1. Graduate School of Agricultural and Life Sciences, The University of Tokyo, Yayoi 1-1-1, Bunkyo-ku, Tokyo 113-8657, Japan, National Institute of Agrobiological Resources (NIAR), 2-1-2 Kannondai, Tsukuba, Ibaraki 305-8602, Japan, and Graduate School of Bioresource and Bioenvironmental Sciences, Kyushu University, 6-10-1, Hakozaki, Fukuoka, 812-8581, Japan