[1] Univ Aberdeen, Sch Biol Sci, NERC Mol Genet Ecol Initiat, Aberdeen AB24 2TZ, Scotland
[2] Univ Exeter, Sch Biol Sci, Hatherly Labs, Exeter EX4 4PS, Devon, England
[3] Environm Agcy, Hampshire & Isle Wight Area Off, Winchester SO21 1WP, Hants, England
[4] Ctr Ecol & Hydrol Banchory, Banchory AB31 4BW, Aberdeen, Scotland