[1] Hokkaido Univ, Fac Environm Earth Sci, Sapporo, Hokkaido 060, Japan
[2] Univ Plymouth, Sch Marine Sci & Engn, Plymouth PL4 8AA, Devon, England
[3] Bayworld Ctr Res & Educ, Cape Town, South Africa
[4] Univ Cape Town, Marine Res Inst, ZA-7700 Rondebosch, South Africa
[5] Plymouth Marine Lab, Plymouth, Devon, England
[6] Hokkaido Univ, Fac Fisheries Sci, Sapporo, Hokkaido 060, Japan
[7] NOAA, Pacific Isl Fisheries Sci Ctr, Silver Spring, MD USA
[8] Japan Sci & Technol Agcy, CREST, Tokyo, Japan
[9] Japan Agcy Marine Earth Sci & Technol JAMSTEC, Res Inst Global Change, Tokyo, Japan