1. Graduate School of Environmental Science, Hokkaido University, 20-5 Benten-cho, Hakodate, Hokkaido 040-0051, Japan
2. Wakkanai Fisheries Research Institute, Hokkaido Research Organization, 4-5-15 Suehiro, Wakkanai, Hokkaido 097-0001, Japan
3. Field Science Center for Northern Biosphere, Hokkaido University, 1 Aikappu, Akkeshi-cho, Akkeshi-gun, Hokkaido 088-1113, Japan
4. Field Science Center for Northern Biosphere, Hokkaido University, 20-5 Benten-cho, Hakodate, Hokkaido 040-0051, Japan