[1] Univ Calif Irvine, Dept Earth Syst Sci, Irvine, CA 92697 USA
[2] Univ Illinois, Dept Biol Sci, Ecol & Evolut Grp, Chicago, IL 60607 USA
[3] Marine Biol Lab, Ctr Ecosyst, Woods Hole, MA 02543 USA
[4] Univ Stirling, Sch Nat Sci, Stirling FK9 4LA, Scotland