[1] ARC, Ctr Excellence Coral Reef Studies, St Lucia, Qld 4072, Australia
[2] Univ Queensland, Sch Biol Sci, St Lucia, Qld 4072, Australia
[3] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld 4011, Australia
[4] James Cook Univ, Sch Marine & Trop Biol, Townsville, Qld 4011, Australia
[5] Woods Hole Oceanog Inst, Dept Geol & Geophys, Woods Hole, MA 02543 USA