[1] Michigan State Univ, Kellogg Biol Stn, Hickory Corners, MI 49060 USA
[2] Tech Univ Denmark, Natl Inst Aquat Resources, Ctr Ocean Life, DK-2920 Charlottenlund, Denmark
[3] Univ Calif San Diego, Scripps Inst Oceanog, Integrat Oceanog Div, San Diego, CA 92093 USA