[1] Univ Stellenbosch, Evolutionary Genet Grp, Dept Bot & Zool, ZA-7602 Matieland, South Africa
[2] Univ Pretoria, Mammal Res Inst, Dept Zool & Entomol, ZA-0002 Pretoria, South Africa
[3] Univ Cambridge, Ctr Vet Sci, Cambridge, England
[4] Queen Mary Univ London, Sch Biol & Chem Sci, London, England
[5] Univ Montpellier 2, Inst Sci Evolut, UMR5554, Montpellier, France