[1] Bournemouth Univ, Sch Appl Sci, Ctr Conservat Ecol & Environm Sci, Poole BH12 5BB, Dorset, England
[2] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld 4811, Australia
[3] Ctr Ecol & Hydrol, Wallingford OX10 8BB, Oxon, England