[1] MIT, Dept Civil & Environm Engn, Cambridge, MA 02139 USA
[2] MIT, Woods Hole Oceanog Inst, Joint Program Biol Oceanog, Cambridge, MA 02139 USA
[3] MIT, Dept Biol, Cambridge, MA 02139 USA
[4] Woods Hole Oceanog Inst, Dept Biol, Woods Hole, MA 02543 USA