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Authors
Luo, Jiali
Li, Jinlong
Sun, Ying
Degen, Abraham Allan
Wang, Xiaoting
Xiong, Junlan
Aqeel, Muhammad
Xie, Shubin
Tang, Dong
Li, Fan
Hu, Weigang
Dong, Longwei
Peng, Jie
Hou, Qingqing
Deng, Yan
Ran, Jinzhi
Deng, Jianming
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Funding
Funding for this research was provided by:
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
National Key Research and Development Program of China (2023YFF0805602)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Natural Science Foundation of Gansu Province (21JR1RA138, 22JR5RA525, 23JRRA1157)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Key Research and Development Programs in Gansu Province (23ZDKA0010, 23ZDNA009, 24ZD13NA016)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
Research Grant Support from Lanzhou City (127000-563224111)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
National Natural Science Foundation of China (32225032, 32271597)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
the National Natural Science Foundation of China (32501402)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
Gansu Provincial Youth Science and Technology Fund (25JRRA710)
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Article History
Received: 23 June 2025
Accepted: 19 December 2025
First Online: 3 January 2026
Declarations
:
: The collection of leaf samples was approved by the Forestry Bureaus of Gansu Province and Shaanxi Province. Field studies and sample collection strictly complied with Chinese law as well as local legislations of the collection areas. All collection activities were conducted in a sustainable and non-destructive manner to minimize disturbance to natural populations and their habitats. No species listed under the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) were collected in this study. Plant specimens were formally identified by Ying Sun and Jiali Luo, and voucher specimens have been deposited in the Herbarium of the School of Life Sciences, Lanzhou University under accession number LZU98859. We confirm that all aspects of the study comply with institutional, national, and international guidelines.
: Not applicable.
: The authors declare no competing interests.
