[1] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, QLD 4811, Australia
[2] James Cook Univ, Sch Marine & Trop Biol, Townsville, QLD 4811, Australia
[3] Newcastle Univ, Sch Marine Sci & Technol, Newcastle Upon Tyne NE1 7RU, Tyne & Wear, England
[4] Marine Programs, Wildlife Conservat Soc, Bronx, NY 10460 USA