1. Department of Genetics, Evolution and Environment, Centre for Biodiversity and Environment Research; University College London; Gower Street London WC1E 6BT UK
2. Institute of Zoology; ZSL; Regent's Park London NW1 4RY UK
3. Ecology, Evolution, and Natural Resources; Rutgers University; New Brunswick NJ 08901 USA
4. School of Biological Sciences; University of Adelaide; Adelaide SA 5005 Australia