1. 1Conservation Programmes, Zoological Society of London, Regent's Park, London, NW1 4RY, United Kingdom; King Khalid Wildlife Research Centre, National Commission for Wildlife Conservation and Development, P.O. Box 61681, Riyadh 11575, Kingdom of Saudi Arabia
2. 2University of Frankfurt, Department of Ecology and Evolution, Frankfurt/M., Germany
3. 3Conservation Programmes, Zoological Society of London, Regent's Park, London, NW1 4RY, United Kingdom; King Khalid Wildlife Research Centre, National Commission for Wildlife Conservation and Development, P.O. Box 61681, Riyadh 11575, Kingdom of Saudi Arabia
4. 4Conservation Programmes, Zoological Society of London, Regent's Park, London, NW1 4RY, United Kingdom; King Khalid Wildlife Research Centre, National Commission for Wildlife Conservation and Development, P.O. Box 61681, Riyadh 11575, Kingdom of Saudi Arabia