[1] Hokkaido Univ, Fac Environm Earth Sci, Kita Ku, North 10 West 5, Sapporo, Hokkaido 0600810, Japan
[2] Japan Sci & Technol, CREST, Kita Ku, North 10 West 5, Sapporo, Hokkaido 0600810, Japan
[3] Tokai Univ, Dept Marine Biol & Sci, Minami Ku, Sapporo, Hokkaido 0058601, Japan
[4] Natl Inst Polar Res, 10-3 Midori Cho, Tachikawa, Tokyo 1908518, Japan
[5] Ishinomaki Senshu Univ, Dept Biol Sci, Ishinomaki, Miyagi 9868580, Japan
[6] Hokkaido Univ, Inst Low Temp Sci, Pan Okhotsk Res Ctr, Kita Ku, North 19 West 8, Sapporo, Hokkaido 0600819, Japan