1. School of Biology, Faculty of Biological Sciences, University of Leeds, Leeds LS2 9JT, UK
2. Groningen Institute for Evolutionary Life Sciences, University of Groningen, 9747 AG Groningen, The Netherlands
3. Wildlife Conservation Research Unit, Department of Zoology, University of Oxford, The Recanati-Kaplan Centre, Abingdon, Oxfordshire OX13 5QL, UK