[1] Univ Toulouse III Paul Sabatier, CNRS, ENFA, Toulouse, France
[2] UMR5174 EDB Lab Evolut & Diversite Biol, Toulouse, France
[3] Univ Exeter, Environm & Sustainabil Inst, Coll Life & Environm Sci, Penryn TR10 9FE, Cornwall, England
[4] Univ Toulouse III Paul Sabatier, CNRS, Toulouse, France
[5] Stn Ecol Theor & Expt, UMR 7321, Toulouse, France
[6] Catholic Univ Louvain, Biodivers Res Ctr, Earth & Life Inst, Louvain La Neuve, Belgium
[7] Museum Natl Hist Nat, Inst Systemat Evolut & Biodiversite, Paris 5, France
[8] Univ Aberdeen, Inst Biol & Environm Sci, Zool Bldg,Tillydrone Ave, Aberdeen AB24 2TZ, Scotland