based on material of H. pubescens. Of the two specimens the present author designates 24

Meded. Landbouwhogeschool Wageningen81-2 (1981)

heretheonecollected atCanoungaklay aslectotype. BUCHANAN-HAMILTON also observed that LINNEAUS erroneously based his Nerium antidysentericum on a mixture and that ROXBURGH followed LINNAEUS in this respect. STEWARD & BRANDISdidnotciteanyspecimenwiththeirH.antidysenterica var.glabrawhich isbased on a plant with glabrous leaves.Asthey did not annotate any specimen the type remains unknown. The present author has not come across any specimens of H.pubescens that are entirely glabrous from Asiatic origin, and therefore he hesitates to provide the variety with a neotype.

'H. C O N G O L E N S I S ' H. congolensis Stapf 1898: 306; De Wildeman & Durand 1899: 37-38; Durand & Schinz 1909: 345(except for Gillet 155). Type: Zaïre: Lower Congo, between Boma-Lundi and Tchoa ( = Tshoa), Cabra3(K, holotype; isotype: BR). All plants referred to '//. congolensis'are found in the neighbourhood of the mouth oftheCongo River(Cabinda, Borna,Kinshasa),considerably outsidethe areaofH.pubescens. Someofthem,belongtoeitherH.floribunda (e.g. Gillet 155) or H. pubescens (e.g. Cabra 3, the type of H. congolensis) while some other specimens appear to be rather intermediate between H. floribunda and H. pubescens: ZAÏRE: B a s - Z a ï r e : Kiobo, Luki Reserve (fl. Dec.) Wagemans 1856 (BR, K). K i n s h a s a : Kimuenza (fl.) Gillet anno 1900(BR); Kimpako (fl. Feb.) Vanderijstanno 1909(BR); ibid. (fl. Nov.) Vanderijst 13628(BR). ANGELA:Cabinda:Sumba, Peco (fl.) Gossweiler8689 (K);ibid. (fl. Nov.) Gossweiler8975 (K, LISJC).

The present author supposes that plants of H.pubescens have been cultivated bythePortuguese(SeeFICALHO 1884)atthemouthoftheCongoRiver,and may haveescaped and hybridized with H.floribunda. Evidence for thispoint of view can begleaned from thefacts that H.pubescens wasknown to thePortuguese of Moçambique under thename of 'Quina' asatonicand afebrifuge implicating a reasonable gradeofapplication implyingthenecessityofitsculture.In India the Portuguese referred to H. pubescens as 'Herbas contra camaras' or 'Herba malabarica'. The bark and seeds of H. pubescens were articles of commerce in India aswellasinEuropebecause oftheirantidysentericvalue.Inthis particular region, i.e. the mouth of the Congo River, specimens of H'.pubescens are found outsidethearea ofthespecies,whileoutsidethisregionbutwithinthearea of H. floribunda intermediates or specimensoïH. pubescenshavenever been reported.

NOMINA NUDA H. angustata Pierre ex Schmit 1950: 12. = H. curtisiiKing & Gamble. H. crassifolia Pierre exPlanchon 1894: 196. = H. curtisii King & Gamble. Meded. Landbouwhogeschool Wageningen81-2 (1981)

25

H. crassifolia var. montana Pierre ex Spire 1906: 144. = H.curtisii King & Gamble. H.perotii Spire 1906: 142;Chevalier 1948: 115-120. = H.pubescens (Buch.Ham.) Wall ex G. Don. Specimen cited: Spire 34(K). H.pierrei Spire 1906: 142;Chevalier 1948:115-120.= H. pubescens (Buch.Ham.) Wall, ex G. Don. Specimen cited: Spire 36 (LE). H. villosaAit. ex Loudon 1830: 641. = H. pubescens (Buch.-Ham.)Wall, ex G. Don.

EXCLUDED SPECIES H. affinis Hook. &Arn. 1841: 198. = Anodendron affine (Hook. &Arn.) Druce (Apocynaceae). H. daronensisElmer 1912: 1455(erroneously published as Halorrhena daronensis). = Kibatalia daronensis(Elmer) Woodson (Apocynaceae). H. landolphioides A. DC. 1844: 414. = Alafia scandens (Thonn.) De Wild. (Apocynaceae). H. (?) madagascariensis Baker 1885:424. = MascarenhasiaarborescensDC. (Apocynaceae). H. microterantha Schum. 1903: 183; DeWildeman 1903c: 515. (No type designated. Theplantdescribed isalianawithcoagulatinglatex,socan't belong to Holarrhena). H. pauciflora Ridley 1911: 132. = Spirolobium cambodianum Baill. (Apocynaceae). H.procumbens (Blanco)Merrill 1905:59.= Aganosma acuminata (Roxb.) G. Don (Apocynaceae).

REFERENCES AuBRÉviLLE, A. 1936. L aflore forestière de laCôte d'Ivoire 3.- Paris. AUBRÉVILLE, A. 1950. Flore forestière S o u d a n o - G u i n é e n n e A. O . F .- C a m e r o u n - A. E .F .- Paris. BAKER, J.G. 1885. F u r t h e r contributions t o the flora of M a d a g a s c a r . Second a n d final part. J. Linn. Soc. 2 1 : 4 0 7 - 4 5 5 . BENTHAM, G . 1849. I n : BENTHAM, G .&J.D .HOOKER, Flora Nigritiana. - L o n d o n . BERHAUT, J. 1967. Flore d uSénégal, 2nd ed. - D a k a r . BERHAUT, J. 1971. Flore illustrée d uSénégal 1.- D a k a r . BISWAS, K. 1956. C o m m o n medicinal plants ofDarjeeling and the Sikkim Himalayas. - Alipore. B R O W N , R. 1811 ('1809'). O nt h eAsclepiadaceae, a natural order of plants separated from t h e Apocineae ofJussieu. M e m . Wern. Soc. 1 :12—78. BUCHANAN-HAMILTON, F . 1822. C o m m e n t a r y on the H o r t u s M a l a b a r i c u s 1. T r a n s . Linn.Soc. L o n d o n 1 3 : 521-525. CHEVALIER, A . 1920. Exploration botanique d el'Afrique Occidentale Française 1.- Paris. CHEVALIER, A . 1948. Legenre H o l a r r h e n a etsadistribution géographique. Rev. int. Bot. appl.et Agr. trop. 2 8 : 1 1 5 - 1 2 0 . C O D D , L.E. 1951.Trees and shrubs ofthe Kruger N a t i o n a l P a r k . - Pretoria. C O D D , L. E. 1963. A p o c y n a c e a e . I n : D Y E R , R. A., L. E. C O D D & H . B. R Y C R O F T (eds.), F l o r a of

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Southern Africa 26. - Pretoria. COOKE, T . 1904. F l o r a ofthe presidency ofB o m b a y 2. - L o n d o n . C O W E N , D . V. 1950. Flowering trees and shrubs inIndia. - Bombay. CRAIB, W . G . 1912. C o n t r i b u t i o n s t othe flora ofSiam 57.- L o n d o n . CRAIB, W . G . 1913. C o n t r i b u t i o n s t othe flora ofSiam. Kew Bull. 2 :65-72. CRAIB, W .G . 1939. F l o r a e Siamensis enumeratio. A list ofthe plants k n o w n from Siam 2 (5). Bangkok. D A L E , I.R.&P. J. G R E E N W A Y 1961. K e n y a trees and shrubs. - L o n d o n . DALZIEL, J.M .1937. T h e useful plants ofWest Tropical Africa. A n appendix t othe flora of West Tropical Africa. - L o n d o n . D A T T A , P. C. & R. K. M A T T I 1972. Relationships of Plumiereae (Apoc.) 1.Based on karyology. Bull. Bot. Soc. Bengal 26 (1): 9-19. D E CANDOLLE, A. 1844. P r o d r o m u s systematis naturalis regni vegetabilis 8. - Paris. D E LA MENSBRUOGE, G. 1966. La germination etles plantules des essences arborées de la foret dense humide de laCôte d'Ivoire. - N o g e n t - s u r - M a r n e . D E WILDEMAN, E. 1903a. Etudes sur laflore du Bas- etd uM o y e n - C o n g o . A n n . M u s . C o n g o 5 ( 1 ) . D E WILDEMAN, E. 1903b. Etudes sur laflore d uK a t a n g a . - Bruxelles. D E WILDEMAN, E. 1903c. Bull. Soc. Etudes Colon. 10 (8). D E W I L D E M A N , E . & T . D U R A N D 1899. M u s . C o n g o A n n . B o t . 2 (1) pt. 1.

D O N , G . 1834. Ageneral system ofgardening and b o t a n y 3.- L o n d o n . D O N , G . 1837. Ageneral system ofgardening and b o t a n y 4. - L o n d o n . D O N , G . 1840-1843. I n : Wight, R., Icônes p l a n t a r u m Indiae orientalis 2(5): pi. 439. - M a d r a s . D U R A N D , T .&H . S C H I N Z 1896. Etudes sur laflore de l'état indépendant d uC o n g o 1. - Bruxelles. D U R A N D , T .&H. S C H I N Z 1909. Sylloge Florae C o n g o l a n a e (Phanerogamae). - Bruxelles. ELMER, A. D .E, 1912. N e w Apocynaceae. Leaflets ofPhilippine Botany 4 : 1445-1467. ENGLER, A . 1910. Die Pflanzenwelt Afrikas. I n : Die Vegetation der Erde 9(1)1.- Berlin. FICALHO, C.DE, 1884. Plantas Uteis d aAfrica Portugueza. - Lisboa. H A L L E , F .&R.A .A . OLDEMAN 1970. Essai surl'architecture etlad y n a m i q u e d ecroissance des arbres tropicaux. - Paris. H A L L E , F . ,R. A .A . OLDEMAN &P . B. TOMLINSON 1978. Tropical Trees a n dForests. - BerlinHeidelberg-New Y o r k . HASSKARL, J.K. 1857. F l o r a 7. HASSKARL, J.K. 1859. H o r t u s Bogoriensis descriptis 2. Bonplandia 7(19). H O O K E R , J.D . 1882. F l o r a ofBritish India 3 .- L o n d o n . H O O K E R , W . J. & G. A. W A L K E R A R N O T T 1841. The b o t a n y of Captain Beechey's voyage.- L o n d o n . H U B E R , H . 1962. K e w Bull. 15. H U B E R , H. 1963.Apocynaceae. I n : H E P P E R , F . N . (ed.), F l o r a of West Tropical Africa, 2nd ed., 2. London. H U B E R , H . 1973. Apocynaceae. I n : F l o r a ofCeylon 1(1).- Peradeniya. K E A Y , R. W . J. & C. F . A . O N O C H I E & D . P . STANFIELD 1964: N i g e r i a n Trees 2.

K I L L I C K , D . J.B. 1977. The flowering plants of Africa 44. K I N G , G .&S. GAMBLE 1908. Materials for aflora ofthe Malay Peninsula. Straits Branch As. Soc. Bengal 7 4 : 387-626. K L O T Z S C H , F . 1862. In : PETERS, W . C. H., Naturwissenschaftliche Reise nach M o s s a m b i q u e : 1-304. LINNAEUS, C. 1753. Species P l a n t a r u m 1.- Stockholm. L O U D O N , J.C. 1830. A catalogue ofall the plants. Supplement 2. - L o n d o n . M A R K G R A F , F . 1950. N e u e Diagnosen. Mitt. Bot. Staatss. M ü n c h e n 1(1): 2 6 - 3 1 . M E R R I L L , E.D . 1905. A review ofthe identification of the species described in Blanco's F l o r a de Filipinas. Bureau ofG o v e r n m e n t Laboratories 2 7 : 1-194. MiÈGE, J. & N . JOSSERAND 1972. N o m b r e s chromosomiques d'espèces africaines et malgaches. Candollea 2 7 : 283-292. M O R R I S (?) 1896. T w o African H o l a r r h e n a s . Kew Bull. 110. T h e a u t h o r s h i p ofMorris is uncertain. N G A N , P .T . 1965.A revision ofthe genus Wrightia (Apocynaceae). A n n . Miss. Bot. G a r d . 52(2): 114-175. OLIVER, D . 1875. T h e b o t a n y ofthe Speke and G r a n t expedition. T r a n s . Linn. Soc. 29(3). Meded. Landbouwhogeschool

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OLIVER, D . 1877. Flora ofTropical Africa 3 . - L o n d o n . PALGRAVE, K.C . 1957. Trees ofCentral Africa. - Salisbury. PALGRAVE, K.C . 1977. Trees ofSouthern Africa. - J o h a n n e s b u r g . PICHON, M . 1948. Classification desApocynacées: I X Rauvolfiées, Alstoniées, Allamandées et T a b e r n é m o n t a n o i d é e s . M e m . M u s . N a t . Hist. Nat., Nouvelle série 27(6): 153-252. PICHON, M . 1950. Classification des Apocynacées: X X V Echitoïdées et X X V I I I Supplementa u x Plumérioidées. M e m . M u s . N a t . Hist. Nat., Série B, Bot. 1(1): 1-174. P I TARD, J. 1933. Apocynacées. I n : LECOMTE, H .&H. H U M B E R T , Flore générale de lTndo-Chine3 : 1087-1262.-Paris. PLANCHON, L. 1894. P r o d u i t s fournis à lamatière médicale par lafamille des Apocynacées. Thèse Agrég. P h a r m . - Paris. RAGHAVAN, R.S. 1957. C h r o m o s o m e numbers ofIndian medicinal plants. Proc. Indian Acad. Sei. 45(B): 294-298. RIDLEY, H .N . 1911. A naccount ofa botanical expedition t olower Siam. J. Straits Branch Royal Asiat. Soc. 59. RIDLEY, H .N . 1923. T h e flora ofMalay Peninsula 2.- L o n d o n . R O X B U R G H , W . 1810. I n : F L E M M I N G S , M e d . PI. & D r u g s . A s . R e s . 11.

R O Y T A P A D A R , N . N . 1964. C y t o t a x o n o m i c studies in Apocynaceae and delineation of the different evolutionary tendencies operating within the family. Caryologia 1 7 : 103-138. SCHMIT, A . 1950. Recherches botaniques chimiques et p h a r m a c o d y n a m i q u e s s u r l'Holarrhena floribunda (G. D o n ) D u r . &Schinz. - Paris. SCHUMANN, K. 1895a. I n : ENGLER, A., Die Pflanzenwelt Ost-Afrikas u n d der Nachbargebiete C. Berlin. SCHUMANN, K. 1895b.I nE N G L E R , A. & K. P R A N T L , Die natürlichen Pflanzenfamilien 4(2).- Berlin. SCHUMANN, K . 1898. I n : D U R A N D , T .&E. DEWILDEMAN, Matériaux p o u r la flore d u C o n g o 2. Bull. Soc. Roy. Bot. Belg. 37. SCHUMANN, K. 1903. Tropenpflanzer 7. SEN, N . K.&N . N .R O Y T A P A D A R 1957. Cytology ofH o l a r r h e n a antidysenterica Wall. Proc. 44th Indian Sei. Cong.4 . SPIRE 1906. C a o u t c h o u c s d ' I n d o - C h i n e . - P a r i s . STAPF, O. 1902. I n : THISELTON-DYER, W . T., Flora ofTropical Africa 4(1). - L o n d o n . STEWARD, J.L.&D . BRANDIS 1874. Flora ofNorthwest and Central India. THWAITES, G .H .K.&J.D .H O O K E R 1860. E n u m e r a t i o p l a n t a r u m . - L o n d o n . V A H L , M . 1790. Symbolae Botanicae 3.- K e b e n h a v n . V A N R H E E D E T O T D R A A K E N S T E I N , H . & J. CASEARIUM& A . SYEN 1678. H o r t u s I n d i c u s M a l a b a r i c u s 1.

- Amsterdam. VILLAR, F . 1880. I n : Blanco, P.M., F l o r a de Filipinas 4. - Manilla. W A L L I C H , N . 1829. Anumerical list ofplants n.1-2153. - L o n d o n . W I G H T , R. 1848. Icônes p l a n t a r u m Indiae Orientalis 4. - M a d r a s . WULFSBERG, N . 1880.H o l a r r h e n a Africana D C . eine tropische Apocynacee. Inaugural-Dissertation.

- Göttingen.

ACKNOWLEDGEMENTS I would like tothank everybody atthe laboratory ofPlant Taxonomy and Plant Geography of Wageningen, especially Dr. A. J. M. LEEUWENBERG for his supervision, Ir. J. J. Bos for his correction of the text, Mr. G. BOELEMA for his assistence in correcting the proofs, and Miss J. WILLIAMSON for her fine drawings. Also I am very grateful toMr. L. AKÉ ASSI ofthe Centre Nationalde Floristique of Abidjan, Ivory Coast, and toMr. BOULABA SAVADOGO ofthe Centre Néerlandais of Adiopodoumé, Ivory Coast, formyintroduction to tropical vegetations;to the directors and keepers of the following herbaria for 28

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putting their material at my disposal: A (Cambridge, U.S.A.), ABI (Abidjan), AAU (Aarhus), B (Berlin), BM (London), BP (Budapest), BR (Bruxelles), C (Kobenhavn), CGE (Cambridge, U.K.), COI (Coimbra), E (Edinburgh), EA (Nairobi), FHO (Oxford), FI (Firenze), G (Genève), G-DC (Genève), GH (Cambridge, U.S.A.),HBG (Hamburg),K (Kew),K-WALL(Kew),L(Leiden), LD (Lund), LE (Leningrad), LINN (London), LISC (Lisboa), LISJC (Lisboa), LISU(Lisboa),MAU (Mauritius),M(München),MO(Missouri),MPU (Montpellier), NY (New York), NH (Durban), OXF (Oxford), P (Paris), PRE (Pretoria), S (Stockholm), SAM (Cape Town), SRGH (Salisbury), STE (Stellenbosch), TCD (Dublin), U (Utrecht), UC (Berkely), UCI (Abidjan), UPS (Uppsala), W (Wien), WAG (Wageningen), Z (Zürich). I would especially like to thank everybody whohelped meduringmyvisitstotheherbaria and librariesof Abidjan (ABI, UCI), Bruxelles (BR), Genève (G, G-DC), Kew (K, K-WALL), Leiden (L), London (BM, LINN), Paris (P),Utrecht (U) and Zürich (Z). I recieved a grant from the Wageningen Agricultural University Fund for my visits to Kew, London and Paris.

INDEX OF EXSICCATAE The number in parentheses corresponds with the species-number of the revision. Only numbered collections have been listed. Achten, L., 747(2). Acocks, J. P. H., 16799(4). Adam, J.G., 992,1375,16477,25844,28104(2). Adams, C. D., 2434, 2470, 2838 (2). Adekunle, A.O., FHI 3007(2). Adlard, P. G., 19,95, 372,(4). Adogla, S. G., 4078(2). Aguiar Macêdo, de, see De Aguiar Macêdo. Aikamphon, K., 182(1) Ainslie, 387(2). Allen, C. E. F., 100, 110(4). Allen, J., 7(4). Anderson, 28(4). Anderson, B., 821, 954(4). Anderson, E. F., 4686 (1) Andoh, J. E., 4884(2). Andrada, E. C , 1227, 1643(4). Andrews, F. W., A292, A338(4). Angus, A., 703, 1439, 1557,2077(4). Anstead, R. D., 20101 (4). Armitage, F. B., 155/55(4). Astle, W. L., 1061,4151(4). Aubréville, A., 64,95D, 115C, 158C,661, 1157, 1417, 1710,2732(2). Aubuli Bandula, 785(4).

Audru, J., 4227,4999(2). Autrique, E., 79(4). Azuma, S., 665(4). Badré, F., 301(2). Bainbrïdge, W. R., 184/55(4). Balansa, B., 2114(4). Baldwin J. T.,jr., 9793, 9883, 10102, 13279(2). Bally, P. R. O., 7534, 8850, 11629, 13610(4). Bally, P. R. O. & Carter, 16419(4). Balsinhas, A., 1055(4). Balwanssing, 11(4). Banda, E. A. K., 604, 1195(4). Banne, J. H„ 9(2). Bar, N. L., 2880(4). Barbosa, L. A. G., 827,988, 1232,2573(4). Barbosa, L. A. G. & M. F. Carvalho, 4390(4). Barker, 533(4). Barnard, D. E., M. C. T. 37(4). Barter, C , 3334(2). Bates, G. R. 15722(4). Batten-Poole, W. H., 205(2). Battiscombe, E., 105(4). Batty, M., 663, 1258(4). Bayliss, R. D. A., 4646, 7200(4).

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Beddome, R. H., 5101,5102(4). Bèjeaus, 800(4). Benedicto, J., 99(4). Benoît, M., 403(2). Benson, F. M., 599,624(4). Berghen, C. van den, seeVanden Berghen. Berhaut, J., 718, 3370, 3900, 5332(2). Bernardi, L., 140-(1962) (2), 15507(3). Best, E. B.,63(4). Beusekom, C. F. van, seeVan Beusekom. Biegel, H. M., 1773,4731(4). Bingham, M. G.,'1449(4). Bishan Singh, 67(4). BisRam, 308, 564(4). Biswas, K„ 4794(4). Bj0rnstad, A., 702(4). Boaler, S. B.,329, 342, 500(4). Bock, K., 231(4). Bon, H. F., 4789(4). Boudet, G., 3924, 5002, 18966(2). Boue, A., 19(2). Boulanger, J. P., 1628(4). Bourne, A. G., 3408(4). Bowbrick,J. M.,J158(4). Brandis, D., 435,442, 443, 1514, 3085(4). Brayne, H. C , 21(4). Bredo, H. J. A. E. R., 3396, 3518, 3521, 3522, 3526 (4), 3548 (2), 3549, 3669, 4017, 4231, 4665, 4857, 5869,6316(4). Brenan, J. P. M. &P.J. Greenway, 8160(4). Brenan, J. P. M. & R. W. J. Keay, 8655(2). Breteler, F. J., 2103(2). Brown, J. W., 326,682(2). Bruce, E. M., 390, 537, 846 (4) Brummitt, R. K., 10579(4). Brummit, R. K. & E. A. K. Banda, 8750(4). Buchanan, J., 32,484, 1114(4). Buchanan-Hamilton, F., 735,736(4). Buchinger, 716(4). Bullock, A. A., 1244, 2287, 2316, 3431, 3504, 3512,3514(4). Bunchuai, K., 396(4). Bunnak, 815(4). Burbridge, K. G., 527, 588(2). Burma Forest SchoolHerbarium, 10,12,21,24, 32, 33, 703 (4). Burnett, G. F.,48/38,49/21 (4). Burrows, H., 969(4). Bum, B. D., 79,2909, 3410, 3544, 5399(4). Busse, W. C. O., 85,248, 248a, 485,490, 490a, 508, 1271,2734(4). Butreau, 20(4). Cabra, A. F. F., 3(4). Caille, O., 18063, 18120(2).

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Callens, H., 2149, 4504(2). Cameron, K. J., 4(4). Campbell, 32, 332(4). Cannell, I., 9(4). Carlier, A., 168(2). Carmichael, 1007(4). Carnochan, F. C , 46, 343,343a (4). Carson, H. M. L., 65(4). Central National Herbarium, Botanical Survey oflndial082(4). Central Research Station, Mazabuka, Department of Animal Health 432, 502(4). Chambo, S4(4). Chapman, J. D., 551, 1741(4). Charles, 16(2). Chase, N. C , 296, 809, 1319, 1519, 2224, 2378 (4). Chesters, D. F., 131(2). Chevalier, A., 620, 621,857, 2685, 3548, 4506, 4507, 6860, 6930, 8019, 8119, 8310, 8525, 8567, 8767, 8824, 9250, 9678, 12235, 12320, 12343, 12425, 12885, 13381, 13436, 13895, 13921, 13927, 13934, 13948, 14083, 14100, 15622, 17948, 18292,21226, 22978,23192, 24110, 24610, 24611, 24613, 24666,(2), 31850 (4), 31949 (1), 34019 (2),40569, 40572(4). Chillou, J., 606, 1388, 2628 (2). Chin, 6830(4). Chorley, J. K., 39943(4). Christiaensen, A. R., 778(4). Clarke, C. B.,6783,20306D (4). Cleghorn, W. B., 320,468(4). Clements,J. B., 154,199, 299, 384, 508,672(4). Clutton-Brock, T. H., TCB143,TCB262 (4). Cochet, S., 27(2). Codd, L. E., 4236, 6896(4). Cole, M„ 47(4). Collen, H., 626(4). Collins, D. J., 777,808,854 (4) Compère, P., 691 (2). Cooke, A., 94(4). Corner, E. J. H., 31519(1). Correia, M. F. &A. Marques, 2582(4). Correia, M. F., A. Marques & M. Adela Dinz, 3560(4). Crichlon, G. E., 34(2). Culwick, G. M., 4(4). Curtis, C , 3392(1). Dale, I. R., 1073(4). Dale, M. O., SKF270 (4). Dalziel, J. M., 156, 208, 370, 534, 971, 1346, 8079, 8080, 8402(2). Danigrake, B. O. & Osanyinlusi, FHI 90127,

Meded. Landbouwhogeschool Wageningen81-2 (198!)

FHI 90198 (2). Davies, R. M., 377(4). Davio, C , 54(2). Dawe, M. T., 40,47 (2), 337,367(4). Dawodu, T. B.,259(2). De Aguiar Macêdo, J. M., 4919(4). De Fabrègues, B.P. 3066(2). Deighton, F. C , 4, 1917, 3147,4218, 5930(2). De Koning, J. 5930, 6555(2). DeKruif,A.P. M., 19,25,26,27,237,261,274, 444, 502, 503, 620(2). De Leeuw, P. M , 54(2). Démange, 2087(2). Demeulemeester, A., 19(2). Den Hoed, G. &A. Kostermans, 891(4). De Saegher.H., 113(4). De Troyer, 7(4). Devred, R., 244,412, 2781(2). DeWailly,4615(2). De Wilde, J., 692(4). De Wit, H. C. D., 6047(2). DeWitte,G.F.,42,2910,2952,4170,4505,4620 (4). Dickason, F. G., 6639, 7590, 7842, 9158(4). Din, 123 (4). Dobremez, J. F., 94,276(4) Donis,C., 1582,2427(2). D'Orey, J., 62, 191,202(2). Drummond, J. R., 498,25510, 25511(4). Dubois, R., 75,203(2). Dumas, 18162, 18179(2). Dussaud, 122(4). Eaux, Forêts et Chasses R. C. A., 2003(2). Eberhardt, P. A., 4076,4896, 5026(4). Edwards, F. C. 24(4). Edwardson, T. E., 123(2). Eggeling, W. J., 6262, 6381(4). Enti,A. A., 1375, 1661(2). Ern, H., 3158(2). Espirito Santo,J., 208, 1978,2393(2). Etesse, M., 31(2). Evans, A. C , 14(4). Exell, A. W., F. A. Mendonça & H. Wild, 864 (4). Eyles, F., 2761,3492, 5167, 6276(4). Eynatten, van, see Van Eynatten. Ezanno, 18(2). Fabrègues, de, seeDe Fabrègues. Falconer, H., 789(4). Fanshawe, D. B.,3961,9068,9303(4). Farmar, L., 275(2). Farron, C , 7019(2). Faster, E. W„ 46(2).

Faulkner, H. G., 328, 488, 1426, 3787, 3920, 4304, 4321(4). Fazl Din, 6269(4). Fernandes, J., 19(4). FHI, 3007, 3694, 7571, 9206, 14177, 15523, 17732,21666,22340,25788, 37775,79443(2). Fischer, 3,378,857(4). Flamigny, F., 229, 6192, 7084(2). Fleury, F., 22092 (2), 32556, 32635, 37698 (4). Fotius, G., 1094, 1594, 1961(2). Franck, C. W., 477,478 (1). Fundi, T. H., 26(4). Gaétan Myembe, 240(4). Galpin, E. E., 15029(4). Gamble, J. S., 326A. 591A, 3207A, 7389,8648, 9111, 16532,26896(4). Garcia, J.G., 611(4). Garrett, H. B.G., 28, 146(4),293(1). Garrigues, 596(2). Gathii, S., 120(4). Geerling, C , 5382(2). Geerling, C. &J. Bokdam, 2172, 2509(2). Geerling, C. &Néné, 4980(2). Geesink, R., D. Phanichapol & T. Santisuk, 5838(1), 5925(4). Geesink, R. & C. Phengkhlai, 6148(4). Geilinger, W., 3184, 3186, 3216(4). Geoffray, 444(1). Germain, R., 2016(2). Gibbs Russell, G. E., 2610(4). Gilbert Rogers, C , 283(4). Gillet,J., 155,347,2758(2). Gillett, J. B., 3057, 3087, 17510, 17928, 17994(4). Gillis,W. T., 10936, 11405(4). Gillman, H., 1191, 1233(4). Glanville, R. R., 181,250(2). Glover, P.E., 6316(4). Godefroy-Leboeuf, A., 189(1). Goetze, W., 36(4). Gomes Pedro,J., 4376,4429(4). Gomes e Sousa, A. F., 4702,4733(4). Goodier, R., 97,234,437, 628,665(4). Gordon, T. A. D., 143(4). Gossweiler, J., 6155, 6822 (2), 8689, 8975 (2 x 4), 9062(2). Graham, R. M., 316(4). Greenway, P.J., 2672,6376,6399(4). Greenway, P. J. &Kanuri, 12653, 14627, 14812(4). Griffith, W., 2346(4). Gueho,J., 10543(4). Guigonis, 2564(2). Gurdial Singh 10(4).

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31

Haarer, A. E., 1655,2007(4). Habid Bakhsh, 80(4). Haerdi, F.,6, 178/0(4). Haines, H.H., 2713(4). Hakki,M.,B.Leuenberger&C.Schiers,980(2). Hall-Martin, A. J., 427,715, 1095(4). Hallier, J. G., C65a, C65b (4). Hambier, D. J., 10(2). Haniff, M. &G. M. G. Nurr, 4272(1). Hara, H., H. Kanai, G. Murata, M. Togashi & T. Tuyama, 1508, 1513(4). Hara, H., H. Kanai, S. Kurosawa, G. Murata, M. Togashi &T. Tuyama, 6303484^4). Harley, W. J„ 358, 1136(2). Harmand, F. J., 23, 1107, 1316, 1828B(4). Harris, B.J., 1979,3813,4456(4). Harris, B.J. &T. &S. Poes, 4228(4). Harsukh, 22141/C, 22141/D, 22142 (4). Hayes, W., 509(2). Hemm, G., 483(4). Henderson, M. R., 22935(1). Henkel, J. S., 187(4). Hepburn, I. D., 49(2). Hepper, F. N., 4400(4). Herbarium Peradeniya 493(1). Herklats, 105(4). Heudelot 795(2). Hira Lai, 13402(4). Hodd, K. T. B.,75(4). Hodgson, L. M., H8/49 (4). Hoed, den, seeDen Hoed. Hohenacker, R. F., 2385(4). Holmes,W. D., 494, 1880(4). Holst, C. H. E. W., 8837(4). Honey, T., 785, 789(4). Hornby, A. J. W., 2958(4). Hornby, R. M., 2754(4). Huart, A., 8(4). Huber, H., 34(3). Hucks, M., 539(4). Hügel, K. A. A., 1896,2362, 2759(4), 3786(3). Hutchinson, J. &J. B.Gillett, 3299(4). Ihlenfeldt, H. D., 2278(4). Ire, W. B.,250 (2). Irvine, F. R., 1526, 1917, 3234(2). Jackson, C. H. N., 1 (4). Jackson, G., 316, 1378, 1677,2131(4). Jackson, J. A. D., 537 = FHI 17732(2). Jacobsen, W. B.G., 2608(4). Jacques-Félix, H., 29, 3545(2). Jamieson, D., 942(4). Jarre«, F. M. &T. P. Ramamoorthy 1079(4). Jarrett, F. M., C. Saldanha & T. P. Rama-

32

moorthy, 614(4). Jayasuriya, M., L. H. Cramer & S. Balasubramium 793(3). Jenkins, F., 3585(4). Joanna, B., 8850(4). Johnston, H. H., 68(4). Jones,A. P.D.,R. W.J. Keay &C. F. Onochie, FHI 14177(2). Jones, M., 55(4). Jorcia,611(4). Jordan, H. D., 226,890,2033(2). Kanai, H., G. Murata &T. Tuyama 1511(4). Kanoth Yeshoda, 146(4). Katz & Schmutz 3056(2). Keay, R. W. J., FHI 7571,FHI 37775(2). Kennedy, J. D., 127(2). Kerr, A. F. G., 1129, 1399(4),2357,2548,3688, 4250(1),4251,4538, 5357(4),5693, 8322(1), 9159,10575,10627,10627a (4), 12266,13094, 13583, 15889,18531,19322(1), 19544,19551, 20344(4),20450,20739(1),20753, 21523(4). Kerr, F. H. W., 2104,2104A, 2104B(4). Kersting, O., 107bis', A361 (2). Khantchai, H., 354(4), 1083(1). King, R. M., 5424, 5506(4). Kingdon Ward, F., 3011, 22191 (4), 37509(1). Koelz, W. N., 18957, 19079, 22825, 24969, 24993, 25013,25143, 30508(4). Koerner, 2270(4). Koning, de, seeDe Koning. Koritschoner, H., 1548, 1632(4). Kruif, de, see De Kruif. Kunstler, H. H., 1402(1). Kuntze, C. E. O., 7267, 7630(4). Lab. de chimie org., Fac. des Sciences, Yaoundé, 16, 30,34, 35, (2). Lace, J. H., 10,2580(4). Ladell,W. R. S., 205(1). Laferrère, M., 102(2). Lakshnakara, M. C , 321 (1),400 (4), 882(1). Lamont, 27983(4). Lane-Poole, C. E„ 61,61 bis, 113(2). Lang, H., 32126(4). Langdale-Brown, L, 2613(2). Larsen, K., 8827(4). Larsen, K., S. S. Larsen, I. Nielsen & T. Santisuk, 31014(1). Larsen, K., T. Smitinand & E. Warncke, 1445

(DLaurent, J., 324(2). Lavieren, van, see Van Lavieren. Lawlor, D. W. & J. B. Hall, 520(2). Lawrence, E., 121(4). Meded. Landbouwhogeschool Wageningen81-2 (1981)

Lawton, R. M., 134,290, 1212, 1345(4). Leach, L. C , 9486A, 9520(4). Leedal, G. P., 2381, 2419, 2474(4). Leeuw, de, see De Leeuw. Leeuwenberg, A. J. M., 3888,4376, 4393,9664 (2), 10709 (3), 10866, 10867 (4), 11046, 11105,11194, 11235, 11341, 11911, 11961, 11990, 12002, 12132(2). Leistner, O. A., 1471(4). Lely, H. V., 122, 311,P502 (2). Leonhardt, 92(4). Lescrauwaet, E., 80(2). LeTestu,G. M.P.C , 122(2),572,577,721,889 (4), 2501,3295, 5053,7803,8540(2). Letouzey, R., 2977,4667, 6836, 14017(2). Lewalle,J., 7971(2). Lewis, W. E., 143(4). Lindeman, H. A., 141(4). Lisowski, S., 55170, 55171 (4), 55199(2). Lovemore, D. F., 322(4). Lowe, T., 1280(2). Ludanga, R. I., 854, 1538(4). Ludanga, R. I. &K. Vollesen, 2469(4). Luja, E. P., 249 ser. 3(4). Lusaka Nat. Hist. Club, 63(4). Macaulay, J., 1053(4). McClounie, J., 70, 116(4). McGregor, G. M., MG/47 (4). Maclaud, C , 236, 327,451 (2). Macrae, J., 52(3). Maesen, van der, see Van der Maesen. Majumber, S. &A. Islam, 60(4). Makin, J., 55,252(4). Mann, G., 884(2). Marcan, A., 375(1),913,951 (4), 1744,1777(1), 2176(4). Marks, S. A., 7(4). Martin, J. P., 401/38(4). Martin, M., 10(1). Mavi, S., 1371(4). Maxwell, J. F., 73-512 (4), 74-372 (1), 74-831, 75-36, 75-225 (4). Meebold, A. K., 2968, 2993,2997, 4698, 14843 (4). Meer, van, seeVan Meer. Meikle, R. D., 869, 1339, 1355, 1450(2). Mendonça, F. A., 477, 738, 1142, 3697, 3795, 3955(4). Menyharth, 850(4). Mfinanga, R. M., 4(4). Mga Hla Maung, 27(4). Mgaza, C. D., 357(4). Michelmore, A. P. G., 706, 1031,1091(4). Michelton, 1158(2).

Miers, J., 15428(4). Migeod, F. W. H., 36, 39, 128, 362, 366, 532, 1014(4). Mildbraed, J., 4616, 8072, 9082, 10717(2). Milien, H., 145, 179(2). Miller, O.B., 3131(4). Milne-Redhead, E., 1250,6265(4). Milne-Redhead, E. &P. Taylor, 7368, 7679(4). Miquel, 34,42(2). Miszewski, K., 36(2). Mitchell, B.L., 2719, 2844(4). Mokim, S., 110,411, 633, 1064(4). Moll, E. J., 354(4). Mongonierma, A., 540(2). Monro, C. F. H„ 809, 1789,2165(4). Mooney, H. F., 430(4). Moor, H. W., 10(2). Moors, D. R., k4(4). Morris Goodall, M. M., 84(4). Morton, J. K., A 489, SL 1296,SL 1351(2). Morton,J. K.&D.Gledhill, SL1164,SL1811(2). Morton, J, K. &A. S. Jarr, SL 3524(2). Mosnier, M., 2576(2). Mukerjee, S. K.,431, 1545(4). Müller, T., 215(4). Mullin, L. J., 77/56 (4). Munch, R. C , 27,222(4). Mundy, H. G., 2802(4). Muri, Y. W., 75(4). Musk, H., 163(4). Mutimushi, J. M., 1645(4). Myembe, G., 240(4). Nandi Singh, R., 1 (4). Neves Rosa, F., 65(4). Newman, J. L., 55(4). Ngameni Kamga, B., 154(2). Nilviset, 12(4). Nodgson, L.W., 48/49(4). Noe,N., 157(1). Nongonierma, A., 540(2). Noronha, A. P., 20(4). Obeng-Darko, K., 509(2). Obermeyer, A. A., 2001(4). Onochie, C , FHI 3694(2). Onwudinjah, D. C , FHI 22340(2). Orey, J. d', see D'Orey. Oyayomi, Flagbemi, Onijamowo, Oguntayo & Arasi, 304 = FHI 79443(2). Pannell, D., 2127(4). Parasweam Datha, 13(4). Pardy.A. A., 11/55(4). Paroisse, G., 11,12,20, 47, 134, 199,218 (2).

Meded. Landbouwhogeschool Wageningen81-2 (1981)

33

Parry, N. E., 503, 1335(4). Patterson, M., 1 (4). Pauwels, L., 4390(2). Pawek, J., 3067, 5088, 6003, 6324, 7734, 12001 (4). Pedro, E., 2104(4). Pedro, E. &Pedragâo, 5386(4). Perrottet, 307,309, 323(4),458(2). Petelot, A., 961, 1444(4), 1696(1), 2422,2437, 4916,5984,5990(4). Peter, A., 31925, 31945, 32592, 33342,33533, 33553, 35851, 36289,45723,45737, 58154, 58287 (4). Phelps, R. J., 5, 173(4). Phengkhlai, C , 141,160(4). Phengkhlai, C. &T. Smitinand, 243(4). Phipps, J. B.,230, 886, 1277(4). Pierre,L., 174 July 1867,174July 1866,174July 1874(1).227,1829Apr. 1866,1829Nov.1866, 1829 July 1877,1829B, 3585,3989,4422,4424 anno 1863, 4424 anno 1874, 4425 (4), 4486, 5243(1). Pierre, N., 1968(2). Pirozynski, K. A., 153,320, 537(4). Plowes, D. C. H., 1522, 1726, 2375, 2507, 3448(4). Pobéguin, H., 14, 1272(2). Poilane,E., 1326,1682,2259,5713(4),11483, 11602, 13782 (1),26092(4),28242(1). Pole-Evans, I. B.,6675,6685(4). Pole-Evans, I. B.&J. Erens, 518(4). Polunin, O., W. R. Sykes & L. H. J. Williams, 5938(4). Porteras, R., 622(2). Prain, 1 (4). Price,J., 23(4). Prittwitz, V., 152(4). Proctor, J. E. A., 84, 300, 1692,2747(4). Put, 1474 (4), 1487, 1681 (1), 2094, 2662, 2843 (4). Pyne, C. T., 10(2). Raizada, M. B.,80(4). Rao, R. S., 7206, 10880, 18407, 39114(4). Rattray, J. M., 1760(4). Rauh, R., 278 (4). Rawlins, S.P., 278(4). Rees, A. F., Tl86(4). Reid, N. R., 550(4). Rensburg, van, seeVan Rensburg. Renvoize, S. A. & R. A. Abdallah, 2370(4). Reporter on EconomicProducts tothe Government of India, 11351, 11489(4). Ribu & Rhomoo, 3281(4). Rich, H. H., 570(4).

34

Richards, H. M , 3589,4054, 5029, 6265,7931, 11470, 11543, 11939, 17990, 18699,20543(4). Richards, M , 26226(4). Richards, M. & S.Arasululu 25964(4). Richards, P.W„ 5093B(2). Ridley, H. N., 14981, 14982, 15864(1). Ritchie 447(4). Robertson,197,236(4). Roberty,G.,2581,6419,6505,6604,10006, 10302,10752,13829,13839,14144,16419, 16441,16442,17195,17326.17532(2). Robijns,W.,137(2). Robinson, E. A., 368,890,2474, 3128(4). Robson,N.K.B.,81,1639(4). Rock, J. F., 1623, 1637(4). Rodgers, W. A. 646, 811(4). Rogers, F. A., 8424, 8941(4). Ross, A. F., 121,136(2). RossTownsend, R.G., seeTownsend, R. G. R.. Rottler, J. P., 22(3). Rounce, N. V., 635(4). Rowland-Jones, M., 1 (4). Royal Forest Department of Thailand, 2504, 11293(4). Ruffock, C. K., 124(4). Ruse, L. F., 92(4). Sacleux,C, 2214,2215(4). Saegher, de, see De Saegher. Saldanha, C. J., 13069, 13273, 16605(4). Salubeni,A. J., 189(4). Samany & Brant, E59(4). Sandwith, C , 38,50,61(4). Sanford,W .W.,6780,6795(2). Sangina,U.,63(4). Sangkachun, B., 566(1). Sangkhachand, B.,969(4). Santapau, H., 454, 11586, 12776(4). Sant Bam Chamla, R28 (4). Savory, C R . , 231(4). Schlagintweit, 14648(4). Schlechter, R., 12296, 12969(2). Schlieben, H. J., 1526, 3124, 3326, 5440, 9333 (4). Schmitz, A., 2105,5029, 6653,6690,6861(4). Schmutz, B6375(2). Schnell, R., 2288(2). Scholes,J., 137(2). Schijff, van der, seeVan der Schijff. Scott Elliot, G. F., 4290a, 4469, 5180 (2), 8419 (4). Sedgewick, L.J. &T. R. D. Bell,6122(4). Semsei, S. R., 560,997, 1033, 1503,1878, 3356(4). Serv. For. Côte d'Ivoire, 406,434(2).

Meded. Landbouwhogeschool Wageningen81-2 (1981)

Serv. For. Sénégal, 54(2). Shabani, R., 63,70,84,257,878(4). Shaw, B. E. B.,97(2). Silva, J. M., 176(3). Simâo, J., 207, 765, 124/48, 162/48,364/48, 682/48, 714/48(4). Simon, B.K., 2295A(4). Sinclair, J., 2922(4). Singh, U., 41,337(4). Sita, P., 1857,2180, 2853(2). Small, D., 149(2). Smith, 51(2). Smitinand, T., 1596, 10887, 10893, 11028(1). Smuts,J. C , 358(4). Smythe, C. W., 4019(2). S0rensen,T.,K. Larsen &B.Hansen 2193,7306 (1). Spichiger, 50(2). Spire, 34, 36(4). Squires, R. W., 778(4). Stainton, J. D. A., 44, 5625,6859(4). Stainton, J. D. A., Sykes & Williams, 5236, 5237, 5875(4). Stevenson, D., 128, 131/30, 144/30, 324/31, 482/32 (4). Stewart, D. R. M., 57(4). Stewart, R. R., 974, 2117, 11465, 13099, 15009 (4). Stinnesbeck, T., 8754(4). Stone,B.C., 11306 = Anderson,E.F.,4686(1). Stopkins, J. B.,6735(4). Strachey, R. &J. E. Winterbottom, 1,2(4). Strid, A., 2284(4). Stuhlmann, 6579(4). Subbarao, G. V.,24214(4). Suksakorn, S., 905(1), 930(4). Swynnerton, C.F. M.,209,209a,209b,990,992, 1002, 1005, 1006(4). Talbot de Malahide, 13,145(1). Tanner, R. E. S., 323,991, 1475(4). Tessier-Yandell, 36(4). Thiébaud, A., 528(4). Thomas, A. S.,D175(2). Thomas, L., 17(4). Thomas, N. W., 106, 167, 420, 1743,2587(4). Thomson, 217, 734(4). Thorel, 1041 annis 1862-1866, 1041 annis 1866-1868(4), 1247, 1403, 1409(1),9127(4). Thornewill, A. S., 42, 51(2). Thulin M. & B.Mhoro, 669, 1256(4). Thwaites, G. H. K., CP 756(3). Thyssen, M. T., 229(2). Tisserant, R. P.C , 843,2501,3295(2). Tixier, P., 13/11/55-05 (1), 18/3/56-07(4). Meded. Landbouwhogeschool Wageningen 81-2 (1981)

Ton, L., 227 (4). Torre,A. R., 927, 1060,3447,3760,4072, 4278, 4787, 5330, 5572, 6140, 6253(4). Torre, A. R., M. F. Carvalho & C. P. Ladeira 18867(4). Torre, A. R. & M. F. Correia, 13020, 13224, 13854, 18069, 18324(4). Torre, A. R. &J. Paiva, 9132, 9272, 11792(4). Toussaint, L., 67,428,2444(2). Toutain, B.,2551(2). Touzet, G., 3(2). Townsend, R. G. R., 63,71,108(4). Trapnell, C. G. 432, 502(4). Trochain, J., 80 bis, 496, 522, 694, 1328,3218, 3316, 3437, 3808 bis,4209(2). Troth, R. G., 757(4). Troupin, G., 44, 507, 1728(2). Troyer, de, see De Troyer. Turner & Shantz, 4147(2). Vaillant, A., 197bis(2). Van Beusekom, C. F. &C. Charenpol, 1742(4). Van Beusekom, C. F. &C. Phengkhlai, 1245(1). Van Beusekom, C.F.,R. Geesink, C. Phenkhlai &B. Wongwan, 3881(4). Vanden Berghen, C , 1869,1882,2456,2768(2). Van der Maesen, L., 3136(4). Van der Schijff, W. P., 1896, 3154, 3337(4). Vanderijst, P. H., 13628 (2 x 4), 13684, 13884 (2). Van der Zon, A. P. M., 510 (2). Van Eynatten, C. L. M., 1425(2). Vanghan, R. E., 10758, 11701(4). Van Lavieren, Bayer & Rees 312, 750(4). VanMeer,P.P.C , 626,679,691,799,865,1052, 1055 (2). Vanpruk, 156,275, 648(1). Van Rensburg, H. G., 2271,2576,2694(4). Vasse, G., 50,304(4). Vaughan, J. H., 2590(4). Verdick, L., 27, 77, 103(4). Verger, 1398(2). Versteegh, C , & R. W. den Outer, 631(2). Vidal,J. E., 1082B(4). Vigne, C , 1648, 1733, 1920,2996(2). Voigt, J. D., 143(4). Vollesen, K., 2469, 3018,3072,4255(4). Vroumsia Tchinaye, 93VT (2). Vuillet. 19, 62,421,(2). Wagemans,J., 856,1078(2), 1856(2 x 4), 1939 (2). Wagh, S. K., 4957(4). Wailly, de, seeDe Wailly. Wallace, C. V., 43, 161, 178(2).

35

Wallace, E. C , 84(4). Wallace, G. B.,265, 370(4). Walker, 27,99(3). Waller, 3803(4). Wallich, herb., 1672.1, 1672.2, 1672.3, 1672.4, 1672E, 1672F, 1672G, 1672H, 1672J, 1672L, 1673.1, 1673.2, 1673.4, 1673.5, 1673.6, 1673.7, 1673H(4). Wang, C. H., 77682, 78076, 78705,80812(4). Ward, 15(4). Warnecke, 134(2). Waterlot, 1122(2). Watkins, G., 539, 2341,2342(4). Watt, G., 2027(4). Welch, J. R., 421(4). West, O., 3455, 3781(4). Whellan.J. A., 474, 2199(4). White, F., 2041,3571,3588 (4), 8296, 8324(2). Whyte, A., 89(4). Wigg, L. T., 1794(4).

36

Wight, 1501 (3), 1874, 1875, 1876, 1877, 2544 (4). Wild, H., 2351,2370, 2620,4372(4). Willan, R. L., 630(4). Williams, L. H. J., 70, 353(4). Williams & Stainton, 8247, 8357,8545(4). Windward, G. R., 1 (4). Winit, K., 237 (4),238, 397, 398(1). Wit, de, see De Wit. Wit, P., 1294 (2). Witt, P. O., 21(4). Witte, de, seeDe Witte. Wood, J. J., 136,187(4). Worthington, T. B.,4539,4625(3). Yuang,44(l). Zimmermann, A., 892,2113, 2152(4). Zoe Brain, 50(4). Zon, van der, seeVan der Zon.

Meded. Landbouwhogeschool Wageningen81-2 (1981)

P H Y T O C H E M I S T R Y O FH O L A R R H E N A R. BR. N. G. BISSET

The bark and seeds ofH. pubescens have long been used inIndia forthe treatment ofamoebic dysentery. They owe their amoebicidal properties to the presence mainly ofconessine. This compound isthemost important representative of a seriesof steroidal alkaloidswhich belongtotheconanine type and which also occur in other species of the genus, both Asian and African. In the leaves,ontheother hand,thesteroidalalkaloidspresentaremostly5oc-pregnane and/or pregn-5-enederivativeswithaminogroupsatthe 3- and/or 20-positions; someofthealkaloidsareidenticalwiththosefound inFuntumiaspecies.The fact thatinmanyofthesesubstancesthereisanoxygenornitrogenfunction atposition 18hasledtoadetailedstudyoftheirchemistryaimedattheirutilizationasstarting materialsfortheproduction ofpharmaceuticalsteroidssuchasthecorticosteroid aldosteroneandrelatedcompounds.Alsopresentintheleavesofcertain speciesH.pubescens, H. mitis, H. curtisii- are various aminoglycosteroids;these substancescontainanamino-sugarandinsomeofthem,e.g.holarosineand holacurtine, the steroidal part of the molecule isacardenolide, such as ismore usually present in the cardiac glycosides of genera like Acokanthera (Plumerioideae), Nerium and Strophanthus (Apocynoideae), and Thevetia (Cerberoideae), and in others, e.g. the holantosines, the oxygenated side-chain israther different. Progesterone andpregnenolone have been isolated from theleaves ofH. floribunda, and thesecompounds areknown tobeinvolved inthebiosynthesisof the steroidal alkaloids and also of the cardenolides. The occurrence of these various steroidal derivatives in Holarrhena is of considerable chemotaxonomic interest, for genera of the Plumerioideaeare known especially fortheir indole alkaloids while genera of the Apocynoideae frequently contain steroidal (cardiac) glycosides and steroidal alkaloids. Holarrhena,currently placed inthe Plumerioideae but having chemical constituents usuallyassociatedwiththeApocynoideae,appearstobeinasomewhat anomalous position. PICHON has pointed tothe fairly close relationship between certain membersofhisNerieaeand ofhisHolarrheninae,andhehasmadethesuggestion that the Holarrheninae could beconsidered ashaving given rise tothe Apocynoideae (Echitoideae) • Thedistribution ofthe steroidal compounds discussed above may also beconsidered to reflect this relationship. The non-alkaloidal constituents ofthe genus have been investigated invery much lessdetail. REFERENCES V. CERNY & F. SORM 1967,in: R. H. F. MANSKE (ed.),The Alkaloids, Chemistry and Physiology, Academic Press, New York and London, vol. 9: pp. 305-374. CHEMICAL SOCIETY SPECIALIST PERIODICAL PEPORTS 1971 - . The Alkaloids, London, vols. 1-.

H. DADOUN &A. CAVÉ 1978.Plantes méd. Phytothérap. 12:225-229. R. GOUTAREL 1964. Les alcaloïdes stéroïdiques desApocynacées, Hermann, Paris. J. KERHARO & J. G. ADAM 1974.Lapharmacopée sénégalaisetraditionnelle, Vigot Frères,Paris,pp. 162-172. M. PICHON 1950. Mém. Mus. nat. Hist, natur. (n.s.) série B.Botanique I