[1] Edith Cowan Univ, Ctr Marine Ecosyst Res, Joondalup, WA 6027, Australia
[2] Univ Western Australia, UWA Oceans Inst, Crawley, WA 6009, Australia
[3] Univ Western Australia, Sch Plant Biol M096, Crawley, WA 6009, Australia
[4] Univ Canterbury, Sch Biol Sci, Marine Ecol Res Grp, Christchurch 1, New Zealand
[5] Australian Inst Marine Sci, Crawley, WA 6009, Australia