[1] INRA, Comportement & Ecol Faune Sauvage UR35, F-31326 Castanet Tolosan, France
[2] UPMC, CNRS, MNHN, Dept Ecol & Gest Biodiversite,UMR 7204, Brunoy, France
[3] Univ Lyon 1, Lab Biometrie & Biol Evolut, Unite Mixte Rech 5558, F-69622 Villeurbanne, France
[4] CNRS, Ctr Rech Cognit Anim, Toulouse, France
[5] Univ Toulouse 3, Ctr Rech Cognit Anim, F-31062 Toulouse, France