[1] Univ Portsmouth, Sch Earth & Environm Sci, Portsmouth PO1 3QL, Hants, England
[2] Univ Southampton, Natl Oceanog Ctr Southampton, Sch Ocean & Earth Sci, Southampton SO14 3ZH, Hants, England
[3] Cefas Lowestoft Lab, Ctr Environm Fisheries & Aquaculture Sci, Lowestoft NR33 0HT, Suffolk, England
[4] Natl Oceanog Ctr Southampton, Southampton SO14 3ZH, Hants, England