1. School of Biology, Faculty of Biological Sciences, University of Leeds, Leeds LS2 9JT, UK
2. Sea Mammal Research Unit, Scottish Oceans Institute, University of St Andrews, St Andrews KY16 8LB, UK
3. School of Mathematics and Statistics, University of Sheffield, Sheffield S3 7RH, UK
4. Scottish Natural Heritage, Battleby, Redgorton, Perth PH1 3EW, UK