[1] Univ Glasgow, Div Environm & Evolutionary Biol, Fish Biol Grp, Inst Biomed & Life Sci, Glasgow G12 8QQ, Lanark, Scotland
[2] Spey Fishery Board Res Off, Knockando AB38 7SD, Morayshire, Scotland
[3] Fisheries Res Serv Freshwater Lab, Pitlochry PH16 5LB, Perthshire, Scotland