[1] Univ Complutense Madrid, Fac Biol, Biodivers Ecol & Evolut Dept, Madrid, Spain
[2] Univ Montpellier, INRAE, Eco&Sols, IRD,CIRAD,Inst Agro Montpellier, Montpellier, France
[3] Univ Vigo, Grp Ecol Anim GEA, E-36310 Vigo, Spain
[4] Univ Montpellier, EPHE, CNRS, CEFE,IRD, Montpellier, France