1. Department of Biological Sciences and NERC Centre for Population Biology, Imperial College London, Silwood Park Campus, Ascot, Berkshire SL5 7PY, United Kingdom
2. Department of Biological Sciences and NERC Centre for Population Biology, Imperial College London, Silwood Park Campus, Ascot, Berkshire SL5 7PY, United Kingdom E‐mail:
3. Department of Ecology and Evolutionary Biology and the Graduate Interdisciplinary Program in Genetics, University of Arizona, Biological Sciences West, Tucson, Arizona 85721
4. Department of Ecology and Evolutionary Biology and the Graduate Interdisciplinary Program in Genetics, University of Arizona, Biological Sciences West, Tucson, Arizona 85721 E‐mail: