[1] Kagoshima Univ, Fac Fisheries, 4-50-20 Shimoarata, Kagoshima 8900056, Japan
[2] Kagoshima Univ, Grad Sch Agr Forestry & Fisheries, 4-50-20 Shimoarata, Kagoshima 8900056, Japan
[3] Hokkaido Univ, Field Sci Ctr Northern Biosphere, Tomakomai Expt Forest, Takaoka, Hokkaido 0530035, Japan
[4] Nagoya Univ, Grad Sch Sci, Sugashima Marine Biol Lab, Toba, Mie 5170004, Japan
[5] Univ Sains Malaysia 11800 USM, Ctr Marine & Coastal Studies, George Town, Malaysia