1. Centre for Biological Diversity, University of St Andrews, Sir Harold Mitchell Building; St Andrews KY16 9TF Scotland UK
2. Department of Biology and CESAM; Universidade de Aveiro, Campus Universitário de Santiago; Aveiro, 3810-193 Portugal
3. College of Science and Engineering, and ARC Centre of Excellence for Coral Reef Studies; James Cook University; Townsville QLD 4810 Australia