1. Key Laboratory of Alpine Ecology, Institute of Tibetan Plateau Research, Chinese Academy of Sciences , Beijing 100101 , China
2. University of the Chinese Academy of Sciences , Beijing 100049 , China
3. College of Science, Tibet University , Lhasa 850000 , China
4. CAS Center for Excellence in Tibetan Plateau Earth Science of the Chinese Academy of Sciences , Beijing 100101 , China
5. Key Laboratory of Adaptation and Evolution of Plateau Biota, Northwestern Institute of Plateau Biology, Chinese Academy of Sciences , Xining 810008 , China