1. Institute of Zoology, Zoological Society of London, Regent's Park, London NW1 4RY, UK
2. Centre for Biodiversity and Environment Research, Department of Genetics, Evolution and Environment, University College London, Gower Street, London WC1E 6BT, UK
3. Oxford Navigation Group, Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK
4. Edward Grey Institute, Department of Zoology, University of Oxford, South Parks Road, Oxford OX1 3PS, UK