[1] Univ Western Australia M470, Sch Environm Syst Engn, Crawley, WA 6009, Australia
[2] Univ Western Australia M470, UWA Oceans Inst, Crawley, WA 6009, Australia
[3] Australian Inst Marine Sci, UWA Oceans Inst M096, Crawley, WA 6009, Australia
[4] CSIRO Marine & Atmospher Res, Hobart, Tas 7001, Australia
[5] Stanford Univ, Hopkins Marine Stn, Pacific Grove, CA 93950 USA
[6] NOAA, Pacific Islands Fisheries Sci Ctr, Natl Marine Fisheries Serv, Honolulu, HI 96822 USA