[1] King Abdullah Univ Sci & Technol, Div Biol & Environm Sci & Engn, Red Sea Res Ctr, Thuwal 23955, Saudi Arabia
[2] James Cook Univ, Australian Res Council, Ctr Excellence Coral Reef Studies, Townsville, Qld 4811, Australia
[3] James Cook Univ, Coll Marine & Environm Sci, Townsville, Qld 4811, Australia
[4] Woods Hole Oceanog Inst, Dept Biol, Woods Hole, MA 02543 USA