[1] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Coll Sci & Engn, Townsville, Qld 4811, Australia
[2] James Cook Univ, Australian Inst Marine Sci, Div Res & Innovat, Townsville, Qld 4811, Australia
[3] Australian Inst Marine Sci, Townsville, Qld 4810, Australia
[4] Max Planck Inst Marine Microbiol, Bremen, Germany