1. Faculty/Graduate School of Fisheries Science, Hokkaido University , 3-1-1 Minato-Cho, Hakodate, Hokkaido 041-8611 , Japan
2. Demonstration Laboratory, Marine Ecology Research Institute Present Address: , 4-7-17 Arahama, Kashiwazaki, Niigata 945-0017 , Japan
3. Arctic Research Centre, Hokkaido University , Kita-21 Nishi-11 Kita-Ku, Sapporo, Hokkaido 001-0021 , Japan
4. Institute of Arctic Climate and Environment, Japan Agency for Marine-Earth Science and Technology , 2-15 Natsushima-Cho, Yokosuka, Kanagawa 237-0061 , Japan
5. Atmosphere and Ocean Research Institute, The University of Tokyo , 5-1-5 Kashiwanoha, Kashiwa-Shi 277-8564 , Japan
6. Polar Biological Oceanography, Alfred Wegener Institute Helmholtz Centre for Polar and Marine Research , Am Handelshafen 12, De-27570 Bremerhaven , Germany