Pleistocene and pre-Pleistocene Begonia speciation in Africa.

This paper presents a historical biogeographic analysis of African Begonia based on combined internal transcribed spacer (ITS) and trnL intron sequences. Age range estimates for Begonia in Africa ranged from only 1.5 Ma for some terminal nodes to 27 Ma for basal nodes when the ages of Réunion (2 Ma) andMayotte (5.4 Ma) were used to date the split between Begonia salaziensis and Begonia comorensis. Assuming a more recent origin age for Begonia salaziensis (2 Ma) provided age estimates in other parts of the phylogeny which agreed with patterns observed in other African organisms. A large proportion of the Begonia diversity seen today in Africa is of pre-Pleistocene origin. Species of Pleistocene origin are concentrated in species-rich groups such as sections Loasibegonia, Scutobegonia, and Tetraphila, which have their centre of diversity in western Central Africa. Phylogenetically isolated taxa such as Begonia longipetiolata, Begonia iucunda, and Begonia thomeana date to the late Miocene, a period of extended aridification on the African continent that had severe effects on African rain forest species. A general pattern is identified where phylogenetically isolated species occur outside the main identified rain forest refuges. Endemic species on the island of São Tomé such as Begonia baccata, Begonia molleri, and Begonia subalpestris appear to be palaeoendemics. Of these species, the most recent age estimate is for B. baccata, which is dated at ca. 3 Ma. Therefore, São Tomé appears to have functioned as an important (if previously unrecognised) pre-Pleistocene refuge. On the mainland, areas such as the Massif of Chaillu in Gabon, southern Congo (Brazzaville), and far western areas of Congo (Kinshasa) have played similar roles to São Tomé.