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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Plant Sci.</journal-id>
<journal-title>Frontiers in Plant Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Plant Sci.</abbrev-journal-title>
<issn pub-type="epub">1664-462X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fpls.2024.1470653</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Plant Science</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Comprehensive prediction of potential spatiotemporal distribution patterns, priority planting regions, and introduction adaptability of <italic>Elymus sibiricus</italic> in the Chinese region</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Lu</surname>
<given-names>Huan-Huan</given-names>
</name>
<uri xlink:href="https://loop.frontiersin.org/people/1385025"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Zheng</surname>
<given-names>Yu-Ying</given-names>
</name>
<uri xlink:href="https://loop.frontiersin.org/people/1731768"/>
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<contrib contrib-type="author">
<name>
<surname>Qiu</surname>
<given-names>Yong-Sen</given-names>
</name>
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<contrib contrib-type="author">
<name>
<surname>Tang</surname>
<given-names>Liu-Ban</given-names>
</name>
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<contrib contrib-type="author">
<name>
<surname>Zhao</surname>
<given-names>Yan-Cui</given-names>
</name>
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<contrib contrib-type="author" corresp="yes">
<name>
<surname>Xie</surname>
<given-names>Wen-Gang</given-names>
</name>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
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</contrib-group>
<aff id="aff1">
<institution>State Key Laboratory of Herbage Improvement and Grassland Agro-Ecosystems, Key Laboratory of Grassland Livestock Industry Innovation, Ministry of Agriculture and Rural Affairs, College of Pastoral Agriculture Science and Technology, Lanzhou University</institution>, <addr-line>Lanzhou</addr-line>, <country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Yangong Du, Chinese Academy of Sciences (CAS), China</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Dou Quanwen, Chinese Academy of Sciences (CAS), China</p>
<p>Xing Fan, Sichuan Agricultural University, China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Wen-Gang Xie, <email xlink:href="mailto:xiewg@lzu.edu.cn">xiewg@lzu.edu.cn</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>08</day>
<month>01</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>15</volume>
<elocation-id>1470653</elocation-id>
<history>
<date date-type="received">
<day>25</day>
<month>07</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>13</day>
<month>12</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Lu, Zheng, Qiu, Tang, Zhao and Xie</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Lu, Zheng, Qiu, Tang, Zhao and Xie</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>The natural grassland in China is facing increasingly serious degradation. <italic>Elymus sibiricus</italic> L., as an important native alpine grass, is widely used in the restoration and improvement of natural grassland. In this study, the geographical distribution and environmental data of <italic>E. sibiricus</italic> in China were collected, and the potential spatiotemporal distribution pattern, planting pattern, and introduction adaptability of <italic>E. sibiricus</italic> were comprehensively predicted by using ensembled ecological niche model and Marxan model. The results show that (1) the potential spatial distribution of <italic>E. sibiricus</italic> mainly spans 33&#xb0;&#x2013;42&#xb0;N and 95&#xb0;&#x2013;118&#xb0;E. It was widely distributed in Qilian Mountains (northeast of Qinghai-Tibet Plateau), Taihang Mountains (junction of Loess Plateau and Inner Mongolia Plateau), and Tianshan Mountains; (2) with the passage of time, the suitable distribution regions of <italic>E. sibiricus</italic> generally showed a collapse trend, but its main distribution regions did not show obvious change, and the suitable regions (centroid) generally migrated to the southwest of China by 2.93 km; (3) the spatial distribution of <italic>E. sibiricus</italic> in the current period was significantly affected by the annual range of monthly near-surface relative humidity, mean annual air temperature, annual range of monthly potential evapotranspiration, mean monthly potential evapotranspiration, annual range of monthly climate moisture index, elevation, exchangeable Ca<sup>2+</sup>, available P, mean monthly near-surface relative humidity, exchangeable H<sup>+</sup>, and annual precipitation amount, respectively; (4) the potential planting area of <italic>E. sibiricus</italic> cover 2.059 &#xd7; 10<sup>5</sup> km<sup>2</sup>, which was mainly distributed in Qilian Mountains (northeast of Qinghai-Tibet Plateau), Taihang Mountains (southeast of Inner Mongolia Plateau), the middle part of Tianshan Mountains, and the southeast of Altai Mountains; (5) the introduction adaptability regions of six <italic>E. sibiricus</italic> germplasm (LM01&#x2013;LM06) were all distributed in the high-elevation regions of western China. The study aims to provide an effective theoretical basis for the collection, preservation, and utilization of <italic>E. sibiricus</italic> germplasm resources in China.</p>
</abstract>
<kwd-group>
<kwd>
<italic>Elymus sibiricus</italic> L.</kwd>
<kwd>functional plant ecology</kwd>
<kwd>global warming</kwd>
<kwd>ensemble ecological niche models</kwd>
<kwd>potential spatiotemporal distribution pattern</kwd>
<kwd>priority planting region prediction</kwd>
<kwd>adaptability evaluation of introduction</kwd>
</kwd-group>
<counts>
<fig-count count="5"/>
<table-count count="3"/>
<equation-count count="0"/>
<ref-count count="71"/>
<page-count count="14"/>
<word-count count="7295"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Functional Plant Ecology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Grasslands, covering 40% of the earth&#x2019;s surface and 69% of agricultural land, provide a wide range of ecosystem services for human beings, such as food production, water supply and regulation, carbon storage and climate mitigation, pollination, and a series of economic and cultural services (<xref ref-type="bibr" rid="B1">Bardgett et&#xa0;al., 2021</xref>). The area of all kinds of grassland in China is about 2.65 &#xd7; 10<sup>6</sup> km<sup>2</sup>, accounting for 27.6% of the total land (<xref ref-type="bibr" rid="B17">He et&#xa0;al., 2024</xref>). Due to human activities and natural factors, as many as 50% of grasslands worldwide have experienced varying degrees of degradation, nearly 5% of grasslands have been seriously degraded, and nearly 90% of grasslands in northern China have been degraded to some extent (<xref ref-type="bibr" rid="B21">Li et&#xa0;al., 2022</xref>). In addition, in the recent 130 years, the global average surface temperature has risen by 0.85&#xb0;C, and the surface temperature will continue to rise in the future (<xref ref-type="bibr" rid="B55">Wu et&#xa0;al., 2020a</xref>). The change in global climate will not only change the precipitation and species distribution pattern but also further reduce the coupling degree between grassland structure and function (<xref ref-type="bibr" rid="B36">O&#x2019;Mara, 2012</xref>). Grassland degradation has a profound impact on productivity decline, biodiversity loss, land degradation, and decline in ecosystem services.</p>
<p>Siberian wildrye (<italic>Elymus sibiricus</italic> L.), belonging to the genus <italic>Elymus</italic> of Poaceae, is a perennial, self-pollinated, allotetraploid (StStHH, 2<italic>n</italic> = 4<italic>x</italic> = 28) native alpine grass (<xref ref-type="bibr" rid="B51">Wang et&#xa0;al., 2017b</xref>; <xref ref-type="bibr" rid="B57">Xiong et&#xa0;al., 2021</xref>). As a model plant of <italic>Elymus</italic>, <italic>E. sibiricus</italic> is the dominant and constructive species in alpine meadow grassland in Qinghai-Tibet Plateau (QTP) (<xref ref-type="bibr" rid="B28">Liu et&#xa0;al., 2015</xref>), and its habitat is mainly composed of valley meadow, mountain meadow grassland, sparse forest, shrub, and open space in forest grassland (<xref ref-type="bibr" rid="B62">Yan et&#xa0;al., 2007</xref>). <italic>E. sibiricus</italic> has excellent cold resistance, drought resistance, strong adaptability, and high hay quality (<xref ref-type="bibr" rid="B67">Zhang et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B52">Wang et&#xa0;al., 2017a</xref>), which is widely used in the establishment of high-yield artificial grassland and the restoration of natural grassland in China. According to the Analysis Report on Supply and Demand of Grass Seeds in China in 2024, the total demand for various grass seeds in China in 2024 is 180,000&#x2013;200,000 tons, including 70,000&#x2013;80,000 tons for ecological restoration. In 2023, 16,500 tons of seeds of various ecological grasses were produced in China, among which the <italic>Elymus</italic> seed yield (including <italic>E. sibiricus</italic>) was 10,500 tons, accounting for 63.63% of the national ecological grass seed yield (<ext-link ext-link-type="uri" xlink:href="https://www.forestry.gov.cn/c/www/lcdt/528077.jhtml">https://www.forestry.gov.cn/c/www/lcdt/528077.jhtml</ext-link>). Ecological grass not only plays their ecological restoration functions but also is important forage grass in traditional pastoral areas, and there is a systematic and serious shortage of ecological grass seed (<xref ref-type="bibr" rid="B54">Wu et&#xa0;al., 2020b</xref>; <xref ref-type="bibr" rid="B20">Li et&#xa0;al., 2021</xref>). Therefore, <italic>E. sibiricus</italic> plays an important role in animal husbandry and ecological maintenance, and its utilization and management are in urgent need of scientific guidance.</p>
<p>Ecological niche models (ENMs), known as species distribution models (SDMs), use the known geographical distribution and environmental data of species to estimate the potential ecological niche of species and project it into the prediction regions, presenting the habitat characteristics of species in the form of probabilities (<xref ref-type="bibr" rid="B35">Miller, 2010</xref>), which is one of the important means of predicting the potential distribution of species. With the deepening of mathematical theory in ecological application research, the types of ENMs have gradually increased, such as generalized linear model (GLM), maximum entropy model (MAXENT), and artificial neural network model (ANN) (<xref ref-type="bibr" rid="B45">Schmitt et&#xa0;al., 2017</xref>). Stacked species distribution models integrate multiple ENMs to eliminate deviations and uncertainties among different algorithms and obtain more robust results and insights (<xref ref-type="bibr" rid="B9">Fan et&#xa0;al., 2024</xref>). Marxan model is an intelligent decision-making tool based on the concept of system conservation planning, which constructs a cost-effective optimization network according to the management objectives and costs of research objects to meet the biodiversity conservation objectives (<xref ref-type="bibr" rid="B53">Watts et&#xa0;al., 2017</xref>). ENMs and Marxan models are widely used in the prediction of potential geographical distribution of species, the assessment of biodiversity conservation, and the analysis of ecological characteristics (<xref ref-type="bibr" rid="B34">Merow and Silander, 2014</xref>; <xref ref-type="bibr" rid="B66">Zhang and Li, 2022</xref>).</p>
<p>At present, the research on <italic>E. sibiricus</italic> mainly focuses on biological characteristics (<xref ref-type="bibr" rid="B56">Xie et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B24">Li et&#xa0;al., 2020</xref>), genetic diversity (<xref ref-type="bibr" rid="B32">Ma et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B68">Zhang et&#xa0;al., 2015</xref>, <xref ref-type="bibr" rid="B69">2018</xref>), variety breeding (<xref ref-type="bibr" rid="B70">Zhou et&#xa0;al., 2011</xref>), cultivation, and utilization (<xref ref-type="bibr" rid="B65">Yang et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B30">Liu et&#xa0;al., 2012</xref>), but the integration of potential spatiotemporal distribution pattern and intelligent selection of planting regions of <italic>E. sibiricus</italic> in China was largely unexplored. Based on ecological theory, this study predicted the potential distribution, planting pattern, and introduction adaptability of different <italic>E. sibiricus</italic> germplasm, in order to answer four scientific questions: (1) the potential spatiotemporal distribution pattern characteristics of <italic>E. sibiricus</italic> in China, (2) the potential planting pattern of <italic>E. sibiricus</italic> in China, (3) The main environmental factors affecting the distribution pattern of <italic>E. sibiricus</italic>, and (4) the characteristics of introduction adaptability regions of different <italic>E. sibiricus</italic> germplasm. This study aims to provide theoretical guidance for the utilization, protection, and management of <italic>E. sibiricus</italic> germplasm resources in China.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Compilation and preprocessing of species distribution data</title>
<p>Based on the Chinese and Latin literary names of <italic>E. sibiricus</italic>, the distribution points in China were comprehensively compiled from the existing literature, field investigation, and public online database. Four online databases (GBIF, Ecoengine, iNaturalist, and iDigBio) related to plant distribution information were queried by the R package spocc v1.2.2 (<ext-link ext-link-type="uri" xlink:href="https://cran.R-project.org/package=spocc">https://cran.R-project.org/package=spocc</ext-link>), and the search ended on 1 June 2024. The species name and its classification were queried and corrected through the NCBI Taxonomy database using the R package taxize v0.9.1 (<ext-link ext-link-type="uri" xlink:href="https://CRAN.R-project.org/package=taxizedb">https://CRAN.R-project.org/package=taxizedb</ext-link>). The distribution data were checked for the common spatial and temporal errors using the R package CoordinateCleaner v3.0.1 (<xref ref-type="bibr" rid="B71">Zizka et&#xa0;al., 2019</xref>). To reduce spatial autocorrelation and sampling bias, the Spatially Rarefy Occurrence Data for SDMs tool in SDM Toolbox v2.4 (<xref ref-type="bibr" rid="B2">Brown et&#xa0;al., 2017</xref>) was used to rarefy the species distribution data to ensure that only one distribution point was contained in each environmental unit at 5 arc-min resolution (~10 km).</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Collection and preprocessing of environmental variable data</title>
<p>To effectively explore the potential spatiotemporal distribution pattern of <italic>E. sibiricus</italic>, this study obtained environmental variables (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>) that widely affect the plant growth and distribution, including bioclimatic, climatic, soil, and topographic data. Bioclimatic and climatic data were sourced from Paleoclim website (<ext-link ext-link-type="uri" xlink:href="http://www.paleoclim.org">http://www.paleoclim.org</ext-link>). Current bioclimatic and climatic data (CP, 1979&#x2013;2013) were obtained using the Anthropocene v1.2b dataset, while future bioclimatic and climatic data (FP, 2071&#x2013;2100) were obtained using the CMIP6 dataset under the MPI-ESM1-2-HR climate model (<xref ref-type="bibr" rid="B18">J&#xe4;germeyr et&#xa0;al., 2021</xref>) and the ssp375 economic sharing pathway (<xref ref-type="bibr" rid="B43">Riahi et&#xa0;al., 2017</xref>). These bioclimatic and climatic variables mainly represent long-term annual trends, seasonality, and extreme situations of temperature and precipitation. The biomass of grass roots was mainly distributed in 0&#x2013;30 cm soil layer to absorb soil nutrients, and the fibrous root system in 0&#x2013;10 cm soil layer was extremely important for growth and development (<xref ref-type="bibr" rid="B25">Li et&#xa0;al., 2011</xref>). Soil layer data (i.e., 0&#x2013;0.045 m, 0.045&#x2013;0.091 m) containing nutrients necessary for plant growth were obtained from the World Soil Database (<ext-link ext-link-type="uri" xlink:href="http://globalchange.bnu.edu.cn/research/soilw">http://globalchange.bnu.edu.cn/research/soilw</ext-link>) (<xref ref-type="bibr" rid="B46">Shangguan et&#xa0;al., 2014</xref>), and the average value of different soil layer data was calculated by ArcGIS v10.8 (<ext-link ext-link-type="uri" xlink:href="https://desktop.arcgis.com/">https://desktop.arcgis.com/</ext-link>). Elevation indirectly affects the adaptability of species to the environment by affecting temperature, and the SRTM elevation data was selected from the WorldClim website (<ext-link ext-link-type="uri" xlink:href="https://www.worldclim.org">https://www.worldclim.org</ext-link>). All environmental variables were unified into GCS_WGS_1984, resampled to 5 arc-min resolution and masked to China region by ArcGIS software. The China region was created from the map with review number GS (2019) 1822, and the base map was not modified. In order to avoid the multicollinearity of environmental variables leading to overfitting of the prediction model, the Correlations and Summary Stats tool in SDM Toolbox was used to detect the correlation among environmental variables, retaining only environmental variables with |<italic>r</italic>| &lt;0.80 and significant ecological significance. The reserved environment variables were normalized through Circuitscape tool v1.0.2 (<ext-link ext-link-type="uri" xlink:href="http://www.circuitscape.org">http://www.circuitscape.org</ext-link>) as a reference with elevation data.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Environmental variables for predicting the potential spatiotemporal distribution pattern of <italic>E. sibiricus</italic>.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Environment variable types</th>
<th valign="middle" align="center">Environmental factors</th>
<th valign="middle" align="center">Description</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">Bioclimatic variables</td>
<td valign="middle" align="center">Bio01</td>
<td valign="middle" align="center">Mean annual air temperature</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio02</td>
<td valign="middle" align="center">Mean diurnal air temperature range</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio03</td>
<td valign="middle" align="center">Isothermality</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio04</td>
<td valign="middle" align="center">Temperature seasonality</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio05</td>
<td valign="middle" align="center">Mean daily maximum air temperature of the warmest month</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio06</td>
<td valign="middle" align="center">Mean daily minimum air temperature of the coldest month</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio07</td>
<td valign="middle" align="center">Annual range of air temperature</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio08</td>
<td valign="middle" align="center">Mean daily mean air temperatures of the wettest quarter</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio09</td>
<td valign="middle" align="center">Mean daily mean air temperatures of the driest quarter</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio10</td>
<td valign="middle" align="center">Mean daily mean air temperatures of the warmest quarter</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio11</td>
<td valign="middle" align="center">Mean daily mean air temperatures of the coldest quarter</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio12</td>
<td valign="middle" align="center">Annual precipitation amount</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio13</td>
<td valign="middle" align="center">Precipitation amounts of the wettest month</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio14</td>
<td valign="middle" align="center">Precipitation amounts of the driest month</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio15</td>
<td valign="middle" align="center">Precipitation seasonality</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio16</td>
<td valign="middle" align="center">Mean monthly precipitation amount of the wettest quarter</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio17</td>
<td valign="middle" align="center">Mean monthly precipitation amount of the driest quarter</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio18</td>
<td valign="middle" align="center">Mean monthly precipitation amount of the warmest quarter</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">Bio19</td>
<td valign="middle" align="center">Mean monthly precipitation amount of the coldest quarter</td>
</tr>
<tr>
<td valign="middle" align="center">Climatic variables</td>
<td valign="middle" align="center">GSP</td>
<td valign="middle" align="center">Accumulated precipitation amounts on growing season days TREELIM</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">NGD0</td>
<td valign="middle" align="center">Number of growing degree days</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">GDD0</td>
<td valign="middle" align="center">Growing degree days heat sum above 0&#xb0;C</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">CMI_mean</td>
<td valign="middle" align="center">Mean monthly climate moisture index</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">CMI_range</td>
<td valign="middle" align="center">Annual range of monthly climate moisture index</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">HURS_mean</td>
<td valign="middle" align="center">Mean monthly near-surface relative humidity</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">HURS_range</td>
<td valign="middle" align="center">Annual range of monthly near-surface relative humidity</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">PET_mean</td>
<td valign="middle" align="center">Mean monthly potential evapotranspiration</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">PET_range</td>
<td valign="middle" align="center">Annual range of monthly potential evapotranspiration</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">NPP</td>
<td valign="middle" align="center">Net primary productivity</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">SCD</td>
<td valign="middle" align="center">Snow cover days</td>
</tr>
<tr>
<td valign="middle" align="center">Soil variables</td>
<td valign="middle" align="center">PH</td>
<td valign="middle" align="center">pH Value (H<sub>2</sub>O)</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">SOM</td>
<td valign="middle" align="center">Soil Organic Matter</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">TN</td>
<td valign="middle" align="center">Total N</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">TP</td>
<td valign="middle" align="center">Total P</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">TK</td>
<td valign="middle" align="center">Total K</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">AN</td>
<td valign="middle" align="center">Alkali-hydrolysable N</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">AP</td>
<td valign="middle" align="center">Available P</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">AK</td>
<td valign="middle" align="center">Available K</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">CEC</td>
<td valign="middle" align="center">Cation Exchange Capacity (CEC)</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">EH</td>
<td valign="middle" align="center">Exchangeable H+</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">EAl</td>
<td valign="middle" align="center">Exchangeable Al3+</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">ECa</td>
<td valign="middle" align="center">Exchangeable Ca2+</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">EMg</td>
<td valign="middle" align="center">Exchangeable Mg2+</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">EK</td>
<td valign="middle" align="center">Exchangeable K+</td>
</tr>
<tr>
<td valign="middle" align="center"/>
<td valign="middle" align="center">ENa</td>
<td valign="middle" align="center">Exchangeable Na+</td>
</tr>
<tr>
<td valign="middle" align="center">Topographic variables</td>
<td valign="middle" align="center">Elev</td>
<td valign="middle" align="center">Elevation</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Analysis of potential spatial distribution pattern of species</title>
<p>To ensure the consistency of environmental data and the effectiveness of model predictions, this study constructed three types of environmental datasets: the full environmental dataset under the current period (CP-full dataset), including bioclimatic, climatic, soil, and topographic data; the simplified environmental dataset under the current or future period (CP or FP-simp dataset), including bioclimatic, soil, and topographic data. The CP-full dataset is used to finely predict the potential spatial distribution pattern of <italic>E. sibiricus</italic> in the current period; the CP or FP-simp dataset is used to analyze the temporal pattern dynamic changes of the potential distribution of <italic>E. sibiricus</italic> from the current to future periods. The potential spatial distribution pattern of <italic>E. sibiricus</italic> was modeled and predicted by R package ssdm v0.2.9 (<xref ref-type="bibr" rid="B45">Schmitt et&#xa0;al., 2017</xref>), which integrated different environmental datasets and species distribution points. To obtain more robust prediction results, this study selected nine modeling algorithms, including MAXENT, GLM, generalized additive model (GAM), multiple adaptive regression splines (MARS), generalized boosting model (GBM), classification tree analysis (CTA), random forest (RF), and ANN and support vector machine (SVM).</p>
<p>During the modeling process, the pseudo-occurrence points of the species of <italic>E. sibiricus</italic> were automatically generated, and 75% distribution data were randomly divided into training datasets for model construction, and the remaining were used as testing datasets to evaluate model performance. The independence between training and testing datasets was verified by 10 repeated crossover methods based on holdout strategy. The area under the curve of receiver operating characteristics (AUC) threshold is 0.75 as the standard for ensemble modeling algorithms, and Pearson correlation tests were performed among algorithms. To coordinate the inherent advantages and disadvantages of each algorithm, an integrated scoring system of true skills statistics (TSS), Kappa, and AUC was used to evaluate the predictive performance of the ensemble model (<xref ref-type="bibr" rid="B6">Chau et&#xa0;al., 2023</xref>). The values of TSS, Kappa, and AUC ranged from 0 to 1, 0.85 &#x2264; TSS | Kappa &#x2264; 1.0 or 0.9 &#x2264; AUC &#x2264; 1.0 considered that the model was accurate; 0.70 &#x2264; TSS | Kappa &lt; 0.85 or 0.8 &#x2264; AUC &lt; 0.9, the model predicts well; 0.55 &#x2264; TSS | Kappa &lt; 0.70 or 0.6 &#x2264; AUC &lt; 0.7, the model predicts general; 0.40 &#x2264; TSS | Kappa &lt; 0.55 or 0.5 &#x2264; AUC &lt; 0.6, the model predicts poorly; if TSS | Kappa &lt; 0.40 or AUC &lt; 0.50, the model predicts abnormal. The relative importance of environmental variables was estimated by R package ssdm. Finally, the R package ssdm generates continuity distribution probability with a threshold accuracy of 100.</p>
<p>Based on the natural break classification method, the distribution probability was divided into three levels of suitability (high, low, and unsuitable regions) by using the Reclassify tool in ArcGIS software, and the area of different suitable levels in each administrative region was counted by using the Region analysis tool in ArcGIS software. To obtain the most suitable survival environment conditions for <italic>E. sibiricus</italic> in the current period, the species survival curve was calculated by MaxEnt v3.4.4 (<xref ref-type="bibr" rid="B37">Phillips et&#xa0;al., 2006</xref>). In the parameter settings of MaxEnt software, enable the options of Create response curves, Do jackknife, Random seed, and Write plot data; set Random test percentage to 25%, Regularization coefficient to 1.1, and Max number of background points to 10,000. The operation model was repeated 10 times based on the Bootstrap strategy and defaulted to other parameters.</p>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Analysis of potential temporal distribution patterns of species</title>
<p>The change and migration of species to suitable regions were a response to environmental changes over a long period of time. To explore the temporal distribution pattern of <italic>E. sibiricus</italic> from the current (1979&#x2013;2013) to future (2071&#x2013;2100) period, Wilcoxon test was conducted on the area of suitable levels at different periods using SPSS v25.0 (<ext-link ext-link-type="uri" xlink:href="https://www.ibm.com/spss">https://www.ibm.com/spss</ext-link>). The geometric centroid of suitable regions is used as a general description of the distribution characteristics of species in each period, and the migration of the geometric centroid can more intuitively display the changes of suitable regions of species in different periods. To clarify the dynamic migration path of the suitable region of <italic>E. sibiricus</italic>, the distribution probability was used, the geometric centroid positions of <italic>E. sibiricus</italic> distribution in different periods were located by the region analysis tool in ArcGIS software and R package foreign v0.8 (<ext-link ext-link-type="uri" xlink:href="https://cran.r-project.org/package=foreign">https://cran.r-project.org/package=foreign</ext-link>), and the geographical distance between different centroids was calculated by R package geosphere v1.5 (<ext-link ext-link-type="uri" xlink:href="https://cran.r-project.org/package=geosphere">https://cran.r-project.org/package=geosphere</ext-link>).</p>
</sec>
<sec id="s2_5">
<label>2.5</label>
<title>Analysis of potential planting patterns of species</title>
<p>The potential planting pattern of <italic>E. sibiricus</italic> in the current period was predicted by Marxan v4.0.5 (<xref ref-type="bibr" rid="B11">Game et&#xa0;al., 2013</xref>), which involves the setting of planning unit, target, cost, and parameter optimization. Considering the continuity of the planting region and the simulation ability of Marxan software, a square planning unit (grid) with a side length of about 5 km covering the land of China was created by using the Fishnet tool in ArcGIS software. According to the protection standard of the Guidelines for Management Planning of Protected Areas issued by the International Union for Conservation of Nature (IUCN, <ext-link ext-link-type="uri" xlink:href="https://www.iucn.org/">https://www.iucn.org/</ext-link>), 10% planting coverage rate was selected as the planting target, which does not affect the ecological environment and maintains the basic ecological functions of species. Due to Chinese policy of &#x201c;arable land minimum&#x201d; (<xref ref-type="bibr" rid="B7">Chen et&#xa0;al., 2019</xref>), this study only selects the probability of refined distribution in the current period and the consensus rate of herbaceous vegetation coverage as equally weighted planting cost. The consensus rate of herbaceous vegetation coverage was selected from Herbaceous Vegetation Reduced v1.0 dataset (<ext-link ext-link-type="uri" xlink:href="https://www.earthenv.org/landcover">https://www.earthenv.org/landcover</ext-link>). The parameters such as boundary length (BLM), species penalty factor (SPF), and iteration times were optimized by ArcGIS plug-in ArcMarxan Toolbox v2.0.2 (<ext-link ext-link-type="uri" xlink:href="https://aproposinfosystems.com/">https://aproposinfosystems.com/</ext-link>). Based on the natural break classification method, the potential planting regions were divided into three levels of planting (high, low, and unselected regions) by using the Reclassify tool in ArcGIS software, and the area of different planting levels in each administrative region was counted by using the region analysis tool in ArcGIS software.</p>
</sec>
<sec id="s2_6">
<label>2.6</label>
<title>Prediction of introduction adaptability of different germplasms of <italic>E. sibiricus</italic>
</title>
<p>In this study, two cultivars [<italic>E. sibiricus</italic> cv. &#x201c;Tongde&#x201d; (S1) and &#x201c;Chuancao No. 2&#x201d; (S2)] and six germplasms [LM01 (S3), LM02 (S4), LM03 (S5), LM04 (S6), LM05 (S7), and LM06 (S8)] of <italic>E. sibiricus</italic> were selected to carry out field experiments in the forage breeding base of the Grassland Science Research Institute, Hongyuan County, Sichuan Province (32&#xb0;47&#x2019;N, 102&#xb0;32&#x2019;E, 3460 m) for 3 years. The experiment adopted a randomized block design with three replicates, with a sample plot area of 15 m<sup>2</sup> (3 m &#xd7; 5 m) and a spacing of 0.5 m between plots. Each sample plot was evenly seeded with 33 g of seeds and the row spacing of 0.3 m. To effectively evaluate the growth performance of <italic>E. sibiricus</italic>, 23 agronomic traits (<xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>) for 8 tested materials were measured (<italic>n</italic> = 10) from 2021 to 2023 according to &#x201c;Description Standard and Data Standard of <italic>Elymus sibiricus</italic> Germplasm Resources&#x201d; (<xref ref-type="bibr" rid="B23">Li et&#xa0;al., 2023</xref>). The values of all agronomic traits were normalized by R package vegan v2.6 (<ext-link ext-link-type="uri" xlink:href="https://CRAN.R-project.org/package=vegan">https://CRAN.R-project.org/package=vegan</ext-link>) to unify dimensions. Principal components analysis (PCA) and phenotypic contribution analysis of all tested materials were carried out using R package FactoMineR v2.11 (<ext-link ext-link-type="uri" xlink:href="https://CRAN.R-project.org/package=FactoMineR">https://CRAN.R-project.org/package=FactoMineR</ext-link>). The membership function analysis among tested materials was analyzed by SPSSPRO software (<ext-link ext-link-type="uri" xlink:href="https://www.spsspro.com/">https://www.spsspro.com/</ext-link>) to evaluate the comprehensive properties of different materials. Based on the potential planting pattern of <italic>E. sibiricus</italic>, the top 25 municipal administrative regions were selected according to the decrease of potential planting area with high selected regions. The suitability probability was extracted by using the extraction analysis tool in ArcGIS software under constraints of different municipal administrative regions. The distribution probability was used as the geographical environment standard to measure the suitable distribution of <italic>E. sibiricus</italic>, and the agronomic traits were weighted by the phenotypic contribution degree to measure the adaptability level of different tested materials. The optimal dissimilarity index was screened by R package vegan, and a Mantel test between the top 25 municipal administrative regions and weighted agronomic trait values of six germplasms was carried out by ChiPlot online software (<ext-link ext-link-type="uri" xlink:href="https://www.chiplot.online/">https://www.chiplot.online/</ext-link>) to predict the introduction adaptability of different germplasms.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Measured agronomic traits indicators of <italic>E. sibiricus</italic>.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Character types</th>
<th valign="middle" align="center">Agronomic traits</th>
<th valign="middle" align="center">Description</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" rowspan="2" align="center">Plant height traits</td>
<td valign="middle" align="center">NPH</td>
<td valign="middle" align="center">Natural plant height</td>
</tr>
<tr>
<td valign="middle" align="center">APN</td>
<td valign="middle" align="center">Absolute plant height</td>
</tr>
<tr>
<td valign="top" rowspan="4" align="center">Leaf traits</td>
<td valign="middle" align="center">FLL</td>
<td valign="middle" align="center">Flag leaf length</td>
</tr>
<tr>
<td valign="middle" align="center">FLW</td>
<td valign="middle" align="center">Flag leaf width</td>
</tr>
<tr>
<td valign="middle" align="center">TSLL</td>
<td valign="middle" align="center">Top second leaf length</td>
</tr>
<tr>
<td valign="middle" align="center">TSLW</td>
<td valign="middle" align="center">Top second leaf width</td>
</tr>
<tr>
<td valign="top" rowspan="5" align="center">Stem traits</td>
<td valign="middle" align="center">SD</td>
<td valign="middle" align="center">Stem diameter</td>
</tr>
<tr>
<td valign="middle" align="center">SNN</td>
<td valign="middle" align="center">Stem node number</td>
</tr>
<tr>
<td valign="middle" align="center">RB</td>
<td valign="middle" align="center">Reproductive branch</td>
</tr>
<tr>
<td valign="middle" align="center">NB</td>
<td valign="middle" align="center">Nutrient branches</td>
</tr>
<tr>
<td valign="middle" align="center">TN</td>
<td valign="middle" align="center">Tiller number</td>
</tr>
<tr>
<td valign="top" rowspan="3" align="center">Panicle traits</td>
<td valign="middle" align="center">EL</td>
<td valign="middle" align="center">Ear length</td>
</tr>
<tr>
<td valign="middle" align="center">SN</td>
<td valign="middle" align="center">Spikelet number</td>
</tr>
<tr>
<td valign="middle" align="center">FN</td>
<td valign="middle" align="center">Flower number</td>
</tr>
<tr>
<td valign="top" rowspan="3" align="center">Seed traits</td>
<td valign="middle" align="center">SL</td>
<td valign="middle" align="center">Seed length</td>
</tr>
<tr>
<td valign="middle" align="center">SW</td>
<td valign="middle" align="center">Seed width</td>
</tr>
<tr>
<td valign="middle" align="center">TGW</td>
<td valign="middle" align="center">Thousand grain weight</td>
</tr>
<tr>
<td valign="top" rowspan="6" align="center">Production traits</td>
<td valign="middle" align="center">FTLR</td>
<td valign="middle" align="center">Fresh stem to leaf ratio</td>
</tr>
<tr>
<td valign="middle" align="center">DSLR</td>
<td valign="middle" align="center">Dry stem to leaf ratio</td>
</tr>
<tr>
<td valign="middle" align="center">SY</td>
<td valign="middle" align="center">Seed yield</td>
</tr>
<tr>
<td valign="middle" align="center">HY</td>
<td valign="middle" align="center">Hay yield</td>
</tr>
<tr>
<td valign="middle" align="center">FGY</td>
<td valign="middle" align="center">Fresh grass yield</td>
</tr>
<tr>
<td valign="middle" align="center">DFR</td>
<td valign="middle" align="center">Dry to fresh ratio</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Data preprocessing and prediction performance evaluation of ENMs</title>
<p>A total of 538 distribution data of <italic>E. sibiricus</italic> were compiled in this study. After data quality control and rarefaction, there were 421 effective distribution points (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1A</bold>
</xref>), which were widely distributed in Qilian Mountains (northeast of QTP), Taihang Mountains (junction of Loess Plateau and Inner Mongolia Plateau), and Tianshan Mountains. The correlation of environmental variables showed that the correlation ranges of 46 environmental factors in the CP-full dataset were &#x2212;0.98 to 0.99 (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1B</bold>
</xref>), and the correlation ranges of 25 environmental factors in the CP and FP-simp dataset were &#x2212;0.98 to 0.99 and &#x2212;0. 99 to 0.99 (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure S1A</bold>
</xref>), respectively. Since 17 environmental factors in bioclimatic variables are derived from mean annual air temperature (bio01) and annual precipitation amount (bio12), bioclimatic variables are screened based on bio01 and bio12. After correlation analysis of all environmental factors, 24 environmental factors with low correlation and obvious ecological significance were screened out from the CP-full dataset, including Elev, AK, EAl, AP, ECa, EH, EMg, ENa, PH, TK, TN, TP, Bio01, Bio12, Bio15, Bio02, Bio03, Bio07, CMI_range, GSP, HURS_mean, HURS_range, PET_mean, and PET_range. The same eight environmental factors (Elev, Bio01, Bio12, Bio15, Bio02, Bio03, Bio07, and GSP) were screened from the CP or FP-simp dataset. The correlation analysis of nine modeling algorithms showed that the correlation range based on the CP-full dataset was 0.33 to 0.88 (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1C</bold>
</xref>); the correlation ranges based on the CP and FP-simp datasets were 0.54 to 0.93 (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure S1B</bold>
</xref>) and 0.60 to 0.93 (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure S1C</bold>
</xref>), respectively, indicating that there are significant deviations and uncertainties among different algorithms. The evaluation results of the integrated scoring system for the model prediction performance showed that RF algorithm had the best modeling effect on the spatial distribution of <italic>E. sibiricus</italic> under different environmental datasets, while MAXENT algorithm was the worst. It is worth noting that GLM algorithm cannot effectively model on CP or FP-simp dataset. Overall, the AUC, Kappa, and TSS based on the CP-full dataset were 0.82, 0.70, and 0.75, respectively (<xref ref-type="fig" rid="f1"><bold>Figure 1D</bold></xref>); the AUC, Kappa, and TSS based on the CP and FP-simp dataset were 0.89, 0.67, 0.72 and 0.89, 0.68, 0.72, respectively. The evaluation results of the integrated scoring system indicated that the ensemble model constructed by different algorithms can effectively simulate the potential spatial distribution pattern of <italic>E. sibiricus</italic>.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Refined modeling data processing and ecological niche model evaluation. <bold>(A)</bold> Geographical distribution records and study regions of <italic>E. sibiricus</italic>; <bold>(B)</bold> correlation among environmental variables in the current period; <bold>(C)</bold> the correlation among different algorithms in the current period; and <bold>(D)</bold> evaluation of integrated scoring system of different algorithms in current period. The meanings of abbreviations for environmental factors were shown in <xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>. MAXENT, maximum entropy; GLM, generalized linear model; GAM, generalized additive model; MARS, multiple adaptive regression splines; GBM, generalized boosting model; CTA, classification tree analysis; RF, random forest; ANN, artificial neural network; SVM, support vector machine.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1470653-g001.tif"/>
</fig>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Importance evaluation and optimal survival value for environmental variables</title>
<p>Relative contribution can be used to evaluate the influence of environmental variables upon the spatial distribution of <italic>E. sibiricus</italic>. 24 environmental factors from the CP-full dataset were used to refine and predict the spatial distribution pattern of <italic>E. sibiricus</italic> in the current period. The relative contribution analysis showed that the relative contribution ranges of 24 environmental factors ranged from 1.57% to 11.36% (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2A</bold>
</xref>), among which the total relative contribution of bioclimatic, climatic, soil, and topographic variables was 23.62%, 39.23%, 31.93%, and 5.21%, respectively. Specifically, the cumulative relative contribution of 11 environmental factors reached 69.03%, including the annual range of monthly near-surface relative humidity (HURS_range, 11.36%), mean annual air temperature (Bio01, 9.67%), annual range of monthly potential evapotranspiration (PET_range, 8.52%), mean monthly potential evapotranspiration (PET_mean, 7.53%), annual range of monthly climate moisture index (CMI_range, 6.48%), elevation (Elev, 5.21%), exchangeable Ca<sup>2+</sup> (ECa, 4.94%), available P (AP, 4.27%), mean monthly near-surface relative humidity (HURS_mean, 3.78%), exchangeable H<sup>+</sup> (EH, 3.71%), and annual precipitation amount (Bio12, 3.57%). Overall, the spatial distribution pattern of <italic>E. sibiricus</italic> is obviously responsive to hydrothermal environmental conditions, especially humidity, air temperature, and potential evapotranspiration, and the elevation and soil calcium and phosphorus nutrients also play an important role in its distribution. In addition, based on the survival response curve, the optimum survival environment value under the optimum survival probability of <italic>E. sibiricus</italic> was extracted (<xref ref-type="table" rid="T3">
<bold>Table&#xa0;3</bold>
</xref>). The ranges of optimum survival probability and its coefficient of variation under different environmental factors were 0.08&#x2013;0.93 and 1.85%&#x2013;125.47%, respectively, among which the optimum survival probability and coefficient of variation with GSP were extremely abnormal, indicating that GSP had no significant effect on the potential spatial distribution of <italic>E. sibiricus</italic>. According to the comprehensive analysis of optimum survival environment values for different types, it is concluded that the most suitable habitat of <italic>E. sibiricus</italic> is a semi-humid area with weak acidic soil rich in nitrogen, phosphorus, and potassium, small fluctuation of air temperature, low temperature, and high altitude.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Relative contribution of environmental variables <bold>(A)</bold> and potential spatial distribution pattern <bold>(B)</bold> of <italic>E. sibiricus</italic>. The meanings of abbreviations for environmental factors were shown in <xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1470653-g002.tif"/>
</fig>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>The optimal survival environment values under optimal survival probability of <italic>E. sibiricus</italic>.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Environmental factors</th>
<th valign="middle" align="center">Mean survival probability (<italic>n</italic> = 10)</th>
<th valign="middle" align="center">Coefficient of variation</th>
<th valign="middle" align="center">Optimal survival value</th>
<th valign="middle" align="center">Unit of optimal survival value</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">AK</td>
<td valign="middle" align="center">0.46 &#xb1; 0.08</td>
<td valign="middle" align="center">16.79%</td>
<td valign="middle" align="center">357.31</td>
<td valign="middle" align="center">mg/kg</td>
</tr>
<tr>
<td valign="middle" align="center">AP</td>
<td valign="middle" align="center">0.82 &#xb1; 0.03</td>
<td valign="middle" align="center">3.07%</td>
<td valign="middle" align="center">7.70</td>
<td valign="middle" align="center">mg/kg</td>
</tr>
<tr>
<td valign="middle" align="center">Bio01</td>
<td valign="middle" align="center">0.82 &#xb1; 0.03</td>
<td valign="middle" align="center">3.58%</td>
<td valign="middle" align="center">2.00</td>
<td valign="middle" align="center">&#xb0;C</td>
</tr>
<tr>
<td valign="middle" align="center">Bio02</td>
<td valign="middle" align="center">0.69 &#xb1; 0.04</td>
<td valign="middle" align="center">6.34%</td>
<td valign="middle" align="center">13.65</td>
<td valign="middle" align="center">&#xb0;C</td>
</tr>
<tr>
<td valign="middle" align="center">Bio03</td>
<td valign="middle" align="center">0.74 &#xb1; 0.03</td>
<td valign="middle" align="center">4.35%</td>
<td valign="middle" align="center">0.36</td>
<td valign="middle" align="center">&#xb0;C</td>
</tr>
<tr>
<td valign="middle" align="center">Bio07</td>
<td valign="middle" align="center">0.75 &#xb1; 0.23</td>
<td valign="middle" align="center">30.79%</td>
<td valign="middle" align="center">59.08</td>
<td valign="middle" align="center">&#xb0;C</td>
</tr>
<tr>
<td valign="middle" align="center">Bio12</td>
<td valign="middle" align="center">0.75 &#xb1; 0.02</td>
<td valign="middle" align="center">2.03%</td>
<td valign="middle" align="center">487.66</td>
<td valign="middle" align="center">kg&#xb7;m<sup>&#x2212;2</sup>&#xb7;year<sup>&#x2212;1</sup>
</td>
</tr>
<tr>
<td valign="middle" align="center">Bio15</td>
<td valign="middle" align="center">0.79 &#xb1; 0.29</td>
<td valign="middle" align="center">37.15%</td>
<td valign="middle" align="center">26.68</td>
<td valign="middle" align="center">kg&#xb7;m<sup>&#x2212;2</sup>
</td>
</tr>
<tr>
<td valign="middle" align="center">CMI_range</td>
<td valign="middle" align="center">0.86 &#xb1; 0.03</td>
<td valign="middle" align="center">2.92%</td>
<td valign="middle" align="center">64.94</td>
<td valign="middle" align="center">kg&#xb7;m<sup>&#x2212;2</sup>&#xb7;month<sup>&#x2212;1</sup>
</td>
</tr>
<tr>
<td valign="middle" align="center">EAl</td>
<td valign="middle" align="center">0.76 &#xb1; 0.02</td>
<td valign="middle" align="center">2.73%</td>
<td valign="middle" align="center">0.10</td>
<td valign="middle" align="center">me/100 g</td>
</tr>
<tr>
<td valign="middle" align="center">ECa</td>
<td valign="middle" align="center">0.81 &#xb1; 0.05</td>
<td valign="middle" align="center">6.00%</td>
<td valign="middle" align="center">14.53</td>
<td valign="middle" align="center">me/100 g</td>
</tr>
<tr>
<td valign="middle" align="center">EH</td>
<td valign="middle" align="center">0.73 &#xb1; 0.03</td>
<td valign="middle" align="center">4.52%</td>
<td valign="middle" align="center">0.14</td>
<td valign="middle" align="center">me/100 g</td>
</tr>
<tr>
<td valign="middle" align="center">Elev</td>
<td valign="middle" align="center">0.93 &#xb1; 0.02</td>
<td valign="middle" align="center">2.48%</td>
<td valign="middle" align="center">3355.44</td>
<td valign="middle" align="center">M</td>
</tr>
<tr>
<td valign="middle" align="center">EMg</td>
<td valign="middle" align="center">0.87 &#xb1; 0.02</td>
<td valign="middle" align="center">2.45%</td>
<td valign="middle" align="center">2.79</td>
<td valign="middle" align="center">me/100 g</td>
</tr>
<tr>
<td valign="middle" align="center">Ena</td>
<td valign="middle" align="center">0.73 &#xb1; 0.02</td>
<td valign="middle" align="center">3.05%</td>
<td valign="middle" align="center">0.56</td>
<td valign="middle" align="center">me/100 g</td>
</tr>
<tr>
<td valign="middle" align="center">GSP</td>
<td valign="middle" align="center">0.08 &#xb1; 0.10</td>
<td valign="middle" align="center">125.47%</td>
<td valign="middle" align="center">343597.36</td>
<td valign="middle" align="center">kg&#xb7;m<sup>&#x2212;2</sup>&#xb7;gsl<sup>&#x2212;1</sup>
</td>
</tr>
<tr>
<td valign="middle" align="center">HURS_mean</td>
<td valign="middle" align="center">0.72 &#xb1; 0.01</td>
<td valign="middle" align="center">1.85%</td>
<td valign="middle" align="center">57.90</td>
<td valign="middle" align="center">%</td>
</tr>
<tr>
<td valign="middle" align="center">HURS_range</td>
<td valign="middle" align="center">0.81 &#xb1; 0.03</td>
<td valign="middle" align="center">3.62%</td>
<td valign="middle" align="center">5.87</td>
<td valign="middle" align="center">%</td>
</tr>
<tr>
<td valign="middle" align="center">PET_mean</td>
<td valign="middle" align="center">0.78 &#xb1; 0.03</td>
<td valign="middle" align="center">3.28%</td>
<td valign="middle" align="center">76.66</td>
<td valign="middle" align="center">kg&#xb7;m<sup>&#x2212;2</sup>&#xb7;month<sup>&#x2212;1</sup>
</td>
</tr>
<tr>
<td valign="middle" align="center">PET_range</td>
<td valign="middle" align="center">0.75 &#xb1; 0.01</td>
<td valign="middle" align="center">1.98%</td>
<td valign="middle" align="center">109.66</td>
<td valign="middle" align="center">kg&#xb7;m<sup>&#x2212;2</sup>
</td>
</tr>
<tr>
<td valign="middle" align="center">PH</td>
<td valign="middle" align="center">0.75 &#xb1; 0.02</td>
<td valign="middle" align="center">2.67%</td>
<td valign="middle" align="center">8.09</td>
<td valign="middle" align="center">PH units</td>
</tr>
<tr>
<td valign="middle" align="center">TK</td>
<td valign="middle" align="center">0.73 &#xb1; 0.02</td>
<td valign="middle" align="center">2.84%</td>
<td valign="middle" align="center">2.11</td>
<td valign="middle" align="center">g/100g</td>
</tr>
<tr>
<td valign="middle" align="center">TN</td>
<td valign="middle" align="center">0.69 &#xb1; 0.03</td>
<td valign="middle" align="center">4.08%</td>
<td valign="middle" align="center">0.44</td>
<td valign="middle" align="center">g/100g</td>
</tr>
<tr>
<td valign="middle" align="center">TP</td>
<td valign="middle" align="center">0.71 &#xb1; 0.02</td>
<td valign="middle" align="center">3.53%</td>
<td valign="middle" align="center">0.09</td>
<td valign="middle" align="center">g/100g</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>The meanings of abbreviations for environmental factors were shown in <xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Potential spatial distribution pattern of <italic>E. sibiricus</italic>
</title>
<p>Based on the natural break classification method, the potential spatial distribution of <italic>E. sibiricus</italic> was divided into three levels of suitability (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S1</bold>
</xref>). The results showed that the potential spatial distribution mainly spans 33&#xb0;&#x2013;42&#xb0;N and 95&#xb0;&#x2013;118&#xb0;E (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2B</bold>
</xref>), which was concentrated in Qilian Mountains (northeastern of QTP), Taihang Mountains (the junction of Loess Plateau and Inner Mongolia Plateau), and Tianshan Mountains, while the rest was scattered in Altai Mountains, Kunlun Mountains (northern of QTP), Hengduan Mountains (central Yunnan-Guizhou Plateau), Qinling Mountains (south of Loess Plateau), and Changbai Mountains (southeast of Northeast China). This study predicted an area of 9.3784 million km<sup>2</sup> in China, with high and low suitable regions accounting for 14.67% (1.3758 million km<sup>2</sup>) and 29.44% (2.7613 million km<sup>2</sup>), respectively. High suitable regions were mainly concentrated in western China, while low suitable regions were mainly distributed around high suitable regions. The high and low suitable regions involve 25 and 33 provincial administrative regions, respectively (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S2</bold>
</xref>), among which the top five provincial administrative regions with the suitable area decreases in high suitable regions are Xinjiang (296,500 km<sup>2</sup>), Gansu (193,300 km<sup>2</sup>), Qinghai (125,700 km<sup>2</sup>), Sichuan (112,600 km<sup>2</sup>), and Heilongjiang (103,900 km<sup>2</sup>); the top 5 provincial administrative regions with the largest area of low suitable regions are Inner Mongolia (399,100 km<sup>2</sup>), Xinjiang (372,600 km<sup>2</sup>), Heilongjiang (270,300 km<sup>2</sup>), Qinghai (197,800 km<sup>2</sup>), and Sichuan (185,300 km<sup>2</sup>). Overall, the suitable area for <italic>E. sibiricus</italic> distribution accounts for about 44.11% of the total area in China, and it is concentrated in plateau regions.</p>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Dynamic changes of the potential temporal distribution pattern of <italic>E. sibiricus</italic>
</title>
<p>The dynamic changes of the potential spatial distribution pattern of <italic>E. sibiricus</italic> across multiple periods were analyzed. The results showed that the levels of high, low, and unsuitable regions remained relatively stable across different periods (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3A</bold>
</xref>), but the area changes within each suitable level were significant during the same period (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3B</bold>
</xref>), indicating that the potential suitable pattern of <italic>E. sibiricus</italic> had migrated with the passage of time. Comparing the current period with the future period, the overall suitable area is estimated to grow between &#x2212;2.72% and 4.65 million km<sup>2</sup>. Specifically, the area of high and low suitable regions is expected to increase by &#x2212;10.66% (up to approximately 1.7684 million km<sup>2</sup>) and 2.89% (up to around 2.8817 million km<sup>2</sup>), respectively (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3C</bold>
</xref>). A deeper analysis of the dynamic changes within the different suitable levels revealed that over time, the percentage range of area growth for high and low suitable regions varied from &#x2212;100.00% to 59.39% and from &#x2212;86.56% to 1185.90%, respectively (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3C</bold>
</xref>, <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S3</bold>
</xref>). The regions predicted to experience relatively large expansions in their suitable areas are Guizhou, Yunnan, and Henan (high suitability), as well as Taiwan, Guangxi, Beijing, and Fujian (low suitability). Meanwhile, the areas expected to have relatively large contractions in their suitable regions are Shandong, Tianjin, and Chongqing (high suitability) and Zhejiang, Jiangsu, and Anhui (low suitability). Overall, compared to the present, the forecasted suitable area for <italic>E. sibiricus</italic> demonstrates a general decline; however, the key distribution centers remain relatively stable, such as Qilian Mountains (northeast of QTP), Taihang Mountains (junction of Loess Plateau and Inner Mongolia Plateau), and Tianshan Mountains. Furthermore, the centroid of the suitable habitat is estimated to shift southwestward by approximately 2.93 km (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3C</bold>
</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Dynamic changes in the potential temporal distribution pattern of <italic>E. sibiricus</italic>. <bold>(A)</bold> The area comparison of the same suitable level in different periods; <bold>(B)</bold> the area comparison of the same period in different suitable level; and <bold>(C)</bold> dynamic changes of potential distribution pattern of <italic>E. sibiricus</italic> in different periods. *<italic>p</italic> &lt; 0.05, **<italic>p</italic> &lt; 0.01.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1470653-g003.tif"/>
</fig>
</sec>
<sec id="s3_5">
<label>3.5</label>
<title>Potential planting distribution pattern of <italic>E. sibiricus</italic>
</title>
<p>Based on the natural break classification method, the planting priority of <italic>E. sibiricus</italic> was divided into three levels of planting (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S4</bold>
</xref>). This study only predicted the natural grassland area of 0.349 million km<sup>2</sup> in China. The potential planting regions of <italic>E. sibiricus</italic> were mainly distributed in the Qilian Mountains (northeast of QTP), Taihang Mountains (southeast of Inner Mongolia Plateau), Tianshan Mountains, and Altai Mountains (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). The high and low selection planting areas were 66,600 km<sup>2</sup> and 139,300 km<sup>2</sup>, respectively, among which the high and low selection planting regions involve 21 and 20 provincial administrative regions, respectively (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S5</bold>
</xref>). The top 6 provincial administrative regions with the planting area decreases in high selection regions are Xinjiang (17,752.50 km<sup>2</sup>), Gansu (12,577.50 km<sup>2</sup>), Qinghai (7,582.50 km<sup>2</sup>), Tibet (7,335.00 km<sup>2</sup>), Inner Mongolia (7,155.00 km<sup>2</sup>), and Sichuan (4,275.00 km<sup>2</sup>). The top six provincial administrative regions with the largest area of low selection regions are Inner Mongolia (36,090.00 km<sup>2</sup>), Tibet (28,777.50 km<sup>2</sup>), Qinghai (20,970.00 km<sup>2</sup>), Xinjiang (20,205.00 km<sup>2</sup>), Sichuan (10,755.00 km<sup>2</sup>), and Gansu (6,007.50 km<sup>2</sup>). Overall, the suitable area for planting <italic>E. sibiricus</italic> accounts for 58.99% of the predicted natural grassland and 2.14% of the national area in China.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Distribution pattern of potential planting of <italic>E. sibiricus</italic>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1470653-g004.tif"/>
</fig>
</sec>
<sec id="s3_6">
<label>3.6</label>
<title>Introduction to the adaptability of different germplasm of <italic>E. sibiricus</italic>
</title>
<p>The PCA and contribution analysis of 23 agronomic traits showed that all agronomic traits explained 58.28% of the overall traits (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5A</bold>
</xref>), among which 12 agronomic traits, such as DFR, TGW, FN, SN, NPH, APN, DSLR, FTLR, HY, FGY, FLL, and FLW, contributed significantly to the comprehensive growth evaluation of <italic>E. sibiricus</italic>. The membership function analysis showed that, in comprehensive evaluation, the varieties S3 (0.47), S5 (0.47), S6 (0.53), S7 (0.60), and S8 (0.49) were superior to <italic>E. sibiricus</italic> varieties S1 (0.11) and S2 (0.45), while S4 (0.23) was superior to S1 but inferior to S2 (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5B</bold>
</xref>). The Mantel Test analysis of the introduction adaptability showed that the Pearson correlation range among all sample plots was &#x2212;0.58 to 1.00 (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5C</bold>
</xref>); for example, the correlation among SP07, SP08, and SP09 was greater than 0.65, indicating that the plots had similar environmental conditions for adaptability distribution. Varieties S3, S5, S6, S7, and S8 had a significant positive correlation with SP03; S6, S7 with SP04; S3, S4, S6, S7, and S8 with SP11; and S5 with SP23 (<italic>p</italic> &lt; 0.05). It was indicated that S3, S5, S6, S7, and S8 were suitable for planting in the southeast of the Altai Mountains represented by the Altay region (SP03); S6 and S7 were suitable for planting in the western section of the Hengduan Mountains represented by the Changdu City (SP04); S3, S4, S6, S7, and S8 were suitable for planting in the middle part of the Qilian Mountains represented by the Hainan Tibetan Autonomous Prefecture (SP11); and S5 was suitable for planting in the northern foot of the eastern part of the Qilian Mountains represented by the Wuwei City (SP23). Overall, the introduction adaptation regions of the six germplasms are all distributed in the high-altitude mountain region in western China, which indirectly verifies the rationality of the potential distribution and planting distribution pattern of <italic>E. sibiricus</italic>.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Comprehensive evaluation of different materials and introduction adaptability of <italic>E. sibiricus</italic>. <bold>(A)</bold> PCA and contribution analysis of agronomic traits; <bold>(B)</bold> Membership function evaluation of different tested materials; <bold>(C)</bold> Mantel Test analysis between tested materials and sample plots. The meanings of abbreviations for agronomic traits were shown in <xref ref-type="table" rid="T2">
<bold>Table&#xa0;2</bold>
</xref>; SP01: Ili Kazakh Autonomous Prefecture; SP02: Tacheng region; SP03: Altay region; SP04: Changdu City; SP05: Zhangjiakou City; SP06: Aba Tibetan and Qiang Autonomous Prefecture; SP07: Zhangye City; SP08: Gannan Tibetan Autonomous Prefecture; SP09: Ulanqab City; SP10: Garze Tibetan Autonomous Prefecture; SP11: Hainan Tibetan Autonomous Prefecture; SP12: Dingxi City; SP13: Bayingolin Mongolian Autonomous Prefecture; SP14: Changji Hui Autonomous Prefecture; SP15: Xilingol League; SP16: Bortala Mongolian Autonomous Prefecture; SP17: Shigatse City; SP18: Haidong City; SP19: Shannan City; SP20: Lanzhou City; SP21: Chifeng City; SP22: Haixi Mongol and Tibetan Autonomous Prefecture; SP23: Wuwei City; SP24: Xinzhou City; SP25: Haibei Tibetan Autonomous Prefecture.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fpls-15-1470653-g005.tif"/>
</fig>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<sec id="s4_1">
<label>4.1</label>
<title>Necessity of ensemble ENMs for prediction</title>
<p>In predicting the potential distribution probability of species, the single ENMs tends to overestimate in places with low species richness and underestimate in places with high species richness (<xref ref-type="bibr" rid="B5">Calabrese et&#xa0;al., 2014</xref>). The uncertainty of distribution prediction may distort policy making and planning. In the process of species distribution modeling, previous studies generally rely on multiple evaluation indicators to test the model results constructed by different algorithms based on the same data, selecting the algorithm with the highest statistical accuracy (<xref ref-type="bibr" rid="B22">Li and Wang, 2013</xref>; <xref ref-type="bibr" rid="B61">Xu et&#xa0;al., 2015</xref>). However, there may be inconsistencies among evaluation indicators. For example, in this study, the model with the highest AUC index was inconsistent with the model corresponding to the highest values of other indexes (Kappa and TSS). In addition, the construction of potential spatial distribution pattern of species is based on three prerequisites: (1) dynamic equilibrium of species distribution and environmental demand (<xref ref-type="bibr" rid="B3">Buckley et&#xa0;al., 2011</xref>), (2) idealized species migration ability (<xref ref-type="bibr" rid="B60">Xu et&#xa0;al., 2021</xref>), and (3) relatively conservative species ecological niche (<xref ref-type="bibr" rid="B16">Harris, 2015</xref>). The simulation results of the potential distribution of <italic>E. sibiricus</italic> in this study fails to effectively cover the existing distribution records, such as Wuyi Mountain, Luoxiao Mountain and Nanling Region, which show that the optimal algorithm may not achieve statistical optimum under all modeling conditions, nor may it effectively reduce the common sampling deviation (<xref ref-type="bibr" rid="B41">Randin et&#xa0;al., 2006</xref>). Nevertheless, to improve the accuracy of the model, using ensemble models to predict species distribution has become the trend of research (<xref ref-type="bibr" rid="B12">Grenouillet et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B15">Hao et&#xa0;al., 2020</xref>). The ensemble model maps the main trends of species distribution with the overall changes of all models and integrates the advantages of different algorithm models, such as the importance of variables or model response curve (<xref ref-type="bibr" rid="B4">Buisson et&#xa0;al., 2010</xref>), so as to effectively achieve and explore the prediction range of different algorithms and find consensus in the prediction of different mechanisms.</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Key environmental factors affecting the potential spatial distribution pattern of <italic>E. sibiricus</italic>
</title>
<p>In this study, the key environmental factors affecting the spatial distribution of <italic>E. sibiricus</italic> in the current period are all related to hydrothermal environmental conditions, especially humidity, air temperature, and potential evapotranspiration, which is consistent with the viewpoint that abiotic factors, mainly climate factors, dominated the species distribution pattern on a large scale (<xref ref-type="bibr" rid="B47">Soberon and Peterson, 2005</xref>). The response of plants to different environmental conditions gives them significant morphological plasticity (<xref ref-type="bibr" rid="B49">Sultan, 2000</xref>). Temperature and precipitation are the main climatic factors affecting forage growth (<xref ref-type="bibr" rid="B26">Li et&#xa0;al., 2018</xref>) and also the most important factors affecting genetic differentiation of the <italic>Elymus</italic> population (<xref ref-type="bibr" rid="B64">Yan et&#xa0;al., 2006</xref>). Studies have indicated that Bio01 and Bio12 have significant effects on the phenotype and genetic diversity of <italic>Elymus</italic> (<xref ref-type="bibr" rid="B38">Qi et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B13">Han et&#xa0;al., 2021</xref>); in this study, Bio01 and Bio12 were the main bioclimatic factors that significantly affect the potential spatial distribution of <italic>E. sibiricus</italic>. Compared to precipitation, temperature has a greater influence. Soil conditions can affect the growth and yield of aboveground and underground parts of plants (<xref ref-type="bibr" rid="B29">Liu et&#xa0;al., 2021</xref>). The appropriate dose of Ca<sup>2+</sup> can activate the activity of antioxidant enzymes, alleviate the oxidative damage caused by low temperature to the cell membrane system, and thus improve the cold resistance of <italic>E. nutans</italic> (<xref ref-type="bibr" rid="B39">Qimei et&#xa0;al., 2021</xref>). Application of phosphate fertilizer can not only effectively increase the yield of artificial grassland build by <italic>E. nutans</italic>, but also prevent the loss of soil organic carbon (<xref ref-type="bibr" rid="B42">Renzneg et&#xa0;al., 2020</xref>). In this study, ECa and AP, as important soil factors, significantly affected the spatial distribution pattern. As an alpine plant, <italic>E. sibiricus</italic> obviously used ECa to improve its cold resistance and AP to increase seed yield to expand population size. Altitude is a comprehensive environmental factor, and the genetic diversity of the <italic>Elymus</italic> population increases first and then decreases with altitude, with the maximum value appearing at around 3,300 m (<xref ref-type="bibr" rid="B63">Yan et&#xa0;al., 2009</xref>). The most suitable altitude for <italic>E. sibiricus</italic> in this study is 3,355.44 m, which provides evidence for the hypothesis that alpine plants have higher genetic diversity at the most suitable altitude. In addition, environmental factors such as precipitation, temperature, and radiation intensity will change with different altitudes (<xref ref-type="bibr" rid="B31">Lv et&#xa0;al., 2023</xref>), while the relative contribution of elevation to the potential distribution of <italic>E. sibiricus</italic> in this study is only 5.21%, which indicates that elevation may not have a direct impact on the quality of life of species, but may indirectly affect the habitat suitability by affecting other environmental factors. These findings well prove that the key environmental variables screened in this study have practical significance.</p>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Positive response of <italic>E. sibiricus</italic> population to climate change</title>
<p>The Qinghai-Tibet Plateau, located in southwest China, is the highest and largest plateau in the world, with an average elevation of over 4,000 m. During the uplift between Miocene periods and Quaternary periods, the geographical and climatic characteristics of QTP have undergone significant changes, making it recognized as one of the most important biodiversity hotspots in the world (<xref ref-type="bibr" rid="B33">Mao et&#xa0;al., 2021</xref>). QTP once served as a refuge for Northern temperate plants during the Quaternary period and was recognized as one of the important origin and radiation places of modern Northern temperate plants after the Glacial period (<xref ref-type="bibr" rid="B59">Xiong et&#xa0;al., 2022</xref>). The QTP is considered a potential center of origin for <italic>E. sibiricus</italic> population (<xref ref-type="bibr" rid="B59">Xiong et&#xa0;al., 2022</xref>, <xref ref-type="bibr" rid="B58">2024</xref>). During the warming process from the last glacial period to Holocene, <italic>E. sibiricus</italic> gradually spread to Sichuan, Gansu, and Xinjiang and then further expanded to Qinghai and Inner Mongolia (<xref ref-type="bibr" rid="B14">Han et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B59">Xiong et&#xa0;al., 2022</xref>, <xref ref-type="bibr" rid="B58">2024</xref>), which has formed a wide distribution pattern today. The pattern formed after the expansion of <italic>E. sibiricus</italic> population is similar to the potential distribution pattern predicted by this study, which is distributed in the vicinity of Qilian Mountains, Taihang Mountains, Kunlun Mountains, Hengduan Mountains, Qinling Mountains, and Changbai Mountains (<xref ref-type="bibr" rid="B59">Xiong et&#xa0;al., 2022</xref>, <xref ref-type="bibr" rid="B58">2024</xref>). The potential planting and introduction adaptation areas of <italic>E. sibiricus</italic> predicted in this study yield similar results. In addition, geographical range is usually considered the basic determinant of species diversity and diversification (<xref ref-type="bibr" rid="B44">Ricklefs, 2006</xref>), and isolation (such as geographical, ecological, or behavioral isolation) is an important factor affecting the blocking mechanism of gene flow among populations, which has the possibility of leading to the formation of continent-island model, island model, and isolation-by-distance model (<xref ref-type="bibr" rid="B40">Qu et&#xa0;al., 2004</xref>). The potential distribution pattern predicted by this study may be caused by some historical geological events, thus forming the <italic>E. sibiricus</italic> population to gather and distribute in mountains and its vicinity. However, due to the lack of molecular data support, it is impossible to clarify specific historical geological events. Climate warming is a significant threat to global biodiversity, especially for endemic species in temperature-sensitive alpine areas (<xref ref-type="bibr" rid="B50">Verrall and Pickering, 2020</xref>). Temperature has an important influence on the productivity of alpine meadow plants, but the continuous rise of temperature may have a negative impact on alpine grasslands (<xref ref-type="bibr" rid="B10">Fazlioglu and Wan, 2021</xref>). Although some studies have suggested that many alpine plants in QTP-Hengduan Mountains may not face a high risk of extinction due to habitat contraction caused by climate warming, some species at extremely high altitudes show significant range contraction (<xref ref-type="bibr" rid="B27">Liang et&#xa0;al., 2018</xref>). In this study, with the climate warming and drought intensification in the future, the potential suitable regions of <italic>E. sibiricus</italic> show a collapse trend and migrate to the high-altitude areas in southwest China. The introduction adaptation regions of six <italic>E. sibiricus</italic> germplasm are also distributed in the high-altitude regions of mountains in west China. It is speculated that in response to global warming, the population of <italic>E. sibiricus</italic> will shrink and concentrate in the Qilian Mountains (northeast of QTP), Taihang Mountains (junction of the Loess Plateau and Inner Mongolia Plateau), and Tianshan Mountains. In addition, many studies have recorded the pattern of alpine species migrating to high altitude in response to climate warming (<xref ref-type="bibr" rid="B8">Dullinger et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B19">K&#xf6;rner and Hiltbrunner, 2021</xref>), which supports the &#x201c;mountain-top&#x201d; extinction and &#x201c;uphill contraction&#x201d; hypothesis (<xref ref-type="bibr" rid="B48">Sorte and Jetz, 2010</xref>), indicating that it is necessary to take protective measures for species inhabiting extremely high altitude or narrowly distributed.</p>
</sec>
</sec>
</body>
<back>
<sec id="s5" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Material</bold>
</xref>. Further inquiries can be directed to the corresponding author.</p>
</sec>
<sec id="s6" sec-type="author-contributions">
<title>Author contributions</title>
<p>HL: Conceptualization, Formal analysis, Investigation, Methodology, Software, Visualization, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. YZ: Investigation, Resources, Software, Validation, Visualization, Writing &#x2013; review &amp; editing. YQ: Formal analysis, Investigation, Resources, Validation, Writing &#x2013; review &amp; editing. LT: Data curation, Investigation, Resources, Validation, Writing &#x2013; review &amp; editing. YZ: Investigation, Resources, Writing &#x2013; review &amp; editing. WX: Funding acquisition, Methodology, Project administration, Resources, Supervision, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s7" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research, authorship, and/or publication of this article. This research was funded by the Leading Scientist Project of Qinghai Province (2023- NK -147), Leading Scientist Project of Gansu Province (23ZDKA013), CARS (CARS-34) and Gansu Provincial Science and Technology Major Projects (22ZD6NA007).</p>
</sec>
<sec id="s8" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s9" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s10" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fpls.2024.1470653/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fpls.2024.1470653/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Table1.docx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document"/>
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