[1] Univ Cambridge, Dept Zool, Large Anim Res Grp, Cambridge CB2 3EJ, England
[2] Univ Pretoria, Mammal Res Inst, Dept Zool & Entomol, ZA-0002 Pretoria, South Africa
[3] Univ Edinburgh, Acad Ctr, Ctr Reprod Biol, MRC HRSU, Edinburgh EH16 4SB, Midlothian, Scotland