[1] Museum Natl Hist Nat, Conservatoire Bot Bassin Parisien, F-75005 Paris, France
[2] Inst Natl Agron Paris Grignon, F-75231 Paris 05, France
[3] Univ Calif Santa Barbara, Dept Ecol Evolut & Marine Biol, Santa Barbara, CA 93106 USA
[4] Univ Paris Sud, URA 2154, Lab Ecol Systemat & Evolut, F-91405 Orsay, France