[1] Southampton Oceanog Ctr, James Rennell Div Ocean Circulat, Southampton SO14 3ZH, Hants, England
[2] Southampton Oceanog Ctr, Sch Ocean & Earth Sci, Southampton SO14 3ZH, Hants, England
[3] Inst Marine Biol Crete, Iraklion 71003, Crete, Greece
[4] Natl Ctr Marine Res, Inst Oceanog, GR-16604 Athens, Greece