1. ARC Centre of Excellence for Coral Reef Studies and Centre of Marine Sciences, School of Biological Sciences, The University of Queensland, St Lucia, Queensland 4072, Australia
2. Smithsonian Tropical Research Institute, Naos Marine and Molecular Laboratories, Balboa 0843-03092, Panamá
3. School of Biological Sciences, The University of Queensland, St Lucia, Queensland 4072, Australia
4. School of BioSciences, The University of Melbourne, Parkville, Victoria 3010, Australia