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A Synopsis of the Genus Beilschmiedia (Lauraceae) in Madagascar

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A Synopsis of the Genus Beilschmiedia (Lauraceae) in Madagascar A synopsis of the genus Beilschmiedia (Lauraceae) in Madagascar Henk van der WERFF Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. [email protected] ABSTRACT A review of the species of Beilschmiedia occurring in Madagascar is presented. Nine species are recognized, of which two, B. pedicellata van der Werff and KEY WORDS B. rugosa van der Werff, are described as new. Two previously described Beilschmiedia, species are treated as imperfectly known. Several collections with dimerous Potameia, Lauraceae, flowers typical of the related genus Potameia are accepted as belonging in Madagascar. Beilschmiedia. RÉSUMÉ Révision synoptique du genre Beilschmiedia (Lauraceae) à Madagascar. Une révision des espèces de Beilschmiedia vivant à Madagascar est présentée. MOTS CLÉS Neuf espèces sont reconnues dont deux nouvelles, B. pedicellata van der Werff Beilschmiedia, et B. rugosa van der Werff, décrites ici. Deux espèces demeurent imparfaite- Potameia, Lauraceae, ment connues. Le rattachement à Beilschmiedia de plusieurs spécimens à Madagascar. fleurs dimères, typiques du genre affin Potameia, est proposé. INTRODUCTION poorly represented in areas with a pronounced dry season. KOSTERMANS (1950) wrote a treat- This is the first of a planned series of articles ment of the family for the “Flore de Madagascar presenting revisions of the genera of Lauraceae in et des Comores”, based largely on a manuscript Madagascar. Lauraceae are a rather large, pre- he published in 1939. Several years later, dominantly tropical family of trees and shrubs KOSTERMANS (1957 a,b,c,d; 1958) published with 55 genera and about 3000 species. Cassytha, updates of the various genera with the description a genus of leafless, parasitic vines is the only non- of several new species, but he did not provide woody genus of Lauraceae. The family is well new keys. As a consequence, Lauraceae in represented in Madagascar by an estimated Madagascar remain difficult to identify. 100 species. Lauraceae are common in wet forest Beilschmiedia is a pantropical genus of about from sea level to the highest mountains, but are 250 species best represented in tropical Asia and ADANSONIA, sér. 3 • 2003 • 25 (1) : 77-92 © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. 77 van der Werff H. Africa. Characteristic for the genus are the flow- genera of Lauraceae based on DNA sequence ers with nine 2-celled stamens and the fruits, data (CHANDERBALI et al. 2001), three from which sit fully exposed on the pedicels. The small Madagascar and three from the Neotropics. tepals rarely persist at the base of the fruits. The These six species formed, together with the two inflorescences are cymose-paniculate, but in con- species of Potameia and the single species of trast to many genera of Lauraceae the flowers are Endiandra included in the study, a strongly sup- not arranged in strict cymes; instead, the lateral ported clade. The single species of Apollonias flowers of the cymes are somewhat alternate, not included in the study formed part of a clade con- strictly opposite (VAN DER WERFF 2001). sisting of Persea (represented by 6 species), Two species reported from Madagascar that are Phoebe, Dehaasia and Alseodaphne (each repre- currently placed in Beilschmiedia were originally sented by one species). These results confirm that described as monotypic, endemic genera. The first the Malagasy species are correctly placed in of these was Bernieria madagascariensis Baill. Beilschmiedia and that they have no close affinity (BAILLON 1884). This species was described as to Apollonias. having 6 fertile stamens; otherwise, it agrees with The Malagasy Beilschmiedia species can be Beilschmiedia. The fertile fragments of the holo- divided in three groups. The first includes species type represent immature flowers; more recent col- with opposite, glabrous leaves; its flowers have a lections with mature flowers show that this species short floral tube, with the tepals deciduous in old has 9 fertile stamens and does not differ from flowers (B. moratii, B. opposita, B. pedicellata and Beilschmiedia. The second endemic genus was B. sary). Members of the second group have alter- Thouvenotia, with the single species T. madagas- nate, pubescent leaves, and rather large flowers cariensis Danguy (DANGUY 1920). The generic with spreading tepals that lack a short floral tube diagnosis and species description fit the concept and whose tepals persist in fruit until they of Beilschmiedia. Neither Bernieria nor become damaged and break off (B. sericans and B. Thouvenotia was compared with Beilschmiedia. velutina). Species in the third group also have KOSTERMANS (1939) transferred these two species alternate leaves, but the flowers are small, with to Apollonias and described 4 new species in that erect or incurved tepals. As in the second group genus. Later he considered these 6 species better the flowers lack a short tube and the tepals persist placed in Beilschmiedia and published the at the base of the fruits (B. madagascariensis, required transfers (KOSTERMANS 1952). He did B. microphylla and B. rugosa). Characters of the not give specific reasons for this transfer, but sim- cuticle support this division in three groups and ply stated that he believed these species should be will be presented in a separate publication incorporated in Beilschmiedia. In 1957 (NISHIDA & VAN DER WERFF, in prep.). KOSTERMANS described three additional species of Beilschmiedia and Potameia, a genus of about Beilschmiedia. VAN DER WERFF (1996) published 20 species endemic to Madagascar, are closely one more species, thus bringing the total of pub- related (CHANDERBALI et al. 2001) and can be eas- lished Beilschmiedia species in Madagascar to ten. ily confused in the absence of flowers. When flow- In most species of Beilschmiedia the tepals fall ers are available, the difference between the two is off in the old flowers and leave the pistil fully readily seen: Beilschmiedia has trimerous flowers, exposed on the pedicel. In some Malagasy species whereas Potameia has dimerous flowers. When though the tepals persist at the base of the fruits, tepals persist at the base of fruits, fruiting material a character also found in Apollonias and which can also be identified in conjunction with the fol- may have led KOSTERMANS to place the Malagasy lowing vegetative characters: Potameia species species initially in Apollonias. However, Apollonias have alternate, mostly glabrous leaves and twigs and Beilschmiedia have different inflorescence with a pale bark; their tepals are often persistent in types (VAN DER WERFF 2001) and the Malagasy fruit, and the fruit is seated on a cushion-like base, species have inflorescences typical of whereas in Malagasy Beilschmiedia some species Beilschmiedia. Six species of Beilschmiedia were have opposite, glabrous leaves, young twigs with a included in a recent phylogenetic study of the nearly black bark and deciduous tepals in fruit 78 ADANSONIA, sér. 3 • 2003 • 25 (1) Beilschmiedia (Lauraceae) of Madagascar while others have alternate, pubescent or glabrous case, stamens can be assigned to whorl I or II by leaves, twigs with a brown bark and persistent their position: stamens of whorl I are opposite the tepals in fruit; the fruit is directly attached to the outer tepals and those of whorl II opposite the pedicel, without a base. I had initially assumed inner tepals. that the Malagasy species with opposite or subop- posite, glabrous leaves, dark twigs, dehiscent tepals and fruits not seated on a distinct base all BEILSCHMIEDIA Nees in Wall. belonged to Beilschmiedia, but several collections have come to my attention with these Pl. Asiaticae Rar. 2: 69 (1831). Beilschmiedia characters and dimerous flowers Bernieria Baill., Bull. Mens. Soc. Linn. Paris 1: 434 (1884). — Type: Bernieria madagascariensis Baill. typical of Potameia. These specimens also have the Thouvenotia Danguy, Bull. Mus. Hist. Nat. (Paris) 26: cuticle characters of the Beilschmiedia species with 652 (1920). — Type: Thouvenotia madagascariensis opposite leaves (NISHIDA, pers. comm.). Because Danguy. most of the characters of these specimens point to Apollonias sensu Kosterm., Notul. Syst. (Paris) 8: 67 Beilschmiedia, I have included them in that genus, (1939); Fl. Madagascar, Fam. 81: 2 (1950). where they key to B. pedicellata. This implies that TYPE.—Beilschmiedia roxburghiana Nees. the presence of dimerous flowers (vs. trimerous flowers) can no longer be regarded as a consistent generic character and that it may even vary within Trees or shrubs. Leaves alternate or suboppo- a species. Thus, it is no longer possible to identify site, sometimes grouped at tips of branches, pin- specimens without flowers to either Beilschmiedia nately veined, entire, petiolate, domatia lacking. or Potameia and consequently one should not Inflorescences axillary, few-to many-flowered, describe new species in these genera based on paniculate, the flowers cymosely or racemosely fruiting or sterile type material. Unfortunately, arranged along the ultimate branchlets. Flowers previous authors have described several species bisexual, small, greenish, the tepals frequently based on incomplete material: B. obovata was incurved and flowers thus globose or depressed based on a sterile type; three species (B. cryptocary- globose; rarely tepals spreading and flowers disc- oides, B. microphylla and B. sary) were based on like; tepals 6, in 2 series of 3, very rarely 4, in fruiting types, and two species (B. madagascarien- 2 series of 2, fertile stamens usually 9, in 3 whorls sis and B. opposita) have type material with only of 3, very rarely 6, in 3 whorls of 2, filaments var- buds, leaving only three species described on the iously developed, all anthers 2-celled, the outer 6 basis of flowering material. It is thus not surpris- with cells introrse or rarely lateral, the inner 3 ing that identification of Beilschmiedia collections with cells extrorse or latrorse, and with 2 glands remains difficult. Although several sterile collec- attached at or near the base; staminodia 3, usually tions are cited under specimens studied, their well-developed. Floral tube short or lacking.
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