[1] Univ Aberdeen, Inst Biol & Environm Sci, Aberdeen AB24 2TZ, Scotland
[2] Natl Museum Nat Hist MNCN CSIC, Dept Evolutionary Ecol, Madrid 28006, Spain
[3] Univ Exeter, Ctr Res Anim Behav, Coll Life & Environm Sci, Exeter EX4 4QG, Devon, England
[4] UPMC, Museum Natl Hist Nat, CNRS, CERSP,MNHN,UMR 7204, F-75005 Paris, France
[5] Univ Freiburg, Dept Wildlife Ecol & Management, D-79106 Freiburg, Germany