[1] James Cook Univ, TropWATER, Townsville, Qld 4811, Australia
[2] CSIRO, Brisbane, Qld 4068, Australia
[3] Australian Inst Marine Sci, PMB 3, Townsville, Qld 4810, Australia
[4] Dept Environm & Sci, GPO Box 5078, Brisbane, Qld 4001, Australia
[5] Univ Queensland, Sch Earth & Environm Sci, St Lucia, Qld 4072, Australia
[6] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld 4811, Australia