1. Australian Research Council Centre of Excellence for Coral Reef Studies and School of Marine Biology and Tropical Biology, James Cook UniversityTownsville, Queensland 4811, Australia
2. Australian Institute of Marine SciencePMB No. 3, Townsville MC, Queensland 4810, Australia
3. Institute of Evolutionary Biology, School of Biological Sciences, University of EdinburghKings Buildings, Edinburgh EH9 3JT, UK