Review of the systematics and global diversity of freshwater mussel species (Bivalvia: Unionoida)

Freshwater mussels (Bivalvia: Unionoida) are interesting because of their unique life cycles, global aggregate distribution and ancient origin. They are also of practical importance due to their worldwide, imperiled status. Of utmost utility for their continued study are a modern assessment of global and regional species diversity and a natural classification that reflects phylogenetic patterns. The freshwater malacological community has taken steps toward satisfying the latter of these requirements, but a consensus census of mussel species has not been published since Fritz Haas's revisions of the late 1960s. We set out to describe the species-level diversity of the Unionoida by reviewing the secondary literature and developing a comprehensive taxonomic database. Each valid species was assigned to one or more geographical regions (i.e. Nearctica, Neotropica, Afrotropica, Palearctica, Indotropica and Australasia) and one or more subregions, and each valid genus was assigned to the lowest possible level in a classification derived from our own, recent phylogenetic analyses. Based upon a consensus of numerous regional works, our global estimate of freshwater mussel diversity is 840 species. Regional diversity was determined as follows: Nearctica: 302 spp., Neotropica: 172, Afrotropica: 85, Palearctica: 45, Indotropica: 219 and Australasia: 33. The largest family is the Unionidae, with 674 species. However, the classification of that taxon is currently in flux, and many genera (corresponding to 225 spp.) were assigned to incertae sedis geographical assemblages. Diversity patterns are discussed, and it is suggested that reevaluation of these faunas with modern methods will likely increase recognized species diversity, especially on the southern continents. Our checklist and classification of freshwater mussel species is included as an appendix and mirrored on the MUSSEL Project Web Site (http://www.mussel-project.net/).