1. Institute of Integrative and Comparative Biology, University of Leeds, Leeds LS2 9JT, UK
2. Woodrow Wilson School and Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544, USA
3. Department of Ecology, Evolution, and Environmental Biology, Columbia University, New York, NY 10027, USA
4. School of Biological Sciences, University of East Anglia, Norwich NR4 7TJ, UK