1. School of Life Sciences, University of SussexBrighton BN1 9QG, UK
2. Institut de Recherche sur les Grands Mammifères, INRAToulouse, France
3. School of Psychology, University of St AndrewsSt Andrews KY16 9JU, UK
4. Large Animal Research Group, Department of Zoology, University of CambridgeCambridge CB2 3EJ, UK