Series of Revisions of Apocynaceae XLIV

Transcription

1 WAGENINGEN AGRICULTURAL UNIVERSITY PAPERS 97-2 (1997) Series of Revisions of Apocynaceae XLIV Craspidospermum Boj. ex A. DC, Gonioma E. Mey., Mascarenhasia A. DC, Petchia Livera, Plectaneia Thou., and Stephanostegia Baill. by A.J.M. Leeuwenberg Dateof publication: 12August 1997 Wageningen MM Agricultural University

2 Nrn,,\;!( N i;-. L.NV 1 ihiisaaat Series of Revisions of Apocynaceae XLIV / Craspidospermum Boj. ex A. DC, Gonioma E. Mey., Mascarenhasia A. DC, Petchia Livera, Plectaneia Thou., and Stephanostegia Baill. / A.J.M. Leeuwenberg ISBN NUGI 823 ISSN X Distribution: Backhuys Publishers, P.O.Box 321, 2300 AH Leiden, The Netherlands. Telephone: Fax: backhuys@euronet.nl All rights reserved Printed in The Netherlands

3 Series of Revisions of Apocynaceae XLIV Craspidospermum Boj. ex A. DC, Gonioma E. Mey., Mascarenhasia A. DC, Petchia Livera, Plectaneia Thou., and Stephanostegia Baill. bya.j.m. Leeuwenberg Department of Plant Taxonomy, Wageningen Agricultural University, P.O.Box 8010, 6700 Wageningen, the Netherlands Abstract Six genera of Apocynaceae have been monographed. These six are restricted to Africa, except for Petchia, which also occurs in Sri Lanka with one endemic species. The study is based on herbarium material and living plants, mostly observed and collected by the author in Madagascar. Petchia replaces the well-known genus name of Cabucala as it has priority, so six new combinations have been made here. Petchia africana from Cameroun has been described as new to science. For Carissa verticillata a new name has been proposed to replace Pichon's homonym, C.pichoniana. In all, 23 names have been reduced to synonymy, 14 of which are even synonymsof Plectaneia thouarsii. A key to all Apocynaceous genera indigenous to the Comoro IslandsandMadagascarhasbeenprovided. Petchia erythrocarpa is locally used as medicine. Mascarenhasia Umceolataand M. lisianthiflora axe potentially of economic valueasornamentals. Introduction This publication is composed of monographic revisions of six genera, Craspidospermum, Gonioma, Mascarenhasia, Petchia, Plectaneia and Stephanostegia, three of which, the first and the last two are endemic to Madagascar and two are represented in that country and on the African continent as well. It is illustrated with photographs,line drawings and distribution maps. Wageningen Agricultural University Papers 97-2 (1997) 5

4 These revisions are mainly based on study of herbarium material, while the author had the opportunity to observe living plants,andtocollect material of the monotypic Craspidospermum, of three of the eight species of Mascarenhasia, four of the eight Petchia species, of all three Plectaneia species, and of Stephanostegia capuroniiinthewild and Gonioma kamassi in cultivation. Several species, Mascarenhasia arborescens, Petchia erythrocarpa, P. madagascariensis and Plectaneia thouarsii turned out to be more variable than was supposed by Markgraf (1976). The present author had the opportunity to study living plants in the field in many different localities and to observe the variation within individual plants and within populations. All these species are widespread and the ecology of their habitats is greatly varied. He therefore was obliged toreducemanynamestosynonyms. It has been possible to trace almost all type specimens of the namesand synonymsdiscussed inthepresentpaper. Geographical Distribution Craspidospermum verticillatum, the only species assigned to this genus, is a forest tree which produces highly appreciated timber and is called Vandrika all over the country of origin, Madagascar.As its populations in the wild are vigourously damaged by harvesting, it should betaken into cultivation for timber purposes. Gonioma countstwospecies,onebelongstothecapeflora and oneoccursin southwestern Madagascar. Mascarenhasia has one species, that is very common in Madagascar, and also occurs in East Africa from Kenya south to Mozambique. All other 7 species of Mascarenhasia are endemic to Madagascar. M. lisianthiflora is widespread and common in dry areas. M. lanceolata is only known from the northwest and M. speciosa from the southeast, while the four remaining species, which are much more rarely represented in herbarium collections, are all from eastern areas. M. havetii and M. macrosiphon have recently been collected. M. rubra, only known from the type, and M. tampinensis, only known from a few rather old collections of twolocalities,even mayalready beextinct. The genus Petchia isrepresented by asingleendemic speciesin SriLanka,sixendemic speciesinmadagascarand onespeciesknown from only two specimens collected in the same forest in Cameroun. The last species, P. africana, described in this paper as new to 6 Wageningen Agricultural University Papers 97-2 (1997)

5 science, has been discovered among the unnamed material in the Wageningen herbarium, where it was present for several decades. It is perfectly understandable, that it could have been recognized only with the revisions of all African Apocynaceae. The revisions of the last yet unpublished genera of this family for Africa have been started, and therefore its identity as a new discovery for the African continental floracould be established. The most widespread species of Petchia is P. erythrocarpa, which is represented on the Comoro Islands and distributed almost over entire Madagascar. It only does not occur in the southwest. P. madagascariensis is the second most widespread species.it occurs in northern and eastern Madagascar. P. cryptophlebia is centraleastern of distribution. P. montana is rare and has been collected in forest remainders on the central plateau. P. plectaneiifoliais only known from a few collections in the southeast. Finally P. humbertii inhabits only onelocality inthenorth,asfar as isknown. Plectaneia, is endemic tomadagascar, and has one widespread variable species, P. thouarsii, which must have a great ability to adapt to various ecological conditions. It occurs over almost the entire country. The second species, P. stenophylla, inhabits dry places in the southwest. The third and last species, P. longisepala is only known from two collections made in the coastal forest in the east. The two species of Stephanostegia share Madagascar almost evenly. S. capuronii inhabits the wet eastern forests and S. hildebrandtii the dry west. At least the first suffers from being exploitedlike Craspidospermum. Itswoodisalsoesteemed astimber. Relation to other genera Petchia belongs to the tribe Alyxieae of the subfamily of the Plumerioideae, where it is housed at present in the subtribe Rauvolfiinae. It shows remarkable resemblances to the Asian, Oceanic and Australian genus Alyxia by the moniliform fleshy follicles which contain drupes,by the frequently whorled leaves,and to a lesser extent by the flowers with an almost cylindrical corolla tube usually longer than the lobes. It could be segregated from Alyxia, asthe seedslacktherumination intheendosperm. Duetothe latter character it has been moved within the Alyxieae from the Alyxiinae to the Rauvolfiinae. The present author, who had the opportunity to see most of the Madagascan species of Petchia and a Wageningen Agricultural University Papers 97-2 (1997) 7

6 few of Alyxia in the wild, corroborates this approach. He saw the Prévost model in the Petchiashrubs and trees, while the Alyxia climbers, quite probably grow in the model of Troll. The four genera Craspidospermum, Gonioma, Plectaneia and Stephanostegia constitute together with the Asia genera Dyera and Kamettia, and the American genus Strempeliopsis, the subtribe Craspidospermatinae of the tribe Plumerieae of the subfamily Plumerioideae. Their capsular fruits contain flat winged seeds with thin non-ruminate endosperm. The corolla tube is cylindrical or nearly soand the lobes overlap to the left. The stamens are included. The wings of the seeds vary in shape, but are mostly more or less entire, only in Craspidospermum they are more or less fimbriate giving the dark brown seeds some resemblance to miniature fried plaice. Mascarenhasia belongs tothesubtribe Malouetiinae of thetribe Wrightieae of the subfamily Apocynoideae, in the context of the subdivision of the subfamily that has been kept uptothe present day. The dry follicles, seeds with coma and the corolla lobes overlapping to the right are all characters known from almost all Apocynoideae. As the subdivision of the Apocynoideae may be altered considerably, when more monographs of the genera will be completed, not much more can be said on the latter subject now (see also Leeuwenberg, 1994). Nevertheless, Mascarenhasia is closely allied to the Asian genus Kibatalia by almost all chararcters, e.g. model of Koriba, the shape of the corolla, the dry follicles, the seeds with coma and the corolla lobes overlapping to the right. Kibatalia, however, differs clearly from Mascarenhasia by the seeds which are beaked with the coma recurved instead of erect, and with the apex directed towards the base of the follicle instead of towards the apex. The seeds of Kibatalia strikingly resemble those of Strophantus. Key to the genera of Apocynaceae indigenous to the Comoro Islands and to Madagascar 1. Herbs;corolla usually pink and with avery narrow long tubeand a wideflat limb Catharanthus Plants woody 2 2. Leaves alternate,often confined tothe apices of thebranchlets 3 Wageningen Agricultural University Papers 97-2 (1997)

7 Leavesopposite or whorled 4 3. Plants succulent and with spinesand clear sap Pachypodium Plants not succulent, unarmed, with white latex Cerbera 4. Plants with spines,often inlow numbers and short;corolla white; berries small, mostly globose Carissa Plants unarmed 5 5. Climbers 6 Trees or shrubs Tendrils present; stems not twining; corolla lobes overlapping to the left; fruits baccate 7 Tendrils absent; stems usually twining; corolla lobes overlapping to the left or to the right; fruits capsular, of 2linear follicles Inflorescence of elongate branched terminal panicles, longerthan the leaves; fruits pubescent Ancylobotrys Inflorescence contracted and shorter than the leaves; fruits glabrous 8 8. Wall of corolla tube not thickened above anthers; inflorescence terminal; corolla tube 16 34mm long Saba Wall of corolla tube thickened above anthers; inflorescence terminal and/or axillary; corolla tube usually less than 15mm long Landolphia 9. Petiole glandular above; leaves mostly with domatia; corolla lobes overlapping tothe right; 5corona lobes;seeds withcoma Oncinotis Petiolenot glandular above;leaveswithout domatia;corollalobes overlapping to the left or to the right; corona absent Ocrea widened into clear intrapetiolar stipules (90 turned as to compare with the interpetiolar stipules of Rubiaceae); corolla lobes overlapping tothe left; seeds with wings;follicles syncarpous,often angularand winged Plectaneia Ocrea not widened into clear intrapetiolar stipules; corolla lobes overlapping to the right; seeds with coma;follicles apocarpous, terete Alafia Wageningen Agricultural University Papers 97-2 (1997) 9

8 11(5). Leaves whorled, at least at apices of branchlets 12 Allleaves opposite Corolla lobes much shorter than the tube; stamens inserted in upper half of tube; fruits drupaceous, with fleshy mesocarp, apocarpous or almost so,or syncarpous globose berries 13 Corollalobesabout aslong asthe tube;stamens inserted inlower half of tube; fruits capsular, bivalved, syncarpous Craspidospermnm 13. Only upper leaves of each branchlet and of main axis verticillate, the other opposite; fruits apocarpous, mericarps moniliform; disk absent;plants growing in the model of Prévost Petchia All leaves verticillate, or only those on main axis verticillate and those on lateral branches opposite; plants model of Prévost in some Carissa species All leaves verticillate 15 Leaves on main axis verticillate and those on lateral branches opposite Corolla lobes overlapping to the right; fruits small globose berries, leaves up to 4 cm long Carissa pichoniana Corolla lobes overlapping to the left; fruits halfway syncarpous and obcordate, or apocarpous and mericarps ellipsoid; most leaves more than 5cm long Rauvolfia 16.Plants repeatedly dichotomously branched; fruits of paired, often fleshy, non-moniliform follicles 17 Plants not dichotomously branched; fruits paired, dry follicles, or moniliform and then fleshy follicles, or globose berries Corolla lobes overlapping to the right; follicles dry 18 Corolla lobes overlapping to the left; follicles fleshy; seeds without coma; corolla white or yellow Follicleslinear; seedswith coma;pedicels (7 )12 35mm long; corolla orange. Madagascar Strophanthus Follicles ellipsoid; seeds without coma; pedicels 2 3mm long; corolla yellow or creamy.comoro Islands Schizozygia 10 Wageningen Agricultural University Papers 97-2 (1997)

9 19.Corolla tube about as long as the sepals;calyx shed with corolla; leaves rounded at apex; mericarps dark green, pale greenspotted and -dotted Voacanga Corolla tube mostly at least twice as long as sepals; calyx persistent when corolla is shed, often even present under the fruit;leaves mostly acuminate or acute Tabernaemontana 20. Corolla lobes overlapping to the right; tube mostly more than 5 mmlong, seeds with comaornot,notwinged 21 Corollalobes overlapping totheleft, tube upto 4mmlong;seeds winged Corolla tube funnel-shaped or with a cylindrical basal part that abruptly widens into an upper part; follicles dry; seeds with coma Mascarenhasia Corolla tube cylindrical; berries globose; seeds without coma Carissa 22. Corolla tube about 2.5 x as long as the lobes, without slits. Dry forest in South Madagascar Gonioma Corolla tube slightly longer than the lobes, with 5 longitudinal slits. Mostly wetforest in the East and the Northwest of Madagascar; often big trees (fragments found on the forest floor could be confused with those of Plectaneia which has intrapetiolar stipules not present here) Stephanostegia 1. Craspidospermum Boj. ex A. DC, Prod. 8: 323 (1844); Markgraf, Fl. Mad. fam. 169: 97 (1976). Type species: C. verticillatum Boj.exA.DC Craspidospermum verticillatum Boj. ex A. DC, Prod. 8: 323 (1844):Markgraf, Fl. Mad. fam. 169:97, pi. 13,map 21 (1976). Type: Madagascar, Antananarivo, near Antananarivo and Imamou, Bojeranno 1839 (holotype G-DC;isotypes BM,K, P). Fig. 1, p. 12; map 1, p.13 Heterotypic synonyms: C. verticillatumvar. petiolare A. DC, op. cit. 324, syn. nov. Type: Madagascar, sin. loc., collector unknown, possibly Commerson, "Ochrosia" (holotype G-DC; possibleisotype Commerson 791,MA). Wageningen Agricultural University Papers 97-2 (1997) 11

10 Fig. 1. Craspidospermum verticillatum. 1, habit (x 2/3); 2, flower above (x 2); 3, central part of flower (x 3); 4, opened corolla (x 5); 5, stamen (x 10); 6, pistil with longitudinal section of ovary (x 10); 7, tétradesof pollen grains much enlarged; 8, infructescence (x 2/3); 9, seed (x 2); 10, part of seed edgemuchenlarged. 12 Wageningen Agricultural University Papers 97-2 (1997)

11 0-15 S to 100km I} M y^i Vf /^A(DS) O l J/ V l^ja Map1. Craspidospermum verticillatum. ML r i. \% -20" mi. y * / i if- : ' TV-A -25" <{ JJ^fuFD) 0 IOQ 200 km 45 I 50 E I C. verticillatum var. sessile Markgr. in Adansonia II, 12:219 (1972); op. cit. 102, map 21, syn. nov. Type: Madagascar, Toliara, Manampanihy R. valley, near Ampasimena, Humbert (holotype P). Tree 3 25 m high, glabrous except for the corolla inside, with white latex at least in branchlets, not in bark of trees.trunk up to 50cm in diameter; bark pale grey-brown, rather smooth. Branches lenticellate; branchlets triangular or quadrangular at the apex, often even when dried. Leaves in whorls of 3 4, petiolate, rarely sessile; petiole 5 25mm long; blade coriaceous even when fresh, narrowly elliptic or obovate, x as long as wide, 4 14 x cm, rounded or sometimes apiculate or acute at the apex, cuneate at the base, with pairs of rather straight secondary veins forming an angle of with the costa; tertiary venation reticulate, conspicuous beneath. Inflorescence terminal and in the axils of the upper leaves, 1 4 together, with very many flowers, 4 12 x 4 10cm, dense. Peduncle 10 85mm long; Wageningen Agricultural University Papers 97-2 (1997) 13

12 pedicels mm long. Lower bracts leafy, other sepal-like and 1 2x as long as the sepals. Flowers: Sepals green, connate at the base, subequal, clasping the base of the corolla or in dried flowers often somewhat spreading, ovate or broadly ovate, obtuse or rounded, without colleters. Corolla white or pale pink, or limb white, throat pink or dark red and tube often pink, with an ellipsoid head in the mature bud about half of the bud length, pilose inside from the mouth to the insertion of the stamens and with a pubescent belt below the insertion of the stamens 1 2 mm wide with straight hairs directed downwards, not ciliate on lobes; tube cylindrical, x as long as the calyx, x as long as the lobes, 6 10 mm long, mm wide at the base, widened around the anthers to mm wide, narrowed above to 1 2 mm wide, again widened at the throat; lobes narrowly elliptic, x as long as the tube, x as long as wide, x mm, rounded, entire, suberect to spreading, overlapping to the left. Stamens deeply included, inserted mm from the base of the corolla tube; filaments 0.3 lmm long; anthers very narrowly ovate to oblong, x as long as wide, x mm, apiculate to obtuse at the apex, cordate at the base, entirely fertile. Pistil with apex near base of anthers; ovary subglobose or nearly so, 0.5 lx x mm, 2- celled, rather abruptly narrowed into the short style; pistil head of a basal stigmatic ellipsoid part x mm and an oblong bilobed stigmoid apex. Ovules approximately 30 in each cell. Fruit an oblong woody capsule x cm, rounded at the apex and at the base, bivalved, septicidal; wall about 2 mm thick. Seed dark brown, flat obliquely elliptic, 12 x 4 x 2 mm, resembling a fried plaice, ciliate with irregularly coherent hairs like fins 1 1.5mm long, minutely papillose; endosperm mealy; embryo straight, spathulate, 8 mm long; cotyledons ovate 3x2 mm, rounded at the apex and at the base; rootlet 5x1 mm. Distribution: Endemic to Madagascar. Ecology: Rain forest. Alt m. Uses: Wood used as timber. Geographicalselectionoftheapproximately140specimensexamined: Madagascar. Mahajanga: Tsiafankantiha, Maevatanana, SF (K,P, TEF); nearandriba, McPherson (TAN).Antsiranana:AnalamohitsoForest, upper Bemarivo R., Perrier de la Bâthie 8841 (P).Antananarivo:AnkazobeMts, km 161Antananarivo-Mahajanga Road, Capuron SF224 (P,TEF);Manankazo, 14 Wageningen Agricultural University Papers 97-2 (1997)

13 Harizo RN 1150 (BR, K, P, WAG); ibid., Ambohitantely, SF7881 (P,TEF);27 km N of Ankazobe, Miller & Phillipson 3724 (BR, K, P, TAN, WAG); near Antananrivo, Bojer anno 1839 (BM,G-DC, K, P, type);ibid., Bouton anno 1838 (G-DC); Ankazondandy, Boiteau 85 C (K, P); Angavokely, 30 km E of Antananarivo, Capuron SF 118 (P, TEF); Soalazaina Forest,Andilanatoby, Cours 1517 (P, TAN); Ambohimanga, Leeuwenberg &Rapanarivo (BR, P,TAN, WAG);ibid., Capuron SF (K, P,TEF);ca 14kmNWofAmbohitsaratelo- Bebao,NW of Tsiroamandidy, Dorr et al (K, P,TAN, WAG); Bongolava, W of Tsiroamandidy, Moral 4609 (P, TAN), 4620 (P, TAN); near Ambavomanoina, E of Antananarivo, Boiteau 2132 bis (K, P); E slope of Iharanandriana, CapuronSF (BR, K, P, TEF); Sirabe (= Antsirabe), Hildebrandt 3602 (BM,BREM,K, P,W).Toamasina: île Ste.Marie, Capuron SF 28818bis (P);Antsaralalana, ManakambahinyEst, Ramanantsoavina RN 1775 (K, P, WAG); Imerimandroso, Ambatondrazaka District, RakotovaoRN (P); Ambatosoratra, Rakotovao RN (P);ManakaEst, Ramanantsoavina RN2793 (P, TAN, WAG);Andranobe Forest, Wof Manohilaly, Capuron SF (BR, K, P, TEF); Manakambahiny, Ramanantsoavina RN 1935 (P, TAN, WAG); Belamba, Andilanatoby, SF 5664 (BR, K, P, TEF); Fierenana, Moramanga, SF 117-R-187 (P);Sakala-Ambany, Marovoay, SF25018 (K, P, TEF);Antsahatsaka, Beravina, SF 7545 (BR, K, P, TEF); Antaniditra, Perinet, SF 2914 (P, TAN, TEF); Analamazaotra, Perinet, SF (P, TEF); Mandraka Forest, km 75 Antananarivo-Tamatave Road, Cremers 1282 (BR, P, TAN); Amboasary, Beforona, SF 85-R-147 (P). Fianarantsoa: Mangatabotahangy, SF 7-R-207 (P); Vazampotsy Forest, Ambatofinandrahana, SF 37-R-10 (P); Ambalanimanakena, Ambositra, SF 5216 (P, TEF); Ampitambe,Fianarantsoa, SF 7056 (K, P, TEF); 9 km Eof km 26 Fianarantsoa-Ambositra, on road to Ranomafana, Leeuwenberg & Rapanarivo (BR, P, TAN, WAG); Ranomafana Reserve, S of Namorona R., Schatz & Miller 2452 (BR, K, P, TAN, WAG); ibid., Malcomber et al (BR, P, WAG); ibid., Phillipson2215 (BR, K, P, TAN, WAG); Morafeno, Ifanadiana District, SF26378 (= 87-R-73) (P);Ankafina, Hildebrandt 3936 (BM, K, P, US, W); Sihanaka, Boiteauin Herb. Jard. Bot. Tana (P); W of Ambalavao, Croat30302 (P, TAN, WAG); Ambatomainty, Mahatsinjony, SF (K, P, TEF); Mahazony, Ambalavao, Rakotovao RN 6830 (P); near Zazafotsy, Croat (TAN); Andringitra Reserve, HenriRN 4875 (P, TAN); ibid., SF (P, TEF); Farafangana, SF (P, TEF); Manombo Forest, Capuron SF9208 (BR, K, P,TEF), SF (K, P,TEF);Angavokely, Carion, SF6639 (P,TEF). Toliara: Manampanihy R. valley, near Ampasimena, Humbert (P,type of C. verticillatum var. sessile); VohimavoMt, NofAmpasimena, Humbert20679 (P); Manantenina, Fort-Dauphin, Debray 2016 (P, TAN); Fort- Dauphin, Scott Elliot 2378 (K);Mahatalaky, SF (K, P,TEF);Mt Enivabe, Ranopisa, SF (P, TEF); Ebakika, Fort-Dauphin, Decary (P); Andohahela Reserve, Rakotoniania RN 5947 (BR, K, P, WAG); ibid., Leandri & Saboureau 4304 (BR, K, P, WAG);Behara, TsilizyRN 8356 (P,TAN);ibid., SF (P, TEF); Ivakoany Mts, Humbert 7024 (P). Sin. loc.: Anonym, s.n. (G- DC, type of C. verticillatum var. petiolare); Baron89 (K), 111 (FT), 768 (BM, K), 853 (FT,K, P); Commerson s.n. (MA, P). Cultivated. Madagascar, Antananarivo, Tsimbazaza Park, Dorr & Barnett 3170 (K, P,TAN,WAG);of same tree, Leeuwenberg (WAG). Note. The leaves have petioles that show a continuous range of variation in length. Exceptionally they are sessile as in the type of Wageningen Agricultural University Papers 97-2 (1997) 15

14 var. sessile. As there are no characters to corroborate the distinction of the varieties,these are notmaintained here. 2. Gonioma E. Mey., Comment. PI. Afr. Austr. 188 (1837); Codd, FI. S. Afr. 26: 262 (1963); Markgraf, Fl. Mad. fam. 169: 112 (1976). Type species: G. kamassie.mey. Shrubs or trees, glabrous except for corolla inside. Branches pale brown, lenticellate; branchlets terete. Leavesin whorls of 3 4 or opposite, petiolate; blade subcoriaceous when fresh, papery when dried, narrowly elliptic, acuminate to obtuse at the apex, cuneate at the base, margin revolute in dried leaves, venation obscure. Inflorescence terminal, many-flowered, dense. Lower bracts leafy, other very narrowly elliptic and long-acuminate to sepal-like. Flowers: Sepals connate at the base, subequal, ovate, obtuse or rounded, without colleters. Corolla white, yellow or orange, pilose inside from mouth to 1 2 mm below insertion of stamens, with a broadly ovoid rounded head in the mature bud about one third of the bud length; tube nearly cylindrical, widened around the anthers, mostly narrowed at the mouth; lobes overlapping to the left in bud, suborbicular or ovate, rounded, entire, spreading. Stamens withapex 0 1mm below mouth of corolla tube; filaments very short; anthers ovate or narrowly so, acuminate to obtuse at the apex, cordate at the base, entirely fertile. Pistil with apex near base of anthers; disk absent; ovary subglobose or ovoid, mostly rather abruptly narrowed into the short style, of 2 separate carpels; pistil head of a basal ellipsoid stigmatic part and a bilobed stigmoid apex. Fruit of 2 separate follicles; follicles light brown, narrowly oblong, capsular, adaxially dehiscent,with athickwall. Seedflat, winged. Two species, one at the Cape and one in southwestern Madagascar. Keytothespeciesof Gonioma 1. Leaves obtuse or acute, rarely acuminate; sepals rounded or obtuse; corolla tube mm long; lobes 2 3 x mm; follicles narrowly ellipsoid, 2 6 x as long as wide. SouthAfrica 1. G. kamassi Leaves acuminate; sepalsobtuse;corolla tube4.5mmlong;lobes 16 Wageningen Agricultural University Papers 97-2 (1997)

15 ovate, 2x1 mm; follicles linear, 8 15 x as long as wide. Madagascar 2. G. malagasy Gonioma kamassi E. Mey., Comment. PI. Afr. Austr. 189 (1837); Codd, FI. S. Afr. 26: 262 (1963). - Types: South Africa, Eastern Cape, near Vanstaadesrivier, Drège a (lectotype P, designated here;isolectotypes G, G-DC, HBG,K, L, P, S, TCD, W). Paratype: South Africa, Eastern Cape, near Meulrivier, Drège b (E, G, HBG,K, L,P, S, W). Map 2, p.18 Heterotypic synonyms: Tabernaemontana camassi Eckl in S.A. Quart. Journ 371 (1830), not cited by E. Meyer and therefore not accepted asbasionym by Codd. Type: South Africa, Eastern Cape, Uitenhage, Krakakamma For., Ecklon3.10 (lectotype P, designated here; isolectotypes BP, C, E, F, GH, HBG, L, NY, TCD,UPS,W, Z). G. kamassivar. brachycarpa E. Mey., I.e. Type: South Africa, Eastern Cape,Uitenhage Division,near Galgebosch, Drèges.n. (lectotype P,designated here;isolectotypes E, G, G-DC,HBG, K, L, P,TCD,W). Shrub or tree 3 10 m high, with white latex or clear sap (teste Codd). Trunk up to 20cm in diameter; bark pale brown, rough, deeply and longitudinally fissured, corky. Leaves: petiole 1 7 mm long; blade 3 6x as long as wide, x cm, obtuse, rounded, acute or rarely acuminate at the apex. Inflorescence x 2 4cm. Peduncle 5 10mm long; pedicels 2 6mm long. Flowers fragrant. Sepals light green, x as long as wide, x mm, rounded or obtuse. Corollawhite and often with a greenish tube, or entirely yellow or orange, pilose inside to mm below insertion of stamens; tube x as long as the calyx, 2 3.2x as long as the lobes, mm long, mm wide at the base, 2.2 3mm wide around the anthers; lobes obliquely and transversely elliptic to suborbicular, x as long as the tube, 0.7 lx as long as wide, 2 3 x mm, auriculate at the right side of the base. Stamens with apex 0.5 1mm below mouth of corolla tube, inserted of the length of the corolla tube; filaments 0.3 lmm long; anthers x as long as wide, Wageningen Agricultural University Papers 97-2 (1997) 17

16 Map 2. Gonioma kamassï, Mascarenhasia arborescens x mm. Fruit: follicles narrowly ellipsoid tooblong, 2 6 x as long as wide, 2 6 x 0.8 1cm, rounded or apiculate at the apex, slightly striate. Seed pale brown, grain darker, obliquely oblong, flat, 8 18 x 5 6x 2mm, with a short wing at the truncate end and a much longer obtuse wing at the other; grain in seeds of 15 18mm long 8 10 x 4 5mm, with irregular raised lines; wings with a minute honeycomb-structure; endosperm cartilaginous; embryo straight, spathulate; cotyledons broadly elliptic, about aslongastherootlet. Distribution: South Africa, the two Cape Provinces, Coast from Knysna to Pondoland. Ecology: Forest understorey, mainly near the coast. Alt m. 18 Wageningen Agricultural University Papers 97-2 (1997)

17 Geographical selectionoftheapproximately90specimensexamined: South Africa. Western Cape: Stellenbosch, Jonkershoek, LaughtonSept (L); 17 km NW of Assegaaibosch, Acocks (K, PRE); near George, Burchell6044 (K, L, P), 6071 (GH, K); ibid., Groenkop, Van der Merwe 4 (WAG); Karatara, Schlechter 5884 (BM, BR, G, HBG, K, L, P, PRE, S, STE, UPS, W, Z); Knysna, Laughton anno 1938 (L, U); ibid., Van Steenis (L, WAG); ibid., Whitish 163 (STE);Buffelsnek, nearknysna, Hafstrom & Lindeberg 9 Dec (S);Knysna Forest, Marloth 695 (PRE, STE); Deepwalls F. R., Bos 765 (K, STE,WAG); ibid., Laughton 8407 (FHO),8408 (FHO);PisangR.valley, Kapp 115 (K, P, PRE); Plettenburg Bay, Rogers (G, PRE, Z). Eastern Cape: near Meulrivier, Drège b (E, G, HBG, K, L, P, S, W, paratype); Willowmore, Bloukrans Pass, Goldblatt 5203 (US, WAG); Tsitsikama, Penther 2004 (S, W); Storms R. F.R., Dahlstrand 478 (C, GB, PRE), 1661 (PRE); ibid., Keet 585 (PRE,STE);GrootR.valley, Gillett 1484 (STE); Clarkson, Thode A 960 (GH,K,PRE);between Blauwrivier KrantzandKowiePoort, Burchell 3659 (GH, K, W);near Galgebosch, Uitenhage Division, Drège s.n. (E, G, G-DC, HBG, K, L, P,TCD, W,type of G. kamassi var. brachycarpa); betweenvanstadens R.and Galgebosch, Burchell4676 (K,W);nearVanStadensR., Drège a (G, HBG, K, L, P, S, TCD, W, lectotype); Van Stadensberg, MacOwan 1055 (A, K, PRE); Van Stadens Nat. Park, Wells 3372 (BR, K, PRE, S); Van Stadens Pass, Acocks (K, PRE);KrakakammaForests, Ecklon & Zeyher3.10 (BP, C, E, F, GH, HBG, L, NY, P, TCD, UPS, W, Z); ibid., Zeyher 730 (E, K, S, STE, TCD), 3413 (BP, G, P, S, STE, UPS, W, Z); Suurberg, Bayliss BS 6104 (BR, HBG, NY, UC, WAG, Z); Port Elisabeth, Van Wijk 3186 (PRU); Dossie Krantz, Grahamstown, MaudeApril 1928 (K); Grobbelaar's Kloof, near Grahamstown, MacOwan 534 (F,G, S, TCD);ca 16kmfrom Grahamstown, Story 3238 (K, P); Blauwkrantz Gorge,Albany District, Cheadle 733 (BM, FHO,K, W);Howiesons Poort, Brink 326 (P); Bathurst Division, Burchell 4138 (K); near East London, Leighton 236 (BM, G, GH, HBG, K, P, UPS, US, W, Z); Komga, Kloof near Kei R. rightbank, Flanagan 367 (FHO,G, P);Haven Forest, Mbanyana R., Gray 1756 (E). Kwazulu-Natal: Faircress Estate, Oribi Gorge, Nicholson 1931 (PRE); Butcher's Place, Durban District, Strey 7317 (BR, PRE); Louwsburg, Hilliard & Burtt 8549 (E,K, PRE, S). Swaziland, near Mbabane, Kemp 1312 (PRE); ibid., Gobolo, Dhlamini 18 April 1963 (K,PRE), 6May 1965 (PRE). Cultivated. England, Kew Palm Stove May 1871 (K). South Africa, Bot. Garden, Cape Town, Dümmer 1772 (E); Compagnie's Tuin, Cape Town, Leeuwenberg (WAG) Gonioma malagasy Markgr. & Boiteau in Adansonia II, 12: 225, pi. 1 (1972); Markgraf, Fl. Mad. fam. 169: 113, pi. 16, map 24 (1976). Type: Madagascar, Toliara, km Tulear-Sakaraha Road, W of Andranovory, Capuron SF (holotype P; isotypes BR, G, HBG, K, TEF, WAG). Fig. 2, p. 20; map 11, p. 80 Large tree up to 18 m high. Trunk up to 40 cm in diameter; bark brown, rough. Leaves: petiole 2 12 mm long; blade (3 )5 6x as long as wide, x cm, acuminate at Wageningen Agricultural University Papers 97-2 (1997) 19

18 Fig. 2. Gonioma malagasy. 1, habit (x 2/3); 2, intrapetiolar stipules (x 5); 3, opened corolla (x 7); 4, calyxwithpistil (x 10); 5, ovary (x 10); 6, fruit (x 2/3); 7, seed (x 1.5). 1-5 from Capuron SF Wageningen Agricultural University Papers 97-2 (1997)

19 the apex. Inflorescence 1 2x 1 2 cm. Peduncle 1 4 mm long; pedicels mm long. Flowers: Sepals 1.3 x 1mm, obtuse. Corolla white, pilose inside to 1 mm below insertion of stamens; tube 3.5 x as long as the calyx, 2.2 x as long as the lobes, 4.5 mm long, 1 mm wide at the base, 1.5 mm wide around the anthers, 1 mm wide at the mouth; lobes ovate, 0.4 x as long as the tube, 2 x 1 mm. Stamens with apex 1 mm below mouth of corolla tube, inserted at 2.5 mm from the base of the corolla tube; filaments 0.3 mm long; anthers ovate, 1 x 0.4 mm, acuminate at the apex. Pistil 3 mm long; ovary ovoid, 1.5 x 1 x 1 mm; style 0.5 mm; pistil head: stigmatic part 0.9 x 0.3 mm; stigmoid apex 0.1 x 0.1 mm. Fruit, follicles linear, 4 11 x mm, acuminate or caudate, striate, not lenticellate, often transformed into irregularly subglobose galls. Distribution: Endemic to SW Madagascar. Ecology: Dry forest. Alt. low. Specimensexamined: Madagascar.Toliara:Analafanja, Antseva,SF15072 (P);Lambomakandro, SF (P); Sakaraha, SF (BR, G, K, P), (P, TEF); Ifazoroa, Sakaraha, SF 3856 (P); Zombitsy Forest, Humbert et al (K, P); ibid., Capuron SF (BR, G, K, P, WAG); ibid., SF (P); 15 km Sof Sakaraha, Imbert 50 (P); km 55-65Tulear-Sakaraha Road, Wof Andranovory, Capuron SF20716 (BR,G,HBG,K,P,TEF,WAG,type);km63ofsame road, SF3668 (P);Ihera Forest,Tulear, SF9794 (P);SakamenaR., Beora,SF14364 (P,WAG).Sin.loc., Richard TAI (K). 3. Mascarenhasia A. DC, Prod. 8: 487 (1844); Pichon in Mém. Inst. Sei. Madag. Sér. B, 2: 76 (1949); Markgraf, Fl. Madag. Farn. 169: 248 (1976). Type species: M. arborescens A. DC. (lectotype species designated by Pichon). Heterotypic synonyms: Tsilaitra Baron in Rev. Madag, 16: 250 (1905). Type species: T. micrantha (Bak.) Baron, lectotype species designated here (= M. arborescens A. DC). Lanugia N.E. Br. in Torreya 27: 51 (1927). Type species: L. latifolia N.E. Br. (= M. arborescens A. DC). Echitella Pichon in op. cit. 88. Type species: E. lisianthiflora (A. DC) Pichon (= M. lisianthiflora A. DC). Shrubs or trees, probably all representing growth model of Wageningen Agricultural University Papers 97-2 (1997) 21

20 Koriba or of Prévost, mostly with white latex in all parts (not in M. speciosa; not known of M. macrosiphon and M. tampinensis). Branchlets terete. Leaves opposite, persistent or deciduous, petiolate; (ocreae not or slightly widened into intrapetiolar stipules); blade coriaceous or subcoriaceous. Inflorescence axillary and/or terminal, 1 15-flowered, fasciculate when with more than one flower. Peduncle obsolete or short; pedicels often rather long. Bracts sepallike or nearly so, much smaller than the sepals, persistent. Flowers rather large (corolla tube 5 60mm long), fragrant or not, open during the day. Sepalsoften leafy, green (always?), free, erect or nearly so, subequal or unequal. Corolla white, cream, red or pink or in a combination of these colours; tube funnel-shaped (M. speciosä) or with a narrow almost cylindrical basal part which abruptly widenes just below the insertion of the stamens into acampanulate or narrowly urceolate apical part; lobes overlapping to the right in bud, almost free to half-way connate and with borders folded inwards in bud, ovate, elliptic or suborbicular, spreading or sometimes recurved. Stamens included, inserted about 2mm above abrupt widening of corolla tube or in M. speciosa in widening part of funnel and then slightly or barely included (often exserted in dried flowers); anthers sessile, narrowly triangular, acuminate and sterile at the apex for mm, fertile for about one half of the remaining part and again sterile below, sagittate at the base and with tails about 1 mm below insertion, glabrous except for the mostly pubescent connective which iscoherent with a circularpatch withthe pistil head above its stigmatic part in mature buds and open flowers, free in young buds. Pistil with apex about halfway along anthers; ovary laterally compressed, of 2 separate carpels, surrounded by a glabrous disk, higher or lower than the ovary, 5-parted with crenate, sinuate, lobed or rarely entire lobes; style various; pistil head variously shaped, always with a bilobed stigmoid apex and probably stigmatic only at theextreme base.ovules many ineach carpel. Fruit dark brown, almost black, of 2 linear cylindrical follicles, mostly longitudinally striate when dried, abaxially and longitudinally dehiscent, much paler and brownish inside. Seed grain dark brown, narrowly elliptic, with a raised line at the hilar side, with the hilum in the middle, convex at the other side,dull, minutely papillose,with apex directed to the apex of the fruit; endosperm thin, surrounding the embryo, amylose; embryo straight, spathulate; cotyledons narrowly ovate,about 4 x as long as the rootlet, rounded at the apex, truncate at the base; coma usually 2 3 x as long as the grain, 22 Wageningen Agricultural University Papers 97-2 (1997)

21 medium brown, of a bunch of simple hairs, directed towards the apexof the follicles. Eight species of which seven endemic to Madagascar; one also ineastand Southeast Africa. Keytothespeciesof Mascarenhasia 1. Corolla tube 5 14mm long, widened at or near the middle 1. M. arborescens Corolla tube at least (14 )16mm long and only if as short as (14 )16mm then abruptly widened at of the length (M. lanceolata) 2 2. Corolla tube funnel-shaped, 45 60mm long, gradually widened from the cylindrical basal part; stamens inserted of the length of the corolla tube which is at 36 49mm from the base 7. M. speciosa Corolla tube abruptly widened into a campanulate or narrowly urceolate part from the cylindrical basal part; stamens often deeply included, inserted just above abrupt widening; leaves ratherlargewhen caudate 3 3. Basal part of corolla tube 2 4mm long; entire tube (14 ) mm long 3. M. lanceolata Basal part of corolla tube 7 32mm long; entire tube mmlong 4 4. Basal part of corolla tube x as long as the entire tube and about as long as the sepals 8. M. tampinensis Basal part of corolla tube x as long as entire tube and often longerthan the sepals 5 5. Corolla tube x as long as the calyx 6 Corolla tube x as long as the calyx 7 6. Leaves caudate to acuminate; ovary and style glabrous 2. M. havetii Leaves rounded to obtusely short-acuminate; ovary pubescent and stylepilose 5. M. macrosiphon Wageningen Agricultural University Papers 97-2 (1997) 23

22 7. Leaves at least partly hairy, mostly rather thin; corolla white, often partly red or pink; corolla tube x as long as the sepals. Dry vegetation 4. M. lisianthiflora Leaves entirely glabrous, thick; corolla red; corolla tube at least x as long as the sepals. Wet forest 6. M. rubra Mascarenhasia arborescens A. DC, Prod. 8: 488 (1844); Dubard in Bull. Soc. Fr. 53: 295 (1906) with M. sessilifolia Dubard in syn.; Pichon in Mém. Inst. Madag. sér. B, 2: 76 (1949); Markgraf, Fl. Madag. Farn. 169: 251, pi. 42 (1976); Kupicha in Fl. Zambes. 7, 2: 487, t. 116 (1985). Type: Madagascar, Mahajanga, Bombetoka Bay, Bojer s.n. (holotype G-DC). Figs. 3, p. 25 and 4, p. 26; phot. 1, p. 121; map 3, p. 27 Heterotypic synonyms: M. angustifolia A. D.C., I.e.; Markgraf, op. cit. 257, pi. 43, 1; M. arborescens subsp. angustifolia (A. DC.) Boiteau, Caoutch. Madag. 13 (1943), syn. nov. Type: Madagascar, Antsiranana, Nosy Be, Richard 121 & 181 (composite sheet, holotype G-DC; isosyntypes P). Holarrhena madagascariensis Bak. in Journ. Linn. Soc. Bot. 21: 424 (1885). Type: Central Madagascar, sin. loc, Baron 3242 (holotype K; isotype P). M. micrantha Bak. in op. cit. 25: 335 (1890). Tsilaitra micrantha (Bak.) Baron in Rev. Madag. 16: 250 (1905). Lanugia micrantha (Bak.) N.E. Br. in Torreya 27: 53 (1927); Mascarenhasia arborescens subsp. arborescens var. micrantha (Bak.) Boiteau, I.e. Type: NW Madagascar, sin. loc., Baron 5141 (holotype K; isotype P). M. variegata Britt. & Rendle in Transact. Linn. Soc. II. 4: 26 (1894). Lanugia variegata (Britt. & Rendle) N.E. Br. in Torreya 27: 53 (1927). - Type: Malawi, Southern, Mt Mulanje, Whyte 108 (holotype BM; isotypes G, K, Z; phot, of K sheet B, BR, GB, S). M.fischeri K. Schum. in Engler, Pflanzenw. Ostafrikas C: 318 (1895). Type: Tanzania, sin. loc, Fischer 322 (holotype Bf). M. elastica K. Schum. in Notizbl. Bot. Gart. Berlin 2: 270 (1899), as caustica on p Type: Tanzania, T6, between Dar es Salaam and Mbaffu, near Vikindo, Stuhlmann s.n. (holotype Bf). M. anceps Boiv. ex Jum. in Rev. Cult. Colon. 5: 300 (1899); M. arborescens var. anceps (Boiv. ex Jum.) Lassia, Masca- 24 Wageningen Agricultural University Papers 97-2 (1997)

23 Fig. 3. Mascarenhasia arborescens. 1, 7 & 9 habit (x2/3); 2 & 6, leaves (x2/3); 3 & 8 fruits (x 2/3); 4, seed (x 2/3); 5, transverse section of seed (x 6). 1 from Bosser 8909; 2 from SF 12-R-240; 3-5 from Boivin s.n.; 6 from Perrier de la Bâthie 16555;7-8 from Lavanchie s.n.; 9 from Capuron SF Wageningen Agricultural University Papers 97-2 (1997) 25

24 Fig Mascarenhasia arborescens. 1-2, habit (x 2/3); 3, sepal inside (x 6); 4, corolla inside with stamens and pistil (x 3) M. macrosiphon. 5, habit (x 2/3); 6, corolla inside with stamens and pistil (x 3). 1 from Keraudren 1656; 2-4from Keraudren-Aymonin &Aymonin 25663; 5 from Capuron SF 28431; 6from Capuron SF Wageningen Agricultural University Papers 97-2 (1997)

25 a to < ^la(ds) a to è^j A A ( D S I km % 0 100km % -15 S Ç^ M /I %* -15 S M /"Î -20 Ml Nil y-p T r" -25 <{ ^ 2 r~{~' V \ A \*È N vt*^' Di n#t(fo) 0 W-r r*' /»y M km -20 MlT \ )<? TV-A -25 «{^ 'M Wr Vi'' ^ \ oa(t) ^~!? OA 3L i~>4 X Or/ / "~Nk ;»'M (Tv/ \ÜJT(FD) /T 200 km E 1 Map 3. Mascarenhasia arborescens. Map 4. Mascarenhasia lanceolata; Af. rubra, AAf. speciosa; ir M. tampinensis. renhasia et Landolphia 41 (1927). Type: Madagascar, Antsiranana, Diego-Suarez, Boivin 2461, comm. Richard (lectotype P,designated here). M. longifolia Jum. in Rev. Gen. Bot. 13: 360 (1901); M. arborescensf. longifolia (Jum.) Jum. & Perr. in Agric. prat, pays chauds 7: 283 (1907); M. arborescens var. longifolia (Jum.) Lassia, I.e.; M. arborescens subsp. arborescens f. longifolia (Jum.) Boiteau, I.e. Type: Madagascar, Mahajanga, Lake Anjiafitsatra, near Mt Tsitondroina, Perrier de la Bâthie 966 (holotypep). M. arborea Boiv. ex Dubard in op. cit. 300; M. arborescens subsp. arborescens f. arborea (Boiv. ex Dubard) Boiteau, I.e. Type: Madagascar, Toamasina(?), East coast, Chapelier93 (lectotype P,designatedhere). M. coriacea Dubard in op. cit. 298; M. arborescens var. coriacea Wageningen Agricultural University Papers 97-2 (1997) I 50 E 1

26 (Dubard) Lassia, I.e. Type: Madagascar, Toliara, Fort- Dauphin, Scott Elliot 2703 (lectotype P, designated here; isolectotypes BM, E, K). M. boivinii Dubard in op. cit. 299; M. arborescens var. boivinii (Dubard) Markgr. in Adansonia II, 12: 588 (1972) and op. cit. 255, pi Type: Madagascar, sin. loc., Boivin s.n. (holotype P). M. barabanja Dubard in op. cit Type: Madagascar, sin. loc, Leboeuf anno 1899 (holotype P). M. grandidieri Dubard, I.e. Type: Madagascar, Toamasina, Antsahalaombe Mts, Grandidier s.n., rec. 16 Mar (holotype P). Lanugia latifolia N.E. Br. in Torreya 27: 52 (1927). Type: Cult., Puerto Rico, Mayaguez Agric. Exp. Stat., seeds sent from Mozambique, Britton 8646 (lectotype NY, designated here, isolectotypes GH, K, P). M. arborescens subsp. arborescens f. albicorticis Boiteau, I.e., French description only. Type not traced, not found in P, nor in TAN. M. angustifolia var. keraudreniana Markgr. in Adansonia II, 12: 586 (1972); Fl. Madag. Fam. 169: 260, pi. 43, 2-4 (1976), syn. nov. Type: Madagascar, Antsiranana, near Diego Suarez, Keraudren-Aymonin (not 25663) (holotype P). M. arborescens var. comorensis Markgr. in op. cit. 588 and op. cit. 257, pi. 42, 7-8, syn. nov. Type: Comoro Islands, Anjouan, alt. 600 m, Lavanchie March (lectotype P, designated here); ibid., 700 m, Lavanchie March (P, paratype); ibid., 700 m, Lavanchie April (P, paratype). M. arborescens var. gracilis Markgr. in op. cit. 589 and op. cit. 257, pi , syn. nov. Type: Madagascar, Antsiranana, Mt d'ambre, Makis R., between Roussettes Station and Grande Cascade, Capuron SF (holotype P; isotypes K, TEF, WAG). Tree or shrub, ( 35) m high, with white latex in all parts. Model of Koriba or Prévost. Trunk 3 40( 60) cm in diameter; bark pale or dark brown or pale or medium grey, rather rough or smooth, shallowly and longitudinally fissured or not, often peeling off in large scales, often lenticellate. Branches brown or grey, sparsely lenticellate or not; branchlets glabrous. Leaves: petiole glabrous, 1 8 mm long; blade coriaceous even when fresh, elliptic 28 Wageningen Agricultural University Papers 97-2 (1997)

27 or obovate to very narrowly so, very variable in shape and size (1.5-)2-4.5(-15)x as long as wide, (l-)4-18x (0.5-) 2 7cm, acuminate to retuse at the apex, cuneate at the base or decurrent into the petiole, margin often revolute, glabrous on both sides, with 5 15 pairs of rather straight to upcurved secondary veins forming an angle of 45 80_ with the costa, often greatly varying within one leaf; tertiary venation reticulate, conspicuous beneath or not. Inflorescencefasciculate, 2 4x 2 6cm, 1 15 ( 25)-flowered. Peduncle 2 5 mm long, glabrous; pedicels 7 12 mm long, glabrous to puberulous. Bracts sepal-like, x as long as the sepals. Flowerssometimes fragrant. Sepals pale to dark green, erect, ovate or triangular, 1 2 x as long as wide, x 1.5 4mm, acute or obtuse, ciliate or ciliolate, sparsely pubescent to glabrous outside, glabrous inside or only at the apex sparsely pubescent, with one continuous row of colleters at the base which is 7 10 colleters with each sepal; colleters equal to very unequal, x mm, entire or lobed. Corollacream or white or partly so and then tube greenish or partly red, lobes often red- or purple-striped outside and/or inside and throat often greenish or pale yellow, puberulous or shortly pubescent outside on the lobes and mostly on the upper quarter or half of the tube or less often on the entire tube, pubescent inside on the lobes and in the throat down to the apices of the anthers or even up to as far as the insertion of the stamens, the belt between the apices of the anthers andthethe insertion ofthe stamens even maybecovered by 5stripes of pubescence behind the anthers, then below the insertion of the stamens with 5 stripes of pubescence 1 3 mm long; tube x as long as the calyx, x as long as the lobes, 5 14mm long, widened abovethebaseto mmwide,contracted atornearthe middle to 1 4mm wide which is just below the insertion of the stamens, again widened around the stamens to 2 5 mm wide and contracted at the throat to mm wide; limb spreading to recurved, with lobes free to halfway connate; lobes ovate or broadly ovate, x as long as the tube, 1 1.8x as long as wide, 4 15 x 2 14 mm, apiculate, acuminate or acute. Stamenswith apex mm below mouth of corolla tube, inserted of the length of the corolla tube (at5 8 mm from the base);anthers 3 4x as long as wide, x mm, apex sterile for mm, fertile below for about half of the remaining part; connective sometimes pubescent. Pistil: ovary cylindrical, 1 2x x mm, retuse or occasionally acute at the apex, Wageningen Agricultural University Papers 97-2 (1997) 29

28 pubescent at the apex or occasionally entirely glabrous; disk (1 )2 3 mm high which is usually higher than the ovary; style rather robust, pubescent; pistil head almost conical, ca 2 x 1 mm and with widened base. Ovules approximately in each carpel. Fruit: follicles x x cm. Seed grain x 2 4x 0.8 lmm; embryo mm long; cotyledons x as long as wide, 7 9x mm; rootlet 2 3 x mm; coma mm long. Distribution: East and Southeast Africa, Seychelles, Comoro Islands and Madagascar. Ecology: On river banks in forest or gallery forest. Alt m. Leaves in forest spreading, in open places erect. Geographical selectionoftheapproximately520specimensexamined: Kenya. K7: Kilifi District, Pangani Crossing of Lwandani Stream, on Chonyi-Ribe Road, Fadenet al. 77/532 (BR, K, US, WAG); Kwale District, Marere Forest, Perdue & Kibuwa (BR, FT, K, WAG); Shimba Hills, Magogo1206 (BR, FT, K);ibid., comm. Batiscombe May 1907 (K,P, Z);Wanga District, Dalvel 3 July 1917 (K). Tanzania. T3: E Usambara Mts, Amani, Verdcourt1731 (K); Mwera. Mwanza R., Peter (B,WAG);Pangani District, Bond 78 (BR, K, M).T6 MorogoroDistrict,Turiani Falls, Drummond & Hemsley 1473 (B,BR, FT, K, S) ibid., Milne-Redhead & Taylor 7099 (B,BR, K);Pugu Hill, Hansen 521 (C).T7 UpperRuhudje R.,LupembeDistrict, Schlieben 1468 (B, P).Pemba: Vaughan 114 (K); near Wesha, Vaughan 2051 (BM); Sengenya Dya, R.O. Williams 99 (BR, K). Zanzibar: Lastanno 1909 (BR, F, K, P). Mafia Island: Tondwa, Greenway 5337 (K). Malawi.Southern:Mulanje Mt, Why te 108 (BM,G, K, Z; phot,of Ksheet B, BR, GB, S, type of M. variegata), 155 (TCD); ibid., Likhubala R., Brass (BR, K, NY, PRE, SRGH, UC, US);ibid., Chapman 350 (BR, K,MAL), 8931 (E, K); path from Lichenya Hut to Likhubula Mission, Patel 857 (BR, K, MAL);LikhubulaForest, Salubeni 1643 (K, MAL,SRGH). Mozambique. Cabo Delgado: between Mocimba da Praia and Quiterajo, Barbosa 2115 (LISC). Nampula: Malema, Gomes e Sousa 4270 (COI, K, LISC, PRE, SRGH); Ribaué Mts, Gomes e Sousa 752 (COI, K, LISC, Z); ibid., Torre 654 (COI, LISC), 1110 (COI, LISC); Chinga Mts, Macedo 3090 (WAG); Moçambique (= Porto Mocambo), Campos Andrada 770 (LISC). Zambézia: Sagura, Sacleux 2205 (P); between Gurué and Mocuba, Mendonça 1344 (LISC); Altomoloinè, road to Gilé, Napome R., Campos Andrada 1911 (COI, LISC); Namagua Plantations, Mocuba District, Faulkner 111 (FT, K, S, SRGH), in GH (BR, K, S, SRGH); Lugela, Faulkner 198 (BR, K, NY, PRE, SRGH); ibid.,between Mobede andtacua, Campos Andrada 1494 (COI);between Pebane andmocubela, Torre 4123 (LISC);Maganja dacosta, Sim6021 (PRE);km36Vila de Maganja-Mocuba Road, Torre & Correia (LISC); Pebane, Barbosa & Carvalho 4296 (G, K); near Quelimane, Luja 479 (BR). Sofala: Gorongosa Nat. Park, Tinley 2566 (K, SRGH); ibid., near Cunduè R., Macedo 2385 (WAG); 30 Wageningen Agricultural University Papers 97-2 (1997)

29 between Inhaminga and Savane, Torre 3070 (LISC); Rio Mupa, Simâo783 (K); Beira Region, Chiniziua R., Gomes e Sousa 4358 (B, BR, COI, K, LISC, PRE), 4404 (COI, K, LISC, PRE), 4427 (COI, K, PRE, W); Dondo (= Macuacua), W.H. Johnson 301 (K, P); near Manga, Beira, Correia & Marques 2789 (BR, WAG); Pungwe Swamps, Pole Evans 5875 (PRE); Beira, Swynnerton 558 (BM, K, SRGH). Manica: Chimani Mts, Müller 1121 (K, SRGH); ibid., Musapa Gap, Wild3529 (K, LISC, NY, PRE, SRGH); Dambo, Machango, Pereira & Marques 851 (BR,WAG). Zimbabwe. Manicaland: Pungwe R., Chase6130 (BM, K, LISC, PRE, SRGH), 6447 (K, LISC, PRE, SRGH), 8040 (BM, BR, K, LISC, SRGH); ibid., Wild 5261 (COI, K, M, PRE, SRGH);Stapleford F.R., Mullin184/63 (K, LISC, NY, SRGH); Musapa Gap, Rail 1/55 (K, PRE, S, SRGH); Chimanimani Mts, Martin F.R., Mavi 656 (K, LISC, SRGH); Mt Pesa, Chase2993 (BM, COI, LISC, SRGH);ibid., Wild 3577 (K, LISC,NY,PRE, SRGH);Lusitu R., near its junction withharoni R., Drummond 5002 (K, PRE, SRGH). Seychelles.Mahe:MorneSeychellois Nat Park, Procter4505 (K). ComoroIslands.Moheli:between Fomboni and Drondoni, Bernardi (G,K, L, P, Z);Voundrouvou Forest, Floret 1232 (P,WAG).Anjouan: Lavanchie March (P, lectotype of M. arborescens var. comorensis); Moya Adda-Doueni Road, Rakotozafy 1134bis (TAN). Madagascar. Antananarivo: Carion, Decary 62 (P). Mahajanga: Mamokornita, Andriba, Kasombo SF (P, TEF, WAG); Mahazoma, Descoings 3379 (TAN); ibid., near Tsaratananakely, Descoings 3364 (P, TAN); Ankara Mts, Decary14537 (P), (P); Ambalarano, Maevatanana, Boiteau 1011 (P); Betsiboka R. valley, near Meavatanana, Capuron SF 212 (P, TEF); Tsavongo, Mahavavy, Dybowski 1731 (P); Soalala, Amparihy, SF (TEF); Tongarejy, Morafeno, Ambato-Boeni, SF (P, TEF); Madirotelo, SF 4180 (P, TEF); Antsohihy, Bosser 5724 (P), 5725 (TAN); 24 km N of Ambondramamy, along Route nat. 6, Leeuwenberg & Rapanarivo (BR, P, TAN,WAG);LakeAnjiafitatra, near MtTsitondraina, Perrier de la Bâthie 966 (P, type of M. longifolia); Tsaramandroso, SF8146 (P,TEF, WAG); Ankarafantsika Reserve, SF 35 (P),71(P),77(P);ibid., RN 1119 (P, WAG);Marovoay, Perrier de la Bâthie13435 (P); Ambanjabe Forest, Ambalomoty, Cours3954 (P, TAN, WAG), 3975 (P, TAN, WAG); Namoroka Reserve, SF 4 (NY, P); ibid. RakotovaoRN 5788 (P); near Bekodoka, Decary 8257 (P); Mangobory, Soalala District, SF 3642 (P,TAN,TEF); 15km SEof Mahajanga, Lam & Meeuse 6097 (L, P);Mahajanga, Amborovy Road, Poisson 25 (P);Bombetoka Bay, Bojer anno 1839 (G-DC, type); Marohoga, SF5502 (P, TEF); near Majunga, Humbert 4064 (P), 4065 (P), 4084 (P); ibid., Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); ibid., Antsanitra, SF 3106 (P, TAN, TEF); Ambalakidy, Bosser 5841 (P); 7 km NW of Port Berge, Leeuwenberg & Rapanarivo (TAN, WAG);Tsinjomitondraka, PortBerge (=Boriziny),SF5519(P, TEF); Bongolava, Boina, Perrier de la Bâthie 8846 (P);Ampakobe, Rabesoana SF35-R-142 (P);km 84PortBerge-Antsohihy Road, nearanjiamangirana, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); Befandriana, Herb. Jard. Bot. Tana (TAN); km 82 Antsohihy-Ambanja Road, between Andrafiabe and Andranosamonta, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); Moramandia, Decary1052 (BM, P); Ambodinanave, Analalava, SF (P, TEF,WAG);Maromandia, Decary 1065 (K, P,TAN), 1245 (C, P);ibid.,Radama Peninsula, Decary 1157 (P).Antsiranana: Nosy Be, Richard 127and 181 (G-DC, composite sheet,typesofaf. angustifolia),127 (P,from Nigny Bay)and 181 (P, from Diego-Suarez);LokobeReserve, Bernardi (G, P);ibid., Pervilléanno Wageningen Agricultural University Papers 97-2 (1997) 31

30 1851 in Boivin s.a. (P, paratype of M. arborea); Beraty, Manongarivo R., Jacquemin 556 (B, P); near Ankaramy, km 50 Ambanja-Maromandia Road, Leeuwenberg et al (BR, MO, P, TAN, WAG); Ankarabato, Ambanja, Bernard RN 1282 (P, WAG); Benavony, Ambanja, SF (P, TEF); lower Sambirano R. valley, near Ambanja, Humbert (G, P); km 16 Ambanja- Ambilobe, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); Ampasindava Peninsula, Nof Kongony, Jacquemin 528 (B, P);nearAnanborano, E.I. White 2 Dec (BM); Mataipaka, SF (TEF); Ankarana Reserve, Humbert (K, P, US), (G, P, TAN); ibid., Keraudren 1656 (K, P, WAG); ibid., Leeuwenberg et al (BR, MO, P, TAN, WAG); ibid., Capuron SF (K, P,TEF,WAG);Mtd'Ambre, near Station des Roussettes, Leeuwenberg et al (BR, MO, P, TAN, WAG); ibid., Capuron SF (K, P, TEF, WAG, type of M. arborescens var. gracilis); ibid., near Grand Cascade, Malcomberetal (BR, P, TAN,WAG); neardiego-suarez, Pervillé 169 (P); ibid., Keraudren-Aymonin & Aymonin (P, WAG, paratype of M. angustifolia var. keraudreniana), (P, type of Af. angustifolia var. k.), 2561A (P, WAG, paratype of M. angustifolia var. k.); ibid., Mangoako Road, Debray 1562 (B, P, WAG); Diego-Suarez, Boivin 2461 (P, lectotype of M. anceps); km 10 Diego-Suarez to Orangea Road, Keraudren-Aymonin & Aymonin (P,WAG); Sahafary Forest, Debray 1539 (B,P, WAG, paratype of M. angustifolia var. k.); ibid., Keraudren-Aymonin & Aymonin (P), (P,paratypeofAf. angustifoliavar.ik.),25669 (P,WAG),25676 (P);km 4 Joffreville-Mt d'ambre Road, Harder et al (P,TAN, WAG);Antsahalalina, 12km Eof Daraina, D. Meyers 269 (K, P, TAN);Ampanihy R., Pervillé285 (P); Vohemar, Richard 58 (P), 107 (L, P), 551 (P); Sambava, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); ibid., Moral 2805 (P,TAN); Eof Ambatobiribiry, Capuron SF 876 (P, TEF, WAG); Antalaha, Roberson 13 June 1950 (P); Anjanaharibe Res., Ravelonarivo et al. 35 (TAN, WAG). Toamasina: Nosy Mangabe, Antongil Bay, Leeuwenberg et al (TAN, WAG); ibid., Schatz et al (BR, K, P, TAN, WAG); Maroantsetra, SF 145-R-199 (P); Ivoliona (= Voloina), Dequaire (P); Rantabe R. basin, near Beanana, Capuron SF9062 (P, WAG),9063 (TEF); Fananehana R. basin,androrana Mts, Capuron SF8963 (P,WAG);Masoala Penisula, 1-3 km Sof Ambanizana, Schatz & Modeste3129 (G, K, P, TAN, WAG); just above Antalavia R., Schatzet al (BR, C, K, P, TAN, WAG);Antalavia, Nicollet al. 573 (BR, K, P, TAN, WAG); Antrohy, Mananara, SF (P, TEF, WAG); île Ste. Marie, Boivin 1786 (G, P); Zahamena Res., Randrianjanaka & Arnaud 16 (WAG); Tampolo Forest, 10 km N of Fenerive, Leeuwenberg et al (BR, MO, P, TAN, WAG); ibid., Zarucchi et al (F, P, TAN, WAG); Analalava Forest, near Foulpointe, Leeuwenberg et al (BR, MO, P,TAN, WAG);ibid., Capuron SF (BR, K, P, TEF, WAG); Betampona Reserve, Ambodiriana, Rakotoniania RN 5913 (P, WAG); Ifantsy R., Rakotoniania RN 2467 (P); Ambendratsara, Manaka Est, Ramanatsoavma RN 2350 (P, TAN, WAG); Anjiofotsy, Manakambahiny Est, Ramanantsoavina RN 1779 (BR, K, P, WAG); Mandraka Forest, SF661 (P, TEF,WAG);Marivolanitra, Robin SF32-R-55 (P); Moramanga, Schlieben 8128 (B, BM, BR, G, HBG, K, M, TAN, UC, Z); ibid., Anosibe, SF 2193 (P, TAN, TEF), 2194 (P, TEF); Moramanga-Anosibe Road, Cours 871 (P, WAG);km 53 and 51 same road, SF7665 (P,TEF, WAG),7813 (P, TEF); km 45, Ankazomanitra, SF (P); Beravina, SF 3771 (P, TEF), 4382 (P, TEF); BetweenAntsahalanbe andantananarivo, Grandidier s.n. (P, type of Af. grandidieri); Analamazaotra Forest, Sof Permet, Leeuwenberg et al (BR, K, P,TAN, WAG); ibid., Müler 3768 (BR, K, P, TAN, WAG), 3836 (BR, 32 Wageningen Agricultural University Papers 97-2 (1997)

31 K, P, TAN, WAG); Ambodilazana, Warbur 21 May 1900 (K); Marongolo R., Dequaire (P,TAN);Anjiro Forest, Brickaville District, Vetondrozy SF 12- R-240 (P);nearAmbila-Lemaitso, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); ibid. SF (P, TEF, WAG); Bevakoana, Abraham SF (P,TEF);Sandrangato, Descoings 103 (TAN); ibid., Capuron SF28789 (P, TEF). Fianarantsoa: Ambohimitombo Forest, Forsyth Major331 (G, K); Ampasimpotsy Forest, Antoetra, SF 29-R-168 (P); N of Azozombe, N Imerina, Perrier de la Bâthie (P); Ampasinambo, Nosy Varika, Rakotozafy SF 85-R-116 (P); Marovato Forest, Manaka, Cours3103 (P, TAN, WAG); Mananjary, Geay 7744 (P), 7929 (P), 7930 (P), 7931 (P), 7932 (P); ibid., Ambodinana, SF (P, TEF); Ampasimazava, SF (P, TEF); Ikongo, SF (P, TEF, WAG); Andrambovato,Tolongoina, SF (P, TEF);Ranomafana, SF 4775 (P, TEF); Ranomafana Reserve, Leeuwenberg et al (BR, K, P, TAN, WAG); ibid., Makomber et al (BR, P, TAN, WAG); Antoetra, Saharanga, SF 29-R-168 (P); Ampamaherana, Fianarantsoa, SF 7054 (P, TEF, WAG), (P, TEF, WAG); Isaloa, Bosser (P, TAN, WAG); ibid., Perrier de la Bâthie (P);Isaloa Plateaux, Sof Sahanafo, Humbert (P,WAG);Isaloa, Gorgesdu Rat, Keraudren 410 (P), 433 (P);Canon des Singes, Jacquemin 275 (B, P, WAG); Ranohira, SF (P, TEF);Sakalema R. valley, Decary (P);Andringitra Res., Iantara R., Lewis et al. 760 (BR); ibid., SF 1491 (P, TAN, TEF); upper Iantara R., Humbert 3430 (G, P); Karianga, Decary 5551 (P);Vondrozo, Decary 4896 (P);Anabotany, Vohitrindry, SF6365 (P, TEF);Bevetraka R., Farafangana, SF25854 (P,WAG);ManomboForest, SF9487 (P,WAG);Vangaindrano, Scott Elliot2207 (P). Toliara:ManampanihyR.basin,Fitana Pass, Humbert 6052 (G, P, US); Nof Manantanina, Faber-Langendoen et al (P, TAN);Analava Forest, Manantenina, Guillaumet 3853 (P, TAN); Fort-Dauphin, Scott Elliot 2361 (E), 2440 (K), 2463 (BM), 2703 (BM, E, K, P, lectotype of M. coriacea); ibid., Pic St. Louis, Leeuwenberg et al (BR, K, P, TAN, WAG); Mandena F.R., N of Fort-Dauphin, Dumetz et al. 504 (K, P,TAN, WAG);ibid., Gereau et al.3298 (BR, K, P,TAN, TEF, WAG);ibid., Leeuwenberg et al (BR, K, P,TAN, WAG); Andohahela Reserve, Parcelle 1, Leeuwenberg14187 (BR, K, P, TAN, WAG); ibid., Makomberet al (BR, P,TAN, WAG);ibid., Phillipson 2963 (BR, K, P,TAN, WAG);ibid., Randriampionona 100 (TAN,WAG);Parcelle 2, 2 km S of Sakaravy, Leeuwenberg14160 (BR, K, P, TAN, WAG); Parcelle 3, Phillipson 2681 (K, P, TAN, WAG); N of Fort-Dauphin, along road to Ranomafana, McPherson (K, P, TAN, WAG); upper Mananara R. valley, Decary 9407 (C, GB, P); Tsilitsily, Behara, RakotosonRN 9431 (P); Ivakoany Mts, Humbert12195 (P); Beravy Mts, Hildebrandt 2078 (BM, G, K, M, P, W); Kirindy Forest, 55km NEof Morondava, Noyés et al (BR, P, WAG). Sin. loc: Baron 7>2A2 (K, P,typeof Holarrhena madagascariensis), 51 Al (K, P,typeof M, micrantha), 5767 (P),6157 (K, P),6433 (K, P); Bender 169 (BR, G, P),285 (B, P), 287 (G, P); Boivinannis (P,type of M. boivinii); Chapelier93 (P,lectotypeofM. arborea); Commerson s.n. (BM, FI-W, G, P, P-JU); Herb. Du Petit-Thouars s.n. (P); Godefroy Leboeuf anno 1899 (P, typeof M. barabanjà). Cultivated. U. S. A: Florida, Fort Meyers, Edison Estate Gardens, Brumbach 6634 (HAL, UPS), 7340 (K); Fairchield Tropical Garden, Gillis 9562 (S, USF). Puerto Rico: Mayaguez, Federal Exp. Station, Britton 8646 (GH, K, NY, P, lectotype of Lanugia latifoliä); ibid., Hess 3017 (NY, paratype of L. latifoliä);ibid., McCklland June 1926 (GH, NY, P, paratype of L. latifoliä); ibid., Wagner1145 (A). Brazil: Botanic Garden of Rio de Janeiro, RB (HBG). U.K.: Kew, Palmhouse H 873/66 (K); ibid., Home May 1878 (K); Edinburgh, Palm House, 9217 (E), of same tree C (E). Netherlands, Wageningen Agricultural University Papers 97-2 (1997) 33

32 Wageningen, Greenhouse, cutting received from Edinburgh, Leeuwenberg (WAG);of same tree, VanSetten 835 (WAG).Ghana:Aburi Gardens, Irvine 1905 (E); Togo, near Lomé, Warnecke 481 b (BM, P). Cameroun: Victoria (= Limbe) Botanic Garden, Leeuwenberg & Berg 9834 (BR, C, HBG, P, WAG); ibid., Leeuwenberg (BR, HBG, K, P, WAG). Gabon: Njégoné (?), Pobéguin 95 (P). Zaire: Equateur, Eala, Corbisier-Baland 2074 (BR, C, K, LG, M, P, UPS, WAG). Angola:Cabinda,Mayumbe,PangaMungo, Rio Luango, Gossweiler 6087 (LISU), 6887 (BM); Buco Zau, Gossweiler 7328 (LISU). Tanzania: Amani, Greenway 2285 (K), 6660 (K); ibid., Peter (B, WAG); ibid., Ruffo 1034 (K, WAG). Zimbabwe: Salisbury (= Harare), Bieget 4692 (K). Madagascar: Antananarivo, TsimbazazaPark, Bosser 8909 (P, TAN);ibid., Harizo RN 18Sept (BR, K, P, WAG). Mauritius: Pamplemoussses Bot. Gardens, Guého MAU (MAU); ibid., Vaughan MAU (MAU), (MAU). India: Bot. Garden Calcutta, Gage July 1903 (G); Lucknow Bot. Gardens, Balapure (A). Sri-Lanka: Bot. Garden Peradeniya H. 29. R.B.a, Jayaweera 1154 (A). Singapore: Bot. Garden, Furtado SING (BM, BO, L),17 June 1929 (A, FHO, K). Philippines: Los Banos, Baker 1 Dec (P).Indonesia: Java, Bogor Bot. Garden, rv. A. 131 (BO,K, L, P), 131 a (BO, WAG); LV. A. 131 a, Woejantoro137 (K, L);Tjikeumeuh Bot.Garden Sept (BO), Jan (BO), sin dat. (L ). Notes. The leaves vary greatly in shape and size. In most specimens they are up to 4.5 x as long as wide, but in some specimens collected they may be up to 15 x as long as wide. They are x as long as wide in Keraudren-Aymonin (paratype of M. angustifolia var. keraudreniana), x in Keraudren-Aymonin 2561A (paratype of same taxon), x as long as wide in Debray 1539 (paratype of same taxon), 6 9 x in Debray 1562, 5 10 x in Humbert 19085, 9 11 x in Capuron SF and x in Keraudren The continuity in this variation makes it impossible to maintain both M. angustifolia var. angustifolia and M. a. var. keraudreniana. The shape and size of the flowers vary independently from the leaves. The type and the topotype, Leeuwenberg 14315, of M. arborescens var. gracilis are much more branched and have smaller leaves than most other specimens belonging to this species. However, the flowers are not the smallest found in the species. Plants of M. arborescens, which were isolated by deforestation, have the leaves erect instead of horizontal to reduce irradiation Mascarenhasia havetii A. DC, Prod. 8: 488 (1844); Markgraf, Fl. Madag. Fam. 169: 262, pi (1976). - Type: Madagascar, Toamasina, Betanimena, near Manambonitra, Havet in Bojer s.n. (holotype G-DC; isotypes BM, K, MAU n.v., P). Fig. 5, p. 35; map 6, p Wageningen Agricultural University Papers 97-2 (1997)

33 Fig , Mascarenhasia havetii. 1 & 4, habit (x 2/3); 2, pistil head (x 6); 3, ovary with disk (x 6) Af. speciosa. 5, habit (x 2/3); 6, pistil head (x 6); 7, ovary with disk (x 6); 8,fruit (x 2/3). 1-2 from Capuron SF 9157; 3 from Capuron SF 9478; 4 from Havet in Bojer s.n.; 5-7 from Scott Elliot 2177; 8from Humbert5804. Wageningen Agricultural University Papers 97-2 (1997) 35

34 Map 5. Mascarenhasia lisianthiflora. Map 6. Mascarenhasia havetii; M. macrosiphon. Homotypic synonym: M. speciosa var. havetii (A. DC.) Boiteau, Caoutch.Madag. 8 (1943). Heterotypic synonyms: M. maroanaaug. D.C. in Bull. Herb. Boiss. II, 1: 580 (1901); Markgraf, Fl. Madag. Farn. 169: 261, pi. 44, 1-3 (1976), syn. nov. Type: Madagascar, Toamasina, Maroantsetra, Mocquerys 271 (holotype G;isotype Z). M. thiryana Pierre ex Dubard in Bull. Soc. Bot. Fr. 53: 303 (1906). Type: NEMadagascar, coast, sin. loc, Thiry s.n. (holotype P;isotype K). Shrub or small tree, 4 8 m high, with white latex in all parts. Branches pale or dark brown, lenticellate; branchlets glabrous or puberulous at the apex. Leaves:petiole glabrous or puberulous, 2 5mm long; blade subcoriaceous when dried, elliptic, x as long as wide, x cm, caudate to acuminate at theapex,cuneate atthe baseor decurrent intothepetiole,margin not 36 Wageningen Agricultural University Papers 97-2 (1997)

35 revolute, glabrous on both sides or with a few hairs on the base of the costa beneath, with 5 9 rather straight obscure secondary veins forming an angle of with the costa; tertiary venation inconspicuous. Inflorescence 1 3-flowered. Peduncle 1 3 mm long, puberulous; pedicels slender, mm long, sparsely puberulous or glabrous. Bracts sepal-like and about half as long as them. Flowers: Sepals light green, erect, triangular or ovate, x as long as wide, 1 3x mm, acute, ciliolate, pubescent or sparsely so outside, glabrous inside and with one row of 5 colleters inside at the base; colleters 0.2 x 0.1 mm. Corolla white, puberulous outside on the lobes, mostly only on the part covered in bud and sometimes at the apex of the tube, otherwise glabrous or even entirely glabrous, pilose on 2 mm of the filament ridges (near connective), otherwise glabrous but on lobes pubescent; tube x as long as the calyx, x as long as the lobes, mm long, above the base mm wide, abruptly widened of the length from the base which is at 8 21 mm to 5 7 mm wide, almost remaining at the same width to the mouth and 5 8 mm wide; limb spreading with lobes up to nearly halfway connate; lobes ovate or suborbicular, x as long as the tube, x as long as wide, x 7 18 mm, acuminate or apiculate. Stamens with apex 3 12 mm below mouth of corolla tube, inserted of the length of the corolla tube (at mm from the base); anthers 4 5 x as long as wide, 8 10 x mm, apex sterile for mm, fertile below for about half of the remaining part; connective pubescent. Pistil: ovary ovoid, x x 1 mm, retuse at the apex, glabrous; disk mm high; style rather slender, glabrous; pistil head of a basal entire veil 0.5 x 1 mm and a stigmoid apex 1.5 x 0.5 mm. Fruit only immature ones known; follicles very slender, x 2 mm. Distribution: Eastern Madagascar. Ecology: Forest understorey. Alt. low. Specimensexamined: Madagascar.Toamasina: MasoalaPeninsula, NofAntalavia R., Schatz et al (BR, K, P,TAN,WAG);Maroantsetra, Mocquerys 271 (G,Z,typeof M. maroana); Vohilava R.basin,tributary of Rantabe R., Capuron SF9157 (P, TEF,WAG);MahavohoForest,km9.5 IbandaRoad, Raharimalala 158 (P),198 (P), 200 (P); Mananara Nord, Ibanda Forest, Raharimalala 368 (P); Onilava, Sahatavy, Ramarokoto RN9478 (P,TAN,WAG);Zahamena Reserve, Botoalina RN3166 (P);Betanimena,nearManambonitra, Havet in Bojer s.n. (BM,G-DC, Wageningen Agricultural University Papers 97-2 (1997) 37

36 K, MAU n.v., P, type). Sin. loc: Bowless.n. (K); Lyall 146 (K); Thiry anno 1904 (K,P,typeof M. thiryana). Note. The flowers of the type of M. havetii are larger than in most other specimens of this species seen. In all other characters the specimens examined are perfectly similar, therefore M. maroana is reduced to synonymy Mascarenhasia lanceolata A. DC, Prod. 8: 488 (1844) Markgraf, Fl. Madag. Fam. 169: 272, pi. 46, 6-7 (1976). - Type Madagascar, Antsiranana, Nosy Be, Richard 370 (holotype G-DC isotype P). Fig. 6, p. 39; phot. 2, p. 121; map 4, p. 27 Homotypic synonym: Tsilaitra lanceolata (A. DC.) Baron in Rev. Madag. 16: 250 (1905). Heterotypic synonyms: M. brevituba Vatke in Abh. Naturw. Ver. Bremen 9: 125 (1885). Type: Madagascar, Antsiranana, Nosy Be, Hildebrandt 3299 (lectotype BREM, designated here; isolectotypes BM, G, K, L, M, P, W). M. rosea Bak. in Journ. Linn. Soc. 25: 335 (1890). Type: NW Madagascar, sin. loc., Baron 5840 (holotype K). M. utilis Bak. in Kew Bull. 1895: 199 (1895), nomen; Hooker, Icon. 24: t (1895). Type: N Madagascar, sin. loc, Baron 6370 (holotype K; isotype P). M.parvifolia Dubard in Bull. Soc. Bot. Fr. 53: 307 (1906). Types: Madagascar, Antsiranana, near Diego-Suarez, Hamelin in Godefroy Leboeufarmo 1900 (lectotype P, designated here; isolectotype BR); Nosy Be, Sakatia, Boivin Feb (paratype P). Shrub or small tree 1 6 m high, with white latex in al parts. Model of Koriba or Prévost. Trunk 1 10 cm in diameter; bark smooth, pale grey-spotted or pale grey-brown. Branches pale greybrown, lenticellate; branchlets puberulous. Leaves: petiole, puberulous or glabrous, 1 6 mm long; blade coriaceous even when fresh, elliptic or less often obovate, 1.7 4( 6.5) x as long as wide, 2 8 x cm, acuminate to rounded at the apex, cuneate at the base, margin often revolute, glabrous on both sides or slightly puberulous on base of costa beneath, with 5 12 pairs of rather straight secondary veins forming an angle of with the costa; tertiary venation reticulate, inconspicuous or invisible. Inflorescence 38 Wageningen Agricultural University Papers 97-2 (1997)

37 Fig Mascarenhasia tampinensis. 1, habit (x 2/3); 2, corolla tube inside withstamensandpistil (x 2); 3,fruit (x2/3); 4, seed (x 1). 5. M. rubra. 5, habit (x 2/3) M. lanceolata. 6, habit (x 2/3); 7, corolla inside with stamens and pistil (x 2). 1-2 from RN 5; 3-4 from Humbert & Swingle 5717; 5 from Perrier de la Bâthie 18267; 6 from Bosser 13232; 7 from Cours5647. Wageningen Agricultural University Papers 97-2 (1997) 39

38 1 4-flowered, about as long as the leaves. Peduncle 0 2 mm long, puberulous; pedicels 8 20 mm long, puberulous. Bracts scale-like, about 1 mm long, acute, puberulous. Flowers: Sepals medium green, free, suberect, ovate or oblong, 2 3 x as long as wide, 4 7 x mm, obtuse, ciliolate, puberulous on both sides, with one continuous row of colleters at the base which is 7 10 of them with each sepal; colleters equal or unequal, x mm. Corolla white, sometimes with a yellow throat and partly pink outside, puberulous outside but mostly glabrous or nearly so on the narrow basal part of the tube, puberulous inside on the lobes and villose from the insertion of the stamens to the mouth; tube x as long as the calyx, x as long as the lobes, (14 )16 20 mm long, mm wide above the base, abruptly widened at of the length which is at 2 4mm to mm wide, gradually narrowed towards the mouth to mm wide, again widened at the mouth to 4 5 mm wide; limb spreading or nearly so, with lobes almost free; lobes x as long as the tube, x as long as wide, x 6 17 mm, acuminate. Stamens with apex 5 9 mm below mouth of corolla tube, inserted of the length of the corolla tube (at 5 6 mm from the base); anthers 3 5 x as long as wide, x mm, apex sterile for mm, fertile below for 2 3 mm. Pistil: ovary cylindrical, x 2x 1.5 mm, retuse at the apex, pubescent except for the glabrous base within the disk; disk 0.5 mm high; style rather robust, glabrous; pistil head consisting of a subglobose basal part about lxl mm with which the anthers cohere, or a veil 0.5 x 1 mm and a stigmoid apical part about 1 x 0.5 mm. Ovules approximately 30 in each carpel. Fruit: follicles cylindrical, 9 17 x cm. Seed 7 10 x 1.5 x 0.5 mm; embryo 6.2 mm long; cotyledons 4.7 x 1.4 mm; rootlet 1.5 x 0.5 mm; coma mm. Distribution: NW Madagascar. Ecology: Dry forest or woodland on laterite or limestone. Alt m. Geographicalselectionoftheapproximately75specimensexamined: Madagascar. Antsiranana: Analamera Hills, Humbert 1915 (P);Ankarana Hills, Humbert (P), (G,P);Ankarana Reserve, Leeuwenberg etal (BR,MO, P,TAN, WAG);Andranomena, Debray 1553 (P,WAG); Diego- Suarez, Boivin 2459 (P);AmbohipirakaMt, Humbert & Cours (P, WAG); near Ambatoarana, Descoings 1923 (TAN); Nosy Be, BoivinFeb (P, 40 Wageningen Agricultural University Papers 97-2 (1997)

39 paratype ofm. parvifolia); ibid., Hildebrandt 3299 (BM, BREM, G, K, L, M, P, W,lectotypeofM. brevitubà); ibid., Richard 312 (P), 370 (G-DC, P, type);ibid., Lokobe Reserve, Bernardi (G, K, L, P, Z); ibid., Boivin March 1851 (P); Nosy Be, Amporaha, Bosser (P, TAN); Nosy Be, Mt Passot, Deroin & Badré189 (BR,K, P,WAG);lower SambiranoR. basin, nearambanja, Humbert (K, P, TAN, US, WAG), (P), (G, P, WAG); Ampasindava Peninsula, Jacquemin 479 (P), 535 (B, P); km 11 Ambilobe-Ambanja Road, Leeuwenberg et al (BR, MO, P,TAN, WAG), (BR, MO, P,TAN, WAG); 3km Sof Mahamanina, Leeuwenberg & Rapanarivo (BR, MO,P, TAN,WAG);km 42Ambanja-Moramandia Road, Leeuwenberg et al (BR, MO, P, TAN, WAG); Maromandia, SF (TEF); between Andrafïabe and Andranosamonta, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); Anaborano,Diego-Suarez, E.I. White 2Dec (BM);Ambolibe, Decary \A111 (GB, P, S), (K, P, TAN), (P); ibid., Waterlot 336 (P);Galoka Mts, SWof Ambilobe, Phillipson 2006 (K, P,TAN);Ampondrabe,Ambanja, SF5841 (P,WAG);SambiranoR. source, Last Dec (K);upper SambiranoR., Perrier de la Bâthie15215 (P); Beangona, Harizo RN 1280 (P, TAN); ibid., TsilizyRN 6963 (P,WAG); Marotolana,RN22 (P);ibid., Rakoto 5 Jan (TAN); Wside of base of Manongarivo Mts, Gentry (TAN). Mahajanga: Maromandia, Decary923 (P), 957 (P), 972 (P), 1326 (P); Sambirano, S of Maromandia, Capuron SF (K,P, TEF,WAG);Ankara Mts, Decary (G,L, P). Sin. loc: Baron 5840 (K,typeofM. rosea),6370 (K,P,type of M. utilis); Hamelinin Godefroy Leboeufanno1900 (BR, P,typeof M. parvifolia) Mascarenhasia lisianthiflora A. DC, Prod. 8: 487 (1844); Markgraf, Fl. Madag. Farn. 169: 265, pl. 1 5 (1976). Type: Madagascar, Mahajanga, Bombetoka Bay, Bojer s.n. (lectotype G- DC, designated here, as it has the fruit described; isolectotypes MAU, n.v., P). Fig. 7, p. 42; phots 3, p. 122 and 4, p. 122; map 5. p. 36 Homotypic synonym: Echitella lisianthiflora (A. DC.) Pichon in Mém. Inst. Sei. Madag. sér. B, 2: 90 (1949). Heterotypic synonyms: M. macrocalyx Bak. in Journ. Bot. 20: 219 (1882); M. lisianthiflora subsp. macrocalyx (Bak.) Boiteau, Caoutch. Madag. 9 (1943); Markgraf, op. cit. 266, pl. 45, 6 7. Type: Madagascar, sin. loc, Baron 92 (holotype K; isotype P). M. rutenbergiana Vatke in Abh. Naturw. Ver. Bremen 9: 124 (1885). Type: Madagascar, sin. loc, Rutenberg 2 Nov (holotype BREM n.v.). M. velutina Jum. in Compt. Rend. Acad. Sei. Paris 128: 1351 (1899). Type: Madagascar, Mahajanga, Betsiboka R., Perrier de la Bâthie 420 (lectotype P, designated here). M. lisianthiflora var. pubescens Dubard in Bull. Soc Bot. Fr. 53: 258 (1906). Type: Madagascar, sin. loc, Baron 5787 (lectotype P, designated here; isolectotype K). Wageningen Agricultural University Papers 97-2 (1997) 41

40 ï-^cxij Fig. 7. Mascarenhasia liskmthiflora. 1, 6, 8 & 9 habit (x 2/3); 2 & 7,calices (x 2); 3, sepal baseinside (x 8); 4 & 10, corollainsidewith stamensandpistil (x 2); 5,fruit (x2/3). 1-4from PerrierdelaBâthie8956; 5from SF3125;6-7 from Keraudren 1548; 8from Capuron SF548; 9from Homolle 1542;10 from Cours Wageningen Agricultural University Papers 97-2 (1997)

41 M. lisianthiflora var. baronica Dubard in op. cit Type: Central Madagascar, sin. loc, Baron 4716 (holotype P; isotypes BM, K). M. lisianthiflora var. hybrida Dubard in op. cit Type: Madagascar, sin. loc., Baron s.n. (holotype P; putative isotype Baron 92, K). M. phyllocalyx Dubard in op. cit Type: Madagascar, sin. loc, Grevé 100 (holotype P; isotypes K, TAN, WAG). M. humblotii Dubard in op. cit Type: Madagascar, sin. loc, Humblot 204 (lectotype P, designated here; isolectotype WAG). M. pallida Dubard in op. cit Type: Madagascar, Fianarantsoa, Ihorombe, Catat 4388 (holotype P). M. tenuifolia Dubard in op. cit Type: Central Madagascar, sin. loc, Baron 4575 (holotype P; isotypes E, K). M. kidroa Cost. & Bois, in Compt. Rend. Acad. Sei. Paris 144: 1054 (1907). Type: Madagascar, Antananarivo, District Ankazoabo, Geay 5983 (holotype P). M. geayi Cost. & Bois., I.e.; M. lisianthiflora subsp. geayi (Cost. & Bois.) Boiteau, Caoutch. Madag. 10 (1943); Margraf, op. cit. 269, pi. 45, 8. Type: Madagascar, Toliara, Taheza Mts, E of Tulear, Geay 6007 (holotype P). M. perrieri Lassia, Mascarenhasia et Landolphia Madag. 48 (1927); Markgraf, op. cit. 270, pi. 45, 9 10, syn. nov.; Echitella perrieri (Lassia) Pichon in Mém. Inst. Sei. Madag. sér. B, 2: 90 (1949). Type: Madagascar, Fianarantsoa, Mania R. basin, Perrier de la Bâthie (holotype P; isotypes K, TAN). Shrub or small tree, m high, with white latex in all parts. Trunks up to 25 cm in diameter; bark pale grey, smooth, lenticellate, often peeling off in large scales. Branches pale to dark brown, lenticellate; branchlets pubescent or puberulous. Leaves: petiole pubescent or puberulous, 1 10 mm long; blade subcoriaceous or coriaceous even when fresh, elliptic or narrowly so, 1.1 3( 4.5) x as long as wide, x cm, variable in shape and size, rounded to acuminate at the apex, truncate to cuneate at the base, margin mostly not revolute, pubescent to almost glabrous on both sides, with indumentum usually at least on costa above and main veins beneath; with 4 12 pairs of rather straight to upcurved secondary veins forming an angle of with the costa; tertiary Wageningen Agricultural University Papers 97-2 (1997) 43

42 venation reticulate, conspicuous or not. Inflorescence 1 6-flowered. Peduncle 1 4mm long, pubescent or puberulous; pedicels 5 12mm long, pubescent or puberulous. Bracts scale-like, much smaller than the sepals. Flowers fragrant. Sepalspale green, often suffused with dark red, erect, unequal, obliquely elliptic or oblong, x as long as wide, largest of a flower 6 15 x mm, variable in shape and size,rounded to caudate,ciliolate,pubescent or sparsely soonboth sidesoronly inside sometimes glabrous,withone row of 4 7colleters at the base;colleters variable in shape and size, x mm, when wide lobed. Corolla white, with tube often partly greenish and/or reddish, sometimes greenish or yellowish inthe throat, pubescent or sparsely sooutside, occasionally partly glabrous on tube, pubescent to sparsely puberulous inside on lobes, glabrous or sometimes with pubescent stripes from the insertion of the stamens to the mouth; tube x as long as the calyx, x as long as the lobes, 25 50mm long, 1 2 mm wide at the base and almost cylindrical for of the length which is for 11 32mm, abruptly widened above to 3 7 mm wide, gradually narrowed towards the throat to 2 5mm wide and slightly widened at the mouth; limb spreading, with lobes almost free; lobes x as long as the tube, x as long as wide, x 5 16mm, ovate or elliptic, acute or acuminate, often crenate. Stamens with apex 1 13 mm below mouth of corolla tube, inserted at of the length of the corolla tube (at 13 34mm from the base); anthers 5 6x as long as wide, x mm, apex sterile for 0.5 lmm, fertile below for 3.5 4mm; connective pubescent. Pistil: ovary cylindrical, 1 2x 1 2x mm, retuse, pubescent except for the glabrous base; disk slightly lower to slightly higher than the ovary, 1 1.7mm high; style rather slender, glabrous; pistil head of 4 parts, first a basal ring mm wide, a cylinder which may be narrower in the middle, x mm, again a ring mm wide and a stigmoid apex x mm. Ovules approximately 50 in each carpel. Fruit: follicles 5 18 x 0.5 lcm, glabrous or pubescent. Seed x 1.2 x 0.5 mm; embryo 5.2 mm long; cotyledons 4.2 x 1.4 mm; rootlet lx 0.5mm; coma about 3 x as long as grain, 25mm long. Distribution: Endemic to Madagascar. Ecology: Semideciduous forest, often with Didieraceaeor woodland, often near base of rocky hills. Alt m. 44 Wageningen Agricultural University Papers 97-2 (1997)

43 Geographical selectionoftheapproximately 300specimensexamined: Madagascar. Mahajanga: Miandrivazo, SF 6303 (P, TEF); Andriambe, Bekopaka, SF 3-R-286 (P); Masoarivo, Leandri 528 (NY, P); Bemaraha Mts, Leandri120 (BM, C, P, TAN); Tsimambo Forest, SF 8243 (P, TEF, WAG); Ambereny Forest, Antsalova, Rakotozafy 1029 (TAN); Tsiampihy Forest, near Besaraha, Leandri 371 (K, P, US), 2225 (P, WAG); Ankinana, Leandri 990 (P); near Maintirano, Decary 8273 (P,W);ibid., Ambararatakely, SF25049 (P, TEF); nearmorafenobe, Decary 2288 (P),2298 (P),2318 (P),2326 (P);ibid., SF (P, TEF, WAG); Ambodiroka, Maevatanana, Descoings3302 (TAN); Belambo Forest, 5 km S of Maevatanana, Dorr et al (BR, K, P, TAN, US,WAG); Besafotra R. basin, between Menavava and Ikopa Rs., Perrier de la Bâthie 420ter (P); Besalampy-Maintirano Road, Morat 983 (TAN);near Betsiboka, Capuron SF 153 (P, TEF); Tsitampiky (= Sitampiky), Decary 8188 (P); Ankarafantsika Reserve, Decary12809 (L, NY, P); Analamitsangana, Ankijabe, SF (P, TEF, WAG); Ambato-Boeni, Perrier de la Bâthie (P); near Bekodoka, Decary 8260 (P); Besalampy, Decary (P, WAG), (P, WAG); ibid., Andranomavety, SF 6262 (P, TEF, WAG); Amborometroka, SF (BR, P, TEF, WAG); Namoroka Reserve, Rakotovao RN 3891 (P, TAN), 5088 (P, WAG); Mahabo, SF (P, TEF); Komehevitra, SF 3988 (P, TAN, TEF); Ankarafantsika Reserve, SF69 (P), 70 (GB, P); ibid.,ampijoroa, Phillipson 1930 (BR, K, TAN, WAG);Tsaramandroso, SF4945 (P,TAN, TEF);Maevarano, SF 3125 (P,TAN, TEF); Boina, Perrier de la Bâthie 8955 (P);ibid., upper Bemarivo R., Perrier de la Bâthie8956 (P); Betsiboka R., Perrier de la Bâthie 420 (P, lectotype of M. velutina);soalala-mitsinjo Road, SF4020 (P,TEF); Madirokely, Marovoay, Bosser8440 (P);Sambirano, Soalala, SF4315 (P, TEF);Mitsinjo, SF 4025 (P, TEF); ibid., Masofandroboaka, SF (P, TEF); Bevazaha, Ramamonjisoa RN 1661 (BR, K, P, WAG); Imonty, Ankazoabo, SF 4138 (P, TEF);Ampagony, Peltier 5386 (P);Boanamary, Descoings 3563 (P,TAN); ibid., Peltier 5249 (P, WAG); Bombetoka Bay, Bojers.n. (G-DC, P, lectotype); ibid., Richard528 (P); near Beronono, Boiteau 1054 (P, WAG); near Mandritsara, Bosser16663 (P);Mahajanga, Afzelius 7April 1912 (K, UPS);ibid., SF 1846 (P, TEF); 2 km E of Mahajanga, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); 6 km N of Mahajanga, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); Bongolava, N of Mahajanga, Morat 2662 (P, TAN); Ampakobe, Port-Berge, SF 5874 (P); Antsahanira, Rakotozafy 1861 (TAN); Analamena Forest, Andranomena, SF (P, TEF); Mamoa Forest, Tsarahonena, SF (P, TEF); Misorobe Forest, Analalava, SF 89-R-78 (P); km33ankarika-analalava, Leeuwenberg & Rapanarivo (BR, MO, P,TAN, WAG); 7 km SW of Ambalafamainty, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); Maromandia, Decary1202 (P). Antsiranana: Nosy Be, Decary 8249 (C, P); ibid., Richard 307 (G-DC, L, P, paratype). Antananarivo: Ankazobo District, Geay 5983 (P, type of M. kidroa), 5985 (K, P), 5986 (P); Tsiroamandidy, Catat 1114 (P); Mangarivo, Grevé 41 (K, P, TAN, US); Antsirabe, Poisson 616 (TAN). Toamasina: Antongil Bay, Boivin s.n. (P); Andilamena-Miarinarivo Road, Bosser16669 (P, TAN); Andilameno,Miarinarivo, Morat 226 (TAN);Ampenefanevohe, Bosser8100 (TAN).Fianarantsoa: ManiaR. basin, Perrier de la Bâthie (K, P, TAN, type of M. perrieri); Wof Itremo, Humbert (K, P, WAG);between Mandosonoro and Amborompotsy, Morat 4172 (P); Fianarantsoa, SF 2135 (P, TAN, TEF); Ankafina, Deans Cowan anno 1880 (BM, P); Ankaramena, SF (P, TEF); Ambalavao, Teteza, SF19-R- 239 (P);Andringitra Reserve, Dorr 3926 (BR, K, P, S, TAN, US, WAG); ibid., Rakoto RN 5831 (P); 35 km S of Ambalavao, along nat. road 7, Miller & Wageningen Agricultural University Papers 97-2 (1997) 45

44 Randrianasolo 6267 (K); km 65 Ambalavao-Ihosy Road, 15 km N of Voatavo, Leeuwenberg & Rapanarivo (BR, MO, P,TAN, WAG); Wof Ambalavao, Croat (K, P,TAN, WAG);Iandranbaky, Sof Ambalavao, Bernardi (A, BR, E, FT, G, L, NY, P); near Zazafotsy, Keraudren 308 (P), 320 (P);ibid., Labatet al (P); ibid., Ambahatoazo, Felder 1243 (P, WAG); Masoala, Vohitsaoka, Razafindrakoto RN 2293 (P, TAN); Ihosy, Peltier 2740 (P, TAN, WAG); 1km S of Ihosy, near Ihosy R., Leeuwenberg & Rapanarivo (BR, MO, P,TAN, WAG);Menarahaka R. valley, between Ihosy and Ivohibe, Leandri & Rakoto 3468 (K, P, WAG); near Ivohibe, Armand 85 (P); Ihosy-Analalivry Road, towards Sakasoa, Allorge & Veyret 594 (P);Iantara R. valley, Ivohibe, SF 1460 (P, TAN, TEF); Ihorombe, Cam 4348 (P, type of M. pallida); Lohony, Farafangana, RakotovaoRN7660 (P);northern partofisaloa Mts, Humbert (BR, K, P,WAG);Isaloa Mts, Cours 5024 (P,TAN, WAG);ibid., Decary (P, WAG);ibid., Ranohira, Schlieben 8219 (B, BM, BR, HBG, K, M, TAN, Z), 8220 (B, BM, BR, HBG, K, M, P, TAN, UC, Z), 8242 (B, BM, BR, HBG, K, M, P, TAN, UC, Z); Wof Ranohira, Capuron SF (P, TEF, WAG); ca 10 kmswof Ranohira, Dorr etal.4181 (BR, P,TAN,WAG); Sofkm 11Ranohira- Sakaraha Road, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG), (BR, MO, P, TAN, WAG);Ampandrabe, SF (TEF); Col de Tapia, 59km EofSakaraha, Croat (P, TAN,WAG);ibid., Capuron SF20244 (K, P, TEF, WAG); Ilalaka, Isalo, Peltier 2984 (P), 5789 (P); ibid., SF (P, TEF); Lombomakondro, Perrier de la Bâthie (P); Benonoka, SF (TEF).Toliara: Mendoravy, SF (P,TEF);Fort-Dauphin, Decary 4105 (P), 4312 (P), 4319 (P);ibid., Scott Elliot 2373 (K, P, M. sessilifoliadubard insyn.of M. arborescens in Bull. Soc.Bot.Fr. 53: 295 (1906));Manavy,Ambovombe, SF 9567 (P, TEF, WAG); Andohahela Reserve, Rakotoniania RN 3461 (P, TAN, WAG),6213 (P, WAG);ibid., Parcelle 1, Randriamampionona 85 (TAN,WAG); Parcelle 2, Leewenberg (BR, K, P,TAN, WAG);Parcelle 3, Leeuwenberg (BR, K, P, TAN, WAG); ibid., Randriamampionona 121 (TAN, WAG), 273 (TAN, WAG); upper Mandrare R. valley, Humbert 6488 (B, NY, P, US, WAG);Lavanala For., GrandidierOct (P);Filanjora, Mahabo, SF (P, TEF, WAG);Ejeda, B. Du Puy MB 42 (TAN);nearAnadabolava Forest,middle Mandrare R., Keraudren 1548 (P,WAG);Ambovombe District, Decary 2824 (P); between Tsivory and Anadabolava, Humbert12333 (P); upper Onilahy R. basin, Andranomiforitra R. valley, Humbert 7050 (P); Malio R. basin, tributary of Mangoky R., near Ambalabe, Humbert (BR, K, P, WAG); Betroka, SF 8396 (P, TEF); Vohipolaka Mt, N of Betroka, Humbert11651 (P); Antanimora, Decary 3323 (P);Beloha-Tranoroa Road, Keraudren 921 (P,WAG);Bevoay,near Beraketa, Peltier 2897 (P,TAN, WAG);Ampandrandava, Seyrig21 (P); Bekely, SF 9663 (P, TEF, WAG); Ankilivalo, S of Berenty-Betsiko, Ankazoabo, Morat 2522 (P);Zombitsy For., Willing 36 (TAN);Betroka, Herb.Jard. Bot.Tana.4989 (TAN); near Morondava, Menabe, Exp. Colon. Marseille s.n. (P); Malaimbandy, Morat69 (P, TAN);between Malaimbandy andankilizato, Keraudren-Aymonin & Aymonin (P, WAG); Dabolava, Decary (P); Ankazomanja, Decary (P), (P); Ambatakazo, Miandrivazo District, Decary (L, P); Beraboka Forest, Miandrivazo District, SF (P, TEF); Ampasimandroatse Forest,Belo/Tsiribihina, SF 54-R-247 (P); Morondava, Grevé204 (K, P, S,TAN, WAG, lectotype of M. humblotii); ibid., Anjenanja, SF (P, TEF, WAG); Manja, Rauh898 (TAN);Ankilizato, Bosser4591 (TAN); Betoboro, Befasy, SF (P, TEF); between Morondava and Bemaraha Mts southern base hills, Humbert11367bis (P); Wof Ankilizato, Keraudren-Aymonin & Aymonin (P, WAG); Beroroho Road, Morat2463 (P, TAN); between Antanimieva and 46 Wageningen Agricultural University Papers 97-2 (1997)

45 Befandriana, Poisson 635 (P); Beravy Mts, Hildebrandt 3099 (BM, K, P, W); upperfiherananar. basin, Perrier de la Bâthie (P);LambomakandroForest, E of Sakaraha, Capuron SF548 (P, TEF);Zombitsy Forest, Bosser (P, TAN), (P,TAN),13981 (P,TAN);km18Sakaraha-IhosyRoad, Müler & Keating 4526 (K,P,TAN,WAG);nearFanjahira, Humbert2745 (P);OnilahyR. valley,neartongobory, Humbert 2111 (P,TAN);Hazoroa,Tulear, SF3396 (P, TAN,TEF);TahezaR., Geay6003 (BM),6004 (P),6005 (K), 6006 (TAN), 6007 (P, type of M. geayi); Beza Mahafaby Reserve, near Betioky, Parcelle 1, Phillipson 2640 (BR, C, K, P, TAN, WAG); Parcelle 2, Phillipson 1828 (P, TAN);Mahafaly Plateau, near Itampolo, Perrier de la Bâthie 4407 bis (P); Eof Ampotaka, Geay 6345 (P). Sin. loc: Baron 92 (K, P,type of M. macrocalyx), 4562 (K, P),4575 (E, K, P, type of M. tenuifolia), 4716 (BM, K, P, typeof M. lisianthiflora var. baronica), 5787 (K, P,typeofAf. /. var. pubescens), 6769 (K), 6874 (K), 6869 (K), s.n. (= 92?) (P, type of M. I. var. hybrida); Commerson s.n. (C, P, P-JU, paratype of M. I.var. pubescens); Grevé 100 (K, P, TAN, WAG, type of M. phyllocalyx); Humblot 204 (P, WAG, type of M. humblotii); Richard239 (FI-W,TCD), 284 (P),845 (P). Cultivated. U. S. A.: Florida, Subtrop. Hort. Research Station, Miami, Avery 1585 (USF); Plant Introduction Station, Miami, Gillis 9453 (P). Madagascar Bojer anno 1833 (G-DC,paratype);TsimbazazaPark,Antananarivo, Rakoto RN 1769 (P).Réunion,BotanieGarden, Pierre 244 (P).Mauritius: Ayr es s.n. (GH); Bojers.n. (K).India:BotanieGardenCalcutta,anonym,s.n. (L,P)and Pierre3584,April 1863 (P). Notes. The leaves of the lectotype and paratype of M. lisianthiflora are 6 14 x 3 5 cm, subcoriaceous and acuminate, and they have conspicuous tertiary venation. Those of the type of M.perrieri are x cm, coriaceous and obtuse or rounded and they have invisible tertiary venation. The flowers of the latter are similar in most details, but they are less hairy. Intermediates between these two and the other forms kept together here exist without any correlation between the differentiating characters. A good series to show why M. perrieri is not kept up as a species has been collected by Schlieben at Ranohira in the Isalo Mts, 8242 with the same leaves as the type of M. perrieri but more hairy all over, 8220 perfectly intermediate, first named M. lisianthiflora and later changed by Markgraf to M. perrieri, and finally 8219 a perfect M. lisianthiflora of which only the leaves are smaller than those of its type. The range of leaf sizes is x cm M. macrosiphon Bak. in Journ. Linn. Soc. Bot. 22: ); Markgraf, op. cit. 260, pi. 43, 5 6. Type: Central Madagascar, sin. loc, Baron 3840 (holotype K; isotypes BM, P). Fig.4, p. 26; map 6, p. 36 Wageningen Agricultural University Papers 97-2 (1997) 47

46 Heterotypic synonym: M. mangorensis Jum. & Perr. in Agric. prat pays chauds 12:433 (1912). Type:Madagascar, Toamasina, upper Anosivola R., tributary right bank of Mangoro R., Perrier de la Bâthie18625 (lectotype P,designated here). Shrub or tree 5 10 m high. Branches pale or dark greybrown, lenticellate; branchlets puberulous. Leaves: petiole puberulous or glabrous, 2 6mm long; blade subcoriaceous when dried, elliptic or obovate, x as long as wide, x 1 2.5cm, rounded to shortly acuminate with a blunt acumen at the apex, cuneate at the base, margin often revolute, with a few minute hairs on the costa beneath or entirely glabrous, with 4 10 pairs of rather obscure and rather straight secondary veins forming an angle of with the costa; tertiary venation not or hardly visible. Inflorescence 1 2-flowered. Peduncle 0 2 mm long, puberulous or glabrous; pedicels 3 12mm long, puberulous or glabrous. Bracts sepal-like, about half as long as them. Flowers: Sepals green or reddish outside, erect or nearly so, ovate or triangular, x as long as wide, x mm, acute, ciliolate, sparsely puberulous to glabrous outside, glabrous inside and with 1 2 colleters atthe edgeof each sepal (5 10in aflower); colleters 0.2 x 0.2 mm. Corolla white, partly puberulous outside all over to only on the lobes and in stripes on upper part of tube, pilose inside often(?) in stripes from the base to the insertion of the stamens and from the apices of the anthers to the mouth, pubescent on the lobes; tube x as long as the calyx, x as long as the lobes, 26 37mm long, 2mm wide above the base, abruptly widened of the length from the base which is 10 11mm from there to 3 5mm wide, again narrowed gradually towards the mouth to mm wide, widened at the mouth to 5mm wide; limb spreading, with lobes almost free; lobes ovate or nearly so, x as long as the tube, x as long as wide, x 6 8 mm, acuminate. Stamens with apex 9 17mm below mouthof corolla tube, inserted of the length of the corolla tube (at 13 14mm from the base); anthers 4 5 x as long as wide, 6 8 x mm, apex sterile for mm, fertile below for about half of the remaining part. Pistil: ovary cylindrical, 1.5 x 1.5 x 1mm, refuse at the apex, pubescent except for the part covered by the disk; disk 0.5mm high; style rather slender, pilose;pistil headof a basal entire veil 0.5 x 1 mm and a stigmoid apex 1 x 0.5mm. 48 Wageningen Agricultural University Papers 97-2 (1997)

47 Ovules approximately 30 in each carpel. Fruit, follicles linear, 20 x 0.5 cm. Distribution: Eastern Madagascar. Ecology: Forest understorey. Alt. up to 1150 m. Specimensexamined: Madagascar. Antsiranana: Beanajada Mts, N of Masoala Peninsula, Capuron SF 8810 (B,G, K, P,TAN, TEF).Toamasina:Analamazaotra Forest, Louvel 9 (P); ibid., Ramanantoavolana in Thouvenot 97 (K, P, paratype of M. mangorensis); WofAntanandava,km45Moramanga-AnosibeRoad, Capuron SF28431 (G,K,P,TEF);Ankazomanitra,Anosibe,SF26832 (P,TEF, WAG); upperanosivola R., tributary rightbankmangoror., Perrier de la Bâthie (P,lectotypeof M. mangorensis). Fianarantsoa:RanomafanaReserve, Rakoto292 (P, TAN, WAG), 341 (P, TAN, WAG); ibid., Turk & Randrianasolo 582 (P, TAN,WAG).Sin. loc: Baron 3840 (BM,K,P,type) Mascarenhasia rubra Jum. & Perr. in Ann. Mus. Colon. Marseille III, 6: 19 (1918); Markgraf, Fl. Madag. Farn. 169: 274, pi. 46, 5 (1976). Type: Madagascar, Antsiranana, Masoala Mts, near Marambo, Perrier de la Bâthie (holotype P). Fig. 6, p. 39; map 4, p. 27 Shrub or small tree. Branches pale brown; branchlets glabrous. Leaves: petiole 3 10 mm long; blade coriaceous when dried, elliptic, x as long as wide, x cm, acuminate at the apex, cuneate at the base, margin revolute, glabrous on both sides, with 8 9 pairs of upcurved secondary veins forming an angle of with the costa; tertiary venation invisible. Inflorescence probably 3-flowered. Peduncle 3 mm long, glabrous; pedicels 12 mm long, glabrous. Flowers: Sepals green(?), oblong, 8 11 x 2 mm, obtuse, glabrous on both sides, with colleters(?). Corolla red, puberulous outside on apex of tube and on lobes, with stripes of pubescence inside from the insertion of the stamens to the mouth and pubescent on the lobes; tube at least 33 mm long, above the base 2 mm wide and cylindrical for 12 mm, abruptly widened to at least 5 mm wide. Stamens deeply included; anthers 9 mm long, glabrous. Fruit, follicles 22 x 0.5 cm (only one known on type), as those of M. arborescens. Distribution: Only known from the type. Ecology: Forest understorey. Wageningen Agricultural University Papers 97-2 (1997) 49

48 Note. It is possible, that some of the fruiting specimens named M. arborescens,which do not flower at the same time, belong to M. rubra Mascarenhasia speciosa Scott-Elliot in Journ. Linn. Soc. Bot. 29: 32 (1891);Markgraf, Fl. Madag. Fam. 169:264,pi. 44, 5-8 (1976). Type: Madagascar, Fianarantsoa, Vangaindrano, Scott- Elliot 2177 (lectotype K, designated here, cited as holotype by Markgraf;isolectotypes BM,E, P). Fig. 5, p.35;map 4,p.27 Heterotypic synonym: M. speciosa var. dextradubard in Bull. Soc. Bot. Fr. 53: 256 (1906). Type: Madagascar, Toliara, Fort- Dauphin, Chisel 66 (holotype P;isotypeBM). Shrub or small tree, m high, with clear sap in all parts. Branches dark brown, obscurely lenticellate; branchlets puberulous or glabrous. Leaves: petiole 1 5 mm long, puberulous or glabrous; blade subcoriaceous when dried, elliptic, x as long as wide, x cm, caudate to apiculate at the apex with a blunt acumen, cuneate or rounded at the base, glabrous on both sides or puberulous on base of costa beneath; secondary and tertiary venation inconspicuous. Flowers solitary. Peduncle 10 25mm long, glabrous. Bracteoles minute, scale-like, near base of peduncle. Sepals green(?) or red (teste Chisel 66 and 79), suberect in dried flowers, obliquely ovate or elliptic, 2 3 x as long as wide, 5 12 x 2.5 5mm, obtuse or acuminate with a blunt acumen, ciliolate or not, sparsely puberulous to glabrous on both sides, with 2 5 colleters at the base; colleters in groups of 2 3at the edge of the sepals or in a continuous row, x mm, often lobed when large. Corolla entirely white or with tube and part of lobes not covered in bud red or pink outside and white on remaining part of lobes and inside, puberulous outside on the in bud covered part of the lobes,otherwise glabrous, glabrous inside in tube and puberulous on lobes, usually in a central longitudinal line;tube4 11 xaslong as the calyx,4 6.5 x aslong as the lobes, 45 60mm long, funnel-shaped, basal part cylindrical, of the tube length, x mm, gradually widened to 4 6mm wide 2 5mm above, at the level where the stamens are inserted, 12 20mm wide at the mouth; lobes spreading, broadly ovate or nearly so, x as long as the tube, x as long as wide, apiculate. Stamens with apex 50 Wageningen Agricultural University Papers 97-2 (1997)

49 0 9 mm below mouth of corolla tube, often exserted in dried flowers, inserted of the length of the corolla tube (at mm from the base); anthers 4 6 x as long as wide, 8 9 x mm, apex sterile for mm, remaining part fertile for upper half or slightly more, again sterile below; connective pubescent to glabrous. Pistil: ovary ovoid, 1.5 2x 1.5 2x mm, gradually narrowed into the style or obtuse, glabrous; disk lower than ovary, mm high; style slender, from pilose at the base to pubescent at the apex; pistil head of a basal veil about 0.5 x 1 mm and a stigmoid apex about 1.5 x 0.5 mm. Ovules approximately 40 in each carpel. Fruit: follicles x 0.4 cm, with a thin wall. Seed grain x 1.5 x 0.5 mm; embryo almost filling the seed; coma about twice as long as the grain. Distribution: Endemic to southeastern Madagascar. Ecology: Dry forest or bush. Alt m. Geographicalselectionofthe35specimensexamined: Madagascar.Toamasina:ManampotsyR.,SWofVatomandry, Perrierdela Bâthie (P). Fianarantsoa: S of Vangaindrano, Scott Elliot 2155 (K, P, paratype), 2177 (BM, E, K, P, lectotype). Toliara: near Manantenina, Humbert (P, WAG); N of Fort-Dauphin, towards Belavenoka, Decary 1732 (P); Tsingafiafy Forest,between ManambatoandFitamalama Rs., Capuron SF28681 (BR,K,P,TEF,WAG); BemangidyForest, NofMahatalaky, Capuron SF (BR,K,P,TEF,WAG);Ste.Luce, Dumetz 1201 (BR,K,P,WAG); 44km Nof Fort-Dauphin, alongroad to Ste.Luce (Manafiafy), Phillipson etal (TAN, WAG); N of Antorendrika R., Gereau et al (BR, K, P, TAN, WAG); ManantantelyForest, Humbert 5804 (G,L,P,TAN,US,WAG),20354 (BR,K, P, WAG); ibid., Rabevohitra 2423 (P, TAN, TEF, WAG); Mandena Forest, Dumetz etal.585 (BR,K,P,TAN,TEF,WAG);ibid., Rabevohitra 2203 (K,P, TAN,TEF,WAG);Fort-Dauphin, Chisel 66 (BM, P, type of M. speciosa var. dextra), 79 (BM,P);ibid., Decary 4166 (BM,K, P,US);nearVinanibe,SW of Fort-Dauphin, Capuron SF (K, P, WAG), (P, TEF, WAG);Italy Road, Bosser (P,TAN) Mascarenhasia tampinensis Pichon in Not. Syst. ed. Humbert 13: 211 (1948); Markgraf, Fl. Madag. Fam. 169: 274, pl. 46, 1 4 (1976). Type: Madagascar, Toamasina, Tampina, Ursch 55 (holotype P). Fig. 6, p. 39; map 4, p. 27 Shrub or small tree. Branches medium grey, lenticellate; branchlets glabrous or minutely puberulous. Leaves: petiole 1 5 mm long, glabrous; blade coriaceous when dried, elliptic or narrowly so, x as long as wide, 3 6x cm, Wageningen Agricultural University Papers 97-2 (1997) 51

50 acuminate to obtuse at the apex, cuneate at the base, glabrous on both sides, with 4 7 pairs of obscure secondary veins; tertiary venation inconspicuous. Flowers solitary. Peduncle 7 12 mm long, glabrous, with 1 2 bracteoles at the base. Bracteoles sepal-like and x as long as them. Sepals green(?), erect, oblong or nearly so, x as long as wide, 6 9x mm, obtuse, not ciliolate, glabrous on both sides, with 10 colleters inside in one row at the base; colleters 0.2 x 0.1 mm. Corolla tube and part of lobes not covered in bud red, remaining part of lobes and inside white, puberulous outside all over, only on the lobes or even only on the part of the lobes covered in bud, pubescent inside on lobes and pilose in upper 1 cm of tube, otherwise glabrous; tube x as long as the calyx, x as long as the lobes, mm long, above the base 2mm wide, abruptly widened at of the tube length which is at 7 10mm into a campanulate or narrowly urceolate part which is 8 14 mm wide above its base and mostly narrowed at the throat to 8 11mm wide, again widened at the mouth; lobes spreading, ovate or nearly so, x as long as the tube, x as long as wide, x 8 15 mm, apiculate. Stamens deeply included, inserted just above the widening of the tube; anthers x as long as wide, 9 12 x mm, apex sterile for 1 2 mm, fertile for one half of the remaining part and again sterile below; connective pubescent. Pistil: ovary ovoid, x x mm, obtuse, glabrous; disk lower than ovary mm high. Fruit unknown. Distribution: Eastern Madagascar. Ecology: Forest understorey, near river banks. Alt. 0 4 m. Specimensexamined: Madagascar. Toamasina: Tampina Forest, Ursch 55 (P, type); Ambila- Lemaitso,SF 1109 (P,TAN,TEF), 1130 (P),6326 (P,TEF);ibid.,Ambodivily Forest, RakotosihanakaRN19Jan.1949 (P,TAN); ibid., RandrianasoloRN 5, 24 Jan (P,TAN);ibid.,SF2926 (P,TAN,TEF, WAG). Species excluded from Mascarenhasia Mascarenhasia curnowiana Hort. in M.T. Masters (ed.) in Gard. Chron. II, 16: 283 (1881), as Mascarenhaisia. Type: Cult., seeds from Madagascar, (holotype K) = Alafia thouarsii Roem. & Schult. M. gerrardiana Bak. in Journ. Linn. Soc. Bot. 22: 504 (1887). Type: Madagascar, sin. loc, Baron 4652 (lectotype K, designated here) = Alafia thouarsii Roem. & Schult. 52 Wageningen Agricultural University Papers 97-2 (1997)

51 4. Petchia Livera in Ann. Roy. Bot. Gard. Peradeniya 10: 140 (1926); H. Huber in Abeywickrama (ed.), Flora of Ceylon 1. 1: 16 (1973). Type species: P. ceylanica (Wight) Livera. Heterotypic synonym: Caubucala Pichon in Not. Syst. ed. Humbert 13: 202 (1948);Markgraf, Fl. Mad. fam. 169:61 (1976), syn. nov. Type species: C. madagascariensis (A. DC.) Pichon (= Petchia madagascariensis (A.DC.) Leeuwenb.). Shrubs or small trees with white latex (latex not mentioned with P. humbertii and P. plectaneiifolia), glabrous all over except for the sometimes ciliate sepals and corollas inside. Branches with concolourous lenticels; branchlets terete. Leaves at least on some branches, especially where they ramify in whorls of 3 5, otherwise opposite, only in P. africana all opposite, petiolate; blade variously shaped, entire. Inflorescences terminal and often also axillary (see note). Sepalsconnate at the extreme base, subequal, clasping the corolla base or only in dried flowers often more or less spreading, ovate or nearly so, mostly obtuse or rounded, rarely ciliate, without colleters. Corolla white,paleyellow or salmon, pubescent at thebase of the lobes (rarely not in P. cryptophlebia),with a callous ring at the mouth, glabrous from the mouth to the insertion of the stamens or pilose in P. plectaneiifoliaand sometimes in P. cryptophlebia, with a pilose belt below the insertion of the stamens which is more sparse towards the base, not ciliate on lobes; tube mostly narrowly cylindrical, widened around the anthers which is close to the mouth, slightly contracted at the mouth; lobes overlapping to the left in bud, often oblique, ovate, elliptic, oblong or narrowly so, rounded or obtuse, entire, spreading, often recurved later. Stamens mostly with apex about 0.5mm below mouth of corolla tube; filaments very short; anthers ovate or nearly so, entirely fertile, acuminate or apiculate at the apex, cordate at the base. Pistilwith apex mostly about halfway along anthers, sometimes below or above; ovary narrowly ovoid, mostly laterally compressed, usually gradually narrowed into the filiform style, of 2 separate carpels; disk absent; pistil head usually of 4 parts of which the first and the third may be absent, a basal ring, a globe or cylinder, again a similar ring and a bilobed stigmoid apex. Ovules in 2 rows of up to 13 in each carpel. Fruit orange or red, of 2 separate moniliform or torulose follicles which contain each a single series of 1 13drupes; exocarp thin; mesocarp fleshy or pulpy; drupe ellipsoid, laterally compressed, lumpy. Seed one, with a thin smooth testa; endosperm mealy, not ruminate, Wageningen Agricultural University Papers 97-2 (1997) 53

52 surrounding the embryo; embryo straight, spathulate; cotyledons elliptic, rounded at the apex and at the base, about as long as the rootlet. Petchiacounts 8species,oneof which isendemicto SriLanka, 5 are endemic to Madagascar, one is endemic to Madagascar and the Comoro IslandsandonetoCameroun. Notes. When monographing CabucalaPichon the present author compared it with Alyxia from which it had been separated by Pichon. His conclusion could be corroborated by the field studies carried out for this monograph. The small often shrub-like trees of Petchia (= Cabucala, as is explained below) have the model of Prévost as the author observed in the field in plants of P. erythrocarpa, P. madagascariensis and P. montana and in the herbarium of P. plectaneiifolia. The tree top and the apex of each branch ramify inasmany elements asthey haveleavesinmostcases. An element is a branchlet or an inflorescence. Therefore when inflorescences are accompanied by branchlets they are axillary. When awhorl of branchlets (and inflorescences) isfully developed, a vigourous shoot starting from alateral bud just below this whorl will overtop it an form the continuation of the tree or the branches as is usual in the model of Prévost. Alyxia may have the model of Troll, as has been observed by the present author in passing by in the field, and should be verified. Livera (1926) has described Petchia,which he founded on Alyxia ceylanica. Pichon could hardly distinguish his Cabucala (1948) from Petchia. Pichon gave a poor key (1949) to distinguish Cabucala from Petchia in which the only characters were the ratioof the sepals and the shape of the pistil head. The sepals range from 1.2 to4.5 xaslong as wide and notfrom 1.5 to 2 xin Cabucala andthey are ca 2 x in Petchia and do not range from 3 to 3.5 x. The pistil head mayhave abasal veil,present in Cabucala and absent in Petchia (Pichon, 1949). The far richer collections available at the moment show that both possibilities are present in Petchia ceylanica and in several of the Madagascan species also this character was no longer distinctive. F.J.H. van Dilst (pers. comm.) observed in a single specimen of Landolphia mytrifolia, a Madagascan species, has the basal veil onthe pistil head in a mature bud,while in open flowers it is lacking. The open flowers of Leeuwenberg et al of P. erythrocarpa (preserved in liquid preservative) have the four 54 Wageningen Agricultural University Papers 97-2 (1997)

53 parts of the pistil head conform the generic description of Petchia (this paper), and has a far less developed veil in the bud. Therefore the shape of the pistil head only indicates the age of the flower and therefore haslesstaxonomieimportance than wasoften supposed.as analyses carried out again on Petchia ceylanica showed that it is so closely allied to Cabucala erytrocarpa in all characters, that the present author is obliged to reduce Cabucala to a synonym of Petchia. Fortunately thepresent monograph exclusive of P. ceylanica was not yet published when it was discovered that Petchiaand Cabucala were distinguished on too weak characters to keep them separate. Keytothespeciesof Petchia 1. Leaves abruptly narrowed at the apex into a linear tail; sepals narrowly ovate, acuminate.cameroun 1. P. africana Leaves not abruptly narrowed if caudate; sepals ovate, obtuse or rounded.madagascar; or acuminate, Sri Lanka 2 2. Corolla tube 5 5.5mm long. Mountains 5. P. humbertii Corolla tube mm long.lowland or mountains 3 3. Leaves subcordate or rounded at the base, shortly petiolate, rounded to very shortly acuminate with a blunt acumen at the apex 3. P. cryptophlebia Leaves cuneate, or in some blades rounded at the base, never subcordate short- or long-petiolate, mostly acuminate at the apex 4 4. Corolla tube 6.5 9mmlong(9-10mm in 2.P. ceylanica) 5 Corolla tube (11 )13 25 mm long 6 5. Leaves small, 3 6cm long, long-acuminate 8. P. plectaneiifolia Leaves mostly more than 8cm long, if smaller, rounded or obtuse 6. P. madagascariensis 6. Montane shrub (alt m); leaves cm long, obtuse or slightly acuminate; inflorescence few-flowered 7. P. montana Lowland shrub or tree (alt m); leaves often larger, long- Wageningen Agricultural University Papers 97-2 (1997) 55

54 acuminate to rounded; inflorescence few or many-flowered 7 7. Leaves mostly herbaceous when fresh and membranaceous when dried, long-acuminate to caudate, if not clearly so, more than 5 x as long as wide and up to 1.2 cm wide; intersections of fruits mostly about x as wide as compartments 4. P. erythrocarpa Leaves coriaceous even when fresh, rounded to acuminate, acumen never long, blade, if 5 x as long as wide, more than 2 cm wide; intersections of fruits x as thick as compartments 6.P. madagascariensis Notes. The flowers of the various species of Petchia are very similar. The only more or less useful key character is the length of the corolla tube. P. madagascariensis is very variable and closely allied to the less variable P. erythrocarpa. Specimens of which the leaves are up to 1.2 cm wide and more than 5 x as long as wide and not (as is the rule in the species) long-acuminate to caudate are considered here to belong to P. erythrocarpa. Specimens with large leaves rounded or obtuse at the apex and mostly cuneate at the base (at least in some blades of a specimen) are placed here in P. madagascariensis, while those with subcordate or rounded otherwise very similar leaves are placed here in P. cryptophlebia. The inconsequent annotations of both Pichon (1948) and Markgraf (1970, 1972) show that it is very difficult to distinguish most of the Petchia species from each other. P. ceylanica resembles P. erythrocarpa by the leaves, but is less variable in this character. The corolla is white in P. ceylanica and 9 10mm long and pale yellow or salmon and mm long in P. erythrocarpa Petchia africana Leeuwenb., sp. no v. Fig. 8, p. 57 Frutex slivae orae. Folia opposita anguste elliptica vel ovata apice abrupte attenuata cauda lineari utroque latere glabra. Inflorescentiae terminales pauciflorae pedunculis perbrevibus pedicellis multo brevioribus. Sepala viridia angustissime ovata longe acuminata utroque latere glabra eglandulosa. Corolla pallide flava extus glabra annulo oris calloso glabra intus infra insertionem staminum pilosa; tubus anguste cylindraceus fauce circa antheras 56 Wageningen Agricultural University Papers 97-2 (1997)

55 Fig. 8. Petchia africana. 1,habit (x2/3); 2,flower (x 2); 3,opened flower (x 6); 4, fruit (x 2/3); 5, drupe on section (x 4); 6, embryo (x 4). 1 3 from W.de Wilde & B. de Wilde-Duyfjes 2901; 3-6 from W. de Wilde & B. de Wilde-Duyfjes Wageningen Agricultural University Papers 97-2 (1997) 57

56 dilatatus; lobi oblique angusteque elliptici apice rotundati integri patentes.staminainclusaantherisangusteovatisapiceacuminatisbasi cordatis omnino fertilibus glabris. Pistillum glabrum ovario anguste ovoideo carpellis duobus separatis stylo filiformi unico gradatim angustato;caput pistilli basi ellipsoidea stigmaticaetapice stigmoidea parva bilobata. Fructus aurantiaci folliculi duo separati lineares torulosi basi stipitati apice caudati drupas circiter decern continentes. Paries drupae glebosa dura. Endospermium farinosum embryonem cingens. Embryo rectus spathulatus cotyledonibus ellipticis radicula aequilongis. Typus: Cameroun, Centre-Sud, about 8 km N of Kribi, flowering, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyjjes2901 (holotypewag). Shrub m high, with white latex, entirely glabrous except for the corolla tube inside. Branches green. Leaves opposite; petiole 6 10mm long; blade papery when dried, narrowly elliptic or narrowly ovate, x as long as wide, x cm, abruptly caudate at the apex, with a linear tail 10 20mm long, cuneate at the base, with pairs of rather straight secondary veins forming various angles with the costa; tertiary venation reticulate. Inflorescences terminal, 1 4 together, ca 2x 2cm, few-flowered. Peduncle very short, mm long; pedicels 4mm long. Bracts like the sepals and about 0.2 x as long as them. Flowers: Sepals green, narrowly to very narrowly ovate, 4 7 x as long as wide, x 0.5mm, long-acuminate, persistent even under the ripe fruit. Corollapale yellow, with a pilose belt below the insertion of the stamens 5mm wide;tube about 4x as long as the calyx, x as long as the lobes, 13 15mm long, 2mm wide at the base, 2.5mm wide around the anthers, narrowed at the mouth to 2 mm wide; lobes elliptic, x as long as the tube, twice as long as wide, 6x3 mm, rounded at the apex, spreading. Stamens with apex 1 2mm below mouth of corolla tube, inserted 0.7 of the length of the corolla tube (at 9 10mm from the base); filaments 0.5mm long; anthers narrowly ovate, 5 x as long as wide, 3 x 0.6mm. Pistilwith apex about 2mm below base of anthers (in tube of 15 mm long 7.5mm long); ovary narrowly ovoid, laterally compressed, 2 x 1 x 0.7mm, gradually narrowed into the filiform style; pistil head of a basal stigmatic ellipsoid part 1 x 0.3mm and a bilobed stigmoid 58 Wageningen Agricultural University Papers 97-2 (1997)

57 apex 0.2 x 0.2 mm. Fruit, follicles orange, linear, 27 x 1 cm, torulose to almost moniliform, with stipe 5.5cm long and a tail 4cm long, with 9 11 drupes. Drupes ellipsoid, 16 x 5 6x 4mm, acute at both ends; wall 2mm thick; embryo 12mm long; cotyledons 6 x 2.5 mm; rootlet 6 x 0.7 mm. Distribution: Only known from the the two unicates collected onthesamelocality onthesameday. Ecology: Understorey of coastal rain forest. Alt. 2m. Paratype: Cameroun, ibid., fruiting, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 2902 (WAG). Notes. This species is the first of Petchia from the African continent. The genus was only known from Sri Lanka, Madagascar and the Comoro Islands until these specimens from Cameroun were recognized asbelonging to Petchia. Although the collectors found the specimens already more than thirty years ago,on 8August 1964,the material travelled through the herbarium until the author could assign it to the correct genus.the fruiting specimen was erroneously filed under Malouetia and the flowering one under Hunteria. They turned out not to belong there only when the monographers, respectively J. van der Ploeg (1984) and E. Omino-Achola (1995) discovered that they did not belong there.the conclusive remarkson the taxonomie position could only be made now almost all Apocynaceae represented inafrica have been monographed Petchia ceylanica (Wight) Livera in Ann. Roy. Bot. Gard. Peradeniya 10: 140, pi. 1 (1926). Type: Sri Lanka, sin. loc., Walkers.n. in herb. Wights.n. (holotypek). Basionym: Alyxia ceylanicawight, Icon. PI. Ind. Or. 4 (2): 2, t (1848). Shrub 2.50 m high. Branches pale brown. Leavesin whorlsof 3, shortly petiolate; petiole 1 3 mm long; blade membranaceous or papery when dried, ovate or elliptic or narrowly so, x as long as wide, x cm, with 5 7 pairs of rather straight secondary veins forming an angle of withthe costa; tertiary venation minutely reticulate, rather conspicuous or invisible. Wageningen Agricultural University Papers 97-2 (1997) 59

58 Inflorescenceterminal, 1 6-flowered, 2 4x 2 4 cm. Peduncle 0 14mm long; pedicels thin, 5 12mm long. Bracts like the sepals and about half as long as them. Flowers: Sepals ovate, ca 2 x as long as wide, x mm, long-acuminate, not ciliate. Corolla white, with a narrowly ovoid head in the mature bud about of the bud length, pilose belt below the insertion of the stamens 1 mm wide;tube 5 6 x as long as the calyx, xas long as the lobes, 9 10mm long, ca 1mm wide at the base, widened around anthers; lobes narrowly ovate, 6 x 3mm, acuminate, spreading. Stamenswith apex 0.5mm below mouth of corolla tube; filaments ca 0.5mm long; anthers 1 x 0.3mm. Pistil with apex about halfway along the anthers; ovary 1 x 0.3 x 0.3mm, gradually narrowed into the style; pistil head of 3 parts, the first of which may be absent, a basal ring ca 0.2 x 0.4mm, a globe or cylinder ca 4 x 4mm and a stigmoid apex ca 0.3 x 0.2mm. Fruit: follicles orange or bright red, moniliform, of 1 5 ellipsoid parts x 4 9x 3 8mm, wrinkled when dried; drupe ellipsoid, slightly smaller than the parts of the follicle; wall 0.3mm thick. Seed one, medium brown, elliptic; embryo 6mm long; cotyledons 3x3 mm; rootlet 4 x 0.7 mm. Distribution: Endemic to Sri Lanka. Ecology: Forest understorey. Alt m. Specimensexamined: Sri Lanka. Badagama F.R., Kurunegala District, Faden 76/404 (AAU, K); ibid., Sumithraarachchi 660 (K); Bibile, Kostermans25299A (L); Kaduwela, Balasubramankun BSM-4 (WAG); Mt Lavinia. near Colombo, Rechinger 2294 (W); Wega For., Colombo District, Dassanayake 492 (AAU, K); Seeduwa, Amandoluwa, Cramer (L); Kelaniya R., opposite Kitugalle, Kostermans (L);Henaratgoda (=Gampaha), Petch 6 Mar (A); ibid., Worthington 7094 (K); Kalatawewa Catchment, Worthington 3520 (K); near Matara, Worthington 4163 (K); Karawita Kanda, Ratnapura Distri, Waas18 (K); Wasse Kadoerae, Oltmans 87 (L);sin. loc, Champion anno 1851 (K); Thwaites CP 1835 (K,W); Walker s.n. in herb. Wight s.n. (K, type) Petchia cryptophlebia (Bak.) Leeuwenb., comb. nov. Type: Central Madagascar, probably Province Toamasina, sin. loc, Baron1790 (holotype K;isotypes BM, P). Fig. 9, p. 61; map7, p.62 Basionym and homotypic synonyms: Carissa cryptophlebia Bak. in Journ. Linn. Soc. 20: 204 (1883). Jasminonerium crypto- 60 Wageningen Agricultural University Papers 97-2 (1997)

59 -. SouA4 Fig Petchia ayptophlebia. 1,habit (x2/3); 2, flowerbud (x 2); 3, young fruit (x 2/3); 4, fruit (x 2/3); 5 & 6, endocarp two sides (x 1.5) P. montana. 7, habit (x 2/3); 8, flower bud (x 2). 1-2 from Viguier & Humbert 852; 3 from Broin 26 Nov. 1969;4-5 from Perrier de la Bâthie 8925; 6from Humbert 3191; 7from PerrierdelaBâthie Wageningen Agricultural University Papers 97-2 (1997) 61

60 km V AA(DS) -15 S Il/î Mt -20» \- 0 TV-A -25" V 4 y.j.y 2)- i; I C o A \ic / \VyT(FD) ZÜOkm 45 I 50"E Map 7. Petchia cryptophlebia; k P. humbertii. Map8. Petchia erythrocarpa. phlebium (Bak.) O. Kuntze, Rev. 415 (1891).Cabucala cryptophlebia (Bak.) Pichon in Not. Syst. ed. Humbert 13: 205 (1948); Markgraf, Fl. Mad. fam. 169: 68, pi. 10, 9-13, map 17 (1976). Shrub or small tree, 3 8 m high, with white latex. Trunk 5 25 cm in diameter or probably more; bark pale grey-brown, smooth or hardly fissured, with large protruding lenticels. Branches brown. Leaves in whorls of 3 5 or opposite, shortly petiolate; petiole 1 3( 8) mm long; blade coriaceous even when fresh, elliptic or ovate, x as long as wide, 3 16 x cm, rounded to very shortly apiculate at the apex and then with a blunt acumen, subcordate or rounded at the base, with 7 20 pairs of rather straight secondary veins forming an angle of (55 )70 90 with the costa; tertiary venation inconspiculous. Inflorescences terminal, 1 3 together, few- or many-flowered. 3 8x 3 6 cm, 62 Wageningen Agricultural University Papers 97-2 (1997)

61 rather lax. Peduncle 3 45 mm long; pedicels 2 6mm long. Bracts sepal-like and 0.2 lx as long as them. Flowers: Sepals medium green, ovate or broadly ovate, x as long as wide, x mm, not ciliate. Corolla white, with a narrowly ovoid head in the mature bud about 0.25 of the bud length, mostly pubescent at the base of the lobes, glabrous or sometimes pubescent from the mouth to the insertion of the stamens, pilose belt below the insertion of the stamens 4 10 mm wide; tube x as long as the calyx, x as long as the lobes, mm long, just above the base mm wide, slightly narrowed above or not to mm wide, mm wide around the anthers; lobes narrowly ovate, x as long as the tube, x as long as wide, 6 10 x mm, rounded or obtuse, spreading. Stamens with apex 0 1 mm below mouth of corolla tube, inserted of the length of the corolla tube (at mm from the base); filaments mm long; anthers x as long as wide, x mm. Pistil with apex 0 lmm above base of anthers; ovary x 1.3 x 0.8 lmm, gradually narrowed into the style; pistil head of a cylinder 1 x 0.5 mm, which is usually widened at the apex and at the base to 0.6 mm wide, and a stigmoid apex, x mm. Fruit: follicles with 2 13 drupes, 3 4 mm thick between drupes; parts at least 10 x 5 mm. Distribution: Endemic to Madagascar. Ecology: Rain forest. Alt m. Geographicalselectionofthe43specimensexamined: Madagascar.Toamasina: Bemanditra, Razafindradora 53 (P);Anjozorobe, Moramanga, Malcomber et al (BR, P, TAN, WAG); Ambatovy, E of Ampitambe, Capuron SF28390 (K,P,TEF);AnalamazaotraForest, SofPerinet, Leeuwenberg (TAN, WAG); ibid., Perrier de la Bâthie 8925 (P); ibid., Capuron SF (K, P, TEF, WAG); ibid., Viguier & Humbert 852 (K, P, TAN,WAG).Antananarivo: Andrangoloaka, Hildebrandt 3679 (BM,G,K,P,W, Z); km 27 Moramanga-Anosibe Road, Broin 26 Nov (B, P); Ambatotsipihina, Ambatolampy, SF 1002 (P, TAN, TEF). Fianarantsoa: Ambohimitombe, Sof NosyVarika, Forsyth Major 324 (K);Ranomafana Res., NEof Fianarantsoa, Leeuwenberg et al (BR, K, P, TAN, WAG);ibid., Schatzetal.1706 (BR,C,K,TAN, WAG);km50Fianarantsoa-Mananjary Road, across Namorana R., Nicoll208 (K, P, TAN, WAG); E of Tsaratanteraka, Anivorano Canton, Fort-Carnot (= ikongo) District, SF (P); between IvohibeandVondrozo, Capuron SF23549 (K, P,TEF,WAG).Sin. loc., Baron 853 (P),1790 (BM,K,P,type),3234 (BM,K,P),5765 (K), 6018 (K). Wageningen Agricultural University Papers 97-2 (1997) 63

62 4. 4. Petchia erythrocarpa (Vatke) Leeuwenb., comb. nov. Type: Madagascar, Antsiranana, Nosy Komba, Hildebrandt 3232 (holotype BREM; isotypes K, L, P, W). Fig. 10, p. 65; map 8, p. 62 Basionym and homotypic synonyms: Ellertonia madagascariensis Radlk. in Abh. Naturw. Ver. Bremen: 8: 402 (1884). Alyxia erythrocarpa Vatke in Abh. Naturw. Ver. Bremen 9: 124 (1885). Gynopogon erythrocarpus (Vatke) K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4, 2: 151 (1895). Cabucala madagascariensis var. latifolia Pichon in Not. Syst. ed Humbert 13: 203 (1948); Cabucala erythrocarpa (Vatke) Markgr. in Adansonia II, 10: 513: (1970); Fl. Mad. fam. 169: 84, map 11 (1976). Heterotypic synonyms: Alyxia lucida Bak. in Journ. Linn. Soc. 22: 503 (1887). Type: Central Madagascar, sin. loc, Baron 4663 (lectotype K, designated here; isolectotypes BM, E, P). Cabucala madagascariensis var. intermedia Pichon, I.e. C. erythrocarpa var. intermedia (Pichon) Markgr., I.e.; op. cit. 86, map 11, syn. nov. Type: Central Madagascar, sin. loc., Baron 4551 (holotype P; isotype K, paratype of A. lucida). C. madagascariensis var. angustifolia Pichon, I.e. C. erythrocarpa var. angustifolia (Pichon) Markgr., I.e.; op. cit. 87, map 11, syn. nov. Type: Madagascar, Mahajanga, Ambodina, Ikopa R., Perrier de la Bâthie 18 (holotype P). C. monarthron Pichon, I.e.; Markgraf, op. cit. 78, map 13, syn. nov. Type: Madagascar, Antananarivo, Manankazo, NE of Ankazobe, Perrier de la Bâthie 8883 (holotype P). C. striolata Pichon, I.e.; Markgraf, op. cit. 80, map 13, syn. nov. Type: Madagascar, Toamasina ("Centre West"), Upper Bemarivo R., Perrier de la Bâthie 8839 (holotype P). C. caudata Markgr. in op. cit. 512; op. cit. 82, map 12, syn. nov. Type: Madagascar, Est, Fianarantsoa, Manakara, Ifaho- Ambila, Debray 533 (holotype P; isotype B). C. oblongo-ovata Markgr. in Adansonia II, 14: 107, t.l, f.l (1974); op. cit. 83, map 12, syn. nov. Type: Madagascar, Fianarantsoa, near Vohipeno, Debray 1256 (holotype P; isotype B). Small tree m high, with white latex in all parts. Trunk 1 10cm in diameter; bark pale brown, shallowly and longitudinally fissured or smooth, with large lenticels. Branches pale brown. Leaves in whorls of 3 5, shortly petiolate; petiole Wageningen Agricultural University Papers 97-2 (1997)

63 Fig. 10. Petchia erythrocarpa. 1,habit (x 2/3); 2, flowerbud (x 4/3); 3,calyxwith pistil (x 4/3); 4,opened corolla (x 4/3); 5, stamen (x 10/3); 6, pistil head (x 10/3). Wageningen Agricultural University Papers 97-2 (1997) 65

64 ( 8) mm long; blade herbaceous or subcoriaceous when fresh, membranaceous or papery and often striolate when dried, especially when young, very variable in shape and size, ovate or elliptic or narrowly or even very narrowly so, x as long as wide, 2 13 x cm, caudate or acuminate at the apex with a blunt acumen, cuneate or rounded at the base, with 5 15 pairs of rather straight often obscure secondary veins forming an angle of with the costa; tertiary venation minutely reticulate, rather conspicuous or invisble. Inflorescence terminal, 1 13-flowered, 2 4x 2 4 cm. Peduncle 0 12mm long; pedicels thin, 5 15 mm long. Bracts sepal-like and about half as long as them. Flowers: Sepals pale or medium green, ovate, x as long as wide, x 0.8 lmm, obtuse, not ciliate. Corollasalmon or pale yellow, with a narrowly ovoid head in the mature bud about of the bud length, pilose belt below the insertion of the stamens mm wide; tube 8 18 x as long as the calyx, x as long as the lobes, 13 23mm long, mm wide at the base, slightly narrowed above or not to mm wide, 2mm wide around the anthers; lobes narrowly ovate to oblong, x as long as the tube, x as long as wide, 6 16 x mm, rounded or obtuse, spreading and recurved later. Stamenswith apex 0.5mm below mouth of corolla tube, inserted of the length of the corolla tube (at mm from the base); filaments 0.5mm long; anthers 2 3 x as long as wide, x mm. Pistilwith apex from 0.5mm above base of anthers to 0.5mm above mouth of corolla tube; ovary 2 3 x 0.8 lx mm, gradually narrowed into the style; pistil head of 4 parts, the first or the third may be absent, a basal ring x mm, a globe or cylinder x mm, again a ring x mm and a stigmoid apex x mm. Fruit: follicles orange or bright red, moniliform, of 1 9 ellipsoid parts, smooth when fresh, 8 18 x 4 9x 3 8 mm, wrinkled when dried; drupe ellipsoid, up to 17 x 6x4 mm, with an acute edge; wall 0.3mm thick. Seed one, medium brown, elliptic; embryo up to 8.5mm long; cotyledons 4.5 x 2.7mm; rootlet 4 x 0.7 mm. Distribution: Endemicto Madagascar. Ecology: Forest understorey or bush. Alt m. 66 Wageningen Agricultural University Papers 97-2 (1997)

65 Geographical selectionoftheapproximately275specimensexamined: Comoro Islands. Grand Comore: La Gille Forest, near Maoueni, Bernardi (G, K, L, P, Z); ibid., Doutrelepont 1235 (BR, WAG).Moheli: Fomboni, Rakotozafy 1114 (B, P, TAN), 1114bis (TAN); Njoumbadjou Forest, SF (K, P, TEF), (P, TEF); ibid., Jacquemin 859 (B, P). Anjouan: sin. loc., Kirk Aug (K); Lavanchie s.n. (P). Mayotte: sin. loc., Boivin 3202 (G, P); Dapani Forest, Floret 1091 (P), 1092 (P), 1106 (P);Dzoumogné, Pobéguin 60 (K, P,WAG);Ouchoungui Forest, SF (P, TEF). Madagascar. Mahajanga: Bekopaka, SF (K, P, TEF, WAG); ibid., Mijamo, Rakotozafy 1000 (TAN); ibid., Jacquemin 753 (B, P); near Tsiandro, Leandri & Capuron2346 (K, P, WAG); Bemaraha Plateau, near Tsiandro, CapuronSF 6750 (P, TEF); near Ambódiriana, E of Antsalova, Leandri & Saboureau 2606 (P,WAG),2606bis (K, P); Tsingy of Bemaraha, Leandri 123 bis (BM, G, P); Antsalova, Harmelin RN (P); Eof Antsalova, near Bemaraha Reserve, Labat et al (P); Nof Bemonto Forest, SF 39-R-44 (TEF);Makara Forest,nearAmbalabongo, Debray 1512 (B, P); AnalalavaForest,nearMaintirano, SF (K, P, TEF, WAG);near Marofenobe, Decary 22% (BM, P);ibid., SF (=90-R-44) (P, TEF); Tampotratsa,km 180Antananarivo-Mahajanga Road, Boiteau 3005 (P,WAG);Ambibaka, Andriba Canton, Kasambo SF 12-R-317 (P, TEF); Ambodina, Ikopa R., Perrier de la Bâthie 18 (P, type of C. madagascariensis var. angustifolia); Ankara Mts,Maevatanana District, Descoings 3370 (TAN);near Maevatanana, Morat 882 (P, TAN); ibid., Perrier de la Bâthie 18bis (P),8881 (P);Mafivato, SF4283 (P, TAN, TEF);SitampikyPlateau, Decary7756 (P);Bekarafa, Eof Bekodoka, Decary 8125 (P);nearBekodoka, Decary 221A (P), 8262 (L, P); Besalampy, Decary (K, P), (K, P, WAG); ibid., Ankaboka, SF (K, P, TEF, WAG);Namoroka Reserve, Morat 834 (TAN); ibid., RN 4606 (P, TAN); Anjiajia, Bosser 3267 (P, TAN); Ampijoroa, F.R., Ankaiafantsite, Leeuwenberg (TAN,WAG); ibid., Nicoll371 (K, P,TAN, WAG), 372 (K, P,TAN, WAG);ibid., Phillipson 1917 (BR, K, P,TAN,WAG); ibid., Schatz & Lowry 1438 (BR, K, P,TAN, WAG);ibid., SF 7949 (K, P, TEF, WAG); Ambalabongo, Ambalajakakomby, Peltier 5983 (P); Sainte Marie de Marovoay, Kaudern 7 Sept (S), 20 June 1912 (S); 23 km N of km 19 Mahajanga-Tsaramandroso Road, towards Anjohibe, Leeuwenberg & Rapanarivo (BR, MO,P,TAN, WAG);Besisika, Port Berge, Rabesaona SF65-R-142 (TEF); between Sofia and Mandritsara Rs., Decary (BM, C, P); ibid., Humbert (G, K, P, TAN); km 21 Befandriana Nord-Antsohihy Road, Leeuwenberg & Rapanarivo (BR,MO, P, TAN,WAG);Mahajanga, Decary (P); Borivatra, Bosser 3527 (TAN); 8 km W of Mandritsara, Schofield 4 (K); Befandriana-Nord, Decary (P); Bora Forest, Antsohihy, Rabevohitra SF29558 (TEF); Ankaizina Mts, Perrier de la Bâthie 8944 (P);Andranosamontana, Decary 2233 (P). Antsiranana: Diégo-Suarez, SF 3560 (HBG, K, P, WAG); Mt d'ambre, near Station des Roussettes, Leeuwenberg et al (BR, MO, P, TAN, WAG); ibid., Capuron SF (G, HBG, K, P, TEF, WAG); Beanamalao,SF (= 164-R-416) (TEF);SEofMarotaolana, Debray 1627 (B, P); Ankarana Mts, Humbert32675 (P, WAG); ibid., Leeuwenberg et al (BR, MO, P, TAN, WAG), (BR, MO, P, TAN, WAG); Nosy Komba, Boivin July 1850 (P); ibid., Hildebrandt 3232 (BREM, K, L, P, W, type); ibid., Mahabo, SF3150 (P, TAN, TEF);Nosy Be, Hildebrandt2949 (BM,G,K, P, W); ibid., Lokobe Reserve, Birkinshaw 60 (BR, P, TAN, WAG); ibid., Boivin 2075 (P); ibid., Deroin & Badré 165 (K,P,WAG); Nof LokobeReserve, Leeuwenberg et al (BR, MO, P,TAN, WAG);Analabe Forest, Sof Vohemar, Capuron SF (K, P, TEF); Anjanaharibe Mt, Cours 3881 (P, TAN); Ambilobe, Wageningen Agricultural University Papers 97-2 (1997) 67

66 Waterlot 330 (P); Sambirano R. valley, 7 km after Ambanja, Imbert 125 (K, P, WAG); Manongarivo Mts, base of Beholosy Ridge, Leeuwenberg et al (BR, MO, P,TAN, WAG); Sambirano R. basin, near Beangona, Humbert (G, K, P, TAN); Tsaratanana Mts, Decary (L, P); Ambodivoara Andranomatavy, Sambava Canton, SF (P, TEF);Sambava, Morat 2813 (P, TAN);ibid., Zamani Vato RN 8826 (K, P,WAG);Bemarivo R., of NE, Perrierde la Bâthie 8896 (P); Lohanantsahabe, between Sambava and Andapa, Capuron SF ter (P, TEF); near Andapa, Lokoho R. basin, Humbert & Capuron (K, P, WAG); Fivondronana of Andapa, Ravelonarivo 70 (WAG); Antanandavahely Escarpment, Antalaha, Rakotozafy 506 (TAN). Toamasina: Masoala Peninsula, near Andranobe, Van Nek2066 (WAG); Upper Bemarivo R. (Centre West), Perrier de la Bâthie 8839 (P,type ofc. striolata), 8842 (P); île Ste Marie, NE of Lokintsy, Capuron SF (K, P, TEF, WAG); Kapiloza R. source,ambongo, Perrier de la Bâthie (P); Tampolo, Rabevohitra SF32004 (TEF), (TEF), (TEF); Ambodimanga, Sahatavy, Ramarokoto RN 9481 (K, P, paratype of C. oblongo-ovata); Ambatosoratra Mt, Cours 3257 (P, TAN);ManakaEst, Rakotovao RN (P);BetamponaReserve,nearTamatave, RN 2617 (P, TAN); ibid., Razanaparany RN 8882 (K, P, WAG); Anosibe, Moramanga District, SF2201 (P,paratypeof C. oblongo-ovata); nearankarahara, near Mangoro R. valley, Capuron SF (K, P, TEF, WAG); W of Ambohidray, upper Mangoro R., Capuron SF (K, P, TEF, WAG); Ambatovola, Perrier de la Bâthie18360 (P); N of Ambodimanga, Mangoro R., Capuron SF (P, TEF). Antananarivo: Tsiroamandidy, Boiteau 3026 (P); Ankazobe Mts,NWof Manarinerina, Capuron SF (G, K, P, TEF, WAG); Manankazo, NE of Ankazobe, Perrier de la Bâthie 8883 (P, type of C. monarthron); ibid., SF (P, TEF); km 130 Antananarivo-Mahajanga Road, Bosser 1849 (TAN); 3 km NW of Ambohitsaratelo-Bebao, NW of Tsiroamandidy, Dorr et al (MO,TAN, WAG); Sahagoma, Bevato, SF2-R- 286 (P);Bongolava, Wof Tsiroamandidy, Morat 4744 (P),4579 (P, TAN);Itasy Lake,Anonym. 4 (P).Fianarantsoa: Itremo, Guillaumet4218 (K, P,TAN,WAG); Mananjary, Geay7722 (P); ibid., SF (= 184-R-496) (TEF); Ranomafana Reserve,NEof Fianarantsoa, Leeuwenberg et al (BR, K, P,TAN,WAG); Andrambovato Forest, E of Fianarantsoa, Capuron SF (K, P, TEF); Ambalavao,IngaroMts,SF (P);Ifaha,Ambila, Nof Manakara, Debray533 (B, P, type of C. caudatà);ese of Ambila, Debray1249 (P); near Vohipeno, Debray 1256 (B, P); near Lokomby, Manakara, Capuron SF (K, P, TEF, WAG); Vondrozo-Ivohibe Road, Broin 138 (P), 139 (P); Matitanana R. basin, Perrier de la Bâthie 8928 (P); near Vondrozo, Boiteau 2123 (P, WAG); Anohambovato, SF (P, TEF); Manombo Forest, Boiteau 2119 (K, P, WAG); ibid., Capuron SF (K, P, TEF, WAG); Midongy du Sud, Decary 4967 (P). Toliara: km 17 Miandrivazo-Mandoto Road, Rakotozafy 1620 (TAN); Amboasary, Nof Ankazobe, Decary 7390 (P);Miandrivazo, Decary (P, S); 15km Nof Fort-Dauphin, Den Outer & Van Veenendaal 1224 (WAG);Analalava Forest, Manantenina, Dumetz1366 (P). Sin. Loc, Baron 654 (K, P), 2525 (K), 4551 (K, P, type of C. m. var. intermedia and paratype of Alyxia lucidd), 4663 (BM, E, K, P, lectotype of A. lucida), 4888 (BM, K, P, paratype of A. lucidd), 5916 (K, P),6084 (K),6541 (K,P). Notes. The earlier epitheton madagascariensis Radlk. (1884) is not available, as the even earlier homonym, Alyxia madagascariensis A. DC. (1844) is used for another species. 68 Wageningen Agricultural University Papers 97-2 (1997)

67 P. erythrocarpa, themost widespread speciesof Petchia, has a great variation in the shape and size of the leaves, but it is rather constant in the flowers. The leaves are smooth as in the type of Cabucala oblongo-ovata orstriolateasinthe typesof C. caudata and C. striolata, or both smooth and striolate within one branch, e.g. Humbert They are clearly acuminate to caudate when upto 5 x as long as wide, as in the types of C. madagascariensis var. intermedia, C. monarthron, C. striolata, C. caudataand C. oblongoovata. The follicles are quite variable in length. They may contain 1 13drupesandtherefore theonly character, asingle drupe ineach follicle, onwhich C. monarthron hasbeenbased,lostitsvalue Petchia humbertii (Markgr.) Leeuwenb., comb. nov. Type: Madagascar, Antsiranana, eastern slopes of Marojejy Mts, W of Manantenina R., tributary of Lokoho R., Humbert (holotype P). Fig. 11, p. 70;map 7, p. 62 Basionym: Cabucala humbertii Markgr. in Adansonia II, 10: 511 (1970;Fl. Mad.fam. 169: 77,pi. 11, 3-4, map14 (1976). Shrub. Branches pale brown. Leavesopposite or in whorlsof 3 at apices of branchlets; petiole 1 5 mm long; blade coriaceous when dried, elliptic, x as long as wide, 1 4x cm, obtuse or rounded at the apex, cuneate at the base, with revolute margin in dried leaves, with 4 8 pairs of rather straight obscure secondary veins forming anangleof45 80 withthe costa; tertiary venation invisible. Inflorescence terminal, few-flowered, 1.5 x cm, lax. Peduncle 10 13mm long; pedicels 5 7mm long. Bracts leafy andmuch smaller than theleaves or sepallike andabout half as long asthem. Flowers: Sepals ovate, x as long as wide, x mm, obtuse, not ciliate. Corolla pale yellow, with anovoid obtuse head inthemature budaboutone third ofthebudlength, pilose belt below theinsertion ofthe stamens 1mmwide; tube x as long as the calyx, x as long as the lobes, 5 5.5mm long, mm wide at the base, mm wide around the anthers; lobes ovate or oblong, 0.6 x as long as the tube, x as long as wide, x 1.2mm, rounded, entire, spreading. Stamenswith apex at mouth of corolla tube, inserted 0.7 of the length of the corolla tube (at 3.5 4mm Wageningen Agricultural University Papers 97-2 (1997) 69

68 Fig , Petchiaplectaneiifolia. 1, habit (x 2/3); 2, flower bud (x 3) P.humbertii. 3, habit (x 2/3); 4, flower bud (x 2) P. madagascariensis. 5 & 11, habit (x2/3); 6 & 12, flowerbuds (x 2); 7 & 13,youngfruits (x2/3); 8,immaturefruit (x 2/3); 9 & 14,fruits (x2/3). 10, endocarp two sides (x 1.5). 1-2 from RN 8173; 3-4 from Humbert 22595; 5-6 from Boiteau 2610; 7-8 from Capuron SF 23936; 9-10 from Decary 9768; 11 from Debray 781; 12-13from Bernier 106;14from Perrier delabâthie Wageningen Agricultural University Papers 97-2 (1997)

69 from the base); filaments 0.5 mm long; anthers 1.5 x 0.4 mm. Pistil with apex just below base of anthers; ovary 1.2 x 0.5 x 0.4 mm, gradually narrowed into the style; pistil head of a basal ring 0.2 x 0.4 mm, a globe 0.4 x 0.4 mm and a stigmoid apex about 0.2 x 0.2 mm. Fruit unknown. Distribution: Only known from four collections three of which were made at the same locality. Ecology: Montane forest. Alt m. Specimensexamined: Madagascar. Antsiranana: Lokoho R. valley, Mt Beondroka, N of Maroambihy, Humbert (B,P);Marojejy Res., Rasoavimbahoaka et al.23 (P,TAN,WAG); easternslopesofmarojejy Mts, WofMananteninaR.,tributary of Lokoho R., Humbert (P,type).Toamasina: Foulpointe,Mahatsara,SF (=1600-R-631) (TEF) Petchia madagascariensis (A. DC.) Leeuwenb., comb, nov. Type: Madagascar, North of Madagascar, Richard{7) in Herbier Boissier ex herbario Musaei Parisiensis anno 1843 (holotype G; isotypes G-DC, L and ). Figs. 11, p. 70 and 12, p. 72; phots 5, p. 123; 6, p. 123 and 7, p. 124; map 9, p. 73 Basionym and homotypic synonyms: Alyxia madagascariensis A. DC, Prod. 8: 345 (1844). Pulassarium madagascariense (A. DC.) O. Kuntze, Rev. 2: 417 (1891). Gynopogon madagascariense (A. DC.) K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4, 2: 151 (1895). Cabucala madagascariensis (A. DC.) Pichon in Not. Syst. ed. Humbert 13: 203 (1948); Markgraf, Fl. Mad. fam. 169: 74, pi. 11, 11-14, map 15 (1976) as var. madagascariensis. C. madagascariensis var. latifolia Pichon, I.e., excl. of syn. Ellertonia madagascariensis and its type which belongs to P. erythrocarpa. Heterotypic synonyms: A. polysperma Sc. Elliot in Journ. Linn. Soc. 29: 33 (1891). C. polysperma (Sc. Elliot) Pichon in op. cit. 204; Markgraf, op. cit. 76, pi. 11, 5 10, map 14, syn. nov. Type: Madagascar, Toliara, Fort-Dauphin, Scott Elliot 231A (holotype K; isotype P). C. torulosa Pichon, I.e.; Markgraf, op. cit. 72, pi. 10, 5-8, map 16, syn. nov. Type: Madagascar, Antsiranana, Sambirano, near Ambanja, Perrier de la Bâthie 8855 (holotype P). Wageningen Agricultural University Papers 97-2 (1997) 71

70 Fig. 12. Petchia madagascariensis. 1, habit (x 2/3); 2, flower bud x 4/3); 3,calyx with pistil x 4/3); 4, opened corolla (x 4/3); 5, stamen (x 10/3); 6, pistil head (x 10/3); 7,immaturefruit (x 2/3). 72 Wageningen Agricultural University Papers 97-2 (1997)

71 a to 0-15 S < 100km S M / ^ Ck/ /LA IDS) atoi 0-15 S < 10ûkm ^»/2 ria(ds) oj 1/ ' V Jrt>* t<h\ ML KTTV ^ rr MlL ~\ V oa(t) ^^-y -20 mi- C A S>7-20 c As>a/ é TV-» g- F.Xf M \"' 0 T T^' M F-.F/W M 7. ; Jn\ -25' M^ 'T 1 *-} 'jw 'y- /- 'flijf \QPT(FD) ZOO km -25 X^ ''-/ )- ' 'C (*'/ \_ P/T(FD) km 45" 50"E 1 Map9. Petchia madagascariensis. Map 10. Petchia montana. 45" 1 50"E 1 C. fasciculata Pichon, I.e.; Markgraf, op. cit. 64, pi. 9, 10-14, map 18, syn. nov. Type: Madagascar, Toamasina, Masoala Peninsula, Perrier de la Bâthie 8900 (holotype P). C. glauca Pichon, I.e. Type: Madagascar, Toamasina, île St. Marie, Bernier 106 (holotype P, erroneously cited as type of C. madagascariensis by Markgraf). C. macrophylla Pichon, I.e.;Markgraf, op. cit. 70, pi. 10, 1 2, map 16, as var. macrophylla, syn. nov. Type: Madagascar, Fianarantsoa, Ikongo Mts, Decary5535 (holotype P; isotypes BM,K,TAN). C. intermedia Pichon in op. cit Type: Madagascar, Toamasina, W of Tamatave, Betampona Reserve, near Ambodiriana, Perrier de la Bâthie17488 (holotype P;isotypes K, TAN). C. multiflora Pichon, I.e.; Markgraf, op. cit. 65, pi. 9, 3-7, map 17, syn. nov. Type: Madagascar, Toliara, Belavenoke,WNW Wageningen Agricultural University Papers 97-2 (1997) 73

72 of Fort-Dauphin, Decary (holotype P; isotypes G, K, S, TAN). C. longipes Pichon, I.e. C. madagascariensis var. longipes (Pichon) Markgr. ex Boiteau in Adansonia II, 12: 222 (1972); Markgraf, op. cit. 75, map 15, syn. nov. Type: Madagascar, sin. loc., Humblot 58 (holotype P). C. crassifolia Pichon in op. cit. 206; Markgraf, op. cit. 66, pi. 9, 1 2, map 18, syn. nov. Type: Madagascar, Fianarantsoa, Vondrozo, Decary 5331 (holotype P; isotypes K, TAN). C. macrophylla var. acuta Markgr. in Adansonia II, 10: 511 (1970). C. madagascariensis var. amygdalifolia Markgr. ex Boiteau in Adansonia II, 11: 555 (1971). C. madagascariensis var. acuta (Markgr.) Markgr., FI. Mad. fam. 169: 75, map 15 (1976), syn. nov. Type: Madagascar, Toamasina, Analamaitso Forest, upper Bemarivo R., Perrier de la Bâthie 8840 (holotype P). C. penduliflora Markgr. in Adansonia II, 10: 511 (1970); op. cit. 82, map 13, syn. nov. Type: Madagascar, Toamasina, W of Tamatave, Betampona Reserve, Rakotoniaina RN 4242 (holotype P; isotype K). C. macrophylla var. oxyphylla Markgr. in Adansonia II, 14: 109 (1974); op. cit. 72, map 16, syn. nov. Type: Madagascar, Toliara, near Fort-Dauphin, Manantantely Forest, Humbert 5714 (holotype P; isotypes G, K, S). Shrub or small tree, 1 8( 15) m high, with white latex. Trunk 1 25 cm in diameter; bark pale grey and dark brown, smooth or shallowly and longitudinally and often transversely fissured. Branches pale brown. Leaves in whorls of 3 4 at apices of branchlets, otherwise often opposite; petiole 1 15 mm long; blade coriaceous even when fresh, elliptic or narrowly elliptic, sometimes obovate or orbicular, 1 5x as long as wide, x cm, acuminate to rounded at the apex, cuneate at the base, margin mostly revolute even in fresh leaves, with 5 20 pairs of obscure rather straight secondary veins forming an angle of with the costa; tertiary venation invisible. Inflorescences terminal, often several together, sometimes also axillary, few- to manyflowered, 2 11x2 7 cm, lax or rather lax. Peduncle 3 60mm long; pedicels thin, 2 11 mm long. Bracts sepal-like, x as long as the sepals. Flowers: Sepals light green, ovate or broadly so, x as long as wide, x mm, not or obscurely ciliolate. Corolla white, tube pale green or pale yellow, in 74 Wageningen Agricultural University Papers 97-2 (1997)

73 bud pale yellow, with a narrowly ovoid head in the mature bud about one third of the bud length, pilose belt below the insertion of the stamens mm wide; tube x as long as the calyx, x as long as the lobes, (6.5 )8 25 mm long, 1 2 mm wide at the base, around the anthers mm wide; lobes ovate, narrowly ovate or elliptic, x as long as the tube, x as long as wide, x mm, obtuse or rounded, spreading, often recurved later. Stamens with apex 0 3 mm below mouth of corolla tube, inserted of the length of the corolla tube (at mm from the base); filaments mm long; anthers x as long as wide, x mm. Pistil with apex from 2 mm below to 1 mm above base of anthers; ovary x x 0.6 lmm, gradually narrowed into the style; pistil head of 4 parts, the third of which may be absent, a basal ring 0.1 x mm, a globe or cylinder x mm, again a ring x mm and a stigmoid apex 0.2 x 0.2 mm. Fruit: follicles orange, with 1 13 drupes; intersections x as wide as the compartments; compartments 6 16 x 5 6x 3 4 mm. Distribution: Endemic to Madagascar. Ecology: Forest, mostly near the coast. Alt m. Geographicalselectionoftheapproximately175specimensexamined: Madagascar.Antsiranana:Sambirano, Perrier de la Bâthie 8855 (P,typeof C. torulosa);manongarivoreserve, EofAkaramy, Malcomber2325 (MO,TAN); ibid., Wsideof Beholosy Ridge, Leeuwenberg et al (BR,MO,P,TAN, WAG);ibid., McPherson & Van der Werf16409 (WAG);ibid.,SEofBeraty,trail to summit of Antsatroto, Malcomber et al. 921 (BR, K, P, TAN, WAG); Maromandia, Rabevohitra SF29504 (TEF);Marojejy Reserve,nearManantenina, Randrianasolo44 (TAN);ibid.,MtBeondrokabase, Miller & Randrianasolo4340 (WAG); Sof Analamanara,near Tsaratanana, Capuron SF (K, P, TEF, WAG); Atiala Forest, Deroin & Badré127 (K, P, WAG); Farimay Forest, Ampahana, SF 8-R-305 (P); Ambolo, Perrier de la Bâthie (P); Ambohitralanana,RN8598 (P);Ambatondradama Pass, Nof MasoalaPeninsula, Capuron SF 8780 bis (P); Ampanavoana, TsilizyRN 5490 (P), 5491 (P). Toamasina: Maroantsetra, Mocquerys 290 (G, Z); E of Sahavary, Lowry et al (BR,K,P,TAN,WAG);NosyMangabe,Antongil Bay, Schatzetal.1676 (K, P, TAN, WAG);Masoala Peninsula, Perrier de la Bâthie 8900 (P, typeof C.fasciculata); ibid., Ambanizana, Lowry 4465 (P, TAN, WAG); S of Ambanizana, Van Nek2212 (WAG); EofAmbanizana, Schatz & Lowry 2833 (K, P,TAN,WAG);AnalamaitsoForest,upperBemarivoR., Perrier de la Bâthie8840 (P, type of C. macrophylla var. acuta); île St. Marie, Bernier 106 (P, typeof C. glauca); Soanierana-Ivongo, Perrier de la Bâthie 8889 (P); Menatany Hill, Capuron SF (K, P, TEF, WAG); Tampolo Forest, Leeuwenberg (WAG);ibid., Capuron SF22116 (K,P,TEF,WAG),23835 (P,TEF);between Volotaraina and Ambodirafia, Cours 1725 (P, TAN); Analalava Forest, Wof Wageningen Agricultural University Papers 97-2 (1997) 75

74 Foulpointe, Capuron SF28076 (P, TEF,WAG),23841 (K, P,TEF,WAG);ibid., Leeuwenberg (BR, P, TAN, WAG); Betampona Reserve, near Ambodiriana,NWofTamatave, Debray781 (P);ibid., Leeuwenberg (TAN, WAG); ibid., Perrier de la Bâthie (K,P, TAN, typeof C. intermedia); ibid., Rakotoniania RN 4242 (K, P, type of C.penduliflora), RN 5923 (K, P, TAN, WAG); Analangavo,Tamatave, SF 1376 (P);between DidyandBrickaville, Cours 4938 (K,P,WAG); Analabe,Marovoay,MoramangaDistrict, SF26193 (P, TEF); near Ambila-Lemaitso, Schatz & Armbuster 3157 (K, P, TAN, WAG); ibid., Schatz et al (P, TAN, WAG); ibid., SF 1132 (P, TAN, TEF); between Ambodimanga and Antanambao, Cours 2789 (P, TAN); Betamotamo Forest, Tsaravinany, Rakotozafy 597 (P, TAN); upper Sahadranamby R., Mangoro R. valley, Perrier de la Bâthie 8920 (P);lower Mangoro R. basin, Perrier de la Bâthie (G, P). Fianarantsoa: Ambodihazo-Androrangavola Road, Ifanadiana, Debray1244 (B, P); Ingaro Mts, W of Ambalavao, Cremers 3603 (P, TAN); Ikongo Mts, Decary 5535 (BM, K, P, TAN, typeof C. macrophylla); FortCarnot (= Ikongo), Decary5819 (BM, P); ibid., Maromiandra, SF 7148 (P, TEF); Anorimbatonamboa, Fort Carnot (= Ikongo), SF (K, P, TEF, WAG); Amboatalanina,Mananjary, RazafySF5630 (K, P,TEF), SF6640 (P, TEF);near Vohitromby-Ifaho, 10kmESEofAmbila, Debray 1251 (B,P); Sihanaka, Boiteau & Cours in Herb. Jard. Bot. Tana (P); Andringitra Reserve, Sendrisoa, Razafindrakoto RN 3492 (P, TAN); Bekatra, Manakara, Capuron SF 6537 (P, TEF); Loharano, SF (P, TEF, WAG); near Vohipeno, Broin 140 (B, P); Ilakatra,SF6426 (P, TEF);km 179Vohipeno-Farafangana Road, Debray 1255 (B, P, TAN); lower Matitanana R. basin, Perrier de la Bâthie 8929 (P); between Vondrozo and Ivohibe, Decary5407 (C, P, paratype of C. macrophylla var. oxyphylla); Vondrozo, Decary 5312 (L, P), 5331 (K, P, TAN, type of C. crassifolia), 5418 (P);Amporoforo Forest,betweenFarafangana andvohipeno, Capuron SF (P, TEF); km 20 Farafangana-Manombo Road, Capuron SF (K, P, TEF, WAG); Manombo Forest, Capuron SF (HBG, K, P, TEF, WAG), (P, TEF), (K, P, TEF, WAG). Toliara: Belavenoke, Decary (G,K, P, S,TAN, typeof C. multiflora);bemangidy,manantenina Canton, Boiteau 2546 (P); ibid., Gueneau SF (= 3-R-32) (P,TEF,WAG); Tsingafiafy Mts, Capuron SF (HBG, K, P, TEF, WAG); Fort-Dauphin, Boiteau 2610 (P);ibid., Decary 9909 (BM,K, P, S,TAN);ibid., Scott Elliot2374 (K, P, type of C. poly sperma); Vatombe, Decary (P, paratype of C. macrophyllavar. oxyphylla), (L, P, paratype of C. macrophyllavar. oxyphylla);ebakika, 50 km N of Fort-Dauphin, McWhriter 214 (K, P); ibid., Capuron SF (P, TEF, paratype of" C. macrophylla var. oxyphylla); Andohahela Reserve, Parcelle 1, Leeuwenberg et al (BR, K, P, TAN, WAG); ibid., Phillipson 2982 (K, P, TAN, WAG); ibid., Ramarokoto RN 3414 (K, P, TAN, WAG), 3427 (P, TAN, WAG); ibid., Tanatana Pass, Decary 9768 (C, L, P); Maningotra, SF 4078 (P, TAN); between Ifarantsy and Ranomafana, Capuron SF 8505 (K, P, TEF, WAG); Petriky Forest, ca 15 km WSW of Fort- Dauphin, Gereau & Dumetz 3253 (BR, K, P, TAN, WAG); ibid., Rabevohitra 2108 (K,P,TAN,TEF,WAG);Manantantely Forest, Humbert 5774 (G, K, P, S, typeof C. mac.var. o.); MandenaReserve, Leeuwenberg et al (BR, K, P, TAN, WAG); ibid., Rabevohitraet al (K, P, TAN, WAG); ibid., Razafindrambao 580 (B, P); ibid., Zarucchi et al (P, TAN, WAG); Wof Mandena, Debray 1752 (P, paratype ofc. macrophylla var. oxyphylla).sin. loc, Richard (?)in Herb.Boissier ex Herb.Mus. Paris anno 1843 (G,G-DC, L, type); Humblot 58 (P, typeof C. longipes), 88 (K, P), 657 (K, P); Lantz 30 Sept (P). 76 Wageningen Agricultural University Papers 97-2 (1997)

75 Notes. Petchia madagascariensis is a very variable species.the characters vary independently. The leaves and inflorescences are very variable in shape and size. The flowers vary in size but much lessinshape. The types of P. madagascariensis and Cabucala glauca have very narrowly elliptic leaves which are acuminate at the apex and have a revolute margin. The leaves of the types of C.fasciculata, C. macrophylla and C. peduliflora are wider and have a less revolute margin and slightly more visible venation. In this respect Leeuwenberg is a perfect intermediate. The type of C. multiflorahas smaller leaves than the specimens mentioned above, which resemble those of the types of C.fasciculata and and C. macrophylla. The type of C. madagascariensis var. acutahas leaves as narrow as those of the types of P. madagascariensis and C. glauca but the venation is more conspicuous. The type of C. torulosa has slightly striolate leaves which resemble those of thetype of C. macrophylla but they are much smaller. In both characters, striolation and size, Capuron SF is a perfect intermediate. As for the size they approach those of the type of C. polysperma of which the leaves are somewhat narrower and rounded at the apex. The type of C. crassifoliahas also small rounded leaves which are wider than those of the type of C. polysperma. The type of C. polysperma has been collected near Fort-Dauphin, probably in the Mandena forest from where the type of C. crassifolia comes. In this forest has been collected Leeuwenberg14203, a rich sample of the population whichhasleaves similar tothoseofthetypesof C. crassifolia, C. polysperma as well as C. torulosa. A perfect intermediate between the types of C. polysperma and C. macrophyllais Capuron SF 28671,especially for theleaves. The inflorescences are few-flowered, as e.g. in Leeuwenberg and 13762, to very many-flowered, as e.g. in Decary10615, thetypeof C. multiflora and Decary The fruits are much more constant in shape and size, as they are usually less indented between the drupes than those of most of thoseof P. erythrocarpa. These observations may show why the author is obliged to reduce somanynamestosynonymsof P. madagascariensis. Wageningen Agricultural University Papers 97-2 (1997) 77

76 4. 7. Petchia montana (Pichon) Leeuwenb., comb. nov. Type: Madagascar, Fianarantsoa, Pic d'ivohibe, Bara, Humbert 3191 (holotype P;isotype G). Fig.9, p. 61; map 10, p.73 Basionym: Cabucala montana Pichon in Not. Syst. ed. Humbert 13: 206 (1948); Markgraf, Fl. Mad. fam. 169: 66, pi. 9, 8-9, map 18 (1976). Shrub 3 m high or tree up to 20 m high. Trunk up to 15 cm in diameter or even more; bark dark brown, rather rough, with protruding concolourous lenticels, with few white latex. Branches pale brown; branchlets subtriangular at apex, with much white latex. Leavesin whorls of three or opposite; petiole 1 4 mm long; blade coriaceous when fresh, subcoriaceous when dried, elliptic, x as long as wide, x cm (up to 5.5 x 3.2cm in sucker shoots),rounded orobtuse atthe apex or slightly acuminate and with ablunt acumen, cuneate orrounded atthe base,with 4 10 pairs of obscure rather straight secondary veins forming an angle of with the costa; tertiary venation invisible. Inflorescences terminal, 1 3 together, 1 7-flowered, lax. Peduncle 3 17 mm long; pedicels 4 12mm long. Bracts sepal-like and about half as long as the sepals. Flowers: Sepals green, ovate, x as long as wide, x mm, obtuse. Corolla white,throat yelloworange, with an ovoid head in the mature bud about one third of the bud length, pilose belt below the insertion of the stamens 3 mm wide; tube 6 15 x as long as the calyx, mm long, 1 2 mm wide at the base, 2 4mm wide around the anthers; lobes ovate, x as long as the tube, 2 3 x as long as wide, 6 12 x 2 5.5mm, rounded, spreading. Stamens with apex 0mm below mouth of corolla tube, inserted of the length of the corolla tube (at 9 20mm from the base); filaments 1 mm long; anthers 3x 1mm. Pistil: ovary 2 4x x mm, gradually narrowed into the style; pistil head of 4 parts, a basal ring 0.2 x 0.8mm, a cylinder 0.3 x 0.3mm, again a cylinder 0.5 x 0.5mm and a stigmoid apex 0.3 x 0.3mm. Fruit :follicles with 3 compartments,only known in immature state. Distribution: Endemic to Madagascar. Ecology: Montane forest. Alt m. Specimensexamined: Madagascar. Antsiranana: Tsaratanana Mts, Pender de la Bâthie (P). 78 Wageningen Agricultural University Papers 97-2 (1997)

77 Fianarantsoa: ca 3 km NE of Antoetra, a village 27 km E of Ivato, Jongkind & Rapanarivo 885 (WAG); ibid., Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); Pic d'lvohibe, Bara, Humbert 3191 (G, P,type);upper Rienana R. valley, Humbert 3614 (G, K, P, TAN). Toliara: Andohahela Mts, Humbert 6200 (P). Sin. loc, Baron 2458 (K, P),5000 (K) Petchia plectaneiifolia (Pichon) Leeuwenb., comb. nov. Type: Madagascar, Fianarantsoa, Ivohibe, Decary 5477 (holotype P; isotypes K, S, TAN). Fig. 11, p. 70; map 11, p. 80 Basionym: Cabucala plecteneiifolia Pichon in Not. Syst. ed. Humbert 13: 206 (1948), as plectaneiaefolia; Markgraf, Fl. Mad. fam. 169: 77, pi. 11, 1-2, map 14 (1976). Shrub or small tree up to 10 m high. Trunk cm in diameter. Branches pale brown. Leaves opposite or in whorls of 3; petiole 1 3 mm long; blade subcoriaceous when dried, narrowly elliptic, 3 5x as long as wide, 3 6x cm, long-acuminate at the apex, cuneate at the base, with 5 10 pairs of rather straight obscure secondary veins forming an angle of with the costa; tertiary venation invisible. Inflorescences terminal, 1 3 together, x cm, few- to many-flowered, rather lax. Peduncle mm long; pedicels 2 4 mm long. Bracts sepal-like, about one third as long as the sepals. Flowers: Sepals ovate or broadly ovate, x as long as wide, x 0.5 lmm, obtuse or rounded, not ciliate. Corolla cream or white and with pale yellow tube, with an ovoid blunt head in the mature bud about one third of the bud length, pilose from the mouth to 2 3 mm below the insertion of the stamens; tube x as long as the calyx, x as long as the lobes, mm long, mm wide at the base, mm wide around the anthers; lobes ovate, x as long as the tube, x as long as wide, x mm, rounded, entire, spreading. Stamens with apex 0 mm below mouth of corolla tube, inserted of the the length of the corolla tube (at mm from the base); filaments 0.5 mm long; anthers x as long as wide, x mm. Pistil with apex near base of anthers; ovary x x mm, gradually narrowed into the style; pistil head of a basal ring 0.2 x mm, a globe 0.3 x 0.3 mm and a stigmoid apex 0.2 x 0.2 mm. Fruit unknown. Distribution: Endemic to southeastern Madagascar. Wageningen Agricultural University Papers 97-2 (1997) 79

78 AA(DS) AAIOS) t o 0-15 S 100km % / \V t-f>^>\ M /"Î < km -15 S U B / Î f V/ 1 J^d ML j^mji >0<7 /1 \ /^-V T ML ST, V^oAlT) _ -^ /T -20 3L OA ^>N/ -20 ƒ L A N 9~- o F;* ^ ( SÔ TWI -25 X\ fa L: / y l~ : * ƒ \jyt(fd) km P. % -25 * ( / 1 J.^ ^N ( 1 WyTfFD) 0 M km 45 I 50 E E 1 Map 11. A Petchia plectaneiifolia; Gonioma malagassy. Map 12. A Plectaneia longisepala; P. stenophylla. Ecology: Dense forest, in the mountains. Alt. up to 1700 m. Specimensexamined: Madagascar. Fianarantsoa: Ivohibe, Decary 5387 (FHO, L, P), 5438 (L, P),5477 (BM,K,P,S,TAN,type);Ankidonolava,Vondrozo, Boiteau 2125 (K, P, WAG). Toliara: Kalambatitra Mts, Humbert (C, G, K, P, WAG); Behara, RakotosonRN8173 (P,TAN). Species excluded from Cabucala Cabucala brachyantha Pichon in Not. Syst. ed. Humbert 13: 205 (1948); Markgraf, Fl. Mad. fam. 169: 73, pi. 10, 3-4, map 17 (1976). Type: Madagascar, Toamasina, Andragovalo Mts, SE of Lake Alaotra, Onibe R. basin, Zakamena Reserve, Humbert & Cours (holotype P). = Carissa campenonii (Drake) Palacky. 80 Wageningen Agricultural University Papers 97-2 (1997)

79 5. Plectaneia Thou., Gen. Nov. Madag. 11 (1806); Markgraf, Fl. Madag.fam. 169: 114 (1976). Twining lianas with white latex in all parts. Branches usually dark brown to black and with cream lenticels, smooth; branchlets terete. Leavesopposite, those of a pair equal, petiolate, inserted on conspicuous leaf cushions; ocreae widened into large intrapetiolar stipules; blade mostly pale green and subcoriaceous or coriaceous even when fresh, ovate or elliptic or narrowly so. Inflorescence terminal and often at the same time axillary and then mostly only in the axils of the upper leaves, usually shorter than the leaves and dense. Flowers small, mm long in the mature bud. Sepals probably always pale green, connate at the base, erect or in dried flowers often spreading, equal or unequal, mostly ovate or narrowly so, sometimes broadly triangular, acute or obtuse, glabrous or pubescent outside, often ciliolate, glabrous or with some hairs inside, without colleters. Corolla white, cream or yellow, glabrous or partly puberulous outside, glabrous inside on basal mm, with a hirto-pubescent belt with recurved hairs just above and mostly directly below the the insertion of the stamens, glabrous from the insertion of the stamens to the mouth and puberulous or glabrous on the lobes;tube almost cylindrical or less often funnel-shaped, mostly contracted below the insertion of the stamens, widened around them and again contracted at the throat, not twisted, with 5 thickenings in the throat just above the anthers; lobes ovate to very narrowly so, obtuse or rounded, mostly twisted in bud, often slightly auriculate at the left side of the base, suberect. Stamens included, mostly barely so; filaments very short, mm long, filiform, glabrous; anthers ovate or broadly so, acuminate or less often obtuse at the apex, cordate at the base, glabrous. Pistil glabrous, with apex mostly just below base of anthers;ovary ovoid,usually laterally compressed, 2-celled; style short, rather thick; pistil head or stigma ovoid. Ovules in two rows of 5 8 in each cell. Fruit mostly dark brown, often with whitish indumentum, a 2-celled dry follicle, acuminate to caudate at the apex, rounded at the base, glabrous or pubescent, with 4 mostly undulate wings, septicidally dehiscent, ca seeded, subtended by the persistent calyx; the two valves envelop the seeds. Seedflat, oblong, winged at both ends, glabrous; grain dark brown, obliquely elliptic or rectangular; wings light brown, obliquely rounded; funiculus long and filiform; hilum small, in the middle of one side; endosperm rather hard, thin, diaphanous, surrounding the Wageningen Agricultural University Papers 97-2 (1997) 81

80 embryo; embryo spathulate, straight; cotyledons ovate or narrowly so, rounded at the apex and at the base; rootlet about x as longascotyledons. Thegenuscounts 3species,allendemictoMadagascar. Keytothespeciesof Plectaneia 1. Leaves rather large and with ca 30 rather straight secondary veins,6 11 cm long;sepals 6 xas long as wide, 3 x 0.5 mm. East Madagascar 1. P. longisepala Leaves mostly smaller, with 5 20 pairs of obscure or anastomosingsecondary veins;sepalsup to 4 x as long as wide 2 2. Leaves 5 8x as long as wide, obtuse or rounded at the apex; corolla glabrous outside. South Madagascar 2. P. stenophylla Leaves, if more than 5 x as long as wide, acuminate at the apex and corolla mostly puberulous outside 1. P. thouarsii Plectaneia longisepala Markgr. in Adansonia II, 12: 585 (1972); Fl. Madag. Fam. 169: 122, pi. 17, (1976). - Type: Madagascar, Toamasina, near Ambila-Lemaitso, Decary 6547 (holotype P). Fig. 13, p. 83;map 12, p.80 Liana. Branches pale brown, with concolourous lenticels; branchlets glabrous. Leaves: petiole 6 10mm long, glabrous; blade coriaceous when dried, elliptic, x as long as wide, 6 11 x 2.5 8cm, apiculate at the apex with a blunt acumen, cuneate or rounded at the base, glabrous on both sides, with ca 30 pairs of rather straight secondary veins, (3 5x 1 1.2cm and with inconspicuous venation on lateral branchlets bearing flowers), tertiary venation invisible. Inflorescenceat least 4x2 cm, lax at least in first branchings. Peduncle at least 2cm long, glabrous; pedicels 1 3 mm long or more, glabrous. Bracts leafy or sepallike. Flowers: Sepals pale green, connate at the base for 0.2mm, equal, very narrowly ovate, 3 x mm, acute, glabrous on both sides. Corolla (in bud only), 2 3mm long, glabrous or sparsely puberulous outside, glabrous inside (probably too young to 82 Wageningen Agricultural University Papers 97-2 (1997)

81 ,}. s -"-»>"U"<-- ^ut/u/ Fig. 13. Plectaneia longisepala. 1, habit (x 1.6); 2, flower bud (x 12). 1-2 from Decary6547. Wageningen Agricultural University Papers 97-2 (1997) 83

82 have developed any indumentum); tube 0.5 mm long; lobes narrowly ovate, 1.5 x 0.7 mm, obtuse. Stamens barely included; anthers 0.5 x 0.3 mm. Fruit immature. Distribution: Only collected twice. Ecology: Coastal forest on sand. Alt m. Secondspecimenknown: Madagascar.Toamasina,Analalava Forest,near Foulpointe, Leeuwenberg &Rapanarivo (BR,MO, P,TAN, WAG) Plectaneia stenophylla Jum. in Ann. Mus. Colon. Marseille V, 2: 27 (1934); Markgraf, Fl. Mad. fam. 169: 126, pi. 18, 5-7 (1976). Type: Madagascar, Toliara, Upper Fiheranona R. valley, Perrier de la Bâthie (lectotype P, designated by Markgraf; isolectotype WAG). Fig. 14, p. 85; map 12, p. 80 Small liana, 2 3 m high climbing or more. Branches dark brown, with whitish lenticels; branchlets glabrous. Leaves: petiole glabrous, 1 3 mm long; blade dark green above, pale green beneath, brittle and papery when dried, very narrowly elliptic or ovate, 5 8 x as long as wide, (often wider in leaves near bases of branchlets), 1 5x 0.2 lcm, obtuse or rounded at the apex, cuneate or rounded at the base, glabrous on both sides, venation invisible. Inflorescence terminal and often at the same time in the axils of the upper leaves, x 1 2 cm, many-flowered, rather dense. Peduncle 1 2 mm long, glabrous; pedicels 1 3 mm long, glabrous. Bracts sepal-like and x as long as them. Flowers: Sepals pale green(?), connate at the base for 0.2 mm, ovate or narrowly ovate, x as long as wide, x mm, obtuse or acute, glabrous on both sides, ciliolate. Corolla yellow or greenish-yellow, mm long in the mature bud, glabrous outside, glabrous inside at basal 1 mm, hairy belt mm wide, otherwise glabrous; tube x as long as the longest sepal, x as long as the lobes, mm long, almost cylindrical and mm wide, widened around the stamens to mm wide, contracted at the throat; lobes very narrowly ovate, x as long as the tube, x as long as wide, x mm, obtuse, twisted in bud. Stamens with apex mm below mouth of corolla tube, inserted of the length of the corolla tube, (at mm from the base); anthers 0.5 x mm. Pistil, mm long; ovary laterally compressed, 84 Wageningen Agricultural University Papers 97-2 (1997)

83 I // J- Sa.<JJ> J ia\\~t..a >nt**3 ) Fig. 14. Plectaneia stenophylla. 1,habit (x 1); 2,flower (x 8); 3,openedcorolla (x 10); 4,fruit (x 1).1-2from Humbert 12945;3-4from Bosser9170. Wageningen Agricultural University Papers 97-2 (1997) 85

84 0.8 1 x x mm; style mm long; pistil head or stigma x 0.2 mm. Ovules in rows of ca 7. Fruit dark brown, 6 13 x 0.7 lcm, glabrous; wings more or less undulate. Seed 15 x 3 4 mm; grain almost square, 5x3 4 mm. Distribution: Endemic to southern Madagascar. Ecology: Dry open forest or bush. Alt m (or less). Geographicalselectionoftheapproximately40specimensexamined: Madagascar. Toliara: near left bank Fiherenana R., 32 km from Tulear, Leeuwenberg & Rapanarivo (BR, K, MO, P, TAN, UPS,WAG);upper FiherenanaR.valley, Humbert 5090 (P,WAG);ibid., Perrier de la Bâthie (P, WAG, lectotype); Ankazoabo, Bosser17359 (P, TAN, WAG); Sakaraha, Bosser 9170 (K, P, TAN, WAG); ibid., Lambomakondro, Decary (P), (P);Andranohinaly, SF (P, TEF, WAG);23-43km E of Tulear, alongroadtoandranovory, Croat (K,P,TAN,WAG);30km SofSakaraha, Moral3510 (P,TAN);Sakaraha-Tongobory Path, Morat2451 (P,TAN);km 45Itampolo-EjedaRoad, Liedeetal.2718 (MO, P);Isalo,nearFanjahira, Humbert 2790 (G, P); Itambono Corridor, 15 km W of junction with Nat. Road 10, Phillipson & Rabesihanaka 3128 (BR, K, P, TAN, WAG); lower Onilahy R. basin,nearbetioky, Humbert & Swingle 5257 (P);BezaMahafaly Reserve, near Betioky, Phillipson 1829 (BR,C,K, P,TAN,WAG); 16kmSWofAmpanihy, Labatetal.2071 (K,P);OnilahyR.,nearBenenitra, Perrier de la Bâthie (P, WAG); nearbeloha, Croat (C,K,WAG);ibid. Decary 4Sept (GB, P), 6Sept (P);betweenBekilyandTsivory, Seyrig 106bis (P);Ambovombe, Decary 8489 (BM, K, P, TAN, US), 8586 (P, S); SW of Ambovombe, Kotoala, Decary 9098 (NY, P, UC); Mandrare R. valley, near Ambovombe, Humbert & Swingle 5627 (G,P,US); Antanimora, Decary 2933 (C,G,P),9556 (BM, P, US);Manambolo R. valley, near Isomono, Humbert (G, K, P, TAN, US); SW of Ifotaka, Lam & Meeuse 5479 (L, P); Berenty Reserve, Phillipson 2308 (BR, K, P, TAN, WAG); Manambolo R. valley, NW of Maroamby, Humbert (P, WAG) Plectaneia thouarsii Roem. & Schult, Syst. Veg. 4: 420 (1819); Markgraf, Fl. Madag. Farn. 169: 129, pl. 19, 1-6 (1976). Type: Madagascar, sin. loc, Du Petit-Thouars s.n. (holotype P). Fig. 15, p. 87; phot. 8, p. 124; map 13, p. 88 Homotypic synonym: P. volubilis Poir. in Lamarck, Tabl. Enc. 2: 306 (1819). Heterotypic synonyms: P. hildebrandtii K. Schum. in Engler & Prantl, Nat. Pflanzenf. 4 (2): 144 (1895); Markgraf, op. cit. 118, pl. 17, 1 5, syn. nov. Type: Madagascar, Antsiranana, Pasandava (= Ampasindava) Bay, Kisimani, Hildebrandt 3012 (holotype Bf; lectotype K, designated here; isotypes BREM, G, M, P, W). 86 Wageningen Agricultural University Papers 97-2 (1997)

85 Fig. 15. Plecteneia thouarsii. 1, habit (x 1); 2, intrapetiolar stipules (x 12); 3, flower (x 12); 4, sepal inside (x27); 5,corolla (x 12); 6, openedcorolla (x 12); 7, pistil (x 12); 8, fruit (x 1). 1-2 from Leeuwenberg 14186;3-7 from Leeuwenberg 14177; 8from Humbert Wageningen Agricultural University Papers 97-2 (1997) 87

86 h M km -15 S C^ Y* 7^~ VÄ~-r s^ -20 / ^. % Vn JIIF«M /^4 ^l'w' -?^!jk^s^/ ( jk/ä -^! '1 V* ƒ (XA(DS) ^M ^ \ wh f* 'T fi to < km -I5-S -20 f~^~^-_ U M S"i AA(DS) ôfry^v /-»À! /A % ^ JM'» ;' O/M ^fês^f' ' / l *-/^r^ ^1/ -2 5 =vy^ jmrt'fd' km ^ -25 X^y I C Cl* 'T(FD) km E 1 Map 13. Plectaneia thouarsii. Map 14. A Stephanostegia capuronii; S. hildebrandtii. 45 I 50 E 1??P. pervillei K.Schum., 1. c. Type: Madagascar, Pervillé 509 (holotype not traced). P. rhomboidalis Jum. & Perr. in Le Caoutchouc et la Gutta-Percha 15 Février 1908: 9 (1908); in Ann. Mus. Colon. Marseille II, 6: 46, t. 1 (1908); Markgraf, op. cit. 121, pi. 17, 6-9, syn. nov. Type: Madagascar, Mahajanga, Ankara Plateau, Bemaimbo, Perrier de la Bâthie 8939 (holotype P; isotype K). P. hildebrandtii f. hirsuta Jum. & Perr. in Ann. Mus. Colon. Marseille II, 6: 46 (1908); Markgraf, op. cit. 120, syn. nov. Type: Madagascar, Mahajanga, Menavava R. valley, Perrier de la Bâthie 1130 (holotype P; isotype WAG). P. inutilis Jum. & Perr. in op. cit. 49, f. 2-5 and 6B (1908). P. elastica var. inutilis (Jum. & Perr.) Markgr., op. cit. 134, pi. 20, 6 7, syn. nov. Type: Madagascar, Mahajanga, Boina, surroundings of Ampanihy, near Ampasimentera, Perrier de la Bâthie 8943 (holotype P; isotype K). 88 Wageningen Agricultural University Papers 97-2 (1997)

87 P. inutilis f. latiflora Jum. & Perr. in op. cit. 56. Type: Madagascar, Mahajanga, Upper Bemarivo R., Perrier de Bâthie Nov (holotype P). P. elastica Jum. & Perr. in op. cit. 56, t. 2, f. 6A and 7 9; Markgraf, op. cit. 132, pi. 20, 1 5. Type: Madagascar, Mahajanga, Upper Bemarivo R. basin, Analamahitso Forest, Perrier de la Bâthie 2 (lectotype P, designated here; isolectotype K). P. elastica f. firingalavensis Jum. & Perr. in Ann. Mus. Colon. Marseille II, 6: 64, fig. 1 (1908). P. firingalavensis (Jum. & Perr.) Jum. in Ann. Mus. Colon. Marseille V, 2: 18 (1934); Markgraf, op. cit. 127, pi. 18, 8-9, syn. nov. Type: Madagascar, Mahajanga, Firingalava Forest, Ikopa R., between Maevatanana and Andriba, Perrier de la Bâthie 351 (holotype P; isotypes K, WAG). P. microphylla Jum. & Perr. in Agric. Prat. Pays Chauds 10: 189 (1910); Markgraf, op. cit. 139, pi. 19, 14-17, syn. nov. Type: Madagascar, Antsiranana, Sambirano R. valley, Perrier de la Bâthie (holotype P). P. boivinii Jum. in Ann. Mus. Colon. Marseille V, 2: 22 (1934), as boivinï, Markgraf, op. cit. 122, pi. 18, 1 2., syn. nov. Type: Madagascar, Mahajanga, Upper Isandrano R., tributary of Ikopa R. left bank, Perrier de la Bâthie 8940 (holotype P). P. inutilis var. hirsuta Jum. in op. cit. 36; P. elastica var. inutilis f. hirsuta (Jum.) Markgr. in Adansonia II, 12: 221 (1972) and op. cit. 136, syn. nov. Type: Madagascar, Antsiranana, Ambohipiraka, Perrier de la Bâthie (lectotype P, designated here, cited as holotype by Markgraf (1976)). P. isalensis Jum. in op. cit. 40; Markgraf, op. cit. 137, pi. 19, 9 13, syn. nov. Type: Madagascar, Fianarantsoa, Isalo, Perrier de la Bâthie (lectotype P, designated here, cited as holotype by Markgraf). P. isalensis f. glomerata Jum. in op. cit. 42. Type: Madagascar, Fianarantsoa, W of Fianarantsoa, Perrier de Bâthie (lectotype P, designated here). P. microphylla var. tsaratanensis Jum. in. op. cit. 44; Markgraf, op. cit. 129, pi. 20, P. tsaratanensis (Jum. & Perr.) Pichon in Mém. Mus. natn. Hist. Nat. II, 27: 194 (1949("1948")), syn. nov. Type: Madagascar, Antsiranana, Mt Tsaratanana, Perrier de la Bâthie (lectotype P, designated here; erroneously cited as holotype by Markgraf). Wageningen Agricultural University Papers 97-2 (1997) 89

88 P. macrocarpa Jum. in op. cit. 46. P. thouarsii var. macrocarpa (Jum.) Markgr. in Adansonia II, 12: 222 (1972) and op. cit. 130, pi. 19, 7-8, syn.nov. Type: Madagascar, Fianarantsoa, Lower Matitana R. basin, Perrier de Bâthie 8197 (lectotype P,designated here,cited asholotype by Markgraf). P.lanceolata Pichon in Not. Syst. ed. Humbert 13: 207 (1948). P.firingalavensis var. Umceolata (Pichon) Markgr. in Adansonia II, 12: 221 (1972) and op. cit. 129,pi. 18, 10-11, syn. nov. Type: Madagascar, Toliara, Miandrivazo District, Dabolava, Decary (holotype P;isotypeK). P. breviloba Markgr., I.e. and op. cit. 124, pi. 18, 3 4, syn. nov. Type: Madagascar, Toamasina, Rahobevava, Cours 4276 (holotype P;isotypeTAN). P. firingalavensis f. setulosamarkgr., I.e.; P.firingalavensis var. firingalavensis f. setulosa (Markgr.) Markgr., op. cit. 128, syn. nov. Type: Madagascar, Toliara, Zombitsy Forest, NEof Sakaraha, Leandri & Rakoto 3895 (holotype P). P. elastica var. insularis Markgr., I.e. and op. cit. 137, pi. 20, 8 9, syn. nov. Type: Madagascar, Antsiranana, Nosy Be, Pervillé703 (holotype P;isotypeK). Liana climbing 1 10 m high, 5 20 m long. Trunk 1 6cm in diameter; bark dark grey or dark brown to black, smooth or rough and longitudinally and often at the same time transversely fissured, mostly with cream lenticels. Branches dark brown, with cream lenticels; branchlets glabrous or sometimes only near apex puberulous or pubescent. Leaves: petiole glabrous or sometimes pubescent, 1 9mm long; blade herbaceous to coriaceous when fresh, membranaceous to coriaceous when dried, ovate, elliptic or narrowly so, variable in shape and size, x as long as wide, x cm, acuminate or less often obtuse at the apex, rounded or cuneate at the base, often suborbicular and rounded or retuse at the apex in leaves at the base of the branchlets, glabrous or rarely pubescent on both sides, sometimes with scattered black dots beneath, with 5 20 pairs of secondary veins which are mostly inconspicuous; tertiary venation mostly invisible, otherwise reticulate. Inflorescence paniculate, dense, especially in last branchings, very variable in size, x 1 6 cm, mostly many-flowered. Peduncle 0 5cm long, glabrous, pubescent or puberulous; pedicels 0.5 5mm long, glabrous, pubescent or puberulous. Bracts sepallike and x as long as the sepals. Flowers: Sepals pale green, 90 Wageningen Agricultural University Papers 97-2 (1997)

89 connate at the base for of their length, x as long as wide, 1 3 x mm or forming a more or less tubular calyx x 1.5 mm with lobes unequal, ovate to trianglar or broadly or narrowly so, x as long as wide, x mm, obtuse to acuminate, pubescent, glabrous or sparsely puberulous outside, ciliolate, glabrous or partly puberulous inside, without colleters. Corolla white, cream or pale yellow, tube often greenish or pale green, ( 12) mm long in the mature bud, puberulous or sparsely so outside, less often entirely glabrous, glabrous inside on basal mm, hairy belt mm wide, entirely glabrous from there or from above the insertion of the stamens to the mouth and puberulous or sometimes sparsely so on the lobes; tube x as long as the longest sepal, x as long as the lobes, 2 4 mm long, obconical or almost cylindrical, mm wide above the base, often slightly contracted below the insertion of the stamens, widened around them to mm wide; lobes ovate or narrowly so to even strap-shaped, ( 5)x as long as the tube, 1 6( 12)x as long as wide, 1 8 ( 10) x mm, obtuse or rounded, mostly suberect, twisted in bud. Stamens with apex mm below mouth of corolla tube, inserted of the length of the corolla tube, (at mm from the base); anthers x as long as wide, x mm. Pistil mm long; ovary laterally compressed, 0.6 lx x mm; style mm long; pistil head or stigma x mm. Ovules in rows of 5 8. Fruit dark brown or less often pale grey-brown, 7 37 x cm, glabrous or pubescent, wings mostly undulate. Seed x 3 8 mm; grain obliquely oblong, 5 15 x 3 8 mm; wings equal or unequal; embryo 4 13 mm long; cotyledons 2 10 x 1 4 mm; rootlet x mm. Distribution: Endemic to Madagascar. Ecology: Light, mostly rather dry forest or woodland. Alt m. Geographicalselectionoftheapproximately340specimensexamined: Madagascar. Mahajanga: Betsiriry, Herb. Jard. Bot Tana (TAN); AntsalovaDistrict, RamarokotoRN (P,WAG); 3km NofAndrananidahy, 15km ESE of Antsalova, Viliiers & Badré 5021 (MO, P); near Ambodiriana, Leandri & Saboureau2667 (K, P,WAG);AnkaraPlateau,Bemaimbo, Parier de la Bâthie8939 (K, P, type of P. rhomboidalis);mahatsinjo, Perrier de la Bâthie (P);upperIsandranoR., tributary of IkopaR., Perrier de la Bâthie 8940 (P, typeof P. boivinii);firingalava Forest,IkopaR. rightbank,between Maevatanana Wageningen Agricultural University Papers 97-2 (1997) 91

90 and Andriba, Perrier de la Bâthie 351 (K, P, WAG, type of P. elastica f. firingalavensis), 351 bis (P); Ankara Mts, Decary (K, P, TAN); Amboromalandy, Bosser8457 (P, TAN);Menavava R. valley, Perrier de la Bâthie 1130 (P, WAG, type of P. hildebrandtii f. hirsuta); Namoroka Reserve, Decary (K, P, WAG); Andranomavo, RakotovaoRN 5328 (TAN); Madirovalo, Perrier de la Bâthie8194 (P); Betsiboka R. valley, near Mananika, Humbert & Perrier de la Bâthie 2132 (P);Boina, Perrier de la Bâthie 2351 (P);Ankarafantsika Reserve, Perrier de la Bâthie8837 (P), A (L, P), B (P); ibid., SF 2269 (P, TAN); ibid., Ampijoroa, Boiteau 1030 (P, WAG); ibid., Dorr & Koenders 2956 (K, P,TAN,WAG);upper BemarivoR., Perrier de la Bâthie Nov (P,type of P. inutilisf. latiflora); AnalamaitsoForest, Perrier de la Bâthie 2 (K, P,lectotype of P. elastica),8941 (P), 1 Aug (K),Dec (P);Boina, surroundings of Ampanihy, nearampasimentera, Perrier de la Bâthie 8943 (K, P, typeof P. inutilis); nearmarovoay, Humbert & Perrier de la Bâthie 2356 (A, B,G, K, P, TAN), 8850 (P), (P), (P), A (P), B (P), (P, WAG), (P); Ste. Marie de Marovoay, Kaudern 17 June 1912 (S); Mahavavy R., Perrier de la Bâthie Sept (G);near Bombetoka, Perrier de la Bâthie June 1908 (P);near Mahajanga, Perrier de la Bâthie May 1908 (P), (P, WAG); N of Mahajanga, Barnett et al. 481 (K, MO, P, TAN, WAG); ibid., Dorr 3827 (BR, K, P,TAN, US,WAG);Mahajamba, Bemarivo R., Perrier de la Bâthie 8938 (P);lower Sofia R., Decary (P, TAN);Befandriana, Rauh 912 (TAN); ibid., Ambilomavo, Herb.Jard. Bot. Tana (TAN); 4 km Wof Port Berge, Leeuwenberg & Rapanarivo (BR, MO, P,TAN, WAG); 1km Eof Ambalafamainty, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); 10 km N of km 7 Befandriana Nord-Antsohihy, 1 km S of Andranova, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG);km 17Ankarika- Analalava, Leeuwenberg & Rapanarivo (BR, MO, P,TAN, WAG);km 33 of sameroad, Leeuwenberg & Rapanarivo (BR,MO, P,TAN,WAG);Bora Mts, betweenantsohihy and Antsahabe, SF31610 (= 1157-R-187) (TEF);30km E of Antsohihy, Gentry11544 (K, P, TAN, WAG); km 82 Antsohihy-Ambanja, betweenandrafiabe andandranosamonta, Leeuwenberg & Rapanarivo (BR, MO, P, TAN,WAG);Maromandia, Decary 1254 (P), 1258 (BM, P);ibid., Bejofo, Decary2089 (P). Antsiranana: SambiranoR.valley, Perrier de la Bâthie (P), (P, type of P. microphylla); between Akaramikely and Moramandia, Rabevohitra SF (TEF); Manongarivo Mts, McPherson (TAN); Mt Tsaratanana, Perrier de la Bâthie (P, lectotype of P. microphylla var. tsaratanensis), (P, paratype of P. m. var. tsaratanensis); Pasandava (= Ampasindava) Bay, Hildebrandt 3012 (G, K, M, P, W, type of P. hildebrandtii); Nosy Be, Boivin 17 (P); ibid., Pervillé 440 (L, P), 703 (G, K, P, type of P. elastica var. insularis), 768 (P); ibid., Richard269 (FI-W, TCD), 327 (K, TCD), 337 (P), 343 (P,TAN); ibid., Lokobe Reserve, Bernardi (G, K, L, P, Z); Nosy Be, near Antafondro, Bernardi (G, K, L, P, Z); Ankarana Reserve, Harderet al (P, TAN, WAG), 1765 (P, TAN, WAG); ibid., Humbert18965 (G, P), (G, K, P, TAN); ibid., Leeuwenberg et al (BR,MO, P,TAN,WAG), (BR,MO, P,TAN,WAG);Orangea Forest,SE ofranomena, Leeuwenberg et al (TAN,WAG), (BR,MO, P,TAN, WAG);Diego-Suarez, Perrier de la Bâthie (P);Mt des Français, Keraudren 1647 (P, WAG); ibid., Leeuwenberg et al (BR, MO, P, TAN, WAG), (BR, MO, P, TAN, WAG), (BR, MO, P, TAN, WAG); ibid., Poisson 92 (P); Marovato, Anivorano-Nord, Humbert32411 (P, WAG); upper Loky R. basin, Perrier de la Bâthie 8930 (P, paratype of P. inutilis var. hirsuta); Ambilobe, Waterlot 357 (P, paratype of P. i. var. hirsuta), 402 (P, paratype of 92 Wageningen Agricultural University Papers 97-2 (1997)

91 P. i. var. hirsuta); nearambohipiraka, Perrier de la Mthie (P,lectotypeof P. i. var. hirsuta); between Vohemar andambilobe, Decary (P);Vohemar, Boivin 2463 (P); Marojejy Res., Ravelonarivo et al. 225 (WAG). Antananarivo: Manerinerina, Perrier de la Bâthie (P);Manankazo, Bosser7889 (P); ibid., Harizo RN 1206 (P); ibid., Perrier de la Bathie (P); Ambohitantely, Rakotozafy 1284 (TAN); km 21 Ankazobe-Maevatanana, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); km 44 Antananarivo-Mahajanga Road, Bosser (TAN); Ankitsika Forest, Ambohijanaka, Cows 1580 (P, TAN);Ambohimasina, Wof Betafo, Perrier de Bâthie (P);MtIbity, Perrier de la Bâthie13578 (P). Toamasina: Maroantsetra, Mocquerys 497 (P); Masoala Peninsula, ca 11 km S of Ambanizana, Schatz & Modeste 3097 (K, P, TAN, WAG); Soanierana-Ivongo, SF 2314 (P, TAN, TEF), 2357 (P); Vatomandry District, Guillot 45 (G, K); Lake Alaotra, Maningoiy R. Falls, Homolle 555 (P); Antandrokomby, Peltier965 (P, TAN);Ambatondrazaka, Decary (P, TAN); Rahobevava, Cours 4276 (P, TAN, type of P. brevilobd); île Ste. Marie, Boivin 13 (P); Tampolo Forest, Leeuwenberg (BR, P, WAG), (TAN, WAG); Sahatavy, Ramarokoto RN (P); ibid., Laibosaka RN (P); road to Analalava, Wof Foulpointe, Dorr & Barnett 3318 (BR, C, K, NY, P, S, TAN, US, WAG); Analalava Forest, Leeuwenberg (K, P, WAG), (BR,MO, P,TAN,WAG);nearTamatave, Perrier de la Bâthie8876 (P); Tampina, Perrier de la Bâthie (P); near Ambila-Lemaitso, Leeuwenberg & Ranaivojaona (BR, MO, P, TAN, WAG), (BR, MO, P, TAN, WAG); ibid., Lowry & Randrianasolo 4549 (BR, TAN, WAG); ibid., Schatz & Armbuster 3146 (K, P, TAN, WAG);Analamazaotra Forest, Boiteau 493 (K, P, WAG); 11 km E of Permet, Moral 3891 (TAN), 3892 (TAN); Ampitambe, SF (= 1137-R-631) (TEF); near Mahanoro, Perrier de la Bâthie (P), (P).Fianarantsoa: nearambatofindrahana, Cremers 3648 (P,TAN,WAG); ibid., Decary (P), (P); 39 km N of Ambositra, along Nat. Road 7, Croat29465 (P, TAN); nearkianjadrakefona, Leeuwenberg et al (BR, MO, P, TAN, WAG); Nosy-Varika, SF (P, TEF); Mananjary, Geay8146 (P), 8147 (P);Ranomafana Reserve, Leeuwenberg et al (BR, K, TAN,WAG), (BR, K, TAN, WAG); W of Fianarantsoa, Perrier de la Bâthie12877 (P, lectotypeof P. isalensis f. glomerata); Ramiova,Ambalavao, Rakotovao RN7627 (P); Vohimaso, Mahazony, Rakotovao 540 (P); km 14 Ambalavao-Ihosy Road, near Ampitaha, Leeuwenberg & Rapanarivo (BR, MO, P, TAN, WAG); near Sakasoa, along Ihosy-Analalira Road, Allorge & Veyret 596 (P, WAG); Vohibory Mts, Wopf Ivohibe, Humbert 3127 (G, P); Sendrisoa, Rakotoson RN (P,TAN); Pic d'lvohibe, Humbert 3135 (BM, P);Vohipeno, Beaujard 261 (P);Ampasy,Ivongo, Rakotovao 589 (P);upperRienanaR. valley, Humbert 3592 (G,K, P,WAG);lower Matitanana R. valley, Perrier de la Bâthie 8136 (P),8137 (P), 8197 (P, lectotype of P. macrocarpa); Andringitra Reserve, Rakoto RN 1383 (P); WofAndringitraMts, Perrier de la Bâthie (P); Tsiazomborona,Ivohibe, Boiteau 2064 (P);Itempona R., SWof Ihosy, Mabberley 900 (TAN), 904 (TAN); Ankibory, Ambalavao, Rakotovao RN 7592 (P).Toliara: nearambanihazo, Geay 6774 (P); Mandena Forest, Leeuwenberg et al (BR, K, P, TAN, WAG); ibid., Rabevohitra et al (K, P, TAN, WAG); ibid., Zarucchi et al.7495 (F, P, TAN, WAG); Manantantely Forest, Boiteau 3076 (P, WAG); near Fort- Dauphin, Scott Elliot 2289 (K, P), 2584 (K, P);ibid., Pic St. Louis, Leeuwenberg et al (BR, K, TAN, WAG), (BR, K, P, TAN, WAG); Petriky, Rabevohitra et al (K, P,TAN, WAG); Wof Ranopisa, Leeuwenberg (BR, K, P, TAN, WAG); Andohahela Reserve, Parcelle 2, Leeuwenberg et al (BR, K, P,TAN, WAG), (BR, K, P,TAN, WAG), (BR, K, Wageningen Agricultural University Papers 97-2 (1997) 93

92 P, TAN, WAG), (BR, K, P, TAN, WAG); Parcelle 3, Leeuwenberg etal (BR, K, P,TAN, WAG), (BR, K, P,TAN, WAG);upper Mandrare R. basin, between Vavara Pass and Manambolo R. valley, Humbert 6747 (G, P); NE of Vohitsiombe, Decary 4569 (C, P); Imanombo, Bosser (P, TAN); Sakamalio R. valley, tributary of Manambolo R., Humbert (G, K, P); Ampandrandava, between Bekily and Tsivory, Seyrig 106 (P); Mangoky, Andranomiforitra R.valley, Humbert7068bis (P); 20km NofBetroka,alongNat. Road 13, Phillipson etal.3937 (TAN,WAG);Vohipolaka Mt, Humbert (P, TAN); Sakalalina Road, Descoings 3708 (P, TAN); Onilahy R., near Benenitra, Perrier de la Barbie12716 (P), (P); Bereketa, Peltier 2783 (P, TAN); Antanimora, Decary 8868 (BM, G, P, S, US); Tsiombe-Ambondro Road, Croat (TAN); Marovato-Tsiombe Road, Croat (TAN); Mahafaly Plateau, near Beomby, Keraudren 798 (K, P, WAG); Fiherenana R. basin, Perrier de la Bâthie (P); km 56 Sakaraha-Anjamala Road, Chauvet431 (P, WAG); betweentulearandmiary, Bosser (P,TAN,WAG);Sakaraha, Bosser9171 (P, TAN); ibid., Decary (GB, P); Zombitsy Forest, Bosser (P, TAN), (P, TAN); ibid., Croat30672 (P, TAN); Analamarina Forest, Hazoroa R. valley, S of Sakaraha, Humbert (K, P, WAG); Ankazoabo, Bosser (P, TAN, WAG); N of Tulear, near Lake Ihotry, Phillipson 3076 (BR,K, P,TAN,WAG);Morondava, Humbert & Perrier de la Bâthie 2391 (B, P); ibid., Rakotozafy1087 (TAN);Marofandilia Forest,nearMorondava, Perrier de la Bâthie 8874 (P); Belo, Tsiribihina R., Decary15538 (P); km 5 Miandrivazo- Mandoto Road, Rakotozafy 1709 (TAN);MiandrivazoDistrict, Dabolava, Decary (K, P, type of P. lanceolata). Sin. loc.: Baron258 (K, P), 1177 (K, P), 2501 (K, P),2929 (K, P),4022 (K), 4764 (K, P). Notes. The leaves of the type of Plectaneia thouarsii have rather inconspicuous venation as those of the type of P. breviloba. Leaves of the types of P. boivinii and P. lanceolata have conspicuous venation, while in this respect the type of P. macrocarpa and the specimens Leeuwenberg and Schatz & Modeste 3097 are intermediate. The smallest leaves are in the types of P. microphylla (1 2 cm long) and P. breviloba (1 3 cm long) and the largests in the types of P. boivinii (4 8.5 cm long) and P. lanceolata (4 8 cm long). The greatest variation in leaf sizes has been observed in Leeuwenberg (2 4.7 cm long). The flowers of the above cited specimens and those placed under these names by Markgraf are remarkably uniform in most characters. The sepals are acute or obtuse, pubescent or glabrous outside. They are united at the base for mm, which is of their length. Up to the present specimens with sepals pubescent, connate at the base for only about 0.2 mm and acute or obtuse were named P. isalensis, those with sepals united for 0.1 of their length, glabrous and mostly acute P. elastica var. inutilis and those with sepals united for of their length, pubescent or glabrous and mostly obtuse P. thouarsii. As several specimens are intermediate, e.g. Perrier de la Bâthie with sepals pubescent, united 94 Wageningen Agricultural University Papers 97-2 (1997)

93 for of their length and obtuse, and Humbert with sepals glabrous, united for of their length and acute or obtuse, it was not possible to keep up P. elasticavar. inutilis and P. isalensis as separate taxa. Preceding authors also had problems to name the latter species. Markgraf named it as P. thouarsii and Pichon considered it as intermediate between P. elastica and P. thouarsii. When collecting in southern Madagascar the author supposed that P.firingalavensis could be maintained as a distinct species, but after analysis of all material it turned out to grade into the variable P. thouarsii. Flowering specimens withcorollalobes glabrous outsideandat least twice as long as the tube were identified as P. hildebrandtii, although the type specimen of this name is fruiting and cannot be distinguished from fruiting specimens of P. thouarsii to which were assigned only specimens with corollas at least partly hairy outside. Some specimens are intermediate in the caracters of the corolla, e.g. Perrier de la Bâthie16559, the lectotype of P. isalensishas corolla lobes 1.5 x as long as the tube which are glabrous or nearly so outside. In Afzelius 10Sept. 1912the corolla is glabrous outside and the lobes vary from 1.5 to 2 x as long as the tube. The corolla is hairy outside and the lobes are x as long as wide in Pervillé 440and Richard 269. The leaves are usually glabrous, but in some specimens they are pubescent, especially beneath, e.g. in Perrier de la Bâthie 18749, thetype of P. inutilis var. hirsuta and partly pubescent, especially on thecostain Perrier de la Bâthie 8939,thetypeof P. rhomboidalis. 6. Stephanostegia Baill. in Bull. Mens. Soc. Linn. Paris 1: 748 (1888); Markgraf, Fl. Mad. fam. 169: 102 (1976). - Type species: S. hildebrandtii Baill. Trees 4 25 m high, with white latex. Trunk up to 70cm in diameter. Branches lenticellate; branchlets terete, glabrous. Leaves opposite, petiolate; petiole glabrous; blade coriaceous even when fresh, entire, glabrous on both sides, with numerous often obscure secondary veins, anastomosing into a submarginal veins, forming an angle of with the costa; tertiary venation reticulate and inconspicuous or invisible. Inflorescence terminal and in the axilsof the upper leaves, many-flowered, lax, except for the the large branchings. Peduncle glabrous or pubescent; pedicels pubescent. Wageningen Agricultural University Papers 97-2 (1997) 95

94 Lower bracts leafy, other sepal-like and like bracteoles about aslong as the sepals. Flowers: Sepals connate at the base, subequal, ovate, broadly ovate or suborbicular, obtuse or rounded, pubescent to almost entirely glabrous outside, ciliate, glabrous or nearly soinside, without colleters. Corolla white or lilac, with an ovoid or subglobose head in the mature bud about one third of the bud length, puberulous to almost entirely glabrous outside, basal part enclosed in the calyx always glabrous, ciliate around lobes, pubescent inside at the baseof thelobes and on the callous ring atthe mouth;tube amply cylindrical to obconical, mostly widened around the anthers which are placed in the upper half, with 5 longitudinal slits alternating with the lobes; lobes overlapping to the left in bud, obliquely elliptic, ciliate, entire, spreading. Stamens with apex about 0.5mm below mouth of corolla tube; filaments very short, glabrous; anthers ovate or nearly so, obtuse at the apex, cordate at the base, entirely fertile, glabrous. Pistil with apex about halfway along anthers; ovary subglobose, abruptly narrowed into the short style, of 2 separate carpels; pistil head of a basal globe or cylinder about x mmand an oblong (when lobes not diverging) bilobed stigmoid apex about 0.7 x 0.2 mm. Fruit of 2 separate follicles forming various angles with each other even within a single branch; follicles ellipsoid, woody, adaxially dehiscent. Seed light brown, with grain slightly darker, elliptic, flat, winged all around, striate or punctate, with a linear hilum in the middle; endosperm thin, mealy; embryo straight, spathulate; cotyledons ovate, rounded at the apex and at the base; rootlet mostly slightly longer than cotyledons. Distribution:Two speciesendemictomadagascar. The genus Stephanostegia strikingly resembles the New World genus Aspidosperma by the leaves, inflorescences, flowers and the wingsof theseeds.both generaaredistinguished mainly as follows: Leaves always opposite; follicles ellipsoid, not laterally compressed; seeds small, up to 16mm long; corolla tube with longitudinal slits Stephanostegia Leaves mostly alternate;follicles elliptic ornearly so,usually laterally compressed; seeds much larger; corolla tube without slits... Aspidosperma 96 Wageningen Agricultural University Papers 97-2 (1997)

95 Keytothe speciesof Stephanostegia 1. Corolla white; leaves mostly acuminate; follicles mostly acuminate. Western Madagascar 2. S. hildebrandtii Corolla lilac; leaves mostly obtuse or rounded; follicles obtuse. Eastern Madagascar 1. S. capuronii Stephanostegia capuronii Markgr. in Adansonia II, 12:219 (1972), as capuroni; Fl. Mad. fam. 169: 108, pi. 15, 1 5, map 23 (1976). Type: Madagascar, Antsiranana, Andrakadraka Forest, nearantalaha, Capuron SF24636 (holotype P;isotypes B,BR,G,K, TEF,WAG, Z). Fig. 16, p. 98;map 14, p.88 Heterotypic synonym: S. brevis Markgr. in op. cit. 220; op. cit. Ill, pi. 15, 6 9, map 23, syn. nov. Type: Madagascar, Toamasina, Ambila-Lemaitso, SF31-R-196 (holotypep). Tree 5 20 m high. Trunk 5 60cm in diameter; bark dark brown, rough and longitudinally fissured or smooth in young trees, with white latex. Leaves: petiole 1 5 mm long; blade elliptic, ovate or narrowly so, x as long as wide, 3 15 x 1 7 cm, rounded to shortly acuminate and with a blunt acumen at the apex, cuneate or rounded at the base. Inflorescence 2 9 x 2 7 cm. Peduncle 10 45mm long; pedicels 0.5 3mm long. Flowers: Sepals pink-magenta, 1 2x as long as wide, x mm. Corolla lilac, lobes sometimes(?) maroon, with an ovoid head in the mature bud, puberulous outside, with a pubescent belt below the insertion of the stamen 1 mm wide;tube x as long as the calyx, 1 2x as long as the lobes, 3 4mm long, 1 1.5mm wide at the base, 1 2 mm wide at the throat; lobes elliptic, x as long as the tube, x as long as wide, x mm, rounded or obtuse. Stamenswith apex about 0.5mm below mouth of corolla tube, inserted of the length of the corolla tube; anthers 1.2 x 0.5 mm. Pistil: ovary 0.7 x 0.7 x0.7mm, glabrous. Fruit, follicles x 1 1.5cm, obtuse, roughly lumpy or smooth. Seed x 6 8 mm; grain 3 4x 2 mm. Distribution: Endemic to Eastern Madagascar. Ecology:Wetforest atlow altitude. Wageningen Agricultural University Papers 97-2 (1997) 97

96 Fig Stephanostegia capuronii. 1 & 6, habit (x 2/3); 2 & 7, corolla with longitudinal slits (x 4); 3 & 8, fruits (x2/3); 4 & 9, seeds (x 1); 5, embryo (x 5) S. hildebrandtii. 10, habit (x 2/3); 11, part of leaf blade muchenlarged; 12, corolla (x 4); 13, fruit (x2/3); 14, seed (x 1). 1-2 from Capuron SF 24636; 3-5 from SF 16103; 6-7 from SF 15698; 8-9 from CapuronSF9091; 10-12from Baron 5451; 13-14from Boiteau Wageningen Agricultural University Papers 97-2 (1997)

97 Geographical selectionoftheapproximately 30specimensexamined: Madagascar. Antsiranana: road from Sambava to Antsiranana-Nord, Capuron SF27219 (P, TEF, paratype of 5. brevis); Andrakadraka Forest, near Antalaha, Capuron SF (B,BR,G,HBG, K, P, TEF, WAG, Z, type); S of Antalaha, Capuron SF27787 (P, TEF, WAG, paratype). Toamasina: Sahajinja, MaroantsetraDistrict,RN (?)2091 (P);SarsoalaPeninsula, Schatzet al (BR, C,K,P, TAN,WAG);ManongaR. basin,tributary RantabeR., Capuron SF9091 (G, K, P, TEF, WAG, paratype of S. b.)\ Tampolo Forest, Leeuwenberg ttlal (WAG), (BR, P, TAN, WAG); ibid., SF (K, P, WAG, paratype, also paratype of S. b.), (K, P, TEF,WAG,paratype, also paratype of S. b.), 468-R-107 (P, TEF);ibid., Capuron SF16103 (P, TEF,paratype);3.5 km N of Foulpointe, Noyes et al. 956 (BR, P, WAG); Analalava Forest, Wof Foulpointe, Leeuwenberg (WAG), (BR, P, TAN, WAG); Antetezana, SF4495 (P, TEF,paratype); Andriantantely,Lorihandava,Brickaville, SF735-R-162 (P); Ambila-Lemaitso, SF3205 (P,TAN,TEF,paratypeof 5. b.), 7570 (P, TEF,paratypeof S. b.),8286 (P,TEF,WAG,paratype of S. b.), 8291 (P, TEF, WAG, paratype of 5. b.), 8316 (P, TEF, WAG, paratype), (K, P, TEF, WAG, paratype of S. b.), (P, TEF, WAG, paratype), (P, TEF, paratype), 31-R-196 (P, type of S. b.). Fianarantsoa: Farafangana, Marofamahy, SF (P,TEF,WAG,paratype of S. b.), (P, TEF, WAG, paratype of S. b.)\ Manombo, SF45-R-62 (P,TEF).Toliara: Tsingafiafy Forest, between Manambato andfitamalama Rs., Capuron SF (G, K, P, TEF,WAG,paratypeof 5. b.) Note. The only character on which S. brevis could be distinguished from S. capuronii is the surface of the fruit, smooth in the first and roughly lumpy in the second. Both forms have been collected in the same forest near Ambila-Lemaitso and the character varies greatly also in the second species of the genus Stephanostegia hildebrandtii Baill. in Bull. Mens. Soc. Linn. Paris 1: 748 (1888); Markgraf, Fl. Mad. fam. 169: 110, pi. 15, 10 14, map 23 (1976). Type: Madagascar, Mahajanga, Vavatobe, Hildebrandt 3330 (holotype P; isotypes G, K, L, M, W, Z). Fig. 16, p. 98; map 14, p. 88 Heterotypic synonyms: Rauvolfia celastrifolia Bak. in Journ. Linn. Soc. 25: 335 (1890), as Rauwolfia. Type: Madagascar, Mahajanga, Androna, Baron 5451 (holotype K; isotype P). S. parvifolia Pichon in Not. Syst. ed. Humbert 13: 206 (1948). S. holophaea var. parvifolia (Pichon) Mgf. in Adansonia II, 12: 221 (1972); op. cit. 108, pi. 14, 7 12, map 22, syn. nov. Type: Madagascar, Antsiranana, Analamera Hills, Humbert (holotype P; isotypes G, K, TAN). S. holophaea Pichon, I.e.; Markgraf, op. cit. 106, pi. 14, 1 6, map Wageningen Agricultural University Papers 97-2 (1997) 99

98 22, syn. nov. Type: Madagascar, Antsiranana, Diego- Suarez, Ursch 272 (holotype P). S. megalocarpa Markgr. in op. cit. 220; op cit. 104, pi. 14, 13 17, map 22, syn. nov. Type: Madagascar, Antsiranana, Nosy Be, Lokobe Reserve, Capuron SF holotype P; isotype TEF). Tree 4 25 m high. Trunk up to 70 cm in diameter; bark brownish; wood hard, yellow. Branches pale or dark brown. Leaves: petiole 2 12mm long; blade elliptic, (1.5 )2 4x as long as wide, (2 )4 11.5x (0.7 )1 6 cm, acuminate to obtuse at the apex, cuneate at the base. Inflorescence 4 10 x 4 10 cm. Peduncle cm long; pedicels mm long. Flowers: Sepals greenish-white or yellowish-green. Corolla white, with a subglobose head in the mature bud, villose behind the anthers, pubescent belt below the insertion of the stamens 0.5 mm wide with straight hairs directed downwards; tube x as long as the calyx, 1 2 x as long as the lobes, 2 3 mm long, mm wide at the base, mm wide around the anthers, contracted at the mouth; lobes falcate, x as long as the tube, x as long as wide, obtuse. Stamens inserted of the length of the corolla tube (at 1 2 mm from the base); filaments 0.3 mm long, glabrous; anthers yellow or orange, x 0.3 mm. Pistil: ovary lxlxl mm, puberulous or glabrous. Fruit, follicles dark brown, obliquely ellipsoid, x x cm, acuminate or almost obtuse, smooth or lumpy, puberulous to glabrous. Seed 8 21 x 4 7 mm; grain 5 7 x 2 3 mm. Distribution: Endemic to western Madagascar. Ecology: Dry forest. Alt. low, up to 400 m. Geographicalselectionoftheapproximately35specimensexamined: Madagascar. Antsiranana: Diego-Suarez, Ursch 272 (P, type of S. holophaea);mt d'ambre, CapuronSF (K, P, TEF, WAG); Andrafiamena and upper RodoRs.confluence, Capuron SF (K, P,TEF, WAG);AnkaranaHills, Humbert & Cours (P),32764 (K,P,WAG); ibid., Capuron SF 3040 (K, P, TEF, WAG);Analamera Hills, Humbert (P), (G,K, P,TAN,typeof 5. parvifolia);mtambohipiraka,neofambilobe, Humbert & Cours 5669 (P), (P, WAG),32894 (K, P,WAG);NosyBe, Lokobe Reserve, Capuron SF (P, TEF, type of S. megalocarpa); Mt Ambohibe, Cours 5678 (P); NW of Ampondrabe, Gachet SF 5 bis (P,TEF); Sambirano, Perrier de la Bâthie (P); Ampondrabe-Ambaliha Hills, Nof Ambanja, Capuron SF (P,TEF,WAG).Mahajanga: Moramandia, Decary 1250 (P,WAG), 1372 (P);Vavatobe, Hildebrandt 3330 (G, K, L, M, P, W,Z, 100 Wageningen Agricultural University Papers 97-2 (1997)

99 type); crossing of Antsohihy and Analalava Roads, Capuron SF (P,TEF); Wof Antonibe Peninsula, Capuron SF (P, TEF,WAG); Berivota Plateau, SEof Mahajanga, Capuron SF (K, P, TEF);Amborovy, N of Mahajanga, Capuron SF (K, P, TEF, WAG);Namoroka Reserve, Rakotovao RN 3900 (P,TAN);ibid., SF 5 (K,P,TAN, US);Belambo,nearMaevatanana, Perrierdela Bâthie 944 (BM, P), (P);Nandrojia, SF (P, WAG); Ankara, Capuron SF 9394 (P, TEF, WAG), SF (P, TEF, WAG); Kelifely Mts, Marat 4730 (P,TAN, WAG);Ambatomitsangana, SF 8162 (P,TEF);Antsalova, Dokobe RN (P); near Ambodiriana, Leandri 2341 (K, P, WAG); Bemaraha Mts base, Boiteauin Herb. Jard. Bot. Tana (P, WAG); Antsalova-Tsiandro Road, Capuron SF 6916 (K, P, TEF, WAG), (P, TEF). Toliara: Analamitsiva, Miandrivazo, SF (K, P, TEF, WAG); Angodongodona, 5 km E of Miandrivazo, SF 7-R-243 (P). Sin. loc.: Baron5451 (K, P, type of Rauvolvia celastrifolia). Note. The indumentum on the sepals, on the corolla outside and on the ovary varies more or less gradually and independently. The fruits of the type of S. megalocarpa are somewhat larger than those of the other specimens seen, but the leaves are perfectly similar to those of the type of S. hildebrandtii. Therefore S. hildebrandtii was assigned some new synonyms. A new name to replace a homonym Carissa pichoniana Leeuwenberg, nom. nov. Type: Madagascar, Toamasina, Manampotsy R., SW of Vatomandry, Perrier de la Bâthie (holotype P). Basionym: C. vertillata Pichon in Mém. Inst. Sc. Madag. ser. B, 2: 126, 136 (1949); non Sessé & Mocino, Plantae Novae Hispaniae 29 (1887) which is a synonym of Alstonia longifolia (A. DC.) Pichon. Correction of author's error Lamarck's Tableau Encyclopédique cited in Leeuwenberg, Tabernaemontana 1, The Old World Species (1991, pages 169 and 204) was wrongly cited. He supposed to cite Vol. 1, part 2, but it was Vol. 2, part 6, which came out in 1819 and not in Taxa published there have been published even earlier by Poiret in Lamarck, Enc. respectively 7: 529 (1806), Tabernaemontana pandacaqui and Suppl. 5: 276 (1817); T. populifolia Poir. published on the same page as T. pandacaqui, was already known as Exacum Wageningen Agricultural University Papers 97-2 (1997) 101

100 vaginale Labill., Nov. Holl. PI. Sp. 1: 37, pl. 51 (1805) = Logania vaginalis (Labill.) F. v. Muell. Therefore the combination Logania populifolia (Poir.) Leeuwenberg resulted in an additional synonym for Logania vaginalis. Acknowledgements The author is greatly indebted to the Directors and Curatorsof the following herbaria for putting material athis disposal: A, B,BM, BO, BP, BR, C, COI, E, F, FHO, FI-W, FT, G, GB, GDC, GH, HAL, HBG,K, L, LG, LISC,LISU,M,MA,MAL,MAU,MO,NY, P,PRE,PRU,S, SRGH,STE,TAN,TCD, TEF,UC,UPS, US, USF, W,WAG,WU, Z. Grateful thanks goto MissM. Spitteler and MrH.de Vries for preparing the excellent drawings, to Prof Dr P. Morat for permitting the use of the nice drawings made in the Paris herbarium by Mr Rabarijaona for Boiteau in Adansonia II, 11: (1971) and by Mme Soussotte-Guérez and othersfor the Flore demadagascar etdes Comores. Prof Dr L.J.G. van der Maesen corrected the English text. For this paper a grant was given by the Landbouw-Export Bureau, Wageningen, in order to have it published with colour photographs. The author is preparing a second edition for the Apocynaceae in the Flore de Madagascar et des Comores on the request of Prof Morat. The editors of the Flore de Madagascar kindly provided the mapsof Madagascar, and Mr P. Bamps (BR) put the map of entire Africa at the author's disposal. 102 Wageningen Agricultural University Papers 97-2 (1997)

101 Index of exsiccatae of collections made in Madagascar and the Comoro Islands Craspidospermum verticillatum (Cv), Gonioma malagasy (Gm), Mascarenhasia arborescens (Ma), M. havetii (Mh), M. lanceolata (Mia), M. lisianthiflora (Ml), M. macrosiphon (Mm), M. rubra (mr), M. speciosa (Ms), M. tampinensis (Mt), Petchia cryptophlebia (Pec), P. erythrocarpa (Pee), P. humbertii (Peh), P. madagascariensis (Pern), P. montana (Pemo), P. plectaneiifolia (Pep), Pkctaneia longisepala (Pll), P. stenophylla (pis), P. thouarsii (Pit), Stephanostegia capuronii (Se), S. hildebrandtii (Sh). Onlynumberedspecimenshavebeenlisted.Herethenamesofthe collectors of the series "Reserves naturelles" (= RN),and of "Service forestier" (= SF)have not been mentioned as these series are continuous and the collector names often havebeenomitted. Allorge, L.,549, 578, 594 (Ml), 596,960 (Pit), 961, 962 (Pem). Adrianarisata,M., 12 (Ma). Antilahimena,P.,62 (Sh),80 (Pee), 138 (Pit). Armand,W., 85 (Ml). Barnett L.144 (Pec) 481 (Pit). Baron, 'R.,89 (Cv), 92 (Ml), 111 (Cv), 258 (Pit), 654 (Pee), 768 (Cv),853 (Cv andpec, now 853a), 1177 (Pit),1790 (Pec), 2458 (Pemo), 2501 (Pit),2525 (Pee), 2929 (Pit), 3234 (Pec), 3242 (Ma), 3840 (Mm), 4022 (Pit), 4551 (Pee), 4562, 4575 (Ml), 4663 (Pee), 4716 (Ml),4764(Plt), 4888 (Pee), 5000 (Pemo), 5451 (Sh), 5747 (Ma), 5765 (Pec), 5767 (Ma), 5787 (Ml), 5840 (Mia), 5916 (Pee), 6018 (Pec), 6084 (Pee), 6157 (Ma), 6370 (Mia), 6433 (Ma), 6541 (Pee), 6769, 6869, 6874 (Ml). Beaujard,Ph.,18 (Pem), 257 (Pee), 259, 260 (Pem), 261 (Pit). Benoist,R., 788 (Pem). Bernardi,L., (Ml), (Pee), (Ma), (Pit),11843 (Mia), (Ma), (Pit). Bender,106 (Pem),169, 285, 287 (Ma). Birkinshaw, C, 60 (Pee). Boiteau,P., 85 C (Cv), 117 D (Ma), 358 (Ml),373, 373 B (Pis), 493 (Pit), 504 (Sc), 1009 (Pee), 1011 (Ma), 1013 (Ml), 1030 (Pit),1054, 1092 (Ml),2007, 2019 (Pee), 2064, 2102 (Pit),2116 (Pem), 2119, 2122, 2123 (Pee),2125 (Pep),2132bis (Cv),2155, 2220 (Pit), 2340 (Ma), 2545, 2546 (Pem), 2577 (Pee), 2610 (Pem), 3005 (Pit), 3022, 3026, 3029 (Pee), 3056 (Ms), 3076 (Pit). Boivin,B., 13, 17 (Pit),1786 (Ma), 2075 (Pee), 2459 (Mia),2460,2461 (Ma), 2463 (Pit), 3202 (Pee). Bosser,J., 1050,1716 (Ml), 1849, 3267, 3521, 3527,4591, 4689,4731 (Ml), 5714 (Pee), 5724, 5725, 5841, 5923, 6692 (Ma), 7874 (Pee), 7889 (Pit), 8100 (Ml), 8438 (Ma), 8440 (Ml), 8457, 8458 (Pit),8909 (Ma), 9170 (Pis), 9171 (Pit),9186 (Ml), 9692 (Cv), 10255, (Pit),10654, (Cv), (Pit), 13232, (Mia), (Cv), (Ml),13872, (Pit), 13947, 13976, (Ml), (Pit), (Cv), 14173, (Pit), (Ml), (Ms), (Ml), (Pit),15840 (Ms), (Pit), 16663, (Ml), (Pit), (Pis), (Ml), (Pem), 19091,19109 (Ma). Wageningen Agricultural University Papers 97-2 (1997) 103

102 Broin,A., 138, 139 (Pee), 140 (Pem), 146bis (Pee). Catat, L.D.M, 1114,4348 (Ml). Chapelier, L.A., 93 (Ma). Chauvet,96,431(Plt). Cloisel,J., 66 (Ms), 70 (Pem), 79 (Ms), 80 (Pem),132 (Ma). Cours,G., 871 (Ma), 1517 (Cv), 1580 (Pit),1725 (Pem), 1860 (Ma), 1991 (Pit), 2009 (Cv), 2012 (Pit), 2014 (Ma), 2015, 2016 (Cv), 2022 (Pit),2026 (Ma), 2104 (Pit), 2264 (Ma), 2789, 2968 (Pem), 3103 (Ma), 3231, 3257, 3881 (Pee), 3954,3975 (Ma), 3988 (Ml), 4276 (Pit), 4860 (Ma), 4887, 4938 (Pem), 5024 (Ml), 5100 (Ma), 5437, 5448 (Pee), 5647 (Mla), 5669, 5678 (Sh). Cremers,G., 1282 (Cv), 1334 (Pee),2850 (Ml), 2914 (Ma), 2926 (Cv),3603 (Pem), 3648 (Pit). Croat, T.B., (Pit), (Cv), (Ml), (Cv), (Ml), (Ml), (Pit), 31029, (Pis), 31597, (Pit), (Ml), (Pit), 31753, (Ma), (Ml). Daniels,P.S., 57 (Ma). D'Arcy, W.G.,15367 (Pem). Debray,M., 51 (Cv), 404 (Ma), 533 (Pee),781 (Pem), 841 (Ml), 1210 (Mla), 1244 (Pem), 1249 (Pee), 1251 (Pem), 1253 (Ma), 1255 (Pem), 1256 (Pee), 1269 (Pem), 1512 (Pee), 1537, 1539 (Ma),1553 (Mla), 1557 (Pee), 1562 (Ma), 1586 (Mla), 1588, 1627, 1672 (Pee), 1725 (Pem), 1732 (Ms),1752, 1755, 1802 (Pem),1807 (Ma), 1819 (Pem),1864 (Sc), 2016 (Cv). Decary,R., 62 (Ma), 923, 957, 972 (Mla), 1052,1065, 1157, 1160 (Ma), 1202 (Ml), 1245 (Ma), 1250 (Sh), 1254, 1258 (Pit),1285 (Ma), 1326 (Mla),1372 (Sh), 2089 (Pit), 2233, 2274 (Pee), 2288 (Ml), 2296 (Pee), 2298, 2318, 2326, 2329, 2334 (Ml), 2360 (Pee), 2440 (Ml), 2700 (Pis), 2824 (Ml),2933 (Pis), 3323 (MI), 3512, 3518 (Pis),4056 (Pit),4105 (Ml), 4166 (Ms), 4172 (Pit), 4312, 4319 (Ml), 4569 (Pit), 4855 (Pee), 4967, 5312, 5331 (Pem), 5387 (Pep),5407,5418, 5421 (Pem), 5438,5477 (Pep),5527,5535 (Pem),5551 (Ma), 5792, 5819 (Pem), 6015 (Cv), 6379 (Pit), 6547 (Pll), 6977 (Cv), 7390 (Pee), 7551 (Pit), 7756 (Pee), 7774, 7800 (Ml),8125 (Pee),8188, 8249 (Ml), 8257 (Ma), 8244 (Pee),8260 (Ml), 8262 (Pee),8264,8273 (Ml),8498,8586 (Pis), 8868 (Pit), 9098 (Pis), 9407 (Ma), 9430 (Ml), 9556 (Pis), 9768 (Pem), 9776 (Ma), 9909 (Pem), 9977 (Pit), 10396, 10502, 10578, 10599, 10615, (Pem), 10820, 10836,10845 (Ma), (Cv), (Pem), (Ma), 12809, (Ml), 12834, 12878, (Pee), 12888, 13132,13292 (Pit), (Ml),14101 (Pit), 14290,14298 (Ma),14395, (Pee), (Pit),14471 (Pee),14537 (Ma), (Pit),14553 (Ma), (Mla), (Pit), 14777, 14780, (Mla), (Pit), (Ma), (Pee),15223 (Ml), (Pit), 15257,15274, (Ml), (Ma), 15403, (Pit),15570, (Ml), 15620, 15641, (Pee),15660, (Ml),15746 (Pit),15924 (Ml),15964 (Pis),16275,16324,16337 (Ml), (Pit),17049 (Pee),18739 (Pit),18883 (Pis),18889,18890 (Ml),18892 (Pis),18951 (Ml). Dequaire,J.,10098 (Ma), (Pee), (Ml),24195,24247, (Pis), 27641, 27644, (Ma). Derleth, P, 65 (Pem),141 (Ma), 152 (Pee). Deroin,T.,127 (Pem),165 (Pee),189 (Mla). Descoings,B.,103 (Ma), 256, 271 (Pit), 306 (Pem), 335 (Ma), 1254, 1295, 1408 (Pit), 1670 (Ml), 1922 (Pit),1923 (Mla),2196 (Ml), 3302 (Ml), 3364 (Ma), 104 Wageningen Agricultural University Papers 97-2 (1997)

103 3365 (Pit), 3370 (Pee), 3379 (Ma), 3527 (Pee), 3563, 3636 (Ml), 3708, 3733 (Pit). Dorr, L.J., 2956 (Pit), 3170 (Cv), 3318 (Pit), 3490 (Cv), 3591 (Pee), 3749 (Ml), 3778 (Pee),3827 (Pit),3926 (Ml), 4016,4021 (Pem),4181 (Ml). Doutrelepont, H., 1235 (Pee). Dumetz, N., 504 (Ma), 564 (Pit), 585 (Ms), 735, 739 (Pem), 1201, 1351 (Ms), 1366 (Pee), 1401 (Ma). DuPuy,B.MB 42, 406,2446 (Ml). Edouard, P.,29 (Ma). Evrard, C, (Pec), (Pit). ExpositioncolonialedeMarseille,19 (Pit). ExpositionpermanentedeMadagascar, 176 (Ml). Faber-Langendoen, D.,2984 (Ma). Floret,J.J., 1091,1092, 1106 (Pee), 1232 (Ma). Fosberg, F.R., (Ml),52546 (Ma). Forsyth-Major, C.J., 132 (Ma),324 (Pec),331 (Ma). Friedmann, F., 331, 481 (Ma). Garnier,145 (Pec). Gautier,L., 2392 (Ma), 2456 (Pem), 2588 (Pit), 2629, 2638 (Ma). Geay,F., 5983,5984,5985, 6003,6004,6005,6006, 6007,6011, 6012, 6014, 6015, 6345, 6379, 6380, 6383 (Ml), 6393, 6676, 6677, 6679, 6680,6681, 6682, 6683, 6684, 6685, 6686, 6713, 6714, 6738 (Ma), 6748 (Ms), 6749 (Pit), 6755 (Ml), 6774 (Pit), 6780 (Pem), 7043, 7044 (Pit), 7254 (Ma), 7722 (Pee), 7744 (Ma), 7898, 7899, 7902, 7905 (Pit),7929, 7930, 7931, 7932 (Ma), 8146, 8147 (Pit). Geldermalsen-de Jongh, van,82/8 (Ms). Gentry,A.,11356 (Ma), (Pit), (Pee), 11544, (Pit),11794 (Pee),11799 (Ml), 11872,11894 (Mia),11906 (Pit),11922 (Pee). Gereau,R.E., 3253 (Pem), 3298 (Ma), 3313 (Ms). Gerold,R., 12 (Pem). Grevé, 29 (Pit), 41, 100, 204 (Ml). Guillaumet,J.L., 3853 (Ma), 4218 (Pee). Guillot,J., 7 (Ma), 45 (Pit). Harder,D.K.,1588 (Pee),1603, 1713 (Ma), 1758, 1765 (Pit). Hildebrandt,J.M., 2949 (Pee), 3012 (Pit),3078 (Ma), 3099 (Ml), 3232 (Pee), 3299 (Mia), 3330 (Sh), 3602 (Cv), 3679 (Pec), 3936 (Cv). Hodgkin,H.T., 65 (Cv). Homolle,A.-M., 45 (Pee), 130 (Ma), 161 (Pee),171 (Ma), 174 (Pee), 391 (Pit), 1542 (Ml), 1735 (Pem), 2009 (Cv), 2014 (Ma), 2016 (Cv), 2104 (Pit), 2264 (Ma). Humbert,H., 2033, 2034 (Pit),2085 (Ma), 2132, 2356,2391 (Pit), 2733, 2745 (Ml), 2790 (Pis), 2873 (Ml), 3127, 3135 (Pit), 3191 (Pemo), 3430 (Ma), 3592 (Pit),3614 (Pemo), 3989 (Pee),4028, 4038 (Ml), 4064, 4065, 4084,4372 (Ma), 4921 (Pit), 4926 (Ml), 4927 (Pit),5013 (Ml),5090,5257, 5627 (Pis), 5669 (Sh), 5717 (Ma), 5774 (Pem), 5800 (Ma), 5804 (Ms), 5939, 6052 (Ma), 6200 (Pemo), 6238, 6251 a (Ma), 6488 (Ml),6747 (Pit), 7024 (Cv), 7050 (Ml), 7068bis (Pit),11367 bis (Ml), (Pit),11651 (Ml), 11670,11735 (Pit), (Pep),12195 (Ma),12333 (Ml), 12751, (Pis), 13346, Wageningen Agricultural University Papers 97-2 (1997) 105

104 13399bis (Pit),13764, 13882, (Cv), (Ma), (Ml), (Pee), (Mia), (Pee), 18773,18774 (Mia), 18775, (Ma), 18870, 18871, 18876,18877 (Mia), 18965, (Pit),19025 (Ma),19043 (Mia),19085 (Ma), (Sh), (Mia),19222 (Sh), (Ml),19517 (Ma),19539, (Ml), 19646, (Pit),20354 (Ms), (Pem), (Pit), (Ms), (Cv), (Ms), (Cv), 20740, (Pem), (Pit),22023 (Pee), (Peh), (Ma), (Peh), (Ma), (Pee), 28661, (Ml), (Pit), (Gm), 29835, (Ml), (Ma), 32094, (Pee), (Pit),32570, (Pee), 32676, 32764, (Sh), (Mia), (Sh), (Pee). Humblot, L, 26 (Pee), 58, 88 (Pem), 204 (Ml), 611 (Cv), 657 (Pem),1026 (Pee). Imbert,T., 50 (Gm), 116 (Ma), 125, 127 (Pee). Jacquemin,H., 253 (Pee), 275, 284, 287 (Ma), 311, 326 bis (Ml), 479 (Mia), 528 (Ma), 535 (Mia), 556 (Ma), 753, 859 (Pee), 1163 (Pem), 1198 (Cv),1202, 1203 (Ml),1205, 1229 (Pee). Jongkind,C.C.H.,885 (Pemo). Keraudren,M., 308, 312, 320 (Ml),410, 433 (Ma), 798 (Pit),921 (Ml), 1242 (Pee), 1488 (Ms), 1525 (Pit),1548 (Ml), 1572 (Pee), 1647 (Pit),1656 (Ma), (Ml), 25472, 25475, 25488, 25635, 25663, 25669, 25674, (Ma), 25813,25831, (Ml). Kiener,H., 65 (Sc). Klackenberg,J., (Pee). Kotozafy, A.,8, 348 (Ma). Labat,J.N., 2071 (Pis), 2173 (Pee), 2446 (Ml). Lam,H.J., 5479 (Pis), 6081 (Pee), 6097 (Ma). Leandri,J., 104 bis (Pee), 120 (Ml), 123bis (Pee),371, 372, 528, 947 IV, 990 (MI), 1074, 2062 (Pee),2225 (Ml), 2341 (Sh),2346, 2606,2606bis (Pee), 2633 (Ml), 2667 (Pit),2713, 2814 (Pee),3468 (Ml), 3895 (Pit), 4261 (Ml), 4304, 4330 (Cv), 4403 (Pit), 4404, 4407 (Ml), 4414, 4429, 4574 (Pit). Leeuwenberg,A.J.M.,13713 (Cv),13735 (Pec),17739 (Ma),13746 (Pit), 13747, (Sc), (Pem), (Pit), 13759, (Pem), (Sc), 13773, (Pem), (Pee), 13907, (Ma), 14004, (Pit), (Ma), (Pec), (Pee), (Ma), 14153, 14154, 14158,14159 (Pit),14160 (Ml), (Pit),14170 (Ml), (Pit),14172, 14177, (Pit), (Ml), (Pit), (Ma),14188 (Pem), 14192, 14198, (Pit), (Ma), (Pem), (Pit), (Ma),14218 (Ml), (Pem),14241, (Pee), (Pit), (Ma), 14260, 14261, 14262, (Mia), (Ma), (Pem),14271, 14273, (Pee), (Ma), (Pee),14395 (Pit), (Pee), 14399, 14400, 14402, 14407, (Pit), (Ma), (Pit),14445 (Ma), (Pit),14468 (Sc), (Ma), (Pem), (Pit),14474 (Ma), (PH), (Sc), (Pem), (Pit),14584 (Ma), (Pit), (Pemo), (Pit), 14614,14621 (Ml), (Pis), (Ml),14635 (Pit),14638 (Ml),14642, (Cv), (Pit),14674,14678 (Pee),14683 (Ma), (Ml), (Pit),14689 (Ma), (Ml),14692 (Ma),14694 (Pee), (Ma),14703 (Pee), 14707, 14710, (Pit), (Ml), (Ma), 14723,14724 (Pit),14738 (Ml), 14739,14744 (Pit), 106 Wageningen Agricultural University Papers 97-2 (1997)

105 14745 (Mia), (Ma),14747 (Pit),14749 (Mia),14750 (Ma),14753 (Pee), (Pit), (Mia), (Pit), (Pee), (Ma), 14765, (Pit),14770 (Pee),14772 (Ma). Le Thomas, A.,104 (Pern). Lewis,B., 618 (Pem), 760,873, 1266 (Ma), 1268 (Ml). Liede,S., 2718 (Pis). Louvel, M., 9 (Mm), 56 (Pec), 75 (Cv). Lowiy,P.P., 4237, 4465 (Pem), 4549 (Pit). Lyall,R., 146 (Mh). Mabberley, D.J., 900, 904 (Pit). Malcomber, ST., 921 (Pem), 1266 (Ma), 1347 (Cv), 1634, 1667 (Ma), 1697 (Pec),2325 (Pem),2660 (Pee). McPherson, G., 14197, (Pem), (Pee), 14626, (Ml), (Pit), (Pem), (Cv), (Pit). McWhriter, J.H., 214 (Pem). Meyers, D., 269 (Ma). Miller,J.S., 3724 (Cv), 3768 (Ma), 3803 (Pit),3836 (Ma), 4340 (Pem), 4526 (Ml), 6137 (Pit), 6267 (Ml). Mocquerys, A., 271 (Mh), 290 (Pem), 497 (Pit). Morat,Ph., 69, 226, 236 (Ml),834, 882 (Pee), 922 (Ma), 983 (Ml), 1243 (Ma), 1407 (Pee),1411 (Mia), 2451 (Pis), 2463, 2522, 2583, 2662 (Ml), 2683, 2705 (Pee), 2775, 2805 (Ma),2813, 3085 (Pee), 3510 (Pis),3891, 3892 (Pit), 4172 (Ml), 4579 (Pee), 4609, 4620 (Cv), 4730 (Sh), 4744 (Pee). Nek,F.I.van, 1829 (Pee),1952, 1959 (Ma), 2066 (Pee), 2212 (Pem). Nicoll, M., 208 (Pec), 371, 372 (Pee), 573 (Ma). Noyés,R.D., 956 (Sc), 996, 1080 (Ma). Outer, R.W.den, 1048 (Pit),1214 (Ma),1224 (Pec). Paroisse,G., 21 (Ma). Pascal,O., 310,489 (Ce). Peltier,J. & M., 965 (Pit),1243 (Ml), 1978 (Cv), 2480 (Ml), 2658 (Pis), 2740 (Ml), 2783 (Pit),2897, 2984 (Ml), 3283 (Pee), 4866 (Pit),5249 (Ml),5298 (Pit), 5303, 5386, 5514, 5789 (Ml), 5983 (Pee). Pernet, 85 (Pit). Perrier dela Bâthie 2 (Pit),18, 18 bis (Pee), 74bis (Ma), 75 (Ml),351, 351bis (Pit), 420, 420 bis, 420ter (Ml), 944 (Sh), 966 (Ma), 1130, 2351 (Pit), 4407 bis (Ml), 5691 (Ma), 7062 (Cv), 8136, 8137, 8194, 8197, 8837 (Pit),8839 (Pee), 8840 (Pem), 8841 (Cv), 8842 (Pee), 8846 (Ma), 8850 (Pit), 8854 (Pee), 8855 (Pem), 8860 (Sh), 8874, 8876 (Pit),8881, 8883 (Pee), 8884 (Cv), 8889 (Pem), 8896, 8897 (Pee), 8898 (Mia), 8900, 8920 (Pem), 8922 (Ma), 8925 (Pec), 8928 (Pee), 8929 (Pem), 8930 (Pit),8936 (Pee), 8938, 8939,8940, 8941, 8942, 8943 (Pit), 8944 (Pee),8948 (Cv), 8949 (Ma),8955, 8956 (Ml), (Pit),11313 (Ml), (Pit),12552 (Ml), (Pit),12725 (Pis), (Pit),12930 (Cv), A, B, 13237,13238,13239 A, B, (Pit),13435 (Ma), 13578, 13735, (Pit),14139 (Ms), (Pit),14654 (Sh),14965 (Ma), (Pee),15212 (Pit),15215 (Mia),15463 (Sh), 15507, (Pit), (Pem), (Ma), (Pemo), (Ma), 16559,16563,16599 (Pit), (Ml), (Pit),16669 (Pis),16698 (Ml), 16819,16846, (Pit), (Ma), (Pem), 17775,17802, Wageningen Agricultural University Papers 97-2 (1997) 107

106 17805 (Pit),18058 (Pern),18110 (Pit),18360 (Pee),18450 (Ml), (Pee), (Ml),18625 (Mm), (Pit),18627 (Mr), (Pit). Pervillé,A., 106 (Pee), 169, 285, 287 (Ma), 355, 369, 413 (Pee), 440, 703, 768 (Pit). Phillipson,P.B., 1828 (Ml), 1829 (Pis),1917 (Pee),1930 (Ml), 2006 (Mia), 2215 (Cv), 2308 (Pis), 2640 (Ml),2681, 2963 (Ma), 2982 (Pern), 3076 (Pit), 3128, 3180 (Pis), 3937 (Pit), 3954 (Ms). Pobéguin, H. 60 (Pee). Poisson,H., 24 (Ml), 25 (Ma), 27 (Ml), 55 (Ma), 92 (Pit), 555,616, 635 (Ml). Ponsonby,L.,18 (Ms). Rabevohitra,R., 1765 (Pem), 1795, 1809 (Pit), 1856, 1879,1913, 2065,2108 (Pern),2124, 2142, 2203 (Ms),2244,2348 (Pem), 2423 (Ms),2431 (Pem), 3313 (=SF 28631) (Ms). Raharimalala,F.,158, 198,200 (Mh), 230 (Ml), 368 (Mh). Rakoto,R.,14 (Pee),176 (Pit),181 (Cv), 285 (Pec), 321 (Ma), 292, 341 (Mm). Rakotomalaza,P.J., 64 (Pit), 65 (Pem), 360 (Pee). Rakotovao,G., 114 (Pem), 174 (Ce), 540, 589 (Pit). Rakotozafy,A., 222 (Pit), 260 (Cv),325 (Ma), 339 (Mia),506, 536 (Pee), 597, 834 (Pem), 1000 (Pee),1029 (Ml), 1087 (Pit),1114,1114bis (Pee),1134bis (Ma), 1284 (Pit),1345 (Pem), 1373 (Ma), 1709 (Pit),1758,1784, 1784bis (Pem), 1861, 1979 (Ml). Randriamampionona,B., 85 (Ml), 94 (Pem), 100 (Ma), 102 (Pit),103, 104,105, 106, 107, 108 (Ma), 114 (Pit),121 (Ml), 153, 184 (Pit), 213 (Ma), 273 (Ml), 317 (Cv), 318 (Ma), 377 (Cv), 411 (Pit), 449 (Ma), 476 (Pit), 583 (Cv), 602 (Pit). Randrianasolo, A., 43,44 (Pem),173, 180 (Ml), 268 (Ma). Randrianjanaka,L.M.,16 (Ma). Rasoavimbahoaka,F., 23Pen). Rauh, W., 24 (Pee), 599 (Pem), 898 (Ml), 912 (Pit),1278 (Ml), 1361 (Ms),1373 (Pem), 1505 (Ml),10802 (Pit). Ravelonarivo,D., 35 (Ma), 70 (Pee), 225 (Pit). Razafindradora, 53 (Pec). Razafindrambao,R., 578 (Ma), 580 (Pem), 596 (Mia). RN 5 (Mt), 8, 20 (Pee), 22 (Mia), 504 (Ma), 622 (Pit),1036 (Pee), 1099 (Cv), 1119 (Ma), 1150 (Cv), 1206 (Pit),1233 (Cv), 1280 (Mia), 1282 (Ma), 1300 (Pec), 1301 (Ma), 1379 (Cv), 1383 (Pit),1661, 1769 (Ml), 1775 (Cv), 1779 (Ma), 1799 (Ml), 1935 (Cv),2091(?) (Sc), 2176, 2251 (Ma), 2293 (Ml), 2350 (Ma), 2388 (Ml), 2467 (Ma), 2510 (Ml), 2617 (Pee), 2679 (Cv),2716, 2729 (Pee), 2793 (Cv), 2972 (Ml), 3166 (Mh), 3414, 3427 (Pem), 3428, 3451 (Ma), 3461 (Ml), 3492 (Pem), 3715 (Cv),3891 (Ml), 3900 (Sh), 4068 (Pee), 4077 (Ml), 4242 (Pem), 4606, 4647 (Pee), 4875 (Cv), 4890 (Pee), 4896 (Ma), 5088 (Ml), 5328 (Pit), 5490,5491 (Pem), 5788 (Ma),5831 (Ml), 5913 (Ma), 5923 (Pem), 5947, 5967 (Cv), 6213 (Ml), 6345 (Pit), 6647, 6779 (Pee), 6830 (Cv), 6963 (Mia), 6964 (Pee), 7066 (Ml), 7200 (Pee), 7443 (Pem), 7592, 7627 (Pit), 7660, 7800 (Ml), 7815 (Mia), 7915, 8004 (Ma), 8154 (Ml), 8160 (Pem), 8173 (Pep), 8356 (Cv), 8598 (Pem), 8826, 8882, 8909, 9093 (Pee), 9431 (Ma), 9478 (Mh), 9481 (Pee), (Pem), (Pit),10179 (Ma), (Pee), (Ma), (Cv), (Ml), (Pit),10812 (Ml), (Pee), 10863, (Cv), (Sh), (Ma), (Pit), 11433, (Pee), 11497, (Pit), (Ma), (Mia), (Pee), (Cv), 12224,12229,12323 (Ma), (Pit). 108 Wageningen Agricultural University Papers 97-2 (1997)

107 Richard, A., 440 (Pit), 550 (Ml). Richard,J.M.C.,13, 58, 80, 100, 107, 127, 181 (Ma), 239 (Ml), 247 (Gm), 269 (Pit), 284, 307 (Ml), 312 (Mia), 327, 337, 343 (Pit), 370 (Mia), 528 (Ml), 551 (Ma), 552 (Mia), 845 (Ml). Schatz,G.E., 1438 (Pee), 1676 (Pem), 1706 (Pec), 1852 (Ma), 1921 (Sc),1929 (Ma), 1933 (Mh), 1972, 2025 (Ma), 2452 (Cv), 2833 (Pem), 3097 (Pit), 3129 (Ma), 3146 (Pit), 3157, 3441 (Pem). Schlieben,H.J., 8128 (Ma),8219, 8220, 8242 (Ml). Schofield,J., 4 (Pee). Scott eiiot,g.f., 2155, 2177 (Ms), 2207 (Ma), 2289 (Pit),2361 (Ma),2373 (Ml), 2374 (Pem), 2378 (Cv), 2440, 2463 (Ma), 2584 (Pit), 2703 (Ma). SF 2 (Ce), 3 (Ml), 4 (Ma),5, 5bis (Sh),11 (Pee),13 (Pit),15 (Pee), 24 (Ml), 30 (Pee), 34 (Ml),35 (Ma), 69, 70 (Ml), 71, 75, 77 (Ma), 118 (Cv), 134 (Pee), 153, 183 (Ml), 200 (Pit), 212 (Ma), 224 (Cv), 253, 271 (Ml), 366 (Pem), 380 (Ma), 548 (Ml), 661, 876 (Ma), 1002 (Pec),1005 (Ml),1099 (Cv), 1109 (Mt), 1130 (Mt), 1132, 1376 (Cm), 1460 (Ml), 1491, 1548 (Ma), 1846, 1929 (Ml), 2118 (Ma), 2135 (Ml), 2143, 2193, 2194 (Ma), 2201 (Pee), 2209 (Ml), 2269, 2314, 2357 (Pit), 2589 (Ma), 2676, 2727 (Ml), 2848 (Ma), 2914 (Cv), 2926 (Mt), 3040 (Sh), 3106 (Ma), 3125 (Ml), 3150 (Pee),3205 (Sc),3231 (Ma), 3396, 3483 (Ml), 3560 (Pee), 3642 (Ma), 3668 (Gm), 3697, 3771, 3814 (Ma), 3856 (Gm), 3988, 4020,4025 (Ml), 4078 (Pem), 4138 (Ml), 4180 (Ma), 4253 (Pec), 4254 (Cv), 4283 (Pee), 4315 (Ml), 4382 (Ma), 4404 (Pec), 4421 (Ma), 4422 (Cv), 4495 (Sc), 4645, 4657 (Cv), 4775,4801, 4875 (Ma), 4945 (Ml), 5216, 5241, 5462 (Cv), 5502, 5519, 5605 (Ma), 5630 (Pem), 5664 (Cv), 5720 (Pec), 5738 (Ml), 5783 (Pec), 5833 (Pee), 5841 (Mia), 5874, 6262, 6303 (Ml), 6326 (Mt), 6365 (Ma), 6426,6485,6537 (Pem), 6639 (Cv), 6640 (Pem), 6720 (Cv), 6750 (Pee), 6916 (Sh), 7007, 7054 (Ma), 7056 (Cv), 7148 (Pem), 7181 (Ml), 7188 (Pem), 7390, 7404 (Ma), 7545 (Cv), 7570 (Sc), 7637 (Ml), 7665, 7813 (Ma),7881 (Cv), 7949 (Pee),8146 (Ma), 8162 (Sh),8243 (Ml),8286, 8291, 8316 (Sc),8396 (Ml), 8508 (Pem), 8571 (Ma), 8780bis (Pem), 8810 (Mm), 8963, 9062, 9063 (Ma),9091 (Sc), 9157 (Mh), 9208 (Cv), 9394 (Sh), 9487 (Ma), 9567, 9629, 9663 (Ml), 9794 (Gm), 10165,10166,10168,10263, 10274, (Ma), 10531, (Ml), (Pee), 10771, 10784,11101 (Ma),11170 (Cv), 11388, 11414,10427 (Sh),11591 (Pee), (Ml), 11760,11791 (Ms), (Gm),11988 (Sh), (Cv), (Ml), 12254,12255 (Sc),12402 (Ma), (Sh), (Pit),12486 (Ma),12546 (Cv),12616 (Ma), (Ml), (Gm), (Pis),13261 (Pee),13274 (Ml), (Sc),13373 (Gm), (Sh),13630,13679 (Ma), (Pee), (Ma), (Ml),13922 (Cv), (Pem),13955 (Cv), (Ma), (Pee),14310 (Ma), (Ml), (Gm), (Ma),14786 (Pee), (Pem), 14840, (Ml), (Ma), (Gm), 15145,15233 (Ml), (Ma), (Gm), 15616,15698 (Sc),15705 (Ma), (Cv), (Sc), (Ma),16146 (Cv), 16193, (Sc),16225 (Pem), 16249, (Cv), (Ma), (Pec), (Pee), 16405, (Ml), 16605, 16606, 16771, 17460, (Pee), (Ml), (Sh), (Cv), 18457, (Sh), (Ml), (Cv), (Sh),18893 (Mia),18976 (Ma),19018 (Cv),19057 (Pee),19489,19664 (Ml),19707 (Ma), (Pit), (Pem), (Cv), (Pem), (Ma), (Sh), (Ml), (Pee), 20521, (Ms), (Gm), (Pee),21057,21225,21305 (Cv),21445 (Pem), (Pec),21622,21655 (Ml), (Pee), (Pec), (Ma), (Pee), (Pec), Wageningen Agricultural University Papers 97-2 (1997) 109

108 (Pem), (Ma), (Pec),23604 (Pem), (Pee),23651 (Pern), (Pee), 23820, 23835, 23841, 23936, 23940, (Pem), 24007, (Pee), (Sh), (Sc), 24946ter (Pee), (Cv),25049 (Ml), 25192, 25209, (Ma), (Pec), (Cv), (Pec), 25821, (Ma), (Cv), (Ma), (Pem), (Cv), (Ml), (Cv), (Ma), (Gm), 27170, (Pem), (Sc), (Cv), (Pee), (Sc), 28076, (Pem), (Pee),28342 (Pem), (Pee),28390,28416 (Pec),28431 (Mm), (Cv), (Ms), (Pem), (Sc), (Ms), (Ma), 28818bis (Cv), (Pee), (Ma), 29282, (Mia), (Pee), (Pem), (Pee), (Ma), (Pit), (Ma), (Mia),29755,29769, 29914, 29919, (Pee), (Pem), (Ma), (Ml), 31005, 31052, 31061, (Ma), (Pee), (Ma), 31225, (Pee), 31417, 31425, (Ml), (Pit),31637 (Pee), (Ml), (Ma), (Mia), 32004, 32005, (Pee), (Pem), (Pit),32583,32584,32585,32586 (Ma), 32590,32606 (Pee), (Pem), (Peh),32923 (Sc), (Pit), (Sc), (Ma), (Pem),33038 (Pit),33108,33294,33337 (Pem), (Pee), (Pem), 33688, (Pee), (Ma),671-R-l (Sh), 276-R-6 (Ma),163-R- 7 (Sc), 37-R-10 (Cv), 295-R-19 (Pem), 39-R-44 (Pee), 32-R-55 (Ma), 45-R-62 (Sc),89-R-78,25-R-87,39-R-100 (Ml),43-R-105 (Sc),46-R-106 (Cv), 468- R-107 (Sc),85-R-116 (Ma), 169-R-140 (Sc),35-R-142 (Ma), 65-R-142 (Pee), 138-R-146,85-R-147 (Cv), 735-R-162 (Sc), 29-R-168,826-R-182 (Ma),117- R-187 (Cv), 190-R-187 (Ma), 43-R-193 (Sh),31-R-196 (Sc),145-R-199, 78- R-206 (Ma), 7-R-207 (Cv), 19-R-239 (Ml), 12-R-240 (Ma), 7-R-243 (Sh), 54- R-247 (Ml),34-R-272,2-R-286 (Pee),3-R-286 (Ml),19-R-288 (Cv),19-R- 295,8-R-305 (Pem),12-R-317 (Pee). Seyrig, A., 21 (Ml),106 (Pit),106bis (Pis). Shuffeldt,U.S., 3 (Cv). Stevens, W.D., (Pit). Herb. Jard.Bot. Tana (Pem), 2102, 2220 (Pit),2925 (Cv), 2950 (Pem), 3245 (Cv), 3728 (Ma), 3866,3875 (Cv), 4989 (Ml), 5106 (Pee), 5130 (Pit), 5238 (Pec),5281, 5411 (Ma),5415, 5689 (Pit), 5802 (Ml), 6110 (Cv), 6174 (Sh), 6181 (Pee), 6203 (Pit). Thouvenot,E.,16 (Pec and Rubiaceae), 75 (Ma), 97 (Mm). Turk,D., 212 (Ma), 582 (Mm). Ursch, E., 28 (Pec), 36 (Ma), 55 (Mt), 256 (Ma), 268,269 (Mia), 270 (Ma),272 (Sh), 280 (Ma), 280 a (Mia),284 (Ma), 284 a (Mia). Viguier, R.,415 (Ma), 852, 1042 (Pec), 1108, 1990 (Ma). Villiers, J.F.,4779,4822 (Pee),5021 (Pit). Waterlot, 330 (Pee), 336 (Mia), 339 (Ma), 354 (Pit), 356 (Pee), 357,402 (Pit), 712 (Cv). Werff, H.vander, (Pit). Willing,T., 36 (Ml). Zarucchi,J.L., 7380 (Ma), 7452 (Pem),7495 (Pit). 110 Wageningen Agricultural University Papers 97-2 (1997)

109 Index of exsiccatae of collections made outside Madagascar and the Comoro Islands Gonioma kamassi (Gk), Mascarenhasia arborescens (Ma), M. lisianthiflora (Ml), Petchia ceylanica (Pece). Acocks, J.P.H., 13739, (Gk). Adam, F.H., MAU (Ma). Archbold, M.E., 1472 (Ma). Avery, G.N., 1585 (Ml). Balapure, K.M.,72474 (Ma). Balasabramaniam,K.,BSM-4 (Pece). Barbosa, G., 2115, 4296 (Ma). Barness, R.D., 1/63 (Ma). Barrett, R.L.,31/55 (Ma). Bayliss, R.D.A., BRI.B.475,BS 6104 (Gk). Biegel, H.M.,4692 (Ma). Blackmore,S., 858 (Ma). Bond, J.S., 78 (Ma). Bos, J.J., 765 (Gk). Botha, D.J., 463 (Gk). Bottomley H (Gk). Bradbuine, P.A., 117 (Ma). Brass, L.J., (Ma). Brian, A.,MAU (Ma). Brink, E., 326 (Gk). Britten, L.L., 1329 (Gk). Britton, N.L.,8646 (Ma). Brousse,G., MAU (Ma). Brumbach, W.C.,6634,7340 (Ma). Burchell, W.J., 3659, 4138, 4676,6044, 6071 (Gk). Burtt, B.D., 236,5417 (Ma). Bum Davy,J., (Gk), (Ma). Campos Andrada,E., 770,1494,1911 (Ma). Chapman,J.D., 350, 5876, 6999, 7066, 8136, 8931 (Ma). Chase,N.C., 2993, 6130, 6447, 8040, 8334 (Ma). Cheadle, V. 733 (Gk). Chevalier, A.,27995 (Ma). Comins, D.M., 2005 (Gk). Compton, R.H., (Gk). Corbisier-Baland, A.,2074 (Ma). Correia, M.F.,2789 (Ma). Cramer, L.H., 3413 (Pece). Dahlstrand, K.A., 478, 1661 (Gk). Dale,I.R., 1053 (Ma). Dale,M.O., SFK 494 (Ma). Dassanayake,M.D., 492 (Pece). Dawe,M.T.,487 (Ma). Wageningen Agricultural University Papers 97-2 (1997) 111

110 Drummond, R.B., 1473,5002 (Ma). Dümmer, R., 1772 (Gk). Edinburgh,Royal Bot.Garden, C9217, (Ma). Ekema, S.N., 23 (Ma). English, 1/50 (Ma). Faden, R.B., 70/815, 76/404 (Pece),77/532 (Ma). Faulkner, H., 177, 198, 389,in GH (Ma). Flanagan, H.G., 367 (Gk). Forests,ass.conservator, Kenya, 308 (Ma). Forest Dept.,Kenya,256 (Ma). Furtadon, C.X., SING (Ma). Gerrard, W.T., 1424 (Gk). Gillett, J.B., 1484 (Gk). Gillis,W.T.,8310 (Ma), 9453 (Ml), 9562 (Ma). Goldblatt, P.,5203 (Gk). Gomes e Sousa,A., 752, 760,777, 4270, 4358, 4404,4427 (Ma). Gordon Gray, J.L., 1756 (Gk). Gossweiler, J., 6087,6887,7328 (Ma). Greenway, P.J., 2285,5337, 6660 (Ma). Guého,J., MAU (Ma). Hansen, O.J., 521 (Ma). Hilliard, O.M., 8549 (Gk). Hilner, O., 290 (Gk). Höft, M.G., 8 (Ma). Irvine, F.R., 1905 (Ma). Jayaweera, D.M.A., 1154 (Ma). Johnson, W.H., 275, 301 (Ma). Kapp, L.M., 115 (Gk). Keay, R.W.J., FHI 28663,28670 (Ma). Keet, J.D., 503,585 (Gk). Kemp, E.S., 1312 (Gk). Kostermans, A.J.G.H., 25299A, (Pece). Laughton, F.S., 8407, 8408 (Gk).. Leeuwenberg, A.J.M., 9834, 10568, (Ma), (Gk). Leighton, 236 (Gk). Liebenberg, L.C.C., 7955 (Gk). Luja, 479 (Ma). Luke,Q., 1729, 1840 (Ma). Macedo,2385,3090,3667 (Ma). MacOwan, P.,534, 1055 (Gk). Magogo, F.C., 1206 (Ma). Marloth, R., 695 (Gk). Masterton, G.M.,466 (Ma). 112 Wageningen Agricultural University Papers 97-2 (1997)

111 Mavi, S., 656 (Ma). McGregor, G.M.,42/48 (Ma). Mendonça,F.A., 1344,2478 (Ma). Merwe,C.J. van der, 4 (Gk). Mgaza, CD., 95 (Ma). Milne-Redhead, E., 7099 (Ma). Mogg,A.O.D., 1188 L(Gk). Müller,T.,562, 1121 (Ma). Mullin,L., 184/63 (Ma). Nicholson, H.B., 1931 (Gk). Oltmans,M.,87 (Pece). Omino,E.A., 8 (Ma). Padwa, J.H., 300 (Ma). Paol, S., 5 (Ma). Patel, I.H., 857 (Ma). Paterson, F.V., 2417, (Gk). Penther, 2004 (Gk). Perdue, R.E., (Ma). Pereira,A., 851 (Ma). Peter, A., 39908, 40253, 58123, 58124, 58127, 58129, 58140, 58172, 58288, (Ma). Phillips, J.F.V., 7355 (Gk). Pierre, L., 244,3584d.d (Ml). Pillans, N.S., (Gk). Pobéguin, H.,95 (Ma). Pole Evans, I.B., 5875,5971 (Ma). Procter,4505 (Ma). Rail,R.W., 1/55 (Ma). RB (Ma). Rechinger, K., 2294 (Pece). Rehmann, A., 197,358 (Gk). Robertson, S.R.,5175 (Ma). Rodin, R.J., 1163 (Gk). Rogers, RA., (Gk). Ruffo, 1034 (Ma). Sacleux,2205 (Ma). Salubeni,A.J., 1643 (Ma). Schlechter, R., 5884 (Gk). Schlieben, HJ., 1468 (Ma). Semsei, S.R., 902, 905, 1134, 1522 (Ma). Setten,K.van, 835 (Ma). Sim,T.R., 124 (Gk),6021 (Ma). Simâo,J., 281, 783 (Ma). S0rensen, T., 402 (Gk). Stables,J.H., 3/55 (Ma). Steenis, C.G.G.J. van, (Gk). Story, R., 3238 (Gk). Wageningen Agricultural University Papers 97-2 (1997) 113

112 Strey, R.G., 7317 (Gk). Sumithraarachchi, D.B., 660 (Pece). Swynnerton, C.F.M., 558 (Ma). Taylor, H.C., 2924 (Gk). Thode,J., 571, A 960 (Gk). Thwaites,CP 1835 (Pece). Tinley, K.L., 2566 (Ma). Torre, A.R., 654, 1110,3070, 4007, 4723, 5207 A, 5909, 10519, 14490, 16168, (Ma). Troughton, S.C., 169 (Gk). VanderLinden,L., 169 (Ma). Vaughan, R.E., 88, 774, 2051, MAU 11586,12023 (Ma). Verdcourt, B., 1731 (Ma). Vermoesen,F.,2162 (Ma). Waas, S., 18 (Pece). Wagner, R.J., 1145 (Ma). Warnecke,O.,481 b (Ma). Wells, M.J., 3372 (Gk). Whitish, 163 (Gk). Whyte, A., 108, 155 (Ma). Wild, H., 3529, 3577, 5261, 5722 (Ma). Williams,R.O., 99 (Ma). Winkler, H.,3699 a (Ma). Woejantoro, 137 (Ma). Wood, J., 3352 (Gk). Worthington, T.B., 3520,4163, 7094 (Pece). Wijk, A.E. van, 3186 (Gk). Zeyher, 730, 1055,3413 (Gk). 114 Wageningen Agricultural University Papers 97-2 (1997)

113 Index of scientific names New names in bold face, synonyms in italics. Page numbers of principal entries are in bold face, those of figures and maps in italics. Alafia Thou. 9 thouarsii Roem. &Schult.52 AlstoniaR.Br. longifolia (A.DC.)Pichon 101 Alyxia BanksexR. Br. 7, 8,54 ceylanka Wight54,59 erythrocarpavatke 64 lucida Bak. 64, 68 madagascariensisa.dc. 68, 71 poly sperma ScottElliot 71 AlyxieaeG.Don 7 AlyxiinaePichonexLeeuwenberg 7 AncylobotrysPierre 9 Apocynoideae 8 AspidospermaMart. & Zucc. 96 Cabucala Pichon 5, 53, 54, 55,80 brachyanthapichon80 caudata Markgr. 64,68 crassifolia Pichon 74, 76,77 cryptophlebia (Bak.)Pichon62 erythrocarpa (Vatke)Markgr. 55,64 var. angustifolia (Pichon)Markgr. 64 var. intermedia (Pichon)Markgr. 64 fasciculata Pichon 73, 75, 77 glauca Pichon 73, 75, 77 humbertii Markgr.69 intermedia Pichon 73,76 longipes Pichon 74, 76 macrophylla Pichon 73, 76, 77 var. acuta Markgr. 74, 75,77 var. macrophylla 73 var. oxyphylla Markgr. 74,76 madagascariensis (A.DC.)Pichon 73,71 var. acuta (Markgr.)Markgr.74 var. amygdalifoliamarkgr.exboiteau 74 var. angustifolia Pichon 64,67 var. intermedia Pichon 64, 68 var. latifoliapichon 64 var. longipes (Pichon)Markgr.exBoiteau 74 var. madagascariensis71 monarthron Pichon 64, 68 montana Pichon78 multiflora Pichon 73,76, 77 oblongo-ovata Markgr. 64, 68 Wageningen Agricultural University Papers 97-2 (1997) 115

114 penduliflora Markgr. 74,76 plectaneiifoliapichon79 polysperma (ScottElliot)Pichon 71,76,77 strioûtta Pichon 64, 68 torulosa Pichon 71, 75, 77 Carissa L.9,11 campenonii (Drake)Palacky80 cryptophlebia Bak.60 pichoniana Leeuwenberg 5, 10,101 verticülata Pichon 5,101 CatharanthusG.Don 8 CerberaL. 9 CraspidospermatinaeA.DC. 8 Craspidospermum Boj.ex A. DC. 5,6,7, 8, 10,11 verticillatum Boj.exA.DC. 6,11, 12,13 var. petiolare A. DC. 11, 15 var. sessile Markgr. 13,15 DidieraceaeDrake 44 Dyera Hook.f. 8 Echitella Pichon21 lisianthiflora (A.DC.)Pichon 21, 41 perrieri (Lassia)Pichon 43 EllertoniaWight madagascariensis Radlk.64, 71 ExacumL. vaginale Labill102 Gonioma E.Mey. 5,6, 8, 11,16 kamassi E.Mey.6, 16,17, 75 var. brachycarpa E.Mey. 17,19 malagasymarkgr. & Boiteau 17,19, 20, 80 Gynopogon Forst. erythrocarpus (Vatke)K.Schum. 64 madagascariensis (A.DC.)O.Kuntze71 Holarrhena R.Br. madagascariensis Bak. 24,33 Hunteria Roxb.59 Jasminonerium L.exO.Kuntze cryptophlebium (Bak.)O.Kuntze60 KamettiaKosteletzky 8 KibataliaG.Don 8 Landolphia P.Beauv. 9 myrtifolia (Poir.)Markgr.54 Lanugia N.E. Br. 21 latifolia N.E.Br. 21, 28,33 micrantha (Bak.) N.E.Br. 24 variegata (Britt. &Rendle)N.E.Br. 24 Logania R.Br. 102 populifolia (Poir.)Leeuwenb. 102 vaginalis (Labill.)F.v. Muell.102 MalouetiaA.DC. 59 Malouetiinae (Muell. Arg.)Pichon 8 Mascarenhasia A. DC. 5,6, 8, 11, 21, Wageningen Agricultural University Papers 97-2 (1997)

115 anceps Boiv. ex Jum. 24, 32 angustifolia A. DC. 24, 31, 34 var. keraudreniana Markgr. 28, 32, 34 arborea Boiv. ex Dubard 27, 32, 33 arborescens A. DC. 6,18, 21, 23, 24, 25, 26, 27, 34, 121 var. anceps (Boiv. ex Jum.) Lassia 24 subsp. angustifolia (A. DC.) Boiteau 24 subsp. arborescens 24 f. albicorticis Boiteau 28 f. arborea (Boiv. ex Dubard) Boiteau 27 f. longifolia (Jum.) Boiteau 27 var. micrantha (Bak.) Boiteau 24 var. boivinii (Dubard) Markgr. 28 var. comorensis Markgr. 28, 31 var. coriacea (Dubard) Lassia 27 var. gracilis Markgr. 28, 32, 34 f. longifolia (Jum.) Jum. & Perr. 27 var. longifolia (Jum.) Lassia 27 barabanja Dubard 28, 33 boivinii Dubard 28, 33 brevituba Vatke 38,41 causticak. Schum. 24 coriacea Dubard 27, 33 curnowiana Hort. 52 elastica K. Schum. 24 fischeri K. Schum. 24 geayi Cost. & Bois. 43, 47 gerrardiana Bak. 52 grandidieri Dubard 28, 32 havetii A. DC. 6, 23, 34, 35, 36, 38 humblotii Dubard 43, 46, 47 kidroa Cost. & Bois. 43 lanceolata A. DC. 5, 6, 23, 27, 38, 39, 121 lisianthiflora A. DC. 5, 6, 21, 24, 36, 41, 42, 47, 722 var. baronica Dubard 43, 47 subsp. geayi (Cost. & Bois.) Boiteau 43 var. hybrida Dubard 43,47 subsp. macrocalyx (Bak.) Boiteau 41 var. pubescens Dubard 41, 47 longifolia Jum. 27, 31 macrocalyx Bak.41, 47 macrosiphon Bak. 6, 22, 23, 26, 36, 47 mangorensis Jum. & Perr. 48, 49 maroana Aug. DC. 36, 37, 38 micrantha Bak. 24, 33 pallida Dubard 43, 46 parvifolia Dubard 38,41 perrieri Lassia 43, 45 phyllocalyx Dubard 43, 47 rosea Bak. 38, 41 rubra Jum. & Perr. 6, 24, 27, 39, 49 rutenbergiana Vatke 41 Wageningen Agricultural University Papers 97-2 (1997) 117

116 sessilifolia Dubard 24, 47 speciosa Scott Elliot 6, 22, 23, 27, 35, 5 0 var. dextra Dubard 50, 51 var. havetii (A. DC.) Boiteau 36 tampinensis Pichon 6, 22, 23, 27, 39, 51 tenuifolia Dubard 43,47 thiryana Pierre ex Dubard 36, 38 Mrife Bak. 38, 41 variegata Britt. & Rendle 24, 30 velutina Jum. 41, 45 Oncinotis Benth. 9 Pachypodium Lindl. 9 Petchia Livera 5, 6, 7, 8, 10, 53, 54, 55, 59 africana Leeuwenb. 5, 6, 53, 55, 56, 57 ceylanica (Wight) Livera 53,54,55,56,59 cryptophlebia (Bak.) Leeuwenb. 7, 53, 55, 56, 60, 61, 62 erythrocarpa (Vatke) Markgr. 5, 6, 7, 54, 56, 62, 64, 65, 69, 71, 77 humbertii (Markgr.) Leeuwenb. 7, 53, 55, 62, 69, 70 madagascariensis (A. DC.) Leeuwenb. 6, 7, 53, 54, 55, 56, 70, 71, 72, 73, 77, 123, 124 montana (Pichon) Leeuwenb. 7, 54, 55, 61, 73, 78 plectaneiifolia (Pichon) Leeuwenb. 7, 53, 54, 55, 70, 79, 80 Plectaneia Thou. 5, 6, 7, 8, 9, 81 boivinii Jum. 89,91,94 breviloba Markgr. 90, 93, 94 elastica Jum. & Perr. 89, 92, 95 f. firingalavensis Jum. & Perr. 89, 92 var. insularis Markgr. 90, 92 var. inutilis (Jum. & Perr.) Markgr. 88, 94, 95 f. hirsuta (Jum.) Markgr. 89 firingalavensis (Jum. & Perr.) Jum. 89, 95 var. firingalavensis 90 f. setulosa (Markgr.) Markgr. 90 f. setulosa Markgr. 90 var. lanceolata (Pichon) Markgr. 90 hildebrandtiik. Schum. 86, 92, 95 f. hirsuta Jum. & Perr. 88, 92 inutilis Jum. & Perr. 88, 92 var. hirsutajum. 89, 92, 93, 95 f. latiflora Jum. & Perr. 89, 92 isalensis Jum. 89, 94, 95 f. glomerata Jum. 89, 93 lanceolata Pichon 90, 94 longisepala Markgr. 7, 80, 82, 83 macrocarpa Jum. 90, 93 microphylla Jum. & Perr. 89, 92, 94 var. tsaratanensis Jum. 89, 92 pervillei K. Schum. 88 rhomboidalis Jum. & Perr. 88, 91, 95 stenophylla Jum. 7, 80, 82, 84, 85 thouarsii Roem. & Schuit. 5, 6, 7, 82, 86, 87, 88, 94, 95,124 var. macrocarpa (Jum.) Markgr Wageningen Agricultural University Papers 97-2 (1997)

117 tsaratanensis (Jum. &Perr.) Pichon 89 volubilis Poir. 86 PlumerieaeEndl. 8 Plumerioideae K.Schum.7, 8 PulassariumO.Kuntze madagascariense (A.DC.)O.Kuntze 71 Rauvolfia L.10 celastrifolia Bak. 99,101 Rauvolfiinae Benth. &Hook.f. 7 Rubiaceae A.L.Juss. 9 SabaPichon 9 Schizozygia Baill.10 Stephanostegia Baill. 5,7, 8, 11, 95, % ferevis Markgr- 97, 99 capuroniimarkgr. 6,7, SS, 97, 98, 99 hildebrandtii Baill. 7,SS, 95, 97, 98, 99, 101 holophaea Pichon 99,100 var. parvifolia (Pichon)Markgr.99 megalocarpa Markgr. 100,101 parvifolia Pichon 99,100 Strempeliopsis Benth. 8 Strophanthus DC. 8,10 Tabernaemontana L. 11,101 camassi Eckl.17 pandacaqui Poir. 101 populifolia Poir. 101 Tsüaitra Baron21 lanceolata (A.DC.)Baron 24,38 micrantha (Bak.)Baron 21, 24 VoacangaThou.11 WrightieaeG.Don 8 Wageningen Agricultural University Papers 97-2 (1997) 119

118 Phot. 1. Mascarenhasia arborescens. Phot,and herb. Leeuwenberg (14689). Phot. 2. Mascarenhasia Umceolata. Phot, and herb. Leeuwenberg (14260). Wageningen Agricultural University Papers 97-2 (1997) 121

119 Phot. 3. Mascarenhasia lisianthiflora. Phot,andherb. Leeuwenberg (14685). ] Phot. 4. Mascarenhasia lisianthiflora. Phot, and herb. Leeuwenberg (14690). 122 Wageningen Agricultural University Papers 97-2 (1997)