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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id>
<journal-title>Frontiers in Microbiology</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Microbiol.</abbrev-journal-title>
<issn pub-type="epub">1664-302X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmicb.2025.1509882</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Microbiology</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Impact of climate change on the distribution of <italic>Isaria cicadae</italic> <italic>Miquel</italic> in China: predictions based on the MaxEnt model</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" equal-contrib="yes">
<name><surname>He</surname> <given-names>Zhipeng</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="author-notes" rid="fn0004"><sup>&#x2020;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2864999/overview"/>
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<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/software/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
</contrib>
<contrib contrib-type="author" equal-contrib="yes">
<name><surname>Ali</surname> <given-names>Habib</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="author-notes" rid="fn0004"><sup>&#x2020;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/307042/overview"/>
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</contrib>
<contrib contrib-type="author">
<name><surname>Wu</surname> <given-names>Junhao</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Liu</surname> <given-names>Zhiqian</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Wei</surname> <given-names>Xinju</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
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<contrib contrib-type="author" corresp="yes">
<name><surname>Zhuo</surname> <given-names>Zhihang</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/880998/overview"/>
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<aff id="aff1"><sup>1</sup><institution>College of Life Science, China West Normal University</institution>, <addr-line>Nanchong</addr-line>, <country>China</country></aff>
<aff id="aff2"><sup>2</sup><institution>Department of Agricultural Engineering, Khwaja Fareed University of Engineering and Information Technology</institution>, <addr-line>Rahim Yar Khan</addr-line>, <country>Pakistan</country></aff>
<author-notes>
<fn fn-type="edited-by" id="fn0005">
<p>Edited by: Javier Carballo, University of Vigo, Spain</p>
</fn>
<fn fn-type="edited-by" id="fn0006">
<p>Reviewed by: Zhiqiang Lu, Chinese Academy of Sciences (CAS), China</p>
<p>Rulin Wang, Yibin University, China</p>
<p>Lei Zheng, Huangshan University, China</p>
</fn>
<corresp id="c001">&#x002A;Correspondence: Zhihang Zhuo, <email>zhuozhihang@foxmail.com</email></corresp>
<fn fn-type="equal" id="fn0004"><p><sup>&#x2020;</sup>These authors share first authorship</p></fn>
</author-notes>
<pub-date pub-type="epub">
<day>07</day>
<month>02</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>16</volume>
<elocation-id>1509882</elocation-id>
<history>
<date date-type="received">
<day>11</day>
<month>10</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>20</day>
<month>01</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2025 He, Ali, Wu, Liu, Wei and Zhuo.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>He, Ali, Wu, Liu, Wei and Zhuo</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>
<sec>
<title>Introduction</title>
<p><italic>Isaria cicadae</italic>, a historically valued edible and medicinal fungus in China, has been experiencing a critical decline in abundance due to ecological degradation and overexploitation. Understanding its potential distribution is essential for promoting sustainable harvesting practices.</p>
</sec>
<sec>
<title>Methods</title>
<p>This study utilizes the MaxEnt model, combined with known distribution records and 22 environmental variables, to predict the potential distribution of <italic>I. cicadae</italic> under three representative emission scenarios (CMIP6: SSP1-2.6, SSP2-4.5, and SSP5-8.5) for the 2050s and 2070s.</p>
</sec>
<sec>
<title>Results</title>
<p>The analysis identifies seven key environmental variables influencing the habitat suitability of <italic>I. cicadae</italic>: the mean temperature of the driest quarter (bio09), the mean temperature of the wettest quarter (bio08), precipitation in the wettest month (bio16), the mean diurnal range (bio02), isothermality (bio03), elevation, and slope. Currently, <italic>I. cicadae</italic> is mainly found in the provinces of Yunnan, Sichuan, Hunan, Hubei, Guizhou, Jiangxi, Guangxi, Fujian, Anhui, and Zhejiang, with Yunnan and Sichuan having the largest areas of high suitability at 25.79&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup> and 21.36&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup>, respectively.</p>
</sec>
<sec>
<title>Discussion</title>
<p>Jiangxi, Hunan, Yunnan, Guizhou, Fujian, and the Guangxi Zhuang Autonomous Region are identified as primary regions of high suitability. This study aims to further elucidate the impact of the environment on the distribution of <italic>I. cicadae</italic> from a geographical perspective and provide theoretical insights for the future cultivation and conservation strategies of this species.</p>
</sec>
</abstract>
<kwd-group>
<kwd><italic>Isaria cicadae</italic></kwd>
<kwd>MaxEnt</kwd>
<kwd>climate change</kwd>
<kwd>potential distribution</kwd>
<kwd>environmental</kwd>
</kwd-group>
<counts>
<fig-count count="7"/>
<table-count count="6"/>
<equation-count count="0"/>
<ref-count count="47"/>
<page-count count="11"/>
<word-count count="6440"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Food Microbiology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="sec1">
<label>1</label>
<title>Introduction</title>
<p><italic>Isaria cicad</italic>ae <italic>Miquel</italic> is a fungus of the genus <italic>Isaria</italic>, belonging to the family Cordycipitaceae and the order Hypocreales (<xref ref-type="bibr" rid="ref29">Shiwei and Hongkai, 2023</xref>; <xref ref-type="bibr" rid="ref44">Zhaoying, 2020</xref>). This species originates from the invasion of cicada larvae by fungi, where the fungal hyphae rapidly proliferate, ultimately leading to the death of the larva. The seven main hosts of this species in China are <italic>Platypleura kaempferi</italic>, <italic>Platylomia pieli</italic>, <italic>Cicada flammata</italic>, <italic>Mogannia conica conica</italic>, <italic>Oncotympana ella</italic>, <italic>Cicadatra shaluensis</italic>, and <italic>Hyalessa ronsnana</italic> (<xref ref-type="bibr" rid="ref15">Jiao, 2021</xref>). The resulting mycelial complex, which matures during the late spring and early summer (<xref ref-type="bibr" rid="ref23">Liu et al., 2008</xref>). The species predominantly inhabits bamboo forests, broadleaf forests at altitudes below 2,500 meters, or mixed coniferous-broadleaf forests dominated by species such as <italic>Cyclobalanopsis glauca</italic>, <italic>Castanea henryi</italic>, <italic>Pinus yunnanensis</italic>, and <italic>Abies</italic> spp. (<xref ref-type="bibr" rid="ref36">Wang et al., 2006</xref>; <xref ref-type="bibr" rid="ref42">Zhang et al., 2022</xref>). <italic>I. cicadae</italic> is a highly regarded <italic>Cordyceps</italic> species, similar to the well-known <italic>Ophiocordyceps sinensis</italic>. It is rich in bioactive compounds such as nucleosides, ergosterol, cordycepic acid, and polysaccharides, which have considerable potential for both culinary and medicinal uses (<xref ref-type="bibr" rid="ref47">Zhu et al., 2024</xref>). Research indicates that <italic>I. cicadae</italic> has health benefits similar to those of <italic>O. sinensis</italic> and holds significant market value (<xref ref-type="bibr" rid="ref3">Bian et al., 2017</xref>). However, global climate change and human activities have severely degraded its habitat, leading to a sharp decline in its population and a mismatch between supply and demand. The International Union for Conservation of Nature (IUCN) has classified this species as vulnerable on its Red List (<xref ref-type="bibr" rid="ref20">Li et al., 2020</xref>). Therefore, it is significant to identify and predict regions suitable for the cultivation and conservation of <italic>I. cicadae</italic>.</p>
<p>Species distribution models (SDMs) are empirical tools used to quantify the ecological niches of species within their environments. SDMs are created by integrating occurrence data of species with corresponding environmental variables, identifying associations between them, and using these relationships to estimate the species&#x2019; distribution across the study area (<xref ref-type="bibr" rid="ref45">Zhonglin et al., 2015</xref>). By combining these occurrence data with environmental variables from sampling sites, SDMs model the species-environment relationships to predict potential distribution patterns (<xref ref-type="bibr" rid="ref7">Deng et al., 2022</xref>). Commonly used niche models include CLIMEX, BIOCLIM, GARP, DOMAIN, and MaxEnt (<xref ref-type="bibr" rid="ref37">Wei et al., 2024</xref>). The MaxEnt model is particularly notable for its advantages, such as its effectiveness with small sample sizes, high predictive accuracy, rapid computation, and user-friendliness. In addition, it can use the Jackknife method to conduct significance testing and evaluation on individual environmental variables. The MaxEnt model has been widely applied both domestically and internationally across various fields (<xref ref-type="bibr" rid="ref5">Brambilla et al., 2024</xref>; <xref ref-type="bibr" rid="ref43">Zhao et al., 2024</xref>). In particular, MaxEnt is extensively used in disciplines such as ecology (<xref ref-type="bibr" rid="ref6">Cao et al., 2023</xref>), conservation biology (<xref ref-type="bibr" rid="ref22">Lin et al., 2024</xref>), and geography (<xref ref-type="bibr" rid="ref35">Wang C. et al., 2023</xref>; <xref ref-type="bibr" rid="ref33">Wang Z. et al., 2023</xref>). Its applications include assessing the potential impacts of climate change (<xref ref-type="bibr" rid="ref11">Guo et al., 2017</xref>; <xref ref-type="bibr" rid="ref10">Guo et al., 2019</xref>), predicting the potential distributions of endangered species (<xref ref-type="bibr" rid="ref1">Ab Lah et al., 2021</xref>; <xref ref-type="bibr" rid="ref21">Lin et al., 2022</xref>), evaluating the spread of invasive species (<xref ref-type="bibr" rid="ref4">Bowen and Stevens, 2020</xref>; <xref ref-type="bibr" rid="ref31">Tu et al., 2021</xref>), forecasting changes in ecological habitat ranges (<xref ref-type="bibr" rid="ref34">Wang et al., 2024</xref>), and investigating biomass reserves (<xref ref-type="bibr" rid="ref27">Rodr&#x00ED;guez-Veiga et al., 2016</xref>).</p>
<p>Research on the distribution of <italic>I. cicadae</italic> is currently limited. Existing studies primarily concentrate on its edibility (<xref ref-type="bibr" rid="ref14">Jia et al., 2023</xref>), chemical composition (<xref ref-type="bibr" rid="ref25">Mei et al., 2023</xref>), metabolomic profiling (<xref ref-type="bibr" rid="ref30">Tang et al., 2023</xref>), and the pharmacological potential of its key compounds (<xref ref-type="bibr" rid="ref19">Li Z. et al., 2024</xref>; <xref ref-type="bibr" rid="ref26">Olatunji et al., 2016</xref>). Investigations into suitable habitats for <italic>I. cicadae</italic> are notably scarce. Therefore, this study provides valuable scientific evidence for exploring the potential distribution of <italic>I. cicadae</italic>.</p>
<p>Using known distribution data for <italic>I. cicadae</italic> and WorldClim environmental variables, this study employed ArcGIS 10.8 and MaxEnt 3.4.4 to analyze the environmental adaptability of <italic>I. cicadae</italic>. This study predicts both the current and future potential geographical distribution of <italic>I. cicadae</italic> and examines the trends in centroid movement affecting its distribution. Additionally, this study identifies key environmental factors influencing the distribution of <italic>I. cicadae</italic> and assesses their impact on the fungus, providing theoretical insights for its conservation.</p>
</sec>
<sec sec-type="materials|methods" id="sec2">
<label>2</label>
<title>Materials and methods</title>
<sec id="sec3">
<label>2.1</label>
<title>Species distribution data</title>
<p>This study primarily obtained species distribution data for <italic>I. cicadae</italic> from the Global Biodiversity Information Facility (GBIF),<xref ref-type="fn" rid="fn0001"><sup>1</sup></xref> including 194 occurrence records for <italic>I. cicadae</italic> and 634 host distribution data points. The precise geographic coordinates of the species distribution points were determined using Google Earth.<xref ref-type="fn" rid="fn0002"><sup>2</sup></xref> The collected distribution points for <italic>I. cicadae</italic> were imported into ArcGIS 10.8. Buffer analysis was employed to filter these distribution points, reducing overfitting caused by spatial overlap or proximity. The filtered records were saved in CSV format, resulting in a total of 161 <italic>I. cicadae</italic> occurrence records and 247 host occurrence records, as shown in <xref ref-type="fig" rid="fig1">Figure 1</xref>.</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Geographical distribution points of <italic>I. cicadae</italic> in China.</p>
</caption>
<graphic xlink:href="fmicb-16-1509882-g001.tif"/>
</fig>
</sec>
<sec id="sec4">
<label>2.2</label>
<title>Environmental variables and data processing</title>
<p>This work downloaded 19 climate variables from the WorldClim database.<xref ref-type="fn" rid="fn0003"><sup>3</sup></xref> To predict the potential distribution of <italic>I. cicadae</italic> under future climate conditions, the study employed three scenarios from the Beijing Climate Center&#x2019;s (BCC) advanced climate model BCC-CSM2-MR: SSP1-2.6 (low emission pathway), SSP3-7.0 (medium emission pathway), and SSP5-8.5 (high emission pathway). To address high autocorrelation among environmental variables and to reduce overfitting while improving model accuracy, Pearson correlation coefficients (<italic>r</italic>) were used to identify multicollinearity among the environmental variables (<xref ref-type="table" rid="tab1">Table 1</xref>). By excluding variables with correlation coefficients greater than 0.8, the impact of multicollinearity on overfitting was minimized (<xref ref-type="bibr" rid="ref8">Fu et al., 2024</xref>). This process ultimately determined the key environmental variables required for modeling.</p>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Presents Pearson correlation coefficients among the environmental factors.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th/>
<th align="center" valign="top">slope</th>
<th align="center" valign="top">bio2</th>
<th align="center" valign="top">bio3</th>
<th align="center" valign="top">bio8</th>
<th align="center" valign="top">bio16</th>
<th align="center" valign="top">bio19</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">bio2</td>
<td align="char" valign="middle" char=".">0.0930</td>
<td/>
<td/>
<td/>
<td/>
<td/>
</tr>
<tr>
<td align="left" valign="middle">bio3</td>
<td align="char" valign="middle" char=".">0.0748</td>
<td align="center" valign="middle">0.4152</td>
<td/>
<td/>
<td/>
<td/>
</tr>
<tr>
<td align="left" valign="middle">bio8</td>
<td align="char" valign="middle" char=".">0.1755</td>
<td align="center" valign="middle">0.5345</td>
<td align="center" valign="middle">0.5265</td>
<td/>
<td/>
<td/>
</tr>
<tr>
<td align="left" valign="middle">bio16</td>
<td align="char" valign="middle" char=".">0.1625</td>
<td align="center" valign="middle">0.0872</td>
<td align="center" valign="middle">0.6396</td>
<td align="center" valign="middle">0.4919</td>
<td/>
<td/>
</tr>
<tr>
<td align="left" valign="middle">bio19</td>
<td align="char" valign="middle" char=".">0.0836</td>
<td align="center" valign="middle">&#x2212;0.0984</td>
<td align="center" valign="middle">0.4786</td>
<td align="center" valign="middle">0.2348</td>
<td align="center" valign="middle">0.6506</td>
<td/>
</tr>
<tr>
<td align="left" valign="middle">elev</td>
<td align="char" valign="middle" char=".">0.0775</td>
<td align="center" valign="middle">&#x2212;0.1737</td>
<td align="center" valign="middle">&#x2212;0.2317</td>
<td align="center" valign="middle">&#x2212;0.7654</td>
<td align="center" valign="middle">&#x2212;0.3474</td>
<td align="center" valign="middle">&#x2212;0.2275</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="sec5">
<label>2.3</label>
<title>Model construction and assessment</title>
<p>Distribution data for <italic>I. cicadae</italic> with various environmental variables to assess their impact on the species distribution. Using MaxEnt version 3.4.4, the potential distribution range of the species was modeled. To minimize the uncertainty introduced by random data selection, 75% of the species occurrence data was randomly designated as training samples, with the remaining 25% reserved for testing. The training process was repeated 10 times, keeping all other parameters at their default settings. This study employed the jackknife technique to determine the contribution rates of key environmental variables and utilized response curves to analyze the relationship between <italic>I. cicadae</italic> distribution and climatic variables.</p>
<p>The MaxEnt model quantifies species suitability on a scale from 0 to 1, with values closer to 1 indicating a higher probability of the species&#x2019; presence under specific environmental conditions (<xref ref-type="bibr" rid="ref18">Li R. et al., 2024</xref>). Following the Intergovernmental Panel on Climate Change (IPCC) reporting methodology and considering the habitat suitability for <italic>I. cicadae</italic>, regions with a suitability score below 0.1 were classified as unsuitable. Areas with scores between 0.1 and 0.3 were designated as low suitability, those between 0.3 and 0.5 as moderate suitability, and habitats with scores of 0.5 or higher were considered highly suitable (<xref ref-type="bibr" rid="ref13">Iturbide et al., 2020</xref>).</p>
<p>The model performance is evaluated using receiver operating characteristic (ROC) curve, which is essential for assessing the accuracy of its predictions. A key metric in this evaluation is the area under the ROC curve (AUC), which measures the effectiveness of the MaxEnt model (<xref ref-type="bibr" rid="ref35">Wang C. et al., 2023</xref>; <xref ref-type="bibr" rid="ref33">Wang Z. et al., 2023</xref>). The AUC ranges from 0.5 to 1, with values closer to 1 indicating higher model precision (<xref ref-type="bibr" rid="ref32">Walden-Schreiner et al., 2017</xref>). Specifically, an AUC between 0.5 and 0.6 reflects substandard performance, 0.6 to 0.7 indicates poor performance, 0.8 to 0.9 signifies excellent performance, and values above 0.9 denote high performance (<xref ref-type="bibr" rid="ref2">Bai et al., 2024</xref>).</p>
</sec>
</sec>
<sec sec-type="results" id="sec6">
<label>3</label>
<title>Results</title>
<sec id="sec7">
<label>3.1</label>
<title>Model performance and key environmental variables</title>
<p>The area under the curve (AUC) is a metric that reflects the accuracy of MaxEnt model simulations. In this study, the MaxEnt model achieved an average AUC of 0.984&#x202F;&#x00B1;&#x202F;0.002, as shown in <xref ref-type="fig" rid="fig2">Figure 2</xref>. This high level of accuracy underscores the model&#x2019;s reliability and supports its use as a foundation for the current investigation.</p>
<fig position="float" id="fig2">
<label>Figure 2</label>
<caption>
<p>Receiver operating characteristic curve and AUC result of MaxEnt modeling.</p>
</caption>
<graphic xlink:href="fmicb-16-1509882-g002.tif"/>
</fig>
<p>Using Pearson correlation coefficients, seven key environmental variables were identified and incorporated into the species distribution model. Jackknife analysis revealed the contribution of each variable as follows: bio16 (50.7%), bio19 (20.0%), slope (7.9%), bio03 (7.1%), bio08 (6.2%), bio02 (6.1%), and elevation (1.9%), with a cumulative contribution of 99.9%. In terms of importance, the rankings were: bio09 (42.9%), bio08 (25.7%), bio16 (20.6%), bio02 (5.5%), bio03 (2.6%), elevation (1.5%), and slope (1.2%), with a total importance score of 100% (<xref ref-type="table" rid="tab2">Table 2</xref>). These results indicate that the selected environmental variables effectively simulate the potential distribution of <italic>I. cicadae</italic>.</p>
<table-wrap position="float" id="tab2">
<label>Table 2</label>
<caption>
<p>The percent contribution and permutation importance of climatic variables in the MaxEnt modeling of <italic>I. cicadae</italic>.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Variable</th>
<th align="center" valign="top">Percent contribution</th>
<th align="center" valign="top">Permutation importance</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">bio16</td>
<td align="char" valign="middle" char=".">50.7</td>
<td align="char" valign="middle" char=".">20.6</td>
</tr>
<tr>
<td align="left" valign="middle">bio09</td>
<td align="char" valign="middle" char=".">20</td>
<td align="char" valign="middle" char=".">42.9</td>
</tr>
<tr>
<td align="left" valign="middle">slope</td>
<td align="char" valign="middle" char=".">7.9</td>
<td align="char" valign="middle" char=".">1.2</td>
</tr>
<tr>
<td align="left" valign="middle">bio03</td>
<td align="char" valign="middle" char=".">7.1</td>
<td align="char" valign="middle" char=".">2.6</td>
</tr>
<tr>
<td align="left" valign="middle">bio08</td>
<td align="char" valign="middle" char=".">6.2</td>
<td align="char" valign="middle" char=".">25.7</td>
</tr>
<tr>
<td align="left" valign="middle">bio02</td>
<td align="char" valign="middle" char=".">6.1</td>
<td align="char" valign="middle" char=".">5.5</td>
</tr>
<tr>
<td align="left" valign="middle">elev</td>
<td align="char" valign="middle" char=".">1.9</td>
<td align="char" valign="middle" char=".">1.5</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="sec8">
<label>3.2</label>
<title>The potential distribution of <italic>Isaria cicadae</italic> in the current period</title>
<p>Employing an MaxEnt model, a predictive map of the most suitable habitats for <italic>I. cicadae</italic> was generated (<xref ref-type="fig" rid="fig3">Figure 3A</xref>). The findings suggest that under the current climate scenario, the species&#x2019; suitable habitats are widely distributed across most of China, excluding the provinces of Jilin, Heilongjiang, Inner Mongolia, Qinghai, Ningxia, and Xinjiang. The total area of suitable habitats covers approximately 341.54&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup>, equivalent to 35.53% of China&#x2019;s land area. High-suitability regions are predominantly situated in Yunnan, Sichuan, Hunan, Hubei, Guizhou, Jiangxi, Guangxi, Fujian, Anhui, and Zhejiang, covering an area of roughly 194.34&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup>, which represents 20.22% of China&#x2019;s total landmass. Currently, Yunnan Province boasts the largest area of high suitability, encompassing 25.79&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup>, followed by Sichuan Province with 21.36&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup> and Hunan Province with 19.13&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup>. These regions account for 13.27, 10.99, and 9.85% of the national high-suitability habitat area, respectively. Notably, the high-suitability area in Hunan constitutes 98.90% of the province&#x2019;s total land area, while in Jiangxi and Guizhou, these proportions are even more significant, reaching 99.23 and 99.27%, respectively. Additionally, in Zhejiang, Hubei, and Fujian, the high-suitability areas surpass 80% of their respective provincial and masses (see <xref ref-type="table" rid="tab3">Table 3</xref>).</p>
<fig position="float" id="fig3">
<label>Figure 3</label>
<caption>
<p>Under the current situation, the potential suitable distribution area of <italic>I. cicadae</italic> and host. <bold>(A)</bold> The potential suitable distribution area of <italic>I. cicadae</italic>. <bold>(B)</bold> The potential suitable distribution area of host. <bold>(C)</bold> Suitable area overlay map.</p>
</caption>
<graphic xlink:href="fmicb-16-1509882-g003.tif"/>
</fig>
<p>The potential distribution areas of the primary hosts for <italic>I. cicadae</italic> under the current climate scenario were also predicted (<xref ref-type="fig" rid="fig3">Figure 3B</xref>). High-suitability regions for the hosts span an area of 24.884&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup>, primarily located at the junction of Zhejiang, Jiangsu, and Anhui provinces. Furthermore, when the suitable areas from the <italic>I. cicadae</italic> potential distribution map were overlaid with those of the host&#x2019;s potential distribution (<xref ref-type="fig" rid="fig3">Figure 3C</xref>), it was found that the shared suitable region for both <italic>I. cicadae</italic> and the hosts spans 267.089&#x202F;&#x00D7;&#x202F;10<sup>4</sup>&#x202F;km<sup>2</sup>, representing 78.41% of the suitable area for <italic>I. cicadae</italic> and 99.13% of the suitable area for the hosts.</p>
<table-wrap position="float" id="tab3">
<label>Table 3</label>
<caption>
<p><italic>I. cicadae</italic> top 10 high-suitability regions for habitat.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Province</th>
<th align="center" valign="top">High suitable area (10<sup>4</sup> km<sup>2</sup>)</th>
<th align="center" valign="top">Total (10<sup>4</sup> km<sup>2</sup>)</th>
<th align="center" valign="top">Percentage of highly suitable areas in the province (%)</th>
<th align="center" valign="top">Percentage of highly suitable areas in China (%)</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">Yunnan</td>
<td align="char" valign="middle" char=".">25.79</td>
<td align="center" valign="middle">34.26</td>
<td align="center" valign="middle">75.29%</td>
<td align="char" valign="middle" char=".">13.27%</td>
</tr>
<tr>
<td align="left" valign="middle">Sichuan</td>
<td align="char" valign="middle" char=".">21.36</td>
<td align="center" valign="middle">45.74</td>
<td align="center" valign="middle">46.70%</td>
<td align="char" valign="middle" char=".">10.99%</td>
</tr>
<tr>
<td align="left" valign="middle">Hunan</td>
<td align="char" valign="middle" char=".">19.13</td>
<td align="center" valign="middle">19.35</td>
<td align="center" valign="middle">98.90%</td>
<td align="char" valign="middle" char=".">9.85%</td>
</tr>
<tr>
<td align="left" valign="middle">Hubei</td>
<td align="char" valign="middle" char=".">16.37</td>
<td align="center" valign="middle">17.57</td>
<td align="center" valign="middle">93.15%</td>
<td align="char" valign="middle" char=".">8.42%</td>
</tr>
<tr>
<td align="left" valign="middle">Guizhou</td>
<td align="char" valign="middle" char=".">15.87</td>
<td align="center" valign="middle">15.99</td>
<td align="center" valign="middle">99.27%</td>
<td align="char" valign="middle" char=".">8.17%</td>
</tr>
<tr>
<td align="left" valign="middle">Jiangxi</td>
<td align="char" valign="middle" char=".">15.13</td>
<td align="center" valign="middle">15.25</td>
<td align="center" valign="middle">99.23%</td>
<td align="char" valign="middle" char=".">7.78%</td>
</tr>
<tr>
<td align="left" valign="middle">Guangxi</td>
<td align="char" valign="middle" char=".">12.38</td>
<td align="center" valign="middle">20.89</td>
<td align="center" valign="middle">59.28%</td>
<td align="char" valign="middle" char=".">6.37%</td>
</tr>
<tr>
<td align="left" valign="middle">Fujian</td>
<td align="char" valign="middle" char=".">9.68</td>
<td align="center" valign="middle">10.94</td>
<td align="center" valign="middle">88.45%</td>
<td align="char" valign="middle" char=".">4.98%</td>
</tr>
<tr>
<td align="left" valign="middle">Anhui</td>
<td align="char" valign="middle" char=".">9.51</td>
<td align="center" valign="middle">13.37</td>
<td align="center" valign="middle">71.13%</td>
<td align="char" valign="middle" char=".">4.89%</td>
</tr>
<tr>
<td align="left" valign="middle">Zhejiang</td>
<td align="char" valign="middle" char=".">8.81</td>
<td align="center" valign="middle">9.36</td>
<td align="center" valign="middle">94.14%</td>
<td align="char" valign="middle" char=".">4.53%</td>
</tr>
<tr>
<td align="left" valign="middle">China</td>
<td align="char" valign="middle" char=".">194.34</td>
<td/>
<td/>
<td align="char" valign="middle" char=".">20.22%</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="sec9">
<label>3.3</label>
<title>The potential distribution of <italic>Isaria cicadae</italic> in the future period</title>
<p>Under the climate change scenarios of SSP1-2.6, SSP3-7.0, and SSP5-8.5, the suitable distribution ranges for <italic>I. cicadae</italic> in the periods 2041&#x2013;2060 and 2061&#x2013;2080 are illustrated in <xref ref-type="fig" rid="fig4">Figure 4</xref>. Exceptions for Xinjiang, Qinghai, Ningxia, Inner Mongolia, Heilongjiang, and Jilin Provinces, suitable habitats for <italic>I. cicadae</italic> are found across the rest of China. High-suitability zones are primarily located in Chongqing, Zhejiang, Jiangxi, Hunan, Yunnan, Guizhou, Sichuan, Fujian, Guangdong, and Guangxi. Additionally, there are scattered areas of high suitability in Gansu, Shaanxi, Henan, and Hebei. The projected distribution regions for the 2050s and 2070s show minimal changes compared to the current distribution. In the 2050s, high-suitability areas only decreased under the SSP3-7.0 and SSP5-8.5 scenarios, while other suitability categories expanded. By the 2070s, high-suitability areas decreased under the SSP1-2.6 and SSP3-7.0 scenarios, while moderate-suitability areas shrank under the SSP5-8.5 scenario, with other categories continuing to expand (<xref ref-type="table" rid="tab4">Table 4</xref>).</p>
<fig position="float" id="fig4">
<label>Figure 4</label>
<caption>
<p>A predictive map of potentially suitable areas for <italic>I. cicadae</italic> in China under different climate change scenarios. <bold>(A)</bold> SSP1-2.62050s. <bold>(B)</bold> SSP3-7.02050s. <bold>(C)</bold> SSP5-8.52050s. <bold>(D)</bold> SSP1-2.62070s. <bold>(E)</bold> SSP3-7.02070s. <bold>(F)</bold> SSP5-8.52070s. (Purple: high suitability zone; pink: moderate suitability zone; yellow: low suitability zone; white: unsuitable zone).</p>
</caption>
<graphic xlink:href="fmicb-16-1509882-g004.tif"/>
</fig>
<table-wrap position="float" id="tab4">
<label>Table 4</label>
<caption>
<p>Predicts suitable areas for <italic>I. cicadae</italic> under the current and future climatic conditions.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top" rowspan="2">Decade scenarios</th>
<th align="center" valign="top" colspan="3">Predicted area (10<sup>4</sup> km<sup>2</sup>)</th>
<th align="center" valign="top" colspan="3">Comparison with current distribution (%)</th>
</tr>
<tr>
<th align="center" valign="top">Low suitable</th>
<th align="center" valign="top">Medium suitable</th>
<th align="center" valign="top">High suitable</th>
<th align="center" valign="top">Low suitable</th>
<th align="center" valign="top">Medium suitable</th>
<th align="center" valign="top">High suitable</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">Current</td>
<td align="char" valign="middle" char=".">61.72</td>
<td align="char" valign="middle" char=".">85.49</td>
<td align="char" valign="middle" char=".">194.34</td>
<td/>
<td/>
<td/>
</tr>
<tr>
<td align="left" valign="middle">41-60ssp126</td>
<td align="char" valign="middle" char=".">66.63</td>
<td align="char" valign="middle" char=".">87.00</td>
<td align="char" valign="middle" char=".">195.60</td>
<td align="center" valign="middle">7.96%</td>
<td align="center" valign="middle">1.77%</td>
<td align="center" valign="middle">0.65%</td>
</tr>
<tr>
<td align="left" valign="middle">41-60ssp370</td>
<td align="char" valign="middle" char=".">77.78</td>
<td align="char" valign="middle" char=".">89.28</td>
<td align="char" valign="middle" char=".">189.38</td>
<td align="center" valign="middle">26.02%</td>
<td align="center" valign="middle">4.44%</td>
<td align="center" valign="middle">&#x2212;2.55%</td>
</tr>
<tr>
<td align="left" valign="middle">41-60ssp585</td>
<td align="char" valign="middle" char=".">75.95</td>
<td align="char" valign="middle" char=".">101.35</td>
<td align="char" valign="middle" char=".">187.44</td>
<td align="center" valign="middle">23.06%</td>
<td align="center" valign="middle">18.55%</td>
<td align="center" valign="middle">&#x2212;3.55%</td>
</tr>
<tr>
<td align="left" valign="middle">61-80ssp126</td>
<td align="char" valign="middle" char=".">74.72</td>
<td align="char" valign="middle" char=".">105.73</td>
<td align="char" valign="middle" char=".">176.25</td>
<td align="center" valign="middle">21.07%</td>
<td align="center" valign="middle">23.67%</td>
<td align="center" valign="middle">&#x2212;9.31%</td>
</tr>
<tr>
<td align="left" valign="middle">61-80ssp370</td>
<td align="char" valign="middle" char=".">93.46</td>
<td align="char" valign="middle" char=".">95.72</td>
<td align="char" valign="middle" char=".">171.32</td>
<td align="center" valign="middle">51.43%</td>
<td align="center" valign="middle">11.97%</td>
<td align="center" valign="middle">&#x2212;11.84%</td>
</tr>
<tr>
<td align="left" valign="middle">61-80ssp585</td>
<td align="char" valign="middle" char=".">74.92</td>
<td align="char" valign="middle" char=".">84.86</td>
<td align="char" valign="middle" char=".">206.04</td>
<td align="center" valign="middle">21.39%</td>
<td align="center" valign="middle">&#x2212;0.73%</td>
<td align="center" valign="middle">6.02%</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>The predictions suggest that the suitable habitat area for <italic>I. cicadae</italic> will generally trend upward during the 2050s. However, under the SSP3-7.0 and SSP5-8.5 scenarios, the high suitability region for <italic>I. cicadae</italic> is expected to decrease by 2.55 and 3.55%, respectively. In contrast, the other suitable habitat areas for <italic>I. cicadae</italic> are projected to increased.</p>
<p>The findings suggest that by the 2070s, the suitable habitat for <italic>I. cicadae</italic> is projected to expand overall. However, this expansion is not uniform: significant growth is observed only under the SSP5-8.5 scenario in high-suited areas, with an increase of approximately 6.02%. In contrast, a decrease in suitable habitat area is evident only within the SSP5-8.5 scenario for mid-suited areas, with a reduction of about 0.73%. Across all three scenarios, low-suited areas exhibit an increase in habitat extent.</p>
</sec>
<sec id="sec10">
<label>3.4</label>
<title>The environmental variables influencing the geographical distribution of <italic>Isaria cicadae</italic></title>
<p>This study evaluated three scenarios-&#x201C;only variables,&#x201D; &#x201C;no variables&#x201D; and &#x201C;all variables&#x201D;-to assess the influence of environmental factors on the distribution of <italic>I. cicadae</italic> using MaxEnt software and the jackknife method. The analysis demonstrated the varying impacts of seven environmental variables on the species&#x2019; distribution (<xref ref-type="fig" rid="fig5">Figure 5</xref>). The results exhibit that when using only a single environmental variable, bio09 and bio16 have the most significant effect on the model&#x2019;s performance. This finding highlights that these two factors play a critical role in shaping the distribution of <italic>I. cicadae</italic>.</p>
<fig position="float" id="fig5">
<label>Figure 5</label>
<caption>
<p>The jackknife test assesses the importance of environmental variables for <italic>I. cicadae</italic>.</p>
</caption>
<graphic xlink:href="fmicb-16-1509882-g005.tif"/>
</fig>
<p>Based on the response curves of the <italic>I. cicadae</italic> probability distribution shown in <xref ref-type="fig" rid="fig6">Figure 6</xref>, with a threshold of 0.5, values above 0.5 are considered suitable. The suitable ranges for the seven environmental variables&#x2014;bio02, bio03, bio08, bio09, bio16, elevation, and slope&#x2014;are as follows: bio02: 7.18 to 11.39&#x00B0;C, bio03: 24.45 to 53.28&#x00B0;C, bio08: 16.05 to 25.62&#x00B0;C, bio09: 0.23 to 13.70&#x00B0;C, bio16: 352.78 to 936.53&#x202F;mm, elevation: less than 2,990&#x202F;m, and slope: greater than 1.71 (<xref ref-type="table" rid="tab5">Table 5</xref>).</p>
<fig position="float" id="fig6">
<label>Figure 6</label>
<caption>
<p>The probability of the presence of <italic>I. cicadae</italic> and the response curve of key environmental variables. <bold>(A)</bold> bio02. <bold>(B)</bold> bio03. <bold>(C)</bold> bio08. <bold>(D)</bold> bio09. <bold>(E)</bold> bio16. <bold>(F)</bold> elev. <bold>(G)</bold> slope. The red curve represents the average of 10 replicates; blue margins represent the standard deviation.</p>
</caption>
<graphic xlink:href="fmicb-16-1509882-g006.tif"/>
</fig>
<table-wrap position="float" id="tab5">
<label>Table 5</label>
<caption>
<p>The optimal range of environmental variables corresponds to the potential distribution of <italic>I. cicadae</italic>.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Environmental variables</th>
<th align="center" valign="top">Suitable range</th>
<th align="center" valign="top">Optimum value</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">bio02/&#x00B0;C</td>
<td align="center" valign="middle">7.18&#x2013;11.39</td>
<td align="char" valign="middle" char=".">7.89</td>
</tr>
<tr>
<td align="left" valign="middle">bio03/&#x00B0;C</td>
<td align="center" valign="middle">24.45&#x2013;53.28</td>
<td align="char" valign="middle" char=".">29.88</td>
</tr>
<tr>
<td align="left" valign="middle">bio08/&#x00B0;C</td>
<td align="center" valign="middle">16.05&#x2013;25.62</td>
<td align="char" valign="middle" char=".">22.37</td>
</tr>
<tr>
<td align="left" valign="middle">bio09/&#x00B0;C</td>
<td align="center" valign="middle">0.23&#x2013;13.70</td>
<td align="char" valign="middle" char=".">6.62</td>
</tr>
<tr>
<td align="left" valign="middle">bio16/mm</td>
<td align="center" valign="middle">352.78&#x2013;936.53</td>
<td align="char" valign="middle" char=".">484.88</td>
</tr>
<tr>
<td align="left" valign="middle">elev</td>
<td align="center" valign="middle">&#x003C;2,990</td>
<td align="char" valign="middle" char=".">24.8</td>
</tr>
<tr>
<td align="left" valign="middle">slope</td>
<td align="center" valign="middle">&#x003E;1.71</td>
<td align="char" valign="middle" char=".">14.49</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="sec11">
<label>3.5</label>
<title>The centroid variation in the potential distribution of <italic>Isaria cicadae</italic></title>
<p><xref ref-type="fig" rid="fig7">Figure 7</xref> illustrates the high suitability centroids of <italic>I. cicadae</italic> under different climate scenarios. Currently, the centroid of the high suitability zone is located in Jishou City, Hunan Province. In the SSP1-2.6 scenario, by the 2050s, the centroid is projected to shift northeast by 43.14&#x202F;km to Yuanling County. By the 2070s, it is expected to move southwest by 58.89&#x202F;km, returning to Jishou City. In the SSP370 scenario, by the 2050s, the centroid is anticipated to move northeast by 8.95&#x202F;km into Luxi County. By the 2070s, it is expected to shift southwest by 26.99&#x202F;km, remaining within Luxi County. Under the SSP585 scenario, by the 2050s, the centroid is projected to shift northwest by 26.63&#x202F;km into Baojing County, and by the 2070s, to move southeast by 32.11&#x202F;km into Luxi County. Among the three carbon emission scenarios, centroid displacement is relatively minor under SSP3-7.0 and SSP5-8.5 from 2050 to 2070, while it is more pronounced under SSP1-2.6.</p>
<fig position="float" id="fig7">
<label>Figure 7</label>
<caption>
<p>The change in the centroid of the potential distribution area of <italic>I. cicadae</italic> in China.</p>
</caption>
<graphic xlink:href="fmicb-16-1509882-g007.tif"/>
</fig>
<p>In summary, except for the 2050s under the SSP5-8.5 and SSP1-2.6 scenarios, the potential high suitability areas for <italic>I. cicadae</italic> are predominantly located within Jishou and Luxi Counties (<xref ref-type="table" rid="tab6">Table 6</xref>). This suggests that the distribution of suitable habitats for <italic>I. cicadae</italic> is influenced by the selected climate scenario, with notable shifts anticipated under specific scenarios.</p>
<table-wrap position="float" id="tab6">
<label>Table 6</label>
<caption>
<p>Under climate change scenarios, the centroid displacement trajectory of <italic>I. cicadae</italic> habitats.</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th align="left" valign="top">Scene</th>
<th align="center" valign="top">Period</th>
<th align="center" valign="top">Direction</th>
<th align="center" valign="top">Displacement/km</th>
<th align="center" valign="top">Location</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" rowspan="2">ssp126</td>
<td align="left" valign="middle">Contemporary to 2050s</td>
<td align="left" valign="middle">Northeast</td>
<td align="char" valign="middle" char=".">40.14</td>
<td align="left" valign="middle">Yuanling county</td>
</tr>
<tr>
<td align="left" valign="middle">2050s to 2090s</td>
<td align="left" valign="middle">Southwest</td>
<td align="char" valign="middle" char=".">58.89</td>
<td align="left" valign="middle">Jishou city</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="2">ssp370</td>
<td align="left" valign="middle">Contemporary to 2050s</td>
<td align="left" valign="middle">Northeast</td>
<td align="char" valign="middle" char=".">8.95</td>
<td align="left" valign="middle">Luxi county</td>
</tr>
<tr>
<td align="left" valign="middle">2050s to 2090s</td>
<td align="left" valign="middle">Southwest</td>
<td align="char" valign="middle" char=".">26.99</td>
<td align="left" valign="middle">Luxi county</td>
</tr>
<tr>
<td align="left" valign="middle" rowspan="2">ssp585</td>
<td align="left" valign="middle">Contemporary to 2050s</td>
<td align="left" valign="middle">Northwest</td>
<td align="char" valign="middle" char=".">26.63</td>
<td align="left" valign="middle">Baojing county</td>
</tr>
<tr>
<td align="left" valign="middle">2050s to 2090s</td>
<td align="left" valign="middle">Southeast</td>
<td align="char" valign="middle" char=".">32.11</td>
<td align="left" valign="middle">Luxi county</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
</sec>
<sec sec-type="discussion" id="sec12">
<label>4</label>
<title>Discussion</title>
<p>The MaxEnt model is widely used in species distribution modeling. In this study, MaxEnt version 3.4.4 was used in combination with 19 climatic variables and 3 topographic factors to model and predict species distribution under current and potential future climatic conditions. ArcGIS version 10.8 was then integrated to analyze the spatial distribution of <italic>I. cicadae</italic>. While some studies, such as those by <xref ref-type="bibr" rid="ref17">Kumar et al. (2014)</xref>, focus exclusively on climatic variables and omit topographic factors, this study adopts a combined approach that integrates both climatic and topographic variables for a more comprehensive analysis. The findings indicate that <italic>I. cicadae</italic> is less influenced by topographic variables and is more significantly affected by climatic factors. MaxEnt model predictions reveal a concentrated distribution of suitable habitats for <italic>I. cicadae</italic>, predominantly in Southwestern, Southern, Central, and Eastern China, with highly suitable habitats concentrated in provinces surrounding the Yangtze River basin. In future climate scenarios, the range of suitable habitats for <italic>I. cicadae</italic> shows slight expansion, with an increase in areas classified as low and moderate suitability, while the extent of high suitability decreases. A trend toward habitat expansion at higher latitudes is observed, while some low-latitude regions experience a reduction in suitability or even a loss of suitable habitats. Currently, Yunnan Province hosts the largest high-suitability area, but future projections suggest a decrease in the high-suitability area within Yunnan, accompanied by an expansion of the moderate-suitability region. Analysis of <xref ref-type="fig" rid="fig4">Figure 4</xref> and <xref ref-type="table" rid="tab4">Table 4</xref> indicates that, over time, the high-suitability area for <italic>I. cicadae</italic> is increasing and extending northward, remaining concentrated in Yunnan, Sichuan, Jiangxi, Hubei, Hunan, Chongqing, and Guizhou provinces.</p>
<p>The MaxEnt model, utilizing a combination of climate, topographic, and <italic>I. cicadae</italic> distribution data, was employed to assess habitat suitability. The model&#x2019;s AUC value is 0.984, which provides robust support for the study. This study employed Pearson correlation coefficients to select seven pivotal environmental variables (bio02, bio03, bio08, bio09, bio16, elevation, and slope) as significant influencers on the distribution of <italic>I. cicadae</italic>. <xref ref-type="bibr" rid="ref16">Jiaojiao et al. (2018)</xref> demonstrated that the suitable habitat of <italic>I. cicadae</italic> is influenced by factors such as temperature and light. In parallel, <xref ref-type="bibr" rid="ref46">Zhu et al. (2017)</xref> investigated the relationship between Cordycipitaceae biomass and climatic conditions, revealing that the yield of Cordycipitaceae is, to some extent, influenced by precipitation levels. The results of this study further support this conclusion. In their phylogenetic study of parasitic fungi, <xref ref-type="bibr" rid="ref9">Gorczak and Trigos-Peral (2021)</xref> observed that infection typically occurs during the warm, humid season when temperatures are favorable. Infection happens when older larvae active in shallow soil layers come into contact with soil contaminated with fungal spores, leading to the development of mature forms. Additionally, the jackknife test was utilized to assess the significance of these seven environmental variables in influencing the distribution of <italic>I. cicadae</italic>. In ranking the significance of the seven key environmental variables, the mean temperature during the wettest quarter (bio08), the rainfall during the wettest season (bio16), and the mean temperature of the driest quarter (bio09) occupied the top three positions, with their importance increasing sequentially. This alignment with the findings of the three preceding researchers suggests the reliability of our results, indirectly validating their credibility.</p>
<p>Based on the overlap analysis of the potential distributions of the hosts and <italic>I. cicadae</italic>, the potential distribution of <italic>I. cicadae</italic> largely overlaps with that of its primary hosts, with the hosts&#x2019; suitable areas covering 78.41% of <italic>I. cicadae&#x2019;s</italic> suitable areas, and vice versa, 99.13%. This provides strong validation of the model&#x2019;s predictive accuracy. The smaller suitable area for the hosts compared to <italic>I. cicadae</italic> is attributed to the presence of other hosts (<xref ref-type="bibr" rid="ref40">Xu et al., 2025</xref>). These results emphasize the potential impact of climate change on species distribution, particularly the shared adaptability of hosts and parasitic species, which could lead to species expansion or contraction in specific regions in the future (<xref ref-type="bibr" rid="ref38">Wei et al., 2023</xref>).</p>
<p>Under the three classic climate scenarios of SSP126, SSP370, and SSP585, the area of suitable habitats is projected to increase by the 2050s and 2070s. The predictive results indicate that high-suitability areas will continue to be concentrated in regions such as Guizhou, Hunan, and Jiangxi. However, there is a decrease in the size of these high-suitability areas, a trend that is particularly pronounced in the 2050s under the SSP126 scenario. Comparing the predicted results across different climate scenarios, it is observed that under the SSP126 scenario, the increase in suitable areas is less pronounced in the 2050s than under the other two scenarios. From 2050 to 2070, the increase in suitable habitat area for <italic>I. cicadae</italic> is more pronounced than in other scenarios, suggesting a potential enhancement in the species&#x2019; adaptability. Currently, the center of mass for the species is situated in Jishou City, Hunan Province (110&#x00B0;0&#x2032;27&#x2033; E, 28&#x00B0;20&#x2032;45&#x2033; N). Projections suggest that, except for the 2050s under the SSP585 and SSP126 scenarios, the center of mass for <italic>I. cicadae</italic> will remain within the coordinates of 109&#x00B0;48&#x2032;37&#x2033; E to 110&#x00B0;3&#x2032;6&#x2033; E and 28&#x00B0;14&#x2032;52&#x2033; N to 28&#x00B0;24&#x2032;59&#x2033; N, covering the areas of Jishou and Luxi counties. The negligible displacement of the center of mass suggests that there is no significant directional change in habitat suitability for the species.</p>
<p>When employing species distribution models for predicting potential distributions, it is common practice to consider the complex interplay of multiple biotic factors and environmental variables that influence species distribution (<xref ref-type="bibr" rid="ref24">Maurya et al., 2023</xref>). The MaxEnt model used in this study integrates data on the known distribution of the target species with environmental variables to estimate the probability of species distribution. While MaxEnt demonstrates strong predictive capabilities, it has notable limitations, including the restriction to temperature, precipitation, and topography as environmental variables, which may overlook other potential determinants. Despite its widespread use for predicting species distributions, MaxEnt has inherent limitations (<xref ref-type="bibr" rid="ref41">Xu et al., 2019</xref>). The response curve illustrates the impact of a single environmental factor without considering the complex interactions among multiple variables. Given the practical limitations of incorporating all environmental factors comprehensively in model development and analysis, it is more appropriate to use this model as a foundational niche model to accurately capture ecological principles (<xref ref-type="bibr" rid="ref21">Lin et al., 2022</xref>; <xref ref-type="bibr" rid="ref39">Weiyao et al., 2019</xref>). As a fungal species, the survival of <italic>I. cicadae</italic> is influenced not only by its biological traits but also by environmental factors such as host distribution, light conditions, and human activities (<xref ref-type="bibr" rid="ref12">He et al., 2024</xref>; <xref ref-type="bibr" rid="ref28">Shi et al., 2015</xref>). However, this study has certain limitations, particularly regarding the potential impact of anthropogenic factors and soil variables on species distribution. Future research should incorporate additional factors, particularly biotic factors such as host types and abiotic factors such as human activities, to provide a more comprehensive analysis and improve the accuracy of the model&#x2019;s predictions.</p>
</sec>
<sec sec-type="conclusions" id="sec13">
<label>5</label>
<title>Conclusion</title>
<p>In this study, distribution data for <italic>I. cicadae</italic> were integrated with three topographic and 19 bioclimatic variables using the MaxEnt model. This integration successfully modeled the potential geographic distributions of <italic>I. cicadae</italic> under three carbon emission scenarios (SSP126, SSP370, and SSP585) for both current conditions and projections for the 2050s and 2070s. Under current climatic conditions, outside of the Xinjiang Uyghur Autonomous Region, Qinghai Province, Ningxia Hui Autonomous Region, Inner Mongolia Autonomous Region, Heilongjiang Province, and Jilin Province, the species&#x2019; suitable areas extend across the remaining regions of China. Among these areas, Yunnan, Sichuan, Hunan, Hubei, Guizhou, Jiangxi, Guangxi, Fujian, Anhui, and Zhejiang are identified as having high suitability for <italic>I. cicadae</italic>. The most primary factor influencing the species&#x2019; distribution is temperature, followed by precipitation, with key variables including the mean temperature of the driest quarter (bio9), the mean air temperature of the wettest quarter (bio08), and the total precipitation during the wettest season (bio16). This study aims to further elucidate the distribution patterns and environmental influences on <italic>I. cicadae</italic> from a geographical perspective, enhancing its potential as an agricultural crop and providing theoretical insights for future cultivation and conservation strategies.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="sec14">
<title>Data availability statement</title>
<p>The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found in the article/supplementary material.</p>
</sec>
<sec sec-type="author-contributions" id="sec15">
<title>Author contributions</title>
<p>ZH: Formal analysis, Methodology, Software, Writing &#x2013; original draft. HA: Supervision, Writing &#x2013; review &#x0026; editing. JW: Data curation, Methodology, Writing &#x2013; review &#x0026; editing. ZL: Data curation, Writing &#x2013; review &#x0026; editing. XW: Data curation, Investigation, Writing &#x2013; review &#x0026; editing. ZZ: Conceptualization, Supervision, Writing &#x2013; review &#x0026; editing.</p>
</sec>
<sec sec-type="funding-information" id="sec16">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. This research was supported by the Fundamental Research Funds of China West Normal University (20A007, 20E051, 21E040 and 22kA011).</p>
</sec>
<sec sec-type="COI-statement" id="sec17">
<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 sec-type="ai-statement" id="sec18">
<title>Generative AI statement</title>
<p>The authors declare that no Gen AI was used in the creation of this manuscript.</p>
</sec>
<sec sec-type="disclaimer" id="sec19">
<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>
<fn-group>
<fn id="fn0001"><p><sup>1</sup><ext-link xlink:href="https://www.gbif.org/" ext-link-type="uri">https://www.gbif.org/</ext-link></p></fn>
<fn id="fn0002"><p><sup>2</sup><ext-link xlink:href="http://www.earthol.com/" ext-link-type="uri">http://www.earthol.com/</ext-link></p></fn>
<fn id="fn0003"><p><sup>3</sup><ext-link xlink:href="http://www.worldclim.org/" ext-link-type="uri">http://www.worldclim.org/</ext-link></p></fn>
</fn-group>
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