1. School of Biosciences, University of Sheffield, Sheffield S10 2TN, UK
2. School of Biology, University of St Andrews, St Andrews KY16 9ST, UK
3. Department of Anthropology, Durham University, Durham DH1 3LE, UK
4. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh EH9 3FL, UK
5. Centre of Ecology and Conservation, College of Life and Environmental Sciences, University of Exeter, Penryn TR10 9FE, UK