1. Institute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, Edinburgh, EH9 3JT, United Kingdom
2. Department of Zoology, University of Cambridge, Cambridge, CB2 3EJ, United Kingdom
3. Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9, Canada and
4. Department of Animal Ecology, Lund University, 22362 Lund, Sweden