1. Ecole Normale Supérieure, Laboratoire Ecologie-Evolution, UMR 7625 UPMC-ENS-CNRS, 46 rue d'Ulm, 75005 Paris, France
2. Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA
3. CNRS, Laboratoire Ecologie-Evolution, UMR 7625 UPMC-ENS-CNRS, 46 rue d'Ulm, 75005 Paris, France