[1] Hokkaido Univ, Grad Sch Environm Sci, Sapporo, Hokkaido, Japan
[2] Tohoku Natl Fisheries Res Inst, Shiogama, Japan
[3] Univ Tokyo, Atmosphere & Ocean Res Inst, Kashiwa, Chiba, Japan
[4] Univ Tokyo, Grad Sch Agr & Life Sci, Tokyo, Japan
[5] Nagasaki Univ, Fac Fisheries & Environm Sci, Nagasaki, Japan
[6] Hokkaido Univ, Inst Low Temp Sci, Sapporo, Hokkaido, Japan
[7] Hokkaido Univ, Fac Fisheries Sci, Hakodate, Hokkaido, Japan
[8] Hokkaido Tokai Univ, Dept Marine Biol & Sci, Sapporo, Hokkaido, Japan
[9] Bowling Green State Univ, Dept Biol Sci, Bowling Green, OH 43403 USA
[10] Hokkaido Univ, Fac Environm Earth Sci, Sapporo, Hokkaido, Japan
[11] Hokkaido Univ, Field Sci Ctr Northern Biosphere, Akkeshi Marine Stn, Akkeshi, Hokkaido, Japan