1. Marine Biology and Aquaculture, College of Science and Engineering, James Cook University, Townsville, Queensland 4811, Australia
2. ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, Queensland 4811, Australia
3. School of Biological Sciences, Life Sciences Building, University of Bristol, Bristol BS8 1TQ, UK
4. Coastal Marine Field Station, School of Science, University of Waikato, Tauranga 3110, New Zealand