1. Institute of Biodiversity, Animal Health and Comparative Medicine, University of Glasgow, Glasgow, G12 8QQ, UK
2. Evolutionary Ecology, Lund University, SE-223 62 Lund, Sweden
3. Biological Sciences Department, North Dakota State University, Fargo, ND 58108, USA
4. RSPB Centre for Conservation Science, The Lodge, Sandy, Bedfordshire, SG19 2DL, UK