[1] Univ Calif San Diego, Ctr Marine Biodivers & Conservat, Scripps Inst Oceanog, 9500 Gilman Dr, San Diego, CA 92093 USA
[2] Smithsonian Trop Res Inst, Apartado Balboa, Ancon 084303092, Panama
[3] Calif State Univ Northridge, Dept Biol, 18111 Nordhoff St, Northridge, CA 91330 USA
[4] Univ Western Australia, Sch Earth & Environm, Crawley, WA 6009, Australia
[5] Univ Western Australia, ARC Ctr Coral Reef Studies, Crawley, WA 6009, Australia
[6] Southern Cross Univ, Sch Environm Sci & Engn, Mil Rd, Lismore, NSW 2480, Australia