[1] Swansea Univ, Fac Sci & Engn, Dept Biosci, Swansea SA2 8PP, Wales
[2] Univ Zurich, Dept Evolutionary Biol & Environm Studies, CH-8057 Zurich, Switzerland
[3] Univ Glasgow, Sch Biodivers, Hlth & Vet Med 1, Glasgow G12 8QQ, Scotland
[4] Univ Cape Town, Inst Communities & Wildlife Africa, Dept Biol Sci, ZA-7701 Rondebosch, South Africa
[5] German Primate Ctr, Endocrinol Lab, D-37077 Gottingen, Germany