1. Biodiversity Assessment and Monitoring, South African National Biodiversity Institute, Silverton, South Africa
2. Amarula Elephant Research Programme, School of Life Sciences, University of KwaZulu-Natal, Durban, KwaZulu-Natal, South Africa
3. Department of Genetics, Evolution and Environment, University College London, University of London, London, United Kingdom
4. Finnish Centre of Excellence in Metapopulation Biology, Department of Biosciences, University of Helsinki, Helsinki, Finland
5. Department of Geosciences and Geography, University of Helsinki, Helsinki, Finland