[1] Univ Toulouse 3, CNRS, UMR Evolut Diversite Biol EDB 5174, F-31062 Toulouse 9, France
[2] Univ Toulouse 3, Lab Immunogenet Mol EA 3034, F-31062 Toulouse, France
[3] Cardiff Univ, Sch Biosci, Biodivers & Ecol Proc Grp, Cardiff, Wales
[4] Univ Malaysia Sabah, Inst Trop Biol & Conservat, Kota Kinabalu, Sabah, Malaysia