1. School of Animal, Plant and Environmental Sciences, University of the Witwatersrand , Johannesburg , South Africa
2. South African National Biodiversity Institute, Kirstenbosch Research Centre , Claremont , South Africa
3. Centre for Ecological Genomics and Wildlife Conservation, University of Johannesburg , Johannesburg , South Africa
4. Department of Biology, Center for Computational and Integrative Biology, Rutgers University–Camden , Camden, New Jersey , USA