1. ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, Queensland 4811, Australia
2. College of Science and Engineering, James Cook University, Townsville, Queensland 4811, Australia
3. Function, Evolution and Anatomy Research (FEAR) Lab and Palaeoscience Research Centre, School of Environmental and Rural Science, University of New England, Armidale, New South Wales 2351, Australia