1. Akkeshi Marine Station, Hokkaido University, Akkeshi, Hokkaido, Japan
2. Faculty of Fisheries Science, Hokkaido University, Hakodate, Hokkaido, Japan
3. Department of Biological Sciences, Hokkaido University, Sapporo, Hokkaido, Japan
4. Kiritappu Wetland National Trust, Hamanaka, Hokkaido, Japan
5. Graduate School of Environmental Science, Hokkaido University, Sapporo, Hokkaido, Japan