[1] Univ So Calif, Dept Biol Sci, Marine Environm Biol Sect, Los Angeles, CA 90089 USA
[2] MIT WHOI Joint Program Oceanog Appl Ocean Sci & E, Woods Hole, MA USA
[3] Woods Hole Oceanog Inst, Dept Marine Chem & Geochem, Woods Hole, MA 02543 USA
[4] Woods Hole Oceanog Inst, Dept Biol, Woods Hole, MA 02543 USA