1. Harvard University 1 Department of Organismic and Evolutionary Biology , , Cambridge, MA 02138 , USA
2. Harvard University 2 Museum of Comparative Zoology , , Cambridge, MA 02138 , USA
3. University of Illinois at Urbana-Champaign 3 Department of Evolution, Ecology, and Behavior , , Champaign, IL 61820 , USA
4. University of Connecticut 4 Department of Ecology and Evolutionary Biology , , Storrs, CT 06269 , USA