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A Synopsis of the Genus Disperis Sw. (Orchidaceae)

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A Synopsis of the Genus Disperis Sw. (Orchidaceae) A synopsis of the genus Disperis Sw. (Orchidaceae) Hubert KURZWEIL Compton Herbarium, South African National Biodiversity Institute, 7735 Claremont (South Africa) Current Address: Singapore Botanic Gardens, 1 Cluny Road, 259569 Singapore [email protected] John C. MANNING Compton Herbarium, South African National Biodiversity Institute, 7735 Claremont (South Africa) [email protected] ABSTRACT An infrageneric classification is provided for the genus Disperis Sw. (Orchidaceae, Orchidoideae, Diseae), with a complete enumeration of all species. The genus comprises 74 species of deciduous herbs that grow largely terrestrially in forest, scrub and grassland. Two subgenera are recognised in this study.Subgenus Dryorkis comprises 44 species found largely in the trop- ics and subtropics of Africa, Madagascar and Asia and is defined by a deeply bilobed lip appendage and possibly by apomorphic, baculate pollen. In addi- tion, in many species the lateral sepals are basally fused and the rostellum arms are simple and not twisted. Subgenus Disperis comprises 30 species mainly in southern Africa and is defined by an entire lip appendage and usu- ally twisted rostellum arms. Within these subgenera some well defined groups of morphologically similar and presumably related species can be distin- guished but the existence of several unusual species makes it premature to propose them as formal taxa. They are consequently recognised as “informal KEY WORDS groups”. Subgenus Disperis appears to be a monophyletic group but the phy- Orchidaceae, Disperis , logenetic status of subgenus Dryorkis is not clear at present. The taxonomic subgeneric classification. value of several vegetative and floral structures is briefly discussed. RÉSUMÉ Synopsis du genre Disperis Sw. (Orchidaceae). Une classification infragénérique est présentée pour le genre Disperis Sw. (Orchidaceae, Orchidoideae, Diseae) avec une énumération complète de toutes les espèces. Le genre comprend 74 espèces d’herbacées décidues princi- palement terrestres croissant en milieux forestiers, dans des fourrés ou en savane. Deux sous-genres sont reconnus. Le sous-genre Dryorkis comprend 44 espèces, principalement de régions tropicales et subtropicales d’Afrique, de Madagascar et d’Asie. Il est défini par un appendice profondément bilobé et un pollen baculé vraisemblablement apomorphique. De plus, chez plusieurs ADANSONIA, sér. 3 • 2005 • 27 (2) : 155-207 © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. 155 Kurzweil H. & Manning J.C. espèces les sépales latéraux sont fusionnés à la base et les bras du rostellum sont simples et non tordus. Le sous-genre Disperis comprend 30 espèces sur- tout d’Afrique du Sud. Il est défini par un appendice entier et un rostellum avec des bras habituellement tordus. Dans les deux sous-genres on peut reconnaître certains groupes bien définis d’espèces morphologiquement proches et vraisemblablement apparentées. Cependant il existe plusieurs «formes» singulières qu’il semble prématuré de reconnaître formellement en tant que taxa. Nous considérons qu’elles forment des «groupes informels». MOTS CLÉS Le sous-genre Disperis est apparemment un groupe monophylétique, mais le Orchidaceae, Disperis , statut phylogénétique du sous-genre Dryorkis reste incertain. La valeur taxo- classification infragénérique. nomique de plusieurs structures végétatives et florales est brièvement discutée. TABLE OF CONTENTS Introduction ..............................................................................................................................156 Characters of taxonomic value .................................................................................................... 157 Vegetative features ................................................................................................................ 157 Perianth ................................................................................................................................ 162 Gynostemium ...................................................................................................................... 164 Pollen .................................................................................................................................. 165 Classification and phylogeny ...................................................................................................... 165 Systematics ................................................................................................................................ 167 Disperis Sw. .......................................................................................................................... 167 Key to subgenera, informal groups and species of Disperis .............................................. 168 Disperis Sw. subgen. Dryorkis (Thouars) Kurzweil & Manning, stat. nov. ........................ 172 Disperis Sw. subgen. Disperis , subgen. nov. ...................................................................... 188 Doubtful names .............................................................................................................. 202 Excluded name ................................................................................................................ 202 References .................................................................................................................................. 202 INTRODUCTION their lip structure. Several synapomorphies, nota- bly the spurred lateral sepals, a linear and clawed The genus Disperis was established by S WARTZ lip, a large, membranous central rostellum lobe (1800) to accommodate a number of species covering the anther on top and sides, and the of terrestrial orchids from the Western Cape of cartilaginous rostellum lobes suggest that South Africa with highly distinctive flowers, char- the genus is monophyletic (L INDER 1986; acterised by spurred lateral sepals and a reflexed L INDER & K URZWEIL 1994). lip bearing a more or less complex appendage The genus Disperis comprises 74 species dis- (Fig. 1D-F, H). As a result of its unique floral tributed in sub-Saharan Africa, Madagascar and structure, the genus has almost always been adjacent Indian Ocean islands. A single variable recognised as distinct, and its circumscription has and widespread species occurs in tropical Asia seldom been disputed. An exception is T HOUARS (Fig. 2). Centres of diversity are the humid (1809), who segregated a few of the tropical African tropics, the Cape Floristic Region, and species into the genus Dryorkis on the basis of Madagascar. Tropical African representatives 156 ADANSONIA, sér. 3 • 2005 • 27 (2) A synopsis of the genus Disperis (Orchidaceae) account for some 44% of the species, the L INDER 1992) retrieved two main clades within Madagascan species for ±30%, and the southern the genus, identified as the “tropical” and “south- African species for an additional 35%. At the ern African” clades, respectively. These two species level these three centres are very distinct, groups are well defined on morphological with some overlap between the southern and the grounds and are recognised here as subgenera. tropical African centres, very little between The aim of the present paper is to provide a the African and Madagascan centres, and none complete enumeration of the species currently between Asia and Africa plus Madagascar. recognised, together with a formal classification In contrast to the other species of tribe Diseae into two subgenera, Dryorkis and Disperis . which prefer rather open habitats, Disperis species Within these subgenera several obvious groups of are mostly found in wooded situations, except for morphologically similar and probably phyloge- the southern African representatives which are netically closely related species exist and may be often found in grassland or sclerophyllous scrub appropriately recognised as sections, but the exis- (fynbos). Disperis is an essentially terrestrial tence of several morphologically unusual taxa of genus, but several tropical species may grow as uncertain affinities makes the formal recognition low-level epiphytes and a few are frequently or of sections inadvisable. Additional data sets, predominantly epiphytic up to several meters including anatomical, ultrastructural and molec- above the ground (for example D. kilimanjarica). ular ones are needed to formulate robust phyloge- Many southern African species are pollinated by netic hypotheses. Consequently, we establish oil-collecting bees in the genus Rediviva 10groups of morphologically similar species but (Mellitidae), which collect oil secreted by the lip do not formally propose them (Table 1). The appendages (S TEINER 1989). A notable exception information included in this paper is derived is the South African D. capensis , which is polli- largely from K URZWEIL & L INDER (1991) and nated by deceit (J OHNSON 1994) and appears to M ANNING & L INDER (1992), with additional have secondarily lost the facility to secrete oil. data from other authors referred to under the Pollination is largely unknown in the tropical individual species as well as the World Checklist of African Disperis species but it has been suggested Monocots (G OVAERTS 2004). that among these species the structure of the lip appendage may not be adequate for oil secretion (S TEINER 1989). CHARACTERS OF TAXONOMIC VALUE On morphological, anatomical and ultrastruc- tural evidence, Disperis is usually placed in sub- V EGETATIVE FEATURES tribe Coryciinaeof tribe Diseae within the orchid Plants
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