[1] Univ Washington, Sch Aquat & Fishery Sci, Seattle, WA 98105 USA
[2] Univ Toulouse 3, CNRS, EDB Lab Evolut & Divers Biol UMR5174, ENFA, 118 Route Narbonne, F-31062 Toulouse, France
[3] Univ Toulouse, INP, UPS, EcoLab Lab Ecol Fonct & Environ, 118 Route Narbonne, F-31062 Toulouse, France
[4] CNRS, EcoLab, F-31062 Toulouse, France