[1] Univ Tasmania, Inst Marine & Antarctic Studies, Hobart, Tas, Australia
[2] Australian Inst Marine Sci, Townsville, Qld 4810, Australia
[3] James Cook Univ, Australian Res Council Ctr Excellence Coral Reef, Townsville, Qld 4811, Australia
[4] Carl von Ossietzky Univ Oldenburg, Inst Biol & Environm Sci, D-2900 Oldenburg, Germany