[1] James Cook Univ, Coll Sci & Engn, Townsville, Qld 4811, Australia
[2] James Cook Univ, Res Hub Coral Reef Ecosyst Funct, Townsville, Qld, Australia
[3] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld, Australia
[4] Australian Inst Marine Sci, Townsville, Qld, Australia
[5] Univ New South Wales, Ctr Marine Sci & Innovat, Sch Biol Earth & Environm Sci, Kensington, NSW, Australia