[1] Australian Inst Marine Sci, Townsville, Qld 4810, Australia
[2] James Cook Univ, AIMS JCU, Townsville, Qld 4810, Australia
[3] James Cook Univ, Sch Marine & Trop Biol, Townsville, Qld 4811, Australia
[4] James Cook Univ, ARC Ctr Excellence Coral Reef Studies, Townsville, Qld 4811, Australia
[5] Univ Brest, Univ Europeenne Bretagne, ORPHY, UFR Sci & Tech,EA 4324, F-29238 Brest, France