1. University of Exeter, Centre for Ecology and Conservation, School of Biosciences, Tremough Campus, Penryn, Cornwall TR10 9EZ, UK
2. Bangor University, School of Biological Sciences, Brambell Laboratories E30, Deiniol Road, Bangor, Gwynedd LL57 2UW, UK
3. North Carolina Wildlife Resources Commission, 1507 Ann Street, Beaufort, NC, USA