1. Department of Evolution, Ecology, and Organismal Biology, The Ohio State University, Columbus, OH 48824
2. Ohio Biodiversity Conservation Partnership, The Ohio State University, Columbus, OH 43210
3. Evolution and Behavior Program, Department of Integrative Biology, Ecology, Michigan State University, East Lansing, MI 48824
4. Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI 48109