1 Key Laboratory of Adaptation and Evolution of Plateau Biota, Northwest Institute of Plateau Biology, the Chinese Academy of Sciences, Xining 810001, China 2 Institute of Molecular Ecology, MOE Key Laboratory of Arid and Grassland Ecology, School of Life Science, Lanzhou University, Lanzhou 730000, China 3 Department of Botany, the University Museum, the University of Tokyo, Tokyo 113-0033, Japan 4 Graduate University of the Chinese Academy of Sciences, Beijing 100049, China