1. School of Anthropology and Conservation; Durrell Institute of Conservation and Ecology, University of Kent; CT2 7NR; Canterbury; United Kingdom
2. Biodiversity Information Reporting and Management Division; Ezemvelo KwaZulu-Natal Wildlife, P.O. Box 13053; Cascades; 3202; South Africa
3. Amarula Elephant Research Programme; School of Life Sciences, University of KwaZulu-Natal; Durban; 4041; South Africa
4. Department of Biosciences, Finnish Centre of Excellence in Metapopulation Biology; P.O. Box 65, FI-00014 University of Helsinki; Finland