1. Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn TR10 EZ, UK.
2. School of Biological Sciences, University of East Anglia, Norwich NR4 7TJ, UK.
3. Department of Animal and Plant Sciences, University of Sheffield, Western Bank, Sheffield S10 2TN, UK.
4. Institute of Evolutionary Biology, School of Biological Sciences, Ashworth Laboratories, King's Buildings, University of Edinburgh, Edinburgh EH9 3JT, UK.