1. Institute of Evolutionary Biology, School of Biological Sciences, University of Edinburgh, West Mains Road, EH9 3LF, UK
2. University of Cambridge, Department of Biochemistry, Sanger Building, 80 Tennis Ct Rd, Cambridge CB2 1GA, UK
3. Centre for Ecology and Conservation, University of Exeter, Penryn, Falmouth, Cornwall, TR10 9FE, UK