[1] Natl Oceanog Ctr, Southampton SO14 3ZH, Hants, England
[2] Univ Southampton, Natl Oceanog Ctr Southampton, Southampton SO14 3ZH, Hants, England
[3] Univ Liverpool, Dept Earth Ocean & Ecol Sci, Liverpool L69 3GP, Merseyside, England
[4] Univ E Anglia, Sch Environm Sci, Norwich NR4 7TJ, Norfolk, England