[1] Univ Southampton, Natl Oceanog Ctr Southampton, Sch Ocean & Earth Sdence, European Way, Southampton SO14 3ZH, Hants, England
[2] Bangor Univ, Sch Ocean Sci, Bangor LL57 2DG, Gwynedd, Wales
[3] Ctr Environm Fisheries & Aquaculture Sci, Lowestoft Lab, Pakefield Rd, Lowestoft NR33 0HT, Suffolk, England