1. Institute of Zoology, Zoological Society of London, Regent's Park, London NW1 4RY, UK
2. Department of Ecology and Evolutionary Biology, Princeton University, 117 Eno Hall, Princeton, NJ 08544, USA
3. Department of Biology, Huck Institutes of the Life Sciences, The Pennsylvania State University, 201 Life Sciences Building, University Park, PA 16802, USA