[1] Univ Queensland, Ctr Marine Studies, ARC Ctr Excellence Coral Reef Studies, St Lucia, Qld 4072, Australia
[2] Univ Queensland, Dept Earth Sci, ARC Ctr Excellence Coral Reef Studies, St Lucia, Qld 4072, Australia
[3] Univ Calif San Diego, Ctr Marine Biodivers & Conservat, Scripps Inst Oceanog, La Jolla, CA 92093 USA
[4] Smithsonian Trop Res Inst, Ctr Trop Paleoecol & Archeol, Balboa, Panama