[1] Univ Calif San Diego, Scripps Inst Oceanog, San Diego, CA 92103 USA
[2] Bangor Univ, Sch Ocean Sci, Bangor, Gwynedd, Wales
[3] Moss Landing Marine Labs, Pob 450, Moss Landing, CA 95039 USA
[4] Boston Univ, Dept Biol, 5 Cummington St, Boston, MA 02215 USA