1. Department of Ecology and Evolutionary Biology, Guyot Hall, Princeton University, Princeton, NJ 08540, USA
2. The Roslin Institute and Royal (Dick) School of Veterinary Studies, University of Edinburgh, Roslin EH25 9PS, UK
3. Centre for Ecology and Hydrology, Edinburgh EH26 0QB, UK
4. Department of Migration and Immuno-ecology, Max Planck Institute for Ornithology, Schlossallee 1a, 78315 Radolfzell, Germany