1. Department of Biological Sciences, University of Bristol, 24 Tyndall Avenue, Bristol BS8 1TQ, UK
2. Center for Population Biology, University of California, 1 Shields Avenue, Davis, Davis, CA 95616, USA
3. Department of Evolution and Ecology, University of California, Davis, 1 Shields Avenue, Davis, CA 95616, USA
4. Department of Animal Biology, University of California, Davis, 1 Shields Avenue, Davis, CA 95616, USA
5. Department of Biological Sciences, California State University, 1250 Bellflower Boulevard, Long Beach, CA 90840, USA