1. Centre for Research into Ecological and Environmental Modelling, University of St Andrews, The Observatory, Buchanan Gardens, St Andrews KY16 9LZ, UK
2. School of Biology, Gatty Marine Laboratory, University of St Andrews, St Andrews KY16 8LB, UK
3. Royal Society for the Protection of Birds and Conservation Biology Group, Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK
4. Department of Statistics, University of Auckland, Private Bag 92019, Auckland, New Zealand