1. Department of Ecology and Evolutionary Biology and Peabody Museum of Natural History, Yale University, New Haven, CT 06520, USA
2. Center for Population Biology, University of California, Davis, CA 95616, USA
3. Department of Organismic and Evolutionary Biology & Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138, USA
4. Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, TN 37669, USA