[1] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld, Australia
[2] Univ Queensland, Sch Earth & Environm Sci, St Lucia, Qld 4072, Australia
[3] Univ Queensland, Ctr Biodivers & Conservat Sci, St Lucia, Qld 4072, Australia
[4] Wildlife Conservat Soc, Marine Program, Bronx, NY 10460 USA