[1] Hokkaido Univ, Field Sci Ctr Northern Biosphere, Akkeshi Marine Stn, Akkeshi, Hokkaido 0881113, Japan
[2] Univ Tokyo, Atmosphere & Ocean Res Inst, 5-1-5 Kashiwanoha, Kashiwa, Chiba 2778564, Japan
[3] Acad Sinica, Res Ctr Biodivers, Taipei 115, Taiwan
[4] Kitasato Univ, Sch Marine Biosci, Minami Ku, 1-15-1 Kitasato, Sagamihara, Kanagawa 2520373, Japan
[5] Japan Agcy Marine Earth Sci & Technol, 2-15 Natsushima Cho, Yokosuka, Kanagawa 2370061, Japan
[6] Univ Copenhagen, Dept Biol, Marine Biol Sect, Univ Pk 4, DK-2100 Copenhagen, Denmark