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Acknowledgements We are grateful for Dr. Jens Strauss and the other two anonymous reviewers for their insightful comments on an earlier version of this MS, and appreciate members of the IBCAS Sampling Campaign Teams for their assistance in field investigation. This work was supported by the National Basic Research Program of China on Global Change (2014CB954001 and 2015CB954201), National Natural Science Foundation of China (31322011 and 41371213), and the Thousand Young Talents Program.