1. Department of Biology, University of North Carolina at Greensboro, Greensboro, NC 27412
2. Salmon and Freshwater Fisheries Research Institute, Hokkaido Research Organization, Eniwa, Hokkaido 061-1433, Japan
3. Faculty of Environmental Earth Science, Hokkaido University, Sapporo, Hokkaido 060-0810, Japan
4. National Institute of Polar Research, Tachikawa, Tokyo 190-8518, Japan