[1] Univ E Anglia, Sch Environm Sci, Norwich NR4 7TJ, Norfolk, England
[2] Plymouth Marine Lab, Plymouth PL13 DH, Devon, England
[3] Univ British Columbia, Changing Ocean Res Unit, Vancouver, BC V6T 1Z4, Canada
[4] Ctr Environm Fisheries & Aquaculture Sci, Lowestoft NR33 0HT, Suffolk, England
[5] Princeton Univ, Atmospher & Ocean Sci Program, Princeton, NJ 08544 USA