[1] Univ Tokyo, Grad Sch Agr & Life Sci, Dept Ecosyst Studies, Tokyo, Japan
[2] Hokkaido Univ, Field Sci Ctr Northern Biosphere, Nayoro Stn, 250 Tokuda, Nayoroshi, Hokkaido, Japan
[3] Univ Tokyo, Grad Sch Agr & Life Sci, Dept Ecosyst Studies, Tokyo 1138657, Japan