1. Edward Grey Institute of Field Ornithology, University of Oxford, Oxford OX1 3PS, UK
2. Research School of Biology, Australian National University, Acton, 2600 Australian Capital Territory, Australia
3. Department of Anthropology, University of California, Davis, CA 95616, USA
4. Smithsonian Tropical Research Institute, Ancon, Panama
5. Department of Mathematics, Uppsala University, Box 480, 75106 Uppsala, Sweden