[1] FRS Freshwater Lab, Pitlochry PH16 5LB, Perth, Scotland
[2] Univ Glasgow, Dept Geog & Earth Sci, Glasgow G12 8QQ, Lanark, Scotland
[3] Univ Aberdeen, Sch Geosci, Aberdeen AB4 3UF, Scotland
[4] Macaulay Land Use Res Inst, Aberdeen AB15 8QH, Scotland