[1] Nihon Univ, Dept Marine Sci & Resources, Fujisawa, Kanagawa 2520880, Japan
[2] Hokkaido Univ, Fac Fisheries Sci, Hakodate, Hokkaido 0418611, Japan
[3] Univ Tokyo, Ocean Alliance, Res Ctr Marine Educ, Bunkyo Ku, Tokyo 1130033, Japan
[4] Tokyo Univ Marine Sci & Technol, Dept Ocean Sci, Minato Ku, Tokyo 1088477, Japan