1. Graduate School of Environmental Science Hokkaido University Sapporo 060‐0810 Japan
2. Muroran Marine Station Field Science Center for Northern Biosphere Hokkaido University Muroran 051‐0013 Japan
3. Graduate School of Environmental Earth Science Hokkaido University Sapporo 060‐0810 Japan
4. Sorbonne Universités CNRS Integrative Biology of Marine Models (LBI2M) Station Biologique de Roscoff 29680 Roscoff France