Financial support was provided by the National Natural Science Foundation of China (No. 31700422, 41731176, 41922056, 31872691), Guangdong Basic and Applied Basic Research Foundation (2019A1515011642), and China Postdoctoral Science Foundation (2017M612775). We are grateful to Dr. Weixing Zhu and two anonymous reviewers for invaluable suggestions and comments. We appreciate Yujia Huang, Nianzhuang Liu, Lijie Deng, Juan Huang and Hui Mo for their assistance in laboratory and fieldwork.