1. Division of Biology and the Natural Environment Research Council (NERC) Centre for Population Biology, Imperial College London, Silwood Park, Ascot, Berkshire, SL5 7PY, UK.
2. Department of Zoology, University of Cambridge, Downing Street, Cambridge, CB2 3EJ, UK.
3. Institute of Evolutionary Biology, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.
4. Department of Biological Sciences, Stanford University, Stanford, CA 94305–5020, USA.