1. Large Animal Research Group, Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK
2. School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG, UK
3. Department of Science and Technology/National Research Foundation Centre of Excellence, Percy FitzPatrick Institute, University of Cape Town, Rondebosch 7701, Western Cape, South Africa
4. Department of Brains, Behaviour and Evolution, Macquarie University, New South Wales, Australia