a Shandong Key Laboratory of Eco-Environmental Science for the Yellow River Delta, Shandong University of Aeronautics, Binzhou 256603, China; b Key Laboratory of Vegetation and Environmental Change, Institute of Botany, the Chinese Academy of Sciences, Beijing 100093, China; c School of Life Sciences, Hebei University, Baoding 071002, China; d Jiangxi Provincial Key Laboratory of Carbon Neutrality and Ecosystem Carbon Sink, Lushan Botanical Garden, Jiangxi Province and Chinese Academy of Sciences, Jiujiang, 332900, China; e Soil Zoology Department, Senckenberg Museum of Natural History Görlitz, Am Museum 1, 02826 Görlitz, Germany #These authors contributed equally to this work. *Corresponding author: Jianhui Huang,E-mail: jhhuang@ibcas.ac.cn
We are deeply grateful to the staff of the Erguna Forest-Steppe Ecotone Research Station for their invaluable assistance during our field work. This work was supported by the Young Scientists Fund of the National Natural Science Foundation of China (32501469), National Key Research and Development Program of China (2022YFF1300602), Open Research Fund Program of Shandong Key Laboratory of Eco-Environmental Science for the Yellow River Delta, Shandong University of Aeronautics (2023KFJJ02) and the PhD Research Startup Foundation of Shandong University of Aeronautics (2026Y18).