[1] Univ Southampton, Natl Oceanog Ctr, Southampton SO14 3ZH, Hants, England
[2] Univ Oxford, Dept Earth Sci, Oxford OX1 3PR, England
[3] Cardiff Univ, Sch Earth Ocean & Planetary Sci, Cardiff CF10 3YE, S Glam, Wales
[4] Stn Biol Roscoff, F-29682 Roscoff, France
[5] Univ Washington, Sch Oceanog, Seattle, WA 98195 USA