1. Institute of Tibetan Plateau Research, Chinese Academy of Sciences, Beijing 100101, China
2. Graduate University of Chinese Academy of Sciences, Beijing 100049, China
3. The authors contributed equally to this paper
4. Key Laboratory for Plant Diversity and Biogeography of East Asia, Chinese Academy of Sciences, Kunming 650204, China
5. Plant Germplasm and Genome Center, Germplasm Bank of Wild Species, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, China
6. Institute of Tibetan Plateau Research at Kunming, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, China
7. Qinghai Academy of Animal Husbandry and Veterinary Medicinal Sciences, Xining 810016, China