[1] Univ Johannesburg, Dept Zool, Mol Zool Lab, Johannesburg, South Africa
[2] Univ Montpellier 3, Univ Montpellier, EPHE Lab Biogeog & Ecol Vertebres, CEFE,UMR 5175,CNRS, 1919 Route Mende, F-34293 Montpellier 5, France
[3] CNRS, SMGE, CEFE, UMR 5175, 1919 Route Mende, F-34293 Montpellier 5, France