[1] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld 4811, Australia
[2] James Cook Univ, Sch Earth & Environm Sci, Townsville, Qld 4811, Australia
[3] James Cook Univ, Sch Earth & Environm Sci, Cairns, Qld, Australia
[4] Australian Inst Marine Sci, Townsville, Qld, Australia
[5] Univ Sydney, Sch Earth Sci, Geocoastal Res Grp, Sydney, NSW 2006, Australia