1. Graduate School of Environment and Information Sciences; Yokohama National University; Yokohama Kanagawa 240-8501 Japan
2. Field Science Center for Northern Biosphere; Hokkaido University; Toikanbetsu 131 Horonobe Hokkaido 098-2943 Japan
3. Center for Ecological Research; Kyoto University; Otsu Shiga 520-2113 Japan
4. Department of Ecology, Evolution and Behavior; University of Minnesota; St Paul MN 55108 USA