[1] Ehime Univ, Global Ctr Excellence, Ctr Marine Environm Studies, Matsuyama, Ehime 7908577, Japan
[2] NOAA, NW Fisheries Sci Ctr, Natl Marine Fisheries Serv, Fishery Resource Anal & Monitoring Div, Newport, OR 97365 USA
[3] NOAA, NW Fisheries Sci Ctr, Natl Marine Fisheries Serv, Fish Ecol Div, Newport, OR 97365 USA