[1] Univ Oxford, Dept Zool, Edward Grey Inst, Oxford OX1 3PS, England
[2] Ctr Rech, F-13200 Arles, France
[3] Univ Montpellier 2, Inst Sci Evolut, UMR 5119, CNRS,UM2, F-34095 Montpellier 05, France
[4] UPMC, MNHN, UMR 5173, Conservat Restaurat & Suivi Populat,CNRS, F-75005 Paris, France
[5] Univ Grenoble 1, UMR 5553, CNRS, Lab Ecol Alpine, F-38041 Grenoble 9, France
[6] Univ Montpellier 2, Lab Ecosyst Lagunaires, CNRS, UMR 5119,UM2, F-34095 Montpellier 5, France