1. Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK
2. Centre for Ecology and Conservation, College of Life and Environmental Sciences, University of Exeter, Penryn Campus, Penryn, Cornwall TR10 9FE, UK
3. Department of Statistical Science, Duke University, PO Box 90251, Durham, NC 27708-0251, USA
4. DST-NRF Centre of Excellence at the Percy FitzPatrick Institute, University of Cape Town, Cape Town, Rondebosch 7701, South Africa