Cola Subgenus Disticha subg. nov. (Malvaceae-Sterculiaceae) of tropical Africa, a synoptic taxonomic revision with seven new species

A new subgenus, Subg. Disticha is erected for 14 species of simple-leaved Cola sharing unique characters within the genus which are presumed synapomorphies: distichous phyllotaxy; petioles lacking pulvini; petioles of uniformly short length, <10 mm long; male flowers with short androphores <1(- 2) mm long; stamens 4 - 5; stem indumentum where present, dominated by short simple translucent hairs (except in C. philipi-jonesii where stellate), and in many species dark purple to black stems with contrasting bright white lenticels. All species have fruit with small globose, unsculptured orange mericarps c. 1 cm diam. (where fruits are known). The species of this subgenus range from SE Nigeria in the west to coastal Kenya and Tanzania in the east and Malawi in the south, but are absent from the Congo basin. Species diversity is highest in the Cross-Sanaga and Eastern Arc-Coastal Forest biogeographic areas, consistent with these taxa being Pleistocene refuge species. The species are mainly monoecious, but dioecy cannot yet be ruled out in some species. In two species where sufficient material was available for investigation (Cola chlorantha, C. roy), the simple cymes were found to be bisexual, the terminal flower being female, the two lateral flowers often being male, a pattern previously unreported in the genus. All species are understorey shrubs or small trees of evergreen lowland or submontane forest, except C. uloloma of coastal semi-deciduous forest in E Africa and which is unique in showing xerophilic characteristics. The species can be divided into group A: five species mainly in E Africa with rapidly glabrescent or glabrous stems, conspicuously asymmetric leaves, few-flowered, pedunculate cymes (C. uloloma, C. chlorantha, C. roy sp.nov., C."udzungwa", C. asymmetrica sp. nov.); group B: in Lower Guinea (Atlantic coast of Africa) with hairy stems, symmetric or inconspicuously asymmetric leaves, sessile, fasciculate inflorescences: (Cola mayimbensis, C. philipi-jonesii, C. metallica, C.moussavoui, C. stigmatosa, C. takamanda sp.nov. C. toyota sp. nov, C. "campo-maan", C. zanaga sp. nov.). Of the seven new species to science, two are named informally since the material is so incomplete (sterile), and five are formally named. Species diversity is highest in Cameroon and Tanzania, each with four species, followed by Gabon with three species. It is expected that additional new species will be found in poorly-surveyed surviving evergreen forest habitat in each of these three countries. Conservation assessments are given for each species using the IUCN 2012 standard. All species are considered threatened, with ten species being Critically Endangered (the highest category of threat), each known from a single location, with their forest habitat threatened mainly by clearance for agriculture.