[1] Leiden Univ, Dept Anim Ecol, Inst Biol, NL-2300 RA Leiden, Netherlands
[2] Univ Bern, Inst Zool, Dept Behav Ecol, CH-3012 Bern, Switzerland
[3] EAWAG Ctr Limnol, Dept Fish Ecol & Evolut, CH-6047 Kastanienbaum, Switzerland
[4] Univ Bern, Dept Aquat Ecol & Macroevolut, Inst Zool, CH-3012 Bern, Switzerland
[5] Univ Hull, Dept Biol Sci, Mol & Evolutionary Ecol Grp, Kingston Upon Hull HU6 7RX, N Humberside, England