1. ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, Queensland 4811, Australia
2. Department of Marine Biology and Aquaculture, College of Science and Engineering, James Cook University, Townsville, Queensland 4811, Australia
3. Biosciences, College of Life and Environmental Sciences, University of Exeter, Geoffrey Pope, Stocker Road, Exeter EX4 4QD, UK