[1] Hokkaido Univ, Grad Sch Environm Sci, 20-5 Benten Cho, Hakodate, Hokkaido 0400051, Japan
[2] Hokkaido Univ, Grad Sch Fisheries Sci, 3-1-1 Minato Cho, Hakodate, Hokkaido 0418611, Japan
[3] Sasakawa Peace Fdn, Ocean Policy Res Inst, Minato Ku, I-15-16 Toranomon, Tokyo 1058524, Japan
[4] Tokyo Univ Marine Sci & Technol, Minato Ku, 4-5-7 Konan, Tokyo 1088477, Japan
[5] Japan Agcy Marine Earth Sci & Technol, Inst Arctic Climate & Environm Res, 2-15 Natsushima Cho, Yokosuka, Kanagawa 2370061, Japan
[6] Japan Fisheries Res & Educ Agcy, Natl Res & Dev Agcy, Fisheries Resources Inst, 1-5939-22 Suidou Cho, Niigata, Niigata 9518121, Japan
[7] Hokkaido Univ, Field Sci Ctr Northern Biosphere, 20-5 Benten Cho, Hakodate, Hokkaido 0400051, Japan
[8] Kyoto Univ, Wildlife Res Ctr, Sakyo Ku, 2-24 Tanaka Sekiden Cho, Kyoto 6068203, Japan