1. Atmosphere and Ocean Research Institute, The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa, Chiba 277-8564, Japan
2. Graduate School of Fisheries Sciences, Hokkaido University, Minato-cho 3-1-1, Hakodate 041-8611, Japan
3. Centre for Ecology & Hydrology, Bush Estate, Penicuik, Midlothian EH26 0QB, UK