Environmental heterogeneity predicts species richness of freshwater mollusks in sub-Saharan Africa

Species diversity and how it is structured on a continental scale is influenced by stochastic, ecological, and evolutionary driving forces, but hypotheses on determining factors have been mainly examined for terrestrial and marine organisms. The extant diversity of African freshwater mollusks is in general well assessed to facilitate conservation strategies and because of the medical importance of several taxa as intermediate hosts for tropical parasites. This historical accumulation of knowledge has, however, not resulted in substantial macroecological studies on the spatial distribution of freshwater mollusks. Here, we use continental distribution data and a recently developed method of random and cohesive allocation of species distribution ranges to test the relative importance of various factors in shaping species richness of Bivalvia and Gastropoda. We show that the mid-domain effect, that is, a humpshaped richness gradient in a geographically bounded system despite the absence of environmental gradients, plays a minor role in determining species richness of freshwater mollusks in sub-Saharan Africa. The western branch of the East African Rift System was included as dispersal barrier in richness models, but these simulation results did not fit observed diversity patterns significantly better than models where this effect was not included, which suggests that the rift has played a more complex role in generating diversity patterns. Present-day precipitation and temperature explain richness patterns better than Eemian climatic condition. Therefore, the availability of water and energy for primary productivity during the past does not influence current species richness patterns much, and observed diversity patterns appear to be in equilibrium with contemporary climate. The availability of surface waters was the best predictor of bivalve and gastropod richness. Our data indicate that habitat diversity causes the observed species–area relationship, and hence, that environmental heterogeneity is a principal driver of freshwater mollusk richness on a continental scale