[1] Ehime Univ, Ctr Marine Environm Studies, Matsuyama, Ehime, Japan
[2] Hokkaido Univ, Dept Nat Hist Sci, Grad Sch Sci, Sapporo, Hokkaido, Japan
[3] Nagasaki Univ, Grad Sch Fisheries Sci & Environm Studies, Nagasaki 852, Japan
[4] Kagoshima Univ, Fac Fisheries, Kagoshima 890, Japan
[5] Mie Univ, Climate & Ecosyst Dynam Div, Tsu, Mie 514, Japan
[6] Japan Agcy Marine Earth Sci & Technol, Res Inst Global Change, Yokosuka, Kanagawa 2370061, Japan
[7] Kagoshima Univ, Fac Engn, Kagoshima 890, Japan