[1] Ctr Ecol & Hydrol, Penicuik EH26 0QB, Midlothian, Scotland
[2] Univ E Anglia, Sch Environm Sci, Norwich NR4 7TJ, Norfolk, England
[3] Univ British Columbia, Dept Earth & Ocean Sci, Vancouver, BC V6T 1Z4, Canada
[4] Natl Oceanog Ctr, Southampton SO14 3ZH, Hants, England