1. Department of Human Evolutionary Biology, Harvard University , 11 Divinity Avenue, Cambridge, MA 02138 , USA
2. Greater Mahale Ecosystem Research and Conservation Project , Box 120, Mpanda , Tanzania
3. School of Biological and Environmental Sciences, Liverpool John Moores University , James Parsons Building, Byrom Street, Liverpool L33AF , UK
4. Department of Anthropology, University College London , 14 Taviton Street, London WC1H OBW , UK