The Genus Antidesma (Euphorbiaceae) in Madagascar and the Comoro Islands

The Genus Antidesma (Euphorbiaceae) in Madagascar and the Comoro IslandsAuthor(s): Petra HoffmannSource: Kew Bulletin, Vol. 54, No. 4 (1999), pp. 877-885Published by: Springer on behalf of Royal Botanic Gardens, KewStable URL: http://www.jstor.org/stable/4111165 .

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KEW BULLETIN 54: 877 - 885 (1999)

The genus Antidesma (Euphorbiaceae) in Madagascar and the Comoro Islands

PETRA HOFFMANN1

Summary. The genus Antidesma (Euphorbiaceae-Phyllanthoideae) in Madagascar and the Comoro Islands is revised. The two endemic species as well as their three varieties and two forms hitherto recognised were found to be conspecific with A. madagascariense, a species earlier considered to be endemic to the Mascarenes. A. madagascariense var. hildebrandtii, comb. nov., is accepted as the only infraspecific taxon. It differs from the type variety by its tomentose floral disc and is confined to northern Madagascar.

INTRODUCTION

Antidesma Burm. ex L. is a genus of dioecious shrubs and trees commonly found in the understorey of tropical rain forests as well as in shrubby vegetation. Its distribution ranges from West Africa to the Pacific Islands and from the Himalayas to Northern Australia, with the centre of diversity in the Malesian region. No

convincing infrageneric classification has yet been proposed for the c. 100 currently recognised species.

The genus belongs to the subtribe Antidesminae in the subfamily Phyllanthoideae. The most notable character of this subtribe is the elongated, U-shaped connective of the anthers, which is found nowhere else in the Euphorbiaceae. Other members of this subtribe in Africa are Thecacoris A. Juss. and the Madagascan endemic Leptonema A. Juss. In addition to its peculiar anthers, Antidesma is distinguished by racemose inflorescences, apetalous flowers with a floral disc in both sexes and unilocular

drupes with a characteristically foveolate endocarp. The endocarp sculpturing and its resemblance to some genera of the Icacinaceae caused the segregation of Antidesma as the monogeneric family Stilaginaceae by Airy Shaw (in Willis 1966: 1076). This segregation cannot be maintained in view of the evidence available

today. Morphological (Hoffmann, in prep.) and molecular studies (Wurdack & Chase, ined.) show that Antidesma is well placed in the subfamily Phyllanthoideae (cf. Webster 1994: 51).

The high morphological and ecological variability of the genus Antidesma has

always presented considerable taxonomic difficulties. This is also the case in Madagascar, the Comores and the Mascarenes, as is indicated by the number of

species and infraspecific taxa described from these islands and the discrepancies between the existing taxonomic treatments (Pax & K. Hoffmann 1922: 112 - 167; Leandri 1937: 23 - 28 and 1958: 14-21). The general lack of consistent

Accepted for publication May 1999. 1 Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, U.K., [email protected]

877



878 KEW BULLETIN VOL. 54(4)

morphological features led to the use of vague vegetative characters such as leaf shape, texture and colour for species delimitation; these characters are highly variable in the genus. The great variability, on the other hand, resulted in the adoption of narrow species concepts, which are unsuited for this genus of remarkable ecological amplitude (cf. Baker et al. 1998: 255).

In the Flore des Mascareignes, Coode (1982: 106 - 109) remarked on the high variability in Mascarene Antidesma in characters such as leaf shape and size, calyx lobes, indumentum of the inflorescences and presence or absence of leaf domatia. However, this variation was found to be continuous and morphological extremes were growing in the same place. He consequently recognised only one taxon, A. madagascariense Lam., in the Mascarenes and also commented that the material from the Mascarenes, Madagascar and the Comores might be conspecific.

A. madagascariense had originally been described by Lamarck (1783: 206) from Madagascar, but for some inexplicable reason Leandri (1937: 23 - 28 and 1958: 14 - 21) applied the name exclusively to plants from the Mascarenes. There is, however, no morphological reason to suspect that the only extant authentic specimen of A.

madagascariense was not collected in Madagascar. In the most recent taxonomic treatment of Antidesma in Madagascar and the Comoro Islands, Leandri (1958: 14 - 21) recognised two species (A. hildebrandtii Pax & K. Hoffm. and A. petiolare Tul.) with three varieties and two forms, some based on the type specimen alone. These taxa are distinguished by overlapping and inconsistent characters, making it difficult to name specimens.

In the present study, material of Antidesma from the Mascarenes was compared with material from Madagascar and the Comoro Islands. Collections from the Mascarenes more often tend to have coriaceous leaves and longer staminate pedicels (0.3 - 1.5 mm), compared to (0 -)0.2 - 0.5(- 1) mm measured in the collections from Madagascar and the Comoros. The petioles are 'on average shorter in the Mascarenes but range within the same limits. These differences are not clear enough to deserve taxonomic recognition.

When examining the abundant herbarium material of Antidesma from Madagascar and the Comoros, the indumentum of the floral disc was found to be the only consistent diagnostic character. The disc is glabrous in the majority of the

specimens, but distinctly tomentose in a number of collections from the area between Nosy Be and the Masoala Peninsula in northern Madagascar, including the type of A. hildebrandtii var. hildebrandtii. However, the disc indumentum is not correlated with any other discrete morphological difference and the two forms are

sympatric (e.g. Keraudren 1583 and Deroin & Badri 186, both from Nosy Be). Therefore this taxon is recognised here at the level of variety.

As a result, only one species, A. madagascariense Lam. is recognised here for

Madagascar, the Comores and the Mascarenes. The necessary new combination is made for the single infraspecific taxon, var. hildebrandtii. Five species are recognised in the recent treatments of Antidesma in continental Eastern Africa (Radcliffe-Smith 1988: 572- 576; Leonard 1995: 16 - 40; Radcliffe-Smith 1996: 105- 110). With the exception of A. laciniatum Muill. Arg. these species seem to be closely related to A. madagascariense.



ANTIDESMA IN MADAGASCAR & THE COMORO ISLANDS 879

TAXONOMY

Antidesma Burm. ex L., Sp. PI.: 1027 (1753); Gen. PI., ed. 5: 451 (1754); Lamarck, Encycl. 1: 206 (1783); Tulasne, Ann. Sci. Nat. Bot., Ser. 3: 182 (1851); Baillon, Etude

Euphorb.: 601 (1858); in DC., Prodr. 15(2): 247 (1866); Benth. & Hook. f., Gen. pl. 3: 284 (1880); Pax & K. Hoffm. in Engl., Pflanzenr. 81: 107 (1922); Leandri, Notul.

Syst. (Paris) 6: 23 (1937), in Humbert, Fl. Madag. 111 (1): 14 (1958); Coode in Bosser et al., Fl. Mascareignes 160: 106 (1982); Radcliffe-Smith in Polhill, F.T.E.A.:

Euphorb. 2: 572 (1988); Webster, Ann. Missouri Bot. Gard. 81: 52 (1994); Leonard in Fl. Afr. Centr., Euphorb. 2: 16 (1995); Radcliffe-Smith in Pope, Fl. Zambesiaca 9

(4): 105 (1996).

Type: A. alexiteria L. (from India)

Dioecious trees or shrubs with a simple indumentum. Leaves alternate, petiolate, simple, eglandular, blades symmetrical, entire, pinnately veined, not decurrent.

Stipules caducous. Inflorescences racemose, axillary. Bracts 1 per flower. Pedicel short to absent, inarticulate. Sepals 3 - 5, imbricate in bud, fused to various degrees. Petals absent. Disc in staminate flowers various, cushion-shaped (enclosing the bases of the stamens and pistillode), dissected to various degrees or extrastaminal- annular, in pistillate flowers annular. Stamens 3(- 4), usually episepalous, filaments free, anthers extrorse in bud, versatile, connective elongated, U-shaped, thecae

resembling swollen ends of the 'U', raised at anthesis, longitudinally dehiscing. Pistillode present. Ovary 1-locular. Stigmas 3 - 8, acute. Ovules 2, anatropous, with individual obturators. Fruits drupaceous, edible, red to black, ellipsoid to lens-

shaped, laterally compressed, base symmetrical, style terminal, rarely subterminal.

Endocarp lignified, foveolate, the sculpture usually obvious through the dried

mesocarp. Seeds ecarunculate, 1 per fruit (rarely two, then fruits twice as big). Endosperm in mature seeds present, cotyledons thin, flat, several times wider and

longer than radicle.

A palaeotropical genus of about 100 species, with 7 species in continental Africa.

Antidesma madagascariense Lam., Encycl. 1: 206 (1783) as 'madagascariensis'; Baker, Fl. Mauritius: 305 (1877); Pax & K. Hoffm. in Engl., Pflanzenr. 81: 134

(1922); Leandri, Notul. Syst. (Paris) 6: 24 (1937); Coode in Bosser et al., Fl.

Mascareignes 160: 106 (1982). Type: Madagascar(?), 'bois de mafoutre', Sonnerat(?) s.n. (lectotype P-LA! (chosen here), microfiche IDC 6207: LM 588/12).

As discussed by Coode (1982: 106), the lectotype designated here is the only specimen eligible for typification of this name to be found in the Paris herbarium

(both general and historical collections). It bears the vernacular name 'bois de mafoutre' which was mentioned by Lamarck in the protologue, but no collector, number or locality. It corresponds with the protologue in having pistillate flowers and fruits.

Shrub or tree, up to 20 m, clear bole up to 10 m, diameter up to 70 cm. Bark smooth. Wood very hard. Young twigs terete, striate or obtusely angled, glabrous to




pubescent, greyish brown. Leaves persistent or deciduous; lamina (2 -)4 - 10(- 20) cm long, (1 -)2 - 5(- 8) cm wide, 1.5 - 5 times longer than wide, elliptic, ovate, obovate or oblong, apically acuminate-mucronate to rounded or slightly retuse, basally acute to rounded, membranaceous to chartaceous, more rarely coriaceous, glabrous, or pilose to pubescent along the midvein adaxially and/or along the major veins abaxially, sometimes also along the margin, sometimes sparsely pilose all over abaxially, moderately shiny, midvein impressed to flat, rarely slightly raised adaxially, tertiary venation reticulate, in larger leaves weakly percurrent, drying olive-green to reddish brown, discolourous or concolourous, domatia in axils of midvein and secondary veins usually present, hairy or glabrous, more often absent in coriaceous leaves. Petioles channelled adaxially, 2 - 15 x 0.5 - 1.5 (- 2) mm, the

longer ones rarely basally and apically slightly swollen, sparsely pilose to pubescent, glabrescent when older. Stipules caducous, linear to deltoid, apically acute, (1 -)4 - 8 x 0.5 - 1 mm, pilose to pubescent. Staminate inflorescences 2 - 8 cm long, axillary, erect, simple, sometimes up to 15 racemes clustered at the leafless tip of branches and giving the appearance of a branched inflorescence, axis pilose to pubescent. Bracts ovate to lanceolate, apically acute, 0.5 - 1 x 0.3 - 0.4 mm, pilose to pubescent. Flowers 2 mm x 1.5 - 2.5 mm. Pedicels (0 -)0.2 - 0.5(- 1) mm long, glabrous to pilose. Calyx 0.5 - 1 x c. 1 mm, cup-shaped, sepals 3 (- 4), fused for half their length or more, apically acute to rounded, glabrous outside, glabrous inside but often with long hairs at the base, margin entire to erose, sometimes fimbriate. Disc cushion-shaped (enclosing the bases of the filaments and the pistillode) or extrastaminal-annular, lobed towards the centre, the lobes filling the space between the filaments, or sometimes irregularly dissected, glabrous or densely tomentose. Stamens 3(- 4), 1 - 2 mm long, 0.8 - 1.5 mm long exserted from the

disc, anthers 0.3 - 0.5 x 0.4 - 0.7 mm. Pistillode cylindrical, conical or ovoid, 0.3 - 0.8 x 0.2 - 0.7 mm, inserted between the filaments inside the extrastaminal or dissected disc, or on top of the cushion-shaped disc, glabrous to pilose. Pistillate inflorescences 1.5 - 5 cm long, axillary, suberect, simple, sometimes up to 3 racemes clustered at the leafless tip of branches and giving the appearance of a branched inflorescence, sometimes in fascicles of 2, axis pilose to pubescent. Bracts ovate to lanceolate, apically acute, 0.3 - 0.7 x 0.2 - 0.5 mm, pilose to pubescent. Flowers 1.5 - 2 x 0.7 - 1 mm. Pedicels 0.5 - 1 mm long, pilose. Calyx 0.5 - 1 x 0.5 - 1.2 mm, cup- shaped, sepals 3 - 5(- 6), fused for /:3 - '2/ of their length, sometimes unequal, apically acute to rounded, glabrous to pilose outside, glabrous inside but with hairs at the base, margin erose to entire, sometimes fimbriate. Disc 0.2 - 0.3 mm long, not exserted from the calyx, glabrous or tomentose, margin entire to crenulate.

Ovary lens-shaped to ellipsoid, glabrous to pilose, style terminal, rarely subterminal, stigmas 3 - 8. Infructescences 1.5 - 7 cm long, pendulous. Fruiting pedicels terete, 1 - 4 mm long, glabrous to pilose. Fruits ellipsoid to lens-shaped, laterally compressed, basally symmetrical, with a terminal, rarely subterminal style, 4- 7 x 3 - 5 mm, areolate, glabrous, rarely sparsely pilose, white-pustulate or not, light green, maturing through white, red and purple to black.

Specimens with subterminal styles have been collected in the districts of Toamasina (Tamatave) (e.g. 9735-RN) and Analamazaotra (Perinet) (e.g. 14966-SF)




only. They tend to have shiny, coriaceous leaves about 7 x 3 cm in size without domatia. There are, however, collections from this area (e.g. 3319-SF from Perinet) with the same leaf characters and perfectly terminal styles.

Like other species in the genus, A. madagascariense can be subject to galling. In both var. madagascariense (e.g. 9735-RN) and var. hildebrandtii (e.g. 7684-SF) glabrous, ellipsoid galls 5 - 15 x 3 - 7 mm in size are occasionally found along the

petiole and the midvein on the abaxial leaf surface. The deformation of the

pistillate inflorescences which is common in some Antidesma species from continental Africa (Leonard 1995: 17; Radcliffe-Smith 1996: 105, 106) has not been observed in the Madagascan material.

KEY TO VARIETIES

Floral disc glabrous .............................. var. madagascariense Floral disc tomentose .......................... ...... . . var. hildebrandtii

var. madagascariense

A. petiolare Tul., Ann. Sci. Nat. Bot., Ser. 3: 207 (1851) (non Airy Shaw 1972), Pax & K. Hoffm. in Engl., Pflanzenr. 81: 130 (1922), Leandri in Humbert, Fl. Madag. 111 (1): 17 (1958), synon. nov. Type: Madagascar sept., in montibus inter Tananarivo et Itas, Febr. 1840, Bernier 271 (lectotype (here designated) and

isolectotype P!). A. erythroxyloides Tul., Ann. Sci. Nat. Bot., Ser. 3: 208 (1851); Baill. in Grandid., Hist.

Phys. Madagascar, Atlas 2: t. 214 (1892); Pax & K. Hoffm. in Engl., Pflanzenr. 81: 134 (1922). Type: Madagascar, Boivin 2369 (lectotype P! (here designated); isolectotype G-DC, P!).

A. alnifolium Baker, J. Linn. Soc., Bot. 22: 519 (1887), nom. illeg. (non Hook. 1842), as 'alnifolia'. Type: Madagascar, Baron 4666 (holotype K!, isotype P!).

A. brachyscyphum Baker, J. Linn. Soc., Bot. 22: 518 (1887), as 'brachyscypha'; Pax & K. Hoffm. in Engl., Pflanzenr. 81: 166 (1922).

A. petiolare Tul. var. brachyscyphum (Baker) Leandri, Notul. Syst. (Paris) 6: 27 (1937), in Humbert, Fl. Madag. 111 (1): 18 (1958), as 'brachyscypha', synon. nov. Type: Madagascar, Baron 4447 (holotype K!, isotype P!).

A. comorense Vatke & Pax: 529 (1893), Pax & K. Hoffm. in Engl., Pflanzenr. 81: 134 (1922), pro syn.

A. hildebrandtii Pax & K. Hoffm. var. comorense (Vatke & Pax) Leandri in Humbert, Fl.

Madag. 111 (1): 17 (1958), synon. nov. Type: Comoren, Insel Johanna, in feuchten Thdilern des Tieflandes, June -Aug. 1875, fem., Hildebrandt 1669

(lectotype K! (here designated)). Leandri (l.c.) cited var. comorense as 'J. Leand., loc. cit.', however, I could not find an earlier place of publication for this stat. nov. than indicated above.

A. petiolare Tul. var. perrieri Leandri, Notul. Syst. (Paris) 6: 28 (1937), in Humbert, Fl. Madag. 111 (1): 21 (1958), synon. nov. Type: Madagascar, Domaine de l'Est, savoka, bassin inferieur du Faraony, c6te est, 30 m alt., arbuste dioique de 4-5 m, pied femin., Perrier 9752 (holotype P!).




A. petiolare Tul. var. brachyscyphum (Baker) Leandri f. elliotii Leandri in Humbert, Fl.

Madag. 111 (1): 20 (1958), synon. nov. Type: Madagascar (est), Fort Dauphin, bois, juin, Scott Elliot 2814 (lectotype P! (here designated), isolectotype K!).

A. petiolare Tul. var. brachyscyphum (Baker) Leandri f. humbertii Leandri in Humbert, Fl. Madag. 111 (1): 20 (1958), synon. nov. Type: Madagascar (centre), contreforts occidentaux du massif du Marojejy (nord-est), pres du col de Doanyanala, foret

ombrophile, sur laterite de gneiss, 800 - 1200 m, janvier - f6vrier, Humbert 23044

(lectotype (here designated) and isolectotype P!).

For further synonymy (taxa described from the Mascarenes) see Coode 1982: 107.

Floral disc glabrous.

DISTRIBUTION. Throughout Madagascar, Comoro Islands and Mascarenes

(Reunion, Mauritius). ECOLOGY. COMORO ISLANDS: Humid forest on clayey soil, 500 - 700 m.

MADAGASCAR: Common throughout the island in both primary and secondary vegetation, from perhumid to subarid bioclimatic zone, mainly in savannah, xerophyllous bush and dry forest, but also on beaches, at roadsides, in deciduous forest, riparian and gallery forest, open swampy areas, moist montane forest and

mossy forest, on sand or lateritic soil, mostly over sandstone but also over gneiss, limestone orjurassic marl: 0 - 1800 m.

VERNACULAR NAMES. COMORO ISLANDS: Chivissa-Cheo (Mohelian), Kibissa, Mouebemba, M'Poua Soha, Schivissa cheou (Anjouanais). MADAGASCAR: Bois de Mafoutre, Hazoambo, Hazombaroa, Hazombato, Hazondomohina, Hazonovy, Hoditrova, Hoditrovy, Hoditrovy vavy, Karambito, Karepokala, Longotrafotsy, Menafony, Mizorotanty, Oditrovy (Betsimisaraka dialect), Taimboalavo, Taindalita, Taindalitra, Tainidalitra, Tsirivodrivotra, Tsirivodrovitra, Varana, Varoana, Varodalitra, Varona (Tsimihety dialect), Varongy, Varotra, Vavaromy, Vavarony, Voafogna, Voafona (Antandroy & Bara dialect), Voafono, Voafony, Voana, Voankarepoka, Voaromoromo, Voaromy, Voarona, Voarony, Vofoana, Vofongo, Vogna, Vofia (Bara dialect), Voromorona, Voroma (Tandroy dialect), Voromy (Tandroy dialect), Vovona (Bara dialect), Vovonana.

USEs. Wood used as fire wood, for construction, fence posts and carpentry. The fruits are eaten by guinea fowl (26616-SF), Lemur catta (Sauther 17) and humans (Leandri 1958: 18, and several specimen labels); their taste resembles Prunus mahaleb L. and they are locally distilled to produce a good 'kirsch' (Perrier de la Bdthie 53). The fruits are also used to gain a red or black dye for fabrics, and an infusion of the bark is used as a remedy for abdominal pain (Herb. Jard. Bot. Tan. 5460). The boiled leaves and stem are said to be effective in the treatment of open wounds (19704-SF).

SELECTED SPECIMENS. COMORO ISLANDS: Anjouan, Sima canton, Boungueni, 9 March 1957, Service des Eaux et Forits 16649-SF (P!); Mayotte, Benara, 29 Feb. 1996, Pascal 398 (K!, P!); Moheli, entre Fomboni et Drondroni, 500 - 700 m, for~t, 6 Dec. 1967, Bernardi 11782 (K!, P!). MADAGASCAR: Antananarivo Province, Presqu'ile d'Amboniara, bord du Lac Itasy, 18 Feb. 1971, Jacquemin 882 (P!); Antsiranana Province, au sud d'Antsiranana, Reserve speciale d'Ankarana, c6te Ouest, 12?49'S,




49001'E, 50 - 409 m, 12 - 20 Oct. 1993, Andrianantoanina 377 (MO, P!); Nosy Be, reserve du Loucoube (Lokobe), March 1962, Keraudren 1583 (P!); Fianarantsoa Province, environs d'Ambatofinandrahana, 1600 - 1800 m, 18 Feb. 1938, Decary 13050 (P!); Ihosy, restes de for&t xerophylle, 6 Aug. 1970, Vovonana 2107 (P!); Mahajanga Province, Presqu'ile Radama, Maromandra, 11 Oct. 1922, Decary 1149 (P!); Andranomavo, 27 Sept. 1930, Decary 8139 (P!); massif de l'Ankarafantsika, 8

Jan. 1938, Decary 12839 (P!); Toamasina Province, Lac Alaotra, Herb. Jard. Bot. Tan. 3896 (P!); Nosy Mangabe, 27 March 1941, Decary 16876 (P!); Toliara Province, For&t Besaka Zombitsy, 15 - 20 km E of Sakaraha, route 7, 730 - 750 m, 4 Feb. 1975, Croat 30674 (MO, P!); Tolafiaro pref. (Fort-Dauphin), plage de Mandena, 0- 10 m, 23 Jan. 1990, Dumetz 1231 (MO, P!); Marosalaza, 50 km N of Morondava, 0 m, closed deciduous forest, 16 Nov. 1973, Petter 17 (P!). More than 300 specimens examined.

ILLUSTRATIONS. Baill. in Grandid., Hist. Phys. Madagascar, Atlas 2: t. 214 (1892) habit, fruit, endocarp, section of fruit and seed; Leandri in Humbert, Fl. Madag. 111 (1): 19, fig. 4.7 - 19 (1958) habit, flowers, fruits; Coode in Bosser et al., Fl. Mascareignes 160: 108, pl. 21.1 - 11 (1982) habit, leaves, flowers, fruits.

var. hildebrandtii (Pax & K. Hoffm.) Petra Hoffm. comb. nov.

A. hildebrandtii Pax & K. Hoffm. in Engl., Pflanzenr. 81: 122 (1922); Leandri in Humbert, Fl. Madag. 111(1): 16 (1958), excl. var. comorense (Pax & Vatke) Leandri. Type: Madagascar, Nosi Komba, Hildebrandt 3254 (lectotype P! (here designated), isolectotypes K!, P!).

Floral disc tomentose.

In the typical variety, there are often hairs at the inner base of the sepals and around the pistillode which can give the wrong impression of a hairy disc. Determinations of var. hildebrandtii should therefore be verified by dissecting a softened flower or fruiting pedicel. The leaf blades in this variety are elliptic or oblong to obovate with an acuminate apex and an acute base, (3.5 -)6 - 12(- 19) x

(1.5 -)2.5 - 4.5(- 7) cm, 1.7 - 3.5 times longer than wide, usually coriaceous and often completely glabrous.

DISTRIBUTION. Madagascar, southern Antsiranana and northern Toamasina

Province, between Nosy Be and the Masoala Penisula. ECOLOGY. Subhumid to perhumid bioclimatic zone of Cornet (1974); no habitat

information available. VERNACULAR NAMES. Taindalitra, Varona, Varodalitra, Varotra. SELECTED SPECIMENS. MADAGASCAR: Antsiranana Province, Nosy Be, Mont

Passot, 13'19'S, 48013'E, versant Nord-Est, pente herbeuse ensoleillee, 5 Dec. 1989, Deroin & Badre 186 (P!); Antalaha distr., Ambohitralanana canton, 15 May 1959, Jaofety Brosy 10837-RN (P!); vallee de Sambirano, Mont Tsitondroina, 1908, Perrier de la Bdthie 9908 (P!); Antalaha distr., Ambohitralanana canton, aux environs d'Amdrahimbazaha, Presqu'ile de Masoala, 15?17'S, 50027'E, O - 60 m, 21 April 1994, Rahajasoa 275 (MO, P!); Parc National de Masoala, Andrombazaha, 15?16'S, 50?29'E, 4 Oct. 1994, Rahajasoa et al. 743 (MO, P!); Ambanja distr., Manarabory, 21




Sept. 1953, Service des Eaux et Forets 7684-SF (P!); Ambanja, Marovato canton, 19

Aug. 1954, Tsiligy 6290-RN (P!); Ambanja, Beangona, 22 Nov. 1954, Tsiligv 6780-RN (P!); Toamasina Province, 2 - 3 km E of Sahavary which is 1 - 2 km E of the

Andranofotsy R. and approx. 15 km NE of Maroantsetra, 15?17'S, 49050'E, 50 - 400 m, 10 Feb. 1988, Schatz et al. 1877 (K!, MO). 15 specimens examined.

ILLUSTRATION. Leandri in Humbert, Fl. Madag. 111 (1): 15, fig. 3.8 - 14 (1958) leaves, flowers, fruits.

EXCLUDED NAME

A. arbutifolium Baker, J. Linn. Soc., Bot. 22: 519 (1887), as 'arbutifolia'; Pax & K. Hoffm. in Engl., Pflanzenr. 81: 121 (1922). Type: Madagascar, Baron 4062

(holotype K!) = Thecacoris madagascariensis A. Juss., Euphorb. Gen.: 105, pl. 1.1 (1824).

This name has been treated as incompletely known by Pax & Hoffmann (1922: 116, key). Leandri (1937: 24, 25) listed A. arbutifolium in his key as insufficiently known but assumed that it may be only a form or variety of Antidesma petiolare. Later he synonymised the name with A. petiolare without further comments (Leandri 1958: 17). Examination of the type in pistillate flower at Kew, however, has shown that this is Thecacoris madagascariensis. The two genera are similar both in staminate and

pistillate flower, but differ in the ovary, which is 3-locular and matures into an autochorous capsule in Thecacoris.

ACKNOWLEDGE MENTS

This study was financed by a grant from the Museum National d'Histoire Naturelle, Paris, for which I am thankful to Prof. Ph. Morat, director of the Laboratoire de Phanerogamie. I am indebted to the curators of the herbaria K and P for the opportunity to study their material, and to G. McPherson (MO), M. Coode (K) and S. Armbruster (Department of Botany, Norwegian University of Science and Technology, Trondheim) for their helpful comments on the manuscript.

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The Genus Antidesma (Euphorbiaceae) in Madagascar and the Comoro Islands

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