1. Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK
2. Institute for Evolutionary Biology, University of Edinburgh, Kings Buildings, Ashworth Laboratories, West Mains Road, Edinburgh EH9 3JT, UK
3. School of Life Sciences, Napier University, Edinburgh EH10 5DT, UK
4. Centre for Ecology and Conservation, University of Exeter in Cornwall, Penryn, Cornwall TR10 9EZ, UK