[1] AgroParisTech, INRA, Ecol Populat & Communautes USC2031, F-75005 Paris, France
[2] Univ Paris 06, CNRS, ENS, Ecol & Evolut UMR7625, F-75005 Paris, France
[3] Univ Paris 06, CNRS, Ecole Normale Super, Fonctionnement & Evolut Syst Ecol UMR7625, F-75005 Paris, France
[4] CNRS Moulis, Stn Ecol Expt, Lab Evolut & Diversite Biol, F-09200 Moulis, Saint Girons, France