[1] Univ New South Wales, Sch Biol Earth & Environm Sci, Evolut & Ecol Res Ctr, Sydney, NSW 2052, Australia
[2] Univ Sydney, Coastal & Marine Ecosyst Grp, Sch Life & Environm Sci, Sydney, NSW 2006, Australia
[3] Sydney Inst Marine Sci, Chowder Bay Rd, Mosman, NSW 2061, Australia
[4] Australian Inst Marine Sci, PMB 3, Townsville, Qld 4810, Australia