Diversification of the yellow-shouldered bats, genus Sturnira (Chiroptera, Phyllostomidae), in the New World tropics.

The Yellow-shouldered bats, Genus Sturnira, are widespread, diverse, and abundant throughout the Neotropical Region, but little is known of their phylogeny and biogeography. We collected 4409 bp of DNA from three mitochondrial (cyt-b, ND2, D-loop) and two nuclear (RAG1, RAG2) sequences from 138 individuals representing all but two recognized species of Sturnira and five other phyllostomid bats used as outgroups. The sequence data were subjected to maximum parsimony, maximum likelihood, and Bayesian inference analyses. Results overwhelmingly support the monophyly of the genus Sturnira but not continued recognition of Corvira as a subgenus; the two species (bidens and nana) allocated to that group constitute separate, basal branches on the phylogeny. A total of 21 monophyletic putatively species-level groups were recovered; pairs were separated by an average 7.09% (SD=1.61) pairwise genetic distance in cyt-b, and three of these groups are apparently unnamed. Several well-supported clades are evident, including a complex of seven species formerly confused with S. lilium, a species that is actually limited to the Brazilian Shield. We used four calibration points to construct a time-tree for Sturnira, using BEAST. Sturnira diverged from other stenodermatines in the mid-Miocene, and by the end of that epoch (5.3 Ma), three basal lineages were present. Most living species belong to one of two clades, A and B, which appeared and diversified shortly afterwards, during the Pliocene. Both parsimony (DIVA) and likelihood (Lagrange) methods for reconstructing ancestral ranges indicate that the radiation of Sturnira is rooted in the Andes; all three basal lineages (in order, bidens, nana, and aratathomasi) have strictly or mainly Andean distributions. Only later did Sturnira colonize the Pacific lowlands (Chocó) and thence Central America. Sturnira species that are endemic to Central America appeared after the final emergence of the Panamanian landbridge ~3 Ma. Despite its ability to fly and to colonize the Antilles overwater, this genus probably accompanied the "legions" of South American taxa that moved overland during the Great American Biotic Interchange. Its eventual colonization of the Lesser Antilles and the appearance of two endemic lineages there did not take place until the Pleistocene. Because of its continual residence and diversification in South America, Andean assemblages of Sturnira contain both basal and highly derived members of the genus.