[1] Univ Western Australia, UWA Oceans Inst, Crawley, WA 6009, Australia
[2] Univ Western Australia, Sch Biol Sci, Crawley, WA 6009, Australia
[3] UWA Great Southern Marine Res Facil, Albany, WA 6330, Australia
[4] Univ Southampton, Natl Oceanog Ctr Southampton, Sch Ocean & Earth Sci, Southampton SO17 1BJ, England
[5] Southern Marine Sci & Engn Guangdong Lab Guangzhou, Guangzhou 511458, Peoples R China