[1] James Cook Univ, Australian Res Council, Ctr Excellence Coral Reef Studies, Townsville, Qld, Australia
[2] James Cook Univ, Sch Marine & Trop Biol, Townsville, Qld, Australia
[3] Univ New S Wales, Australian Wetlands & Rivers Ctr, Sch Biol Earth & Environm Sci, Sydney, NSW, Australia