1. Centre for Ecology and Conservation, College of Life and Environmental Sciences, University of Exeter, Penryn TR10 9EZ, Cornwall, UK
2. Centre for Evolutionary Biology, School of Biological Sciences (M092), The University of Western Australia, Crawley 6009, Western Australia, Australia
3. Department of Biological Sciences (E8C 209), Macquarie University, Sydney 2109, New South Wales, Australia