1. School of Fisheries Sciences, Hokkaido University , 3−1−1 Minato-cho, Hakodate, Hokkaido 041−8611, Japan
2. Graduate School of Fisheries Sciences, Hokkaido University , 3−1−1 Minato-cho, Hakodate, Hokkaido 041−8611, Japan
3. School of Marine Biosciences, Kitasato University , 1–15–1 Kitasato, Minami-ku, Sagamihara 252–0373, Japan
4. Japan Agency for Marine-Earth Science and Technology , 2−15 Natsushima-cho, Yokosuka, Kanagawa 237−0061, Japan
5. Faculty of Fisheries Sciences, Hokkaido University , 3−1−1 Minato-cho, Hakodate, Hokkaido 041−8611, Japan
6. Arctic Research Center, Hokkaido University , Kita-21 Nishi-11 Kita-ku, Sapporo, Hokkaido 001−0021, Japan