[1] Univ Toulouse 3, Inst Natl Polytech, Lab Ecol Fonct & Environm, F-31062 Toulouse, France
[2] CNRS, Lab Ecol Fonct & Environm, F-31062 Toulouse, France
[3] Yale Univ, Tell Leilan Project, New Haven, CT 06511 USA
[4] Yale Univ, Sch Forestry & Environm Studies, New Haven, CT 06511 USA