[1] Hokkaido Univ, Inst Low Temp Sci, Sapporo, Hokkaido, Japan
[2] Natl Cent Univ, Grad Inst Hydrol & Ocean Sci, Zhongli, Taiwan
[3] Hokkaido Univ, Fac Fisheries Sci, Grad Sch Fisheries Sci, Hakodate, Japan
[4] Hokkaido Univ, Arctic Res Ctr, Sapporo, Japan
[5] Hokkaido Univ, Field Sci Ctr Northern Biosphere, Hakodate, Hokkaido, Japan