[1] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld 4811, Australia
[2] Australian Inst Marine Sci, Townsville, Qld 4810, Australia
[3] Dept Biodivers Conservat & Attract, Kensington, WA 6151, Australia
[4] Univ Western Australia, Oceans Inst, Crawley, WA 6009, Australia
[5] James Cook Univ, Coll Sci & Engn, Townsville, Qld 4811, Australia
[6] James Cook Univ, Ctr Trop Water & Aquat Syst Res, Townsville, Qld 4811, Australia
[7] Great Barrier Reef Marine Pk Author, Townsville, Qld 4811, Australia