1. Smithsonian Tropical Research Institute, AA 2072, Balboa, Panama
2. The Galton Laboratory, Department of Biology, University College London, London NW1 2HE, UK
3. Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK
4. Department of Biology, Middle Tennessee State University, Murfreesboro, TN 37132, USA