[1] Swiss Fed Res Inst WSL, CH-8903 Birmensdorf, Switzerland
[2] Univ Tromso, Fac Sci, Dept Biol, N-9037 Tromso, Norway
[3] Utah State Univ, Utah Cooperat Fish & Wildlife Res Unit, US Geol Survey, Logan, UT 84322 USA
[4] Univ Grenoble 1, CNRS, UMR 5553, Lab Ecol Alpine, F-38041 Grenoble 9, France
[5] Univ Lausanne, Dept Ecol & Evolut, CH-1015 Lausanne, Switzerland