[1] Univ Washington, Dept Biol, Seattle, WA 98195 USA
[2] Danish Inst Fisheries Res, Dept Inland Fisheries, Vejlsovej, Silkeborg, Denmark
[3] Natl Environm Res Inst, Dept Freshwater Biol, Vejlsovej, Silkeborg, Denmark
[4] Univ Aarhus, Dept Ecol Bot, Risskov, Denmark