1. Australian Research Council Centre of Excellence for Coral Reef Studies, and School of Marine and Tropical Biology; James Cook University; Townsville QLD 4811 Australia
2. Department of Ecology, Evolution, and Marine Biology; University of California; Santa Barbara CA 93106 USA
3. School of Biological Sciences; Monash University; Melbourne VIC 3800 Australia
4. Australian Research Council Centre of Excellence for Coral Reef Studies; Centre for Marine Science, and School of Biological Sciences; The University of Queensland; Brisbane QLD 4072 Australia