Phylogeny and temporal diversification of Calomys (Rodentia, Sigmodontinae): Implications for the biogeography of an endemic genus of the open/dry biomes of South America

A thorough analysis of character evolution and biogeography of a group is only possible with a comprehensive sampling of its diversity. The sigmodontine genus Calomys is particularly interesting for the study of neotropical biogeography as it occurs exclusively in the dry and grassland biomes: Cerrado, Caatinga, Chaco, Pampas, Venezuelan Llanos, Puna, and a diversity of dry forests biomes. Although Brazil encompasses a large part of the geographic distribution of the genus and at least three endemic species, the last published phylogeny of Calomys included only two specimens (both representing the same species) from a single locality in this country. In the present paper we add complete cytochrome b sequences of Brazilian karyomorphs to previously published sequences in order to provide a phylogenetic hypothesis including most of the genus diversity. The main objectives of this study were to clarify taxonomic issues related to Brazilian karyomorphs and to study the diversification processes of the genus by analyzing its biogeography in combination with cladogenesis dates estimated with a molecular clock. The phylogeny indicates that at least six different species inhabit the Brazilian territory, one of them still undescribed. Date estimates indicate that two sequential basal splits, the first separating Andean from lowland species and the second isolating species north and south of the Amazonia, took place in the Pliocene between 3 and 4  Ma. A large-bodied and speciose clade of lowland species associated with dry forests and ecotones of the core Cerrado and Chaco areas with adjacent biomes diversified in the Pleistocene. This indicates the importance of safeguarding ecotones and the Cerrado at a time where this biome is being rapidly destroyed.