1. Graduate School of Environmental Science, Hokkaido University, Akkeshi, Hokkaido 088-1113, Japan
2. Akkeshi Marine Station, Field Science Center for Northern Biosphere, Hokkaido University, Akkeshi, Hokkaido 088-1113, Japan
3. Department of Life Sciences and Innovation and Development Center of Sustainable Agriculture, National Chung Hsing University, Taichung 402, Taiwan
4. Department of Zoology and Biodiversity Research Centre, University of British Columbia, Vancouver, BC V6T 1Z4, Canada