[1] Natl Oceanog Ctr, Southampton, Hants, England
[2] Univ Southampton, Dept Ocean & Earth Sci, Southampton, Hants, England
[3] Univ Sheffield, Dept Anim & Plant Sci, Sheffield S10 2TN, S Yorkshire, England
[4] Ctr Environm Fisheries & Aquaculture Sci, Lowestoft, Suffolk, England
[5] Univ E Anglia, Sch Environm Sci, Norwich NR4 7TJ, Norfolk, England