[1] CNRS, SETE, UMR 5321, Moulis, France
[2] Univ Toulouse III Paul Sabatier, Moulis, France
[3] Katholieke Univ Leuven, Dept Biol, Plant Conservat & Populat Biol, Kasteelpk Arenberg 31, B-3001 Leuven, Belgium
[4] Catholic Univ Louvain, Earth & Life Inst, Louvain, Belgium
[5] Sorbonne Univ, Museum Natl Hist Nat, Inst Systemat Evolut Biodiversite, UMR 7205, Paris, France