[1] James Cook Univ, Australian Res Council Ctr Excellence Coral Reef, Townsville, Qld 4811, Australia
[2] Univ Calif San Diego, Scripps Inst Oceanog, La Jolla, CA 92093 USA
[3] Smithsonian Inst, Dept Paleobiol, Natl Museum Nat Hist, Washington, DC 20013 USA
[4] Natl Ctr Atmospher Res, POB 3000, Boulder, CO 80307 USA
[5] Wageningen Univ, Dept Environm Sci, POB 47, NL-6700 AA Wageningen, Netherlands
[6] Australian Inst Marine Sci, PMB 3, Townsville, Qld 4810, Australia
[7] Stanford Univ, Dept Biol, Hopkins Marine Stn, Pacific Grove, CA 93950 USA