1. Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Penryn Campus, TR10 9FE, UK
2. Groningen Institute for Evolutionary Life Sciences, University of Groningen, PO Box 11103, 9700 CC Groningen, The Netherlands
3. Department of Psychology, Neuroscience and Behaviour, McMaster University, 6 Hamilton, Ontario, Canada