Origin and diversification of dung beetles in Madagascar

Madagascar has a rich fauna of dung beetles (Scarabaeinae and Aphodiinae) withalmost 300 species described to date. Like most other taxa in Madagascar, dung beetles exhibit an exceptionally high level of endemism (96% of the species). Here,we review the current knowledge of the origin and diversification of Malagasy dung beetles. Based on molecular phylogenies, the extant dung beetles originate from eight colonizations, of which four have given rise to extensive radiations. These radiations have occurred in wet forests, while the few extant species in the less successfulradiations occur in open and semi-open habitats. We discuss the likely mechanisms of speciation and the ecological characteristics of the extant communities, emphasizing the role of adaptation along environmental gradients and allopatric speciation in generating the exceptionally high beta diversity in Malagasy dung beetles. Phylogeographic analyses of selected species reveal complex patterns with evidence for genetic introgression between old taxa. The introduction of cattle to Madagascar 1500 years ago created a new abundant resource, onto which a few species haveshifted and thereby been able to greatly expand their geographical ranges.