1. Present address: Division of Evolution, Infection and Genomics, School of Biological Sciences, Faculty of Biology, Medicine and Health, The University of Manchester, Manchester M13 9PT, UK
2. Centre for Ecology and Conservation, University of Exeter, Penryn, TR10 9FE, UK
3. Present address: Department of Biology, University of York, Wentworth Way, York, YO10 5DD, UK