[1] Univ Calif Santa Barbara, Dept Ecol Evolut & Marine Biol, Santa Barbara, CA 93106 USA
[2] Univ Calif Santa Barbara, Inst Marine Sci, Coastal Res Ctr, Santa Barbara, CA 93106 USA
[3] James Cook Univ, Ctr Excellence Coral Reef Studies, Australian Res Council, Townsville, Qld 4811, Australia
[4] James Cook Univ, Sch Marine & Trop Biol, Townsville, Qld 4811, Australia