1. Smithsonian Tropical Research Institute , Apartado 0843-03092, Balboa, Ancón , Republic of Panamá
2. Department of Integrative Biology, University of Texas at Austin , 1 University Station C0930, Austin, TX 78756 , USA
3. Department of Evolution, Ecology and Organismal Biology, The Ohio State University , 318 West 12th Avenue, Columbus, OH 43210 , USA