1. Division of Biology, Faculty of Life Sciences, Imperial College at Silwood ParkAscot, Berks SL5 7PY, UK
2. Institute of Evolutionary Biology, University of EdinburghEdinburgh EH9 3JT, UK
3. Department of Zoology, University of CambridgeDowning Street, Cambridge CB2 3EJ, UK
4. NERC Centre for Population Biology, Imperial College at Silwood ParkAscot, Berks SL5 7PY, UK