1. Graduate School of Bioresource and Bioenvironmental Sciences, Kyushu University, Fukuoka 812-8581, Japan
2. Section of Biological Sciences, Graduate School of Life Sciences, Hokkaido University, Sapporo 060-0810, Japan
3. National Salmon Resources Center, Fisheries Research Agency, Sapporo 062-0922,Japan
4. School of Aquatic and Fishery Sciences, University of Washington, Seattle, WA 98195, USA
5. Northwest Fisheries Science Center, NOAA Fisheries, Seattle, WA 99164,USA
6. Hokkaido National Fisheries Research Institute, Fisheries Research Agency,Kushiro 085-0802, Japan