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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
<issn pub-type="epub">2296-701X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2023.1225466</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Ecology and Evolution</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Effects of clearcutting on species composition and community renewal of <italic>Rhododendron</italic> shrubs in northwest Guizhou Province, China</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Zhang</surname><given-names>Yaoyao</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2311836"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Zhao</surname><given-names>Xuechun</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="author-notes" rid="fn001"><sup>*</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2326505"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname><given-names>Lingjun</given-names>
</name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname><given-names>Zhaoyi</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Shuai</surname><given-names>Honggang</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wang</surname><given-names>Yuefeng</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2302468"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Jin</surname><given-names>Baocheng</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2143298"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Chen</surname><given-names>Chao</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="author-notes" rid="fn001"><sup>*</sup></xref>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup><institution>College of Animal Science, Guizhou University</institution>, <addr-line>Guiyang</addr-line>, <country>China</country></aff>
<aff id="aff2"><sup>2</sup><institution>Guizhou Provincial Key Laboratory for Biodiversity Conservation and Utilization in the Fanjing Mountain Region, Tongren University</institution>, <addr-line>Tongren</addr-line>, <country>China</country></aff>
<aff id="aff3"><sup>3</sup><institution>Institute of Azalea, Baili Dujuan Management Committee</institution>, <addr-line>Bijie</addr-line>, <country>China</country></aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Weiguo Sang, Minzu University of China, China</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Suresh Chandra Subedi, Arkansas Tech University, United States; Yongji Wang, Shanxi Normal University, China</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Xuechun Zhao, <email xlink:href="mailto:xczhao@gzu.edu.cn">xczhao@gzu.edu.cn</email>; Chao Chen, <email xlink:href="mailto:chenc@gzu.edu.cn">chenc@gzu.edu.cn</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>31</day>
<month>08</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>11</volume>
<elocation-id>1225466</elocation-id>
<history>
<date date-type="received">
<day>19</day>
<month>05</month>
<year>2023</year>
</date>
<date date-type="accepted">
<day>17</day>
<month>08</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2023 Zhang, Zhao, Wang, Wang, Shuai, Wang, Jin and Chen</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Zhang, Zhao, Wang, Wang, Shuai, Wang, Jin and Chen</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>Tree base sprouting is the main reproduction and expansion mode of <italic>Rhododendron</italic> plants. By leveraging the plot survey method, the species composition, community renewal, and species diversity in three <italic>Rhododendron</italic> shrub communities in control, and before and after clearcutting (CK, D<sub>3yr</sub>, and D<sub>6yr</sub>, respectively) were studied. Moreover, the dissimilarity of <italic>Rhododendron</italic> communities in CK, D<sub>3yr</sub> and D<sub>6yr</sub> were analyzed. The results showed that there were 26 plant species belonging to 14 families and 22 genera, in 3 communities in total, with 19 species of shrub plants and 7 species of herbaceous plants. The number of species increased from 13 in CK to 23 in D<sub>3yr</sub> and then decreased to 20 in D<sub>6yr</sub>. The height and coverage of D<sub>3yr</sub> and D<sub>6yr</sub> reached 39.3% and 58.9% of that of CK, respectively. The relative height of CK, D<sub>3yr</sub>, and D<sub>6yr</sub> was 43.79%, 65.4%, and 58.54%, respectively. The coverage of D<sub>3yr</sub> and D<sub>6yr</sub> reached 60.8% and 114.70% of that of CK, respectively. The relative coverage of CK, D<sub>3yr</sub>, and D<sub>6yr</sub> was 19.05%, 83.24%, and 77.32%, respectively. The important value of <italic>Rhododendron</italic> plants in the communities increased from 0.42 in CK to 0.74 in D<sub>3yr</sub>, and then decreased to 0.67 in D<sub>6yr</sub>. The &#x3b1; diversity in the shrub layer of D<sub>3yr</sub> and D<sub>6yr</sub> were generally lower than those of CK except Pielou evenness index. The &#x3b2; diversity indicates that the similarity between CK and D<sub>3yr</sub> was lower, that between CK and D<sub>6yr</sub> was moderate, and that between D<sub>3yr</sub> and D<sub>6yr</sub> was higher. The sprouting height and coverage of <italic>Rhododendron</italic> plants was significantly correlated with age and sprouting time. The sprouting ability of <italic>Rhododendron</italic> plants increased first and then decreased with age, while the sprouting ability of <italic>Rhododendron</italic> plants with age of 10&#x2013;12 years was the strongest. Clearcutting measures can improve the dominance of <italic>Rhododendron</italic> plants in the communities, promote the sprouting and renewal of <italic>Rhododendron</italic> population, and accelerate the succession rate of communities.</p>
</abstract>
<kwd-group>
<kwd><italic>Rhododendron</italic> shrub</kwd>
<kwd>clearcutting</kwd>
<kwd>species composition</kwd>
<kwd>species diversity</kwd>
<kwd>community renewal</kwd>
</kwd-group>
<contract-num rid="cn001">QKHPTRC-CXTD[2022]011, &#x94b1;&#x79d1;&#x4f55;&#x5fd7;&#x6210;[2020]1Y076, &#x94b1;&#x53ef;&#x4f55;&#x5fd7;&#x6210;[2021]&#x6613;&#x73ed;503</contract-num>
<contract-sponsor id="cn001">Guizhou Science and Technology Department<named-content content-type="fundref-id">10.13039/501100004001</named-content>
</contract-sponsor>
<counts>
<fig-count count="4"/>
<table-count count="5"/>
<equation-count count="10"/>
<ref-count count="45"/>
<page-count count="10"/>
<word-count count="4670"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Conservation and Restoration 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><italic>Rhododendron</italic> is one of the largest genera of angiosperms with more than 1200 species worldwide, which have important ecological and socio-economic applications of ornamental, cultural, scientific, economic, and medicinal value (<xref ref-type="bibr" rid="B31">MacKay and Gardiner, 2017</xref>; <xref ref-type="bibr" rid="B22">Li et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B1">Ahmad et&#xa0;al., 2021</xref>). The genus is mostly distributed in subtropical evergreen broad-leaved mountain forests, mixed coniferous broad-leaved forests, coniferous forests, and dark coniferous forests at altitudes varying from 1000 to 3800&#xa0;m (<xref ref-type="bibr" rid="B11">Fang and Min, 1995</xref>). It only forms a single species of <italic>Rhododendron</italic> shrub or <italic>Rhododendron</italic> dwarf forest in certain high mountains above the tree line (<xref ref-type="bibr" rid="B36">Sun, 2002</xref>). Guizhou Province is located at the edge of the distribution center of modern <italic>Rhododendron</italic> (Hengduan Mountain region in China) and its transition zone along the eastern distribution range (<xref ref-type="bibr" rid="B10">Dai et&#xa0;al., 2020</xref>). More than 110 species of <italic>Rhododendron</italic> plants are distributed naturally in Guizhou Province (<xref ref-type="bibr" rid="B10">Dai et&#xa0;al., 2020</xref>). Northwest Guizhou Province is a vital area for the distribution of <italic>Rhododendron</italic>, with six subgenera and more than fifty species accounting for approximately 50% of the total <italic>Rhododendron</italic> population in Guizhou. The Baili Rhododendron National Nature Reserve at the junction of Qianxi and Dafang counties is the most representative, with 43 species of <italic>Rhododendron</italic> plants (<xref ref-type="bibr" rid="B40">Wang et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B6">Chen et&#xa0;al., 2013</xref>). It is a typical representative of the largest and contiguously distributed natural wild <italic>Rhododendron</italic> communities in the world (<xref ref-type="bibr" rid="B32">Rong et&#xa0;al., 2009</xref>) which may play an important role in understanding <italic>Rhododendron</italic> shrub ecosystem processes, succession, and shrub management. However, both the quantity of seedlings and the renewal rate of <italic>Rhododendron</italic> plants are low (<xref ref-type="bibr" rid="B43">Yang et&#xa0;al., 2020a</xref>). Meanwhile, it is easy for <italic>Rhododendron</italic> plants to be replaced by highly competitive trees and other shrubs because of succession (<xref ref-type="bibr" rid="B3">Bian et&#xa0;al., 2006</xref>). Coupled with global changes in recent years, <italic>Rhododendron</italic> plants, particularly some narrowly distributed <italic>Rhododendron</italic> species, have begun to decline or even die (<xref ref-type="bibr" rid="B29">Ma et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B25">Liu et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B44">Yu et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B4">Bitayan et&#xa0;al., 2021</xref>).</p>
<p>Sprouting renewal primarily refers to dry base sprouting and underground stem sprouting (<xref ref-type="bibr" rid="B27">Lu et&#xa0;al., 2021</xref>). New plants are formed <italic>via</italic> dormant or adventitious buds from underground stems and stubble of trees to achieve forest renewal (<xref ref-type="bibr" rid="B39">Vesk and Westoby, 2004</xref>). Sprouting plants can use nutrients in the soil more effectively through their strong root system, which usually grows faster than seedlings and has stronger adaptability (<xref ref-type="bibr" rid="B17">Kauffman, 1991</xref>; <xref ref-type="bibr" rid="B7">Chen et&#xa0;al., 2019</xref>). Owing to the weak competitiveness and high canopy density of <italic>Rhododendron</italic> shrub communities in northwest Guizhou, seed renewal of the <italic>Rhododendron</italic> population is difficult, and tree-based sprouting renewal is the main mode of reproduction and expansion (<xref ref-type="bibr" rid="B18">Kong et&#xa0;al., 2019</xref>). At present, the primary measures to promote tree-based sprouting renewal are controlled forest fires, clearcutting and other natural disasters such as hurricane, mudslides (<xref ref-type="bibr" rid="B20">Li, 1992</xref>; <xref ref-type="bibr" rid="B28">Luoga et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B35">Subedi et&#xa0;al., 2019</xref>). However, forest fires are harmful as these reduce the sprouting ability of shrub communities (<xref ref-type="bibr" rid="B37">Tang et&#xa0;al., 2001</xref>). In contrast, planned clearcutting promotes the growth of shrub tillering branches, regenerates and rejuvenates plant clusters, and increases the canopy width and biomass (<xref ref-type="bibr" rid="B33">Shang et&#xa0;al., 2020</xref>). Following clearcutting, sprouting plants usually grow faster than seedlings (<xref ref-type="bibr" rid="B39">Vesk and Westoby, 2004</xref>).</p>
<p>Research on sprouting renewal of <italic>Rhododendron</italic> plants in global started late at present; further, research on sprouting and stress tolerance of wild <italic>Rhododendron</italic> plants is limited, decreasing the promotion and utilization of wild <italic>Rhododendron</italic> plants resources (<xref ref-type="bibr" rid="B30">MacKay and Gardiner, 2016</xref>). Although several studies have reported renewal of <italic>Rhododendron</italic> shrubs by disturbance, altitude, and climate changes (<xref ref-type="bibr" rid="B34">Singh et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B9">Choudhary et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B14">Jia et&#xa0;al., 2021</xref>), there is limited knowledge regarding the sprouting renewal of <italic>Rhododendron</italic> plants by clearcutting. In this study, the effects of rational clearcutting on the sprouting renewal of <italic>Rhododendron</italic> communities were studied to determine the effect of clearcutting&#x2013;sprouting on the role of <italic>Rhododendron</italic> plants in <italic>Rhododendron</italic> shrub communities. The objectives were to (1) Can clearcutting promote the community renewal of <italic>Rhododendron</italic> shrubs? (2) Does age affect the response of <italic>Rhododendron</italic> shrubs to clearcutting? (3) Can the sprouting and regeneration of <italic>Rhododendron</italic> be predicted after clearcutting?</p>
</sec>
<sec id="s2" sec-type="material|methods">
<label>2</label>
<title>Material and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Overview of the study area</title>
<p>The study area is in the Tiaohuapo Scenic Spot in the Baili Rhododendron National Nature Reserve, Guizhou Province, China (27&#xb0;23&#x2032;22&#x2032;&#x2032;N and 105&#xb0;51&#x2032;52&#x2032;&#x2032;E), near the Jiaozi Mountain, at an altitude of 1700&#x2013;1900 m. The climate of the region can be characterized as subtropical humid monsoon climate, with an annual average temperature and precipitation of 11.8&#xb0;C, and 1150.4&#xa0;mm, respectively, a frost-free period of 257 days, and an annual sunshine duration of 1335.5&#xa0;h. Zonal vegetation is dominated by the evergreen broad-leaved mountain forest. The existing vegetation is primarily <italic>Rhododendron</italic> shrub species exhibiting succession and transitional characteristics (<xref ref-type="bibr" rid="B21">Li and Chen, 2005</xref>; <xref ref-type="bibr" rid="B15">Jiang et&#xa0;al., 2015</xref>). Among <italic>Rhododendron</italic> shrubs, <italic>R. delavayi</italic>, <italic>R. annae</italic>, and <italic>Lyonia ovalifolia</italic> are the main dominant species. The soil type is primarily yellow soil with an acidic pH (4.61&#x2013;5.32).</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Plot setting and survey</title>
<p>In August 2015, three 20&#xa0;m &#xd7; 20&#xa0;m fixed quadrats were set in the study area to represent the control community (CK), and three 5&#xa0;m &#xd7; 5&#xa0;m survey quadrats were set diagonally in each quadrat. The species, plant number, branch number, height, and canopy width of the shrubs as well as the species and number of the herbaceous plants in the quadrats were recorded. Each <italic>Rhododendron</italic> shrub in the fixed quadrat was labelled.</p>
<p>In conjunction with the guidelines of Management Office of the Baili Rhododendron National Nature Reserve, all plants in the fixed quadrat were deforested and recovered. The shrubs in the fixed quadrat were deforested in December 2015 and the stubble height was recorded as approximately 15&#xa0;cm. The number of stubbles was also recorded. The age of shrubs was determined based on the rings of the main pile of shrubs. All other plants were deforested and eliminated.</p>
<p>The same method was used to survey and record the sprouting number, height, and canopy width of <italic>Rhododendron</italic> shrubs in August 2018 and August 2021. Moreover, the species number, height, and canopy width of other shrubs, as well as the species and number of herbaceous plants in the quadrats, were also surveyed. The communities surveyed in 2018 and 2021 were denoted as D<sub>3yr</sub> (Community 3 years after clearcutting) and D<sub>6yr</sub> (Community 6 years after clearcutting), respectively.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Data analyses</title>
<sec id="s2_3_1">
<label>2.3.1</label>
<title>Importance value</title>
<p>IV of the shrub layer for plant species in the three annual communities (CK, D<sub>3yr</sub>, and D<sub>6yr</sub>) was calculated using the equation (<xref ref-type="bibr" rid="B41">Wang et&#xa0;al., 2023</xref>)</p>
<disp-formula>
<label>(1)</label>
<mml:math display="block" id="M1">
<mml:mrow>
<mml:mi>I</mml:mi>
<mml:mi>V</mml:mi>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>R</mml:mi>
<mml:mi>H</mml:mi>
<mml:mo>+</mml:mo>
<mml:mi>R</mml:mi>
<mml:mi>C</mml:mi>
<mml:mo>+</mml:mo>
<mml:mi>R</mml:mi>
<mml:mi>D</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo stretchy="false">/</mml:mo>
<mml:mn>3</mml:mn>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>RH</italic> denotes the relative height, <italic>RC</italic> denotes the relative coverage, and <italic>RD</italic> denotes the relative density.</p>
</sec>
<sec id="s2_3_2">
<label>2.3.2</label>
<title>&#x3b1;-Diversity</title>
<p>The Margalef richness index (<italic>R</italic>), Simpson diversity index (<italic>D</italic>), Shannon&#x2013;Wiener diversity index (<italic>H</italic>), and Pielou evenness index (<italic>J</italic>) were determined to measure the &#x3b1;-diversity of the three communities using the equations (<xref ref-type="bibr" rid="B45">Zhang et&#xa0;al., 2017</xref>)</p>
<disp-formula>
<label>(2)</label>
<mml:math display="block" id="M2">
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>S</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo stretchy="false">)</mml:mo>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>l</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>N</mml:mi>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(3)</label>
<mml:math display="block" id="M3">
<mml:mrow>
<mml:mi>D</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
<mml:mo>&#x2212;</mml:mo>
<mml:mstyle displaystyle="true">
<mml:msubsup>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>S</mml:mi>
</mml:msubsup>
<mml:mrow>
<mml:msubsup>
<mml:mi>P</mml:mi>
<mml:mi>i</mml:mi>
<mml:mn>2</mml:mn>
</mml:msubsup>
</mml:mrow>
</mml:mstyle>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(4)</label>
<mml:math display="block" id="M4">
<mml:mrow>
<mml:mi>H</mml:mi>
<mml:mo>=</mml:mo>
<mml:mo>&#x2212;</mml:mo>
<mml:mstyle displaystyle="true">
<mml:msubsup>
<mml:mo>&#x2211;</mml:mo>
<mml:mrow>
<mml:mi>i</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
<mml:mi>S</mml:mi>
</mml:msubsup>
<mml:mrow>
<mml:msub>
<mml:mi>P</mml:mi>
<mml:mi>I</mml:mi>
</mml:msub>
<mml:mtext>&#x2009;</mml:mtext>
<mml:mi>l</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>I</mml:mi>
</mml:mrow>
</mml:mstyle>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(5)</label>
<mml:math display="block" id="M5">
<mml:mrow>
<mml:mi>J</mml:mi>
<mml:mo>=</mml:mo>
<mml:mi>H</mml:mi>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>l</mml:mi>
<mml:mi>n</mml:mi>
<mml:mi>S</mml:mi>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>P<sub>i</sub>
</italic> denotes the relative IV of the i<sup>th</sup> species, <italic>S</italic> denotes the number of species, and <italic>N</italic> denotes the number of individual plants in the quadrat.</p>
</sec>
<sec id="s2_3_3">
<label>2.3.3</label>
<title>&#x3b2;-Diversity</title>
<p>The Cody index (<italic>&#x3b2;<sub>c</sub>
</italic>), Whittaker index (<italic>&#x3b2;<sub>ws</sub>
</italic>), Jaccard index (<italic>C<sub>j</sub>
</italic>), and Sorenson index (<italic>C<sub>s</sub>
</italic>) were determined to measure the &#x3b2;-diversity of the three communities before and after clearcutting using the equations (<xref ref-type="bibr" rid="B2">Baselga, 2010</xref>)</p>
<disp-formula>
<label>(6)</label>
<mml:math display="block" id="M6">
<mml:mrow>
<mml:msub>
<mml:mi>&#x3b2;</mml:mi>
<mml:mi>c</mml:mi>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mo stretchy="false">[</mml:mo>
<mml:mi>g</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>H</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo>+</mml:mo>
<mml:mi>L</mml:mi>
<mml:mrow>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>H</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
<mml:mo stretchy="false">]</mml:mo>
<mml:mo stretchy="false">/</mml:mo>
<mml:mn>2</mml:mn>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(7)</label>
<mml:math display="block" id="M7">
<mml:mrow>
<mml:msub>
<mml:mi>&#x3b2;</mml:mi>
<mml:mrow>
<mml:mi>w</mml:mi>
<mml:mi>s</mml:mi>
</mml:mrow>
</mml:msub>
<mml:mo>=</mml:mo>
<mml:mi>m</mml:mi>
<mml:mi>s</mml:mi>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>m</mml:mi>
<mml:mi>a</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mn>1</mml:mn>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(8)</label>
<mml:math display="block" id="M8">
<mml:mrow>
<mml:mi>C</mml:mi>
<mml:mi>j</mml:mi>
<mml:mo>=</mml:mo>
<mml:mi>j</mml:mi>
<mml:mo stretchy="false">/</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>a</mml:mi>
<mml:mo>+</mml:mo>
<mml:mi>b</mml:mi>
<mml:mo>&#x2212;</mml:mo>
<mml:mi>j</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<disp-formula>
<label>(9)</label>
<mml:math display="block" id="M9">
<mml:mrow>
<mml:mi>C</mml:mi>
<mml:mi>s</mml:mi>
<mml:mo>=</mml:mo>
<mml:mn>2</mml:mn>
<mml:mi>j</mml:mi>
<mml:mo stretchy="false">/</mml:mo>
<mml:mo stretchy="false">(</mml:mo>
<mml:mi>a</mml:mi>
<mml:mo>+</mml:mo>
<mml:mi>b</mml:mi>
<mml:mo stretchy="false">)</mml:mo>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>&#x3b2;<sub>c</sub>
</italic> and <italic>&#x3b2;<sub>ws</sub>
</italic> denote the dissimilarity measures, <italic>C<sub>j</sub>
</italic> and <italic>C<sub>s</sub>
</italic> denote the similarity measures, <italic>g</italic>(<italic>H</italic>) denotes the number of species increased along the time gradient <italic>H</italic>, <italic>L</italic>(<italic>H</italic>) denotes the number of plant species existing in the previous community but lost in the next community, <italic>ms</italic> denotes the total number of species recorded in the three communities, <italic>ma</italic> denotes the average number of species in each community, <italic>a</italic> denotes the number of species in community A, <italic>b</italic> denotes the number of species in community B, and <italic>j</italic> denotes the number of species shared by communities A and B.</p>
</sec>
<sec id="s2_3_4">
<label>2.3.4</label>
<title>Sprouting status</title>
<p>The ratio between the number of <italic>Rhododendron</italic> sprouts and stumps after clearcutting was determined to measure the ability of <italic>Rhododendron</italic> plant to rejuvenate and renew using the equation</p>
<disp-formula>
<label>(10)</label>
<mml:math display="block" id="M10">
<mml:mrow>
<mml:mi>R</mml:mi>
<mml:mi>S</mml:mi>
<mml:mo>=</mml:mo>
<mml:mi>E</mml:mi>
<mml:mi>S</mml:mi>
<mml:mo stretchy="false">/</mml:mo>
<mml:mi>N</mml:mi>
<mml:mi>S</mml:mi>
</mml:mrow>
</mml:math>
</disp-formula>
<p>where <italic>RS</italic> denotes sprouting ability, <italic>ES</italic> denotes the number of sprouts, and <italic>NS</italic> denotes the number of stubbles.</p>
</sec>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Data processing and statistical analyses</title>
<p>SPSS 26.0 software was used to measure the correlation (<italic>Pearson correlation</italic>) and linear regression analysis between the height and coverage as well as clearcutting time and age of <italic>Rhododendron</italic>. Analysis of variance was performed (<italic>p</italic> &lt; 0.05 indicates significant differences) to determine the difference in IV of community species and biodiversity indices between the different stages of <italic>Rhododendron</italic> shrubs. All plots were prepared using SigmaPlot 14.0.</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Effects of clearcutting on <italic>Rhododendron</italic> shrub community characteristics</title>
<p>The three communities comprised 26 species, including nineteen species of shrubs and seven species of herbaceous plants, belonging to 14 families and 22 genera (<xref ref-type="table" rid="T1"><bold>Tables&#xa0;1</bold></xref>, <xref ref-type="table" rid="T2"><bold>2</bold></xref>). Ericaceae, Rosaceae, and Theaceae accounted for the highest number of species, specifically 19.23%, 15.38%, and 11.54% of the total, respectively. Three species of <italic>Rhododendron</italic> plants accounted for 11.54% of the total species.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Effects of clearcutting on species composition of <italic>Rhododendron</italic> shrub communities.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" rowspan="2" align="left">Layer</th>
<th valign="middle" rowspan="2" align="center">Family</th>
<th valign="middle" rowspan="2" align="center">Species name</th>
<th valign="middle" colspan="3" align="center">Community</th>
</tr>    <tr>
<th valign="middle" align="center">CK</th>
<th valign="middle" align="center">D<sub>3yr</sub>
</th>
<th valign="middle" align="center">D<sub>6yr</sub>
</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">Shrub</td>
<td valign="middle" align="center">Ericaceae</td>
<td valign="middle" align="center"><italic>R. delavayi</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>R. annae</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>R. lilacinum</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>Lyonia ovalifolia</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>Pieris japonica</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Rosaceae</td>
<td valign="middle" align="center"><italic>Rubus trianthus</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>R. corchorifolius</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>Cotoneaster horizontalis</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>Pyracantha fortuneana</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Lauraceae</td>
<td valign="middle" align="center"><italic>Litsea pungens</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>Machilus pingii</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">NA</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Theaceae</td>
<td valign="middle" align="center"><italic>Eurya japonica</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>E. loquaiana</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>Camellia oleifera</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Salicaceae</td>
<td valign="middle" align="center"><italic>Populus adenopoda</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>Salix matsudana</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Pinaceae</td>
<td valign="middle" align="center"><italic>Pinus armandii</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">NA</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Fagaceae</td>
<td valign="middle" align="center"><italic>Castanea seguinii</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Smilacaceae</td>
<td valign="middle" align="center"><italic>Smilax china</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="center">Herbaceous</td>
<td valign="middle" align="center">Poaceae</td>
<td valign="middle" align="center"><italic>Miscanthus floridulus</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center"/>
<td valign="middle" align="center"><italic>Fargesia spathacea</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Thelypteridaceae</td>
<td valign="middle" align="center"><italic>Macrothelypteris oligophlebia</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">NA</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Rosaceae</td>
<td valign="middle" align="center"><italic>Duchesnea indica</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Cyperaceae</td>
<td valign="middle" align="center"><italic>Cyperus rotundus</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">NA</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Juncaceae</td>
<td valign="middle" align="center"><italic>Juncus effusus</italic>
</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">NA</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="center">Asteraceae</td>
<td valign="middle" align="center"><italic>Elephantopus scaber</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">NA</td>
</tr>
<tr>
<td valign="middle" align="left">Total</td>
<td valign="middle" align="center">14</td>
<td valign="middle" align="center">26</td>
<td valign="middle" align="center">13</td>
<td valign="middle" align="center">23</td>
<td valign="middle" align="center">20</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>&#x201c;NA&#x201d; indicates that the species did not appear in the community, and &#x201c;1&#x201d; indicates that the species appeared in the community. CK, Community before clearcutting; D<sub>3yr</sub>, Community 3 years after clearcutting; D<sub>6yr</sub>, Community 6 years after clearcutting.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Effects of clearcutting on importance values (IVs) of species in the shrub layer of <italic>Rhododendron</italic> shrub communities.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" rowspan="2" align="center">Species name</th>
<th valign="middle" colspan="3" align="center">IV</th>
</tr>    <tr>
<th valign="middle" align="center">CK</th>
<th valign="middle" align="center">D<sub>3yr</sub>
</th>
<th valign="middle" align="center">D<sub>6yr</sub>
</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center"><italic>R. delavayi</italic>
</td>
<td valign="middle" align="center">0.31</td>
<td valign="middle" align="center">0.53</td>
<td valign="middle" align="center">0.46</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>R. annae</italic>
</td>
<td valign="middle" align="center">0.12</td>
<td valign="middle" align="center">0.19</td>
<td valign="middle" align="center">0.18</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>R. lilacinum</italic>
</td>
<td valign="middle" align="center">0.02</td>
<td valign="middle" align="center">0.02</td>
<td valign="middle" align="center">0.02</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Lyonia ovalifolia</italic>
</td>
<td valign="middle" align="center">0.17</td>
<td valign="middle" align="center">0.04</td>
<td valign="middle" align="center">0.07</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Pieris japonica</italic>
</td>
<td valign="middle" align="center">0.12</td>
<td valign="middle" align="center">0.02</td>
<td valign="middle" align="center">0.03</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Rubus trianthus</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.03</td>
<td valign="middle" align="center">0.04</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>R. corchorifolius</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.01</td>
<td valign="middle" align="center">0.01</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Cotoneaster horizontalis</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.03</td>
<td valign="middle" align="center">0.04</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Pyracantha fortuneana</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.00</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Litsea pungens</italic>
</td>
<td valign="middle" align="center">0.12</td>
<td valign="middle" align="center">0.01</td>
<td valign="middle" align="center">0.02</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Machilus pingii</italic>
</td>
<td valign="middle" align="center">0.09</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">NA</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Eurya japonica</italic>
</td>
<td valign="middle" align="center">0.04</td>
<td valign="middle" align="center">0.04</td>
<td valign="middle" align="center">0.05</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>E. loquaiana</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.02</td>
<td valign="middle" align="center">0.01</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Camellia japonica</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.00</td>
<td valign="middle" align="center">0.01</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Populus adenopoda</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.01</td>
<td valign="middle" align="center">0.01</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Salix matsudana</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.03</td>
<td valign="middle" align="center">0.03</td>
</tr>
<tr>
<td valign="middle" align="center"><italic>Pinus armandii</italic>
</td>
<td valign="middle" align="center">0.01</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">NA</td>
</tr>
<tr>
<td valign="top" align="center"><italic>Castanea seguinii</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.01</td>
<td valign="middle" align="center">0.01</td>
</tr>
<tr>
<td valign="top" align="center"><italic>Smilax china</italic>
</td>
<td valign="middle" align="center">NA</td>
<td valign="middle" align="center">0.01</td>
<td valign="middle" align="center">0.01</td>
</tr>
<tr>
<td valign="top" align="center">Total: 19</td>
<td valign="middle" align="center">9</td>
<td valign="middle" align="center">16</td>
<td valign="middle" align="center">17</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>&#x201c;NA&#x201d; indicates that the species did not appear in the community. CK, Community before clearcutting; D<sub>3yr</sub>, Community 3 years after clearcutting; D<sub>6yr</sub>, Community 6 years after clearcutting.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>CK comprised 13 species belonging to seven families (including Ericaceae, Lauraceae, Theaceae, and Poaceae) and 11 genera. Four species were identified in the herbaceous layer and nine in the shrub layer of which five viz., <italic>R. delavayi</italic>, <italic>L. ovalifolia</italic>, <italic>R. annae</italic>, and <italic>P. japonica</italic> of Ericaceae and <italic>L. pungens</italic> of Lauraceae exhibited IVs &gt; 0.1 (0.31, 0.17, 0.12, 0.12, and 0.12, respectively). <italic>R. delavayi</italic> was the dominant species in the shrub layer of CK, and <italic>L. ovalifolia</italic> was the subdominant species.</p>
<p>D<sub>3yr</sub> comprised the highest number of species, with 23 species from 12 families (including Ericaceae, Rosaceae, Theaceae, and Poaceae) and 19 genera. There were seven species in the herbaceous layer and 16 in the shrub layer of which two, <italic>R. delavayi</italic> and <italic>R. annae</italic> of Ericaceae, exhibited IVs of &gt; 0.1 (0.53 and 0.19, respectively). <italic>R. delavayi</italic> was the dominant species in the shrub layer of D<sub>3yr</sub>, and <italic>R. annae</italic> was the subdominant species.</p>
<p>D<sub>6yr</sub> comprised 20 species belonging to nine families (including Ericaceae, Rosaceae, Theaceae, and Poaceae) and 16 genera. There were three species in the herbaceous layer and seventeen in the shrub layer. Consistent with D<sub>3yr</sub>, two species in the shrub layer, <italic>R. delavayi</italic> and <italic>R. annae</italic>, exhibited IVs &gt; 0.1 (0.46 and 0.18, respectively). <italic>R. delavayi</italic> was the dominant species in the shrub layer of D<sub>6yr</sub>, and <italic>R. annae</italic> was the subdominant species.</p>
<p>In the CK&#x2192;D<sub>3yr</sub>&#x2192;D<sub>6yr</sub> community sequence, the height and coverage of <italic>Rhododendron</italic> shrubs first decreased and then increased (<xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>). The height decreased significantly from 191.87&#xa0;cm in CK to 75.61&#xa0;cm in D<sub>3yr</sub> and then recovered to 116.71&#xa0;cm in D<sub>6yr</sub>. The coverage decreased significantly from 68.46% in CK to 37.89% in D<sub>3yr</sub> and then increased significantly to 73.28% in D<sub>6yr</sub>. The relative height increased from 43.79% in CK to 65.40% in D<sub>3yr</sub> and then decreased to 58.54% in D<sub>6yr</sub>. The relative coverage increased from 49.05% in CK to 83.24% in D<sub>3yr</sub> and then decreased to 77.32% in D<sub>6yr</sub>. The height of <italic>Rhododendron</italic> species in the three communities were significantly different (<italic>p</italic> &lt; 0.05). The coverage in D<sub>3yr</sub> was significantly different from that in CK and D<sub>6yr</sub> (<italic>p&lt;</italic> 0.05). The relative height and coverage in CK were both significantly different from those in D<sub>3yr</sub> and D<sub>6yr</sub> (<italic>p</italic> &lt; 0.05).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Effects of clearcutting on the height and coverage of <italic>Rhododendron</italic> shrubs. Height <bold>(A)</bold>; Relative height <bold>(B)</bold>; Coverage <bold>(C)</bold>; Relative coverage <bold>(D)</bold>. <bold>(A&#x2013;C)</bold> indicate significant difference (<italic>p&lt;</italic> 0.05). CK, control; D<sub>3yr</sub>, community in 2018; D<sub>6yr</sub>, community in 2021.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-11-1225466-g001.tif"/>
</fig>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Effects of clearcutting on the diversity of species in <italic>Rhododendron</italic> shrub communities</title>
<p>The Margalef richness index values of species in the shrub layers of the three communities were significantly different (<italic>p &lt;</italic> 0.05) (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref>). However, the differences in the Simpson diversity, Shannon&#x2013;Wiener diversity, and Pielou evenness indices were not significant (<italic>p</italic> &gt; 0.05). The values of all four indices were the highest in CK and lowest in D<sub>3yr</sub>. The values of the Margalef richness, Simpson diversity, Shannon&#x2013;Wiener diversity, and Pielou evenness indices of species in D<sub>3yr</sub> were 61.15%, 80.30%, 84.79%, and 96.61% of that of species in CK, respectively.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Effects of clearcutting on &#x3b1;-diversity of <italic>Rhododendron</italic> shrub communities. Margalef richness index <bold>(A)</bold>; Simpson diversity index <bold>(B)</bold>; Shannon&#x2013;Wiener diversity index <bold>(C)</bold>; Pielou evenness index <bold>(D)</bold>. <bold>(A&#x2013;C)</bold> significant difference (<italic>p&lt;</italic> 0.05). CK, control; D<sub>3yr</sub>, community in 2018; D<sub>6yr</sub>, community in 2021.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-11-1225466-g002.tif"/>
</fig>
<p>As CK&#x2192;D<sub>3yr</sub>&#x2192;D<sub>6yr</sub> progressed, the dissimilarity coefficients of the three communities showed a decreasing trend (<xref ref-type="table" rid="T3"><bold>Table&#xa0;3</bold></xref>). Both Cody and Whittaker indices were the highest in CK and D<sub>3yr</sub> (3.83 and 1.65, respectively) and the lowest in D<sub>3yr</sub> and D<sub>6yr</sub> (0.83 and 1.20, respectively). The similarity indices, the Jaccard and Sorenson indices, were the highest in D<sub>3yr</sub> and D<sub>6yr</sub> (0.87 and 0.93, respectively) and the lowest in CK and D<sub>3yr</sub> (0.38 and 0.55, respectively). The similarity between CK and D<sub>3yr</sub> was low, that between CK and D<sub>6yr</sub> was moderate, and that between D<sub>3yr</sub> and D<sub>6yr</sub> was high.</p>
<table-wrap id="T3" position="float">
<label>Table&#xa0;3</label>
<caption>
<p>Effects of clearcutting on the &#x3b2;-diversity index of species in <italic>Rhododendron</italic> shrub communities.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Index</th>
<th valign="middle" align="left">Community</th>
<th valign="middle" align="left">CK</th>
<th valign="middle" align="left">D<sub>3yr</sub>
</th>
<th valign="middle" align="left">D<sub>6yr</sub>
</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">&#x3b2;<sub>c</sub>
</td>
<td valign="middle" align="left">CK</td>
<td valign="middle" align="left">0.00</td>
<td valign="middle" align="left">3.83</td>
<td valign="middle" align="left">3.50</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">D<sub>3yr</sub>
</td>
<td valign="middle" align="left">3.83</td>
<td valign="middle" align="left">0.00</td>
<td valign="middle" align="left">0.83</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">D<sub>6yr</sub>
</td>
<td valign="middle" align="left">3.50</td>
<td valign="middle" align="left">0.83</td>
<td valign="middle" align="left">0.00</td>
</tr>
<tr>
<td valign="middle" align="left">&#x3b2;<sub>ws</sub>
</td>
<td valign="middle" align="left">CK</td>
<td valign="middle" align="left">0.00</td>
<td valign="middle" align="left">1.65</td>
<td valign="middle" align="left">1.55</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">D<sub>3yr</sub>
</td>
<td valign="middle" align="left">1.65</td>
<td valign="middle" align="left">0.00</td>
<td valign="middle" align="left">1.20</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">D<sub>6yr</sub>
</td>
<td valign="middle" align="left">1.55</td>
<td valign="middle" align="left">1.20</td>
<td valign="middle" align="left">0.00</td>
</tr>
<tr>
<td valign="middle" align="left">C<sub>j</sub>
</td>
<td valign="middle" align="left">CK</td>
<td valign="middle" align="left">1.00</td>
<td valign="middle" align="left">0.38</td>
<td valign="middle" align="left">0.45</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">D<sub>3yr</sub>
</td>
<td valign="middle" align="left">0.38</td>
<td valign="middle" align="left">1.00</td>
<td valign="middle" align="left">0.87</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">D<sub>6yr</sub>
</td>
<td valign="middle" align="left">0.45</td>
<td valign="middle" align="left">0.87</td>
<td valign="middle" align="left">1.00</td>
</tr>
<tr>
<td valign="middle" align="left">C<sub>s</sub>
</td>
<td valign="middle" align="left">CK</td>
<td valign="middle" align="left">1.00</td>
<td valign="middle" align="left">0.55</td>
<td valign="middle" align="left">0.62</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">D<sub>3yr</sub>
</td>
<td valign="middle" align="left">0.55</td>
<td valign="middle" align="left">1.00</td>
<td valign="middle" align="left">0.93</td>
</tr>
<tr>
<td valign="middle" align="left"/>
<td valign="middle" align="left">D<sub>6yr</sub>
</td>
<td valign="middle" align="left">0.62</td>
<td valign="middle" align="left">0.93</td>
<td valign="middle" align="left">1.00</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>&#x3b2;<sub>c</sub> denotes the Cody index, &#x3b2;<sub>ws</sub> denotes the Whittaker index, C<sub>j</sub> denotes the Jaccard index, and C<sub>s</sub> denotes the Sorensen index. CK, Community before clearcutting; D<sub>3yr</sub>, Community 3 years after clearcutting; D<sub>6yr</sub>, Community 6 years after clearcutting.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Effects of clearcutting on the renewal of <italic>Rhododendron</italic> shrub communities</title>
<p>The height and coverage as well as the sprouting time and age of <italic>Rhododendron</italic> shrubs were significantly and positively correlated (<italic>p</italic> &lt; 0.01). The correlation degree of sprouting time was higher than that of the age of <italic>Rhododendron</italic> shrubs (<xref ref-type="table" rid="T4"><bold>Table&#xa0;4</bold></xref>).</p>
<table-wrap id="T4" position="float">
<label>Table&#xa0;4</label>
<caption>
<p>Correlation between sprouting height and coverage and time and age of <italic>Rhododendron</italic> shrubs.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left"/>
<th valign="top" align="center">Height</th>
<th valign="top" align="center">Coverage</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">Time</td>
<td valign="top" align="center">0.69**</td>
<td valign="top" align="center">0.62**</td>
</tr>
<tr>
<td valign="middle" align="center">Age</td>
<td valign="top" align="center">0.42**</td>
<td valign="top" align="center">0.37**</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>** indicates significant correlation at the 0.01 level.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>The sprouting time (t) and age (a) are the two main factors that affect clearcutting sprouting of <italic>Rhododendron</italic> shrubs. The linear relationships between sprouting height (h), sprouting coverage (c), sprouting time (t), and age (a) of <italic>Rhododendron</italic> shrubs after clearcutting were well fitted (<italic>R</italic>&#xb2; = 0.65, <italic>p</italic> &lt; 0.01 and <italic>R</italic>&#xb2; = 0.50, <italic>p</italic> &lt; 0.01, respectively) (<xref ref-type="table" rid="T5"><bold>Table&#xa0;5</bold></xref>).</p>
<table-wrap id="T5" position="float">
<label>Table&#xa0;5</label>
<caption>
<p>Regression model of sprouting height, sprouting coverage, and sprouting time of <italic>Rhododendron</italic> shrubs.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left"/>
<th valign="top" align="center">Regression equation</th>
<th valign="top" align="center"><italic>R<sup>2</sup>
</italic>
</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">Height</td>
<td valign="top" align="center"><italic>h</italic> = 13.55<italic>t</italic> + 8.15<italic>a</italic> &#x2212; 44.60</td>
<td valign="top" align="center">0.65**</td>
</tr>
<tr>
<td valign="middle" align="center">Coverage</td>
<td valign="top" align="center"><italic>c</italic> = 0.60<italic>t</italic> + 0.37<italic>a</italic> &#x2212; 3.43</td>
<td valign="top" align="center">0.50**</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>** indicates significant correlation at the 0.01 level. h, sprouting height; t, sprouting time; a, age; and c, sprouting coverage.</p>
</fn>
</table-wrap-foot>
</table-wrap>
<p>The height of <italic>Rhododendron</italic> shrubs in CK, D<sub>3yr</sub>, and D<sub>6yr</sub> showed a gradual increasing trend with age (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3</bold></xref>). Compared with CK, the height of <italic>Rhododendron</italic> shrubs in D<sub>3yr</sub> and D<sub>6yr</sub> was lesser and increased slowly. The coverage rates of <italic>Rhododendron</italic> shrubs in the three communities also increased gradually with increasing age. The coverage rates of <italic>Rhododendron</italic> shrubs in D<sub>3yr</sub> and D<sub>6yr</sub> at the early stage were higher than those in CK.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Variation in height and coverage of <italic>Rhododendron</italic> shrubs with age. Height of CK community <bold>(A)</bold>; Coverage of CK community <bold>(B)</bold>; Height of D<sub>3yr</sub> community <bold>(C)</bold>; Coverage of D<sub>3yr</sub> community <bold>(D)</bold>; Height of D<sub>6yr</sub> community <bold>(E)</bold>; Coverage of D<sub>6yr</sub> community <bold>(F)</bold>. CK, control; D<sub>3yr</sub>, community in 2018; D<sub>6yr</sub>, community in 2021.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-11-1225466-g003.tif"/>
</fig>
<p>To determine the sprouting ability at different ages, the age of <italic>Rhododendron</italic> shrubs was divided into three classes (I, II, and III). The sprouting ability of <italic>Rhododendron</italic> shrubs first increased and then decreased with age (<xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4</bold></xref>). Class II <italic>Rhododendron</italic> shrubs had the strongest sprouting ability. Moreover, the sprouting ability of class III <italic>Rhododendron</italic> shrubs was significantly different from that of classes I and II shrubs (<italic>p</italic> &lt; 0.05).</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Effect of age of <italic>Rhododendron</italic> shrubs on sprouting ability. I, 7&#x2013;9 years; II, 10&#x2013;12 years; and III, &gt; 12 years. a and b denote significant difference (<italic>p &lt;</italic> 0.05).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-11-1225466-g004.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>Effects of clearcutting on species composition and community characteristics of <italic>Rhododendron</italic> shrubs</title>
<p>Species composition reflects the structure, dynamic changes, and succession characteristics of a community and plays a decisive role in community diversity (<xref ref-type="bibr" rid="B12">Gilliam, 2007</xref>; <xref ref-type="bibr" rid="B24">Liu et&#xa0;al., 2020</xref>). In this study, 26 species belonging to 14 families and 22 genera were identified in the three communities studied (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>). The number of species in CK was the lowest probably because the canopy density of <italic>Rhododendron</italic> shrubs was high and the litter layer was thick, which reduced the migration and colonisation of <italic>Rubus trianthus</italic>, <italic>Cotoneaster horizontalis</italic>, <italic>Eurya loquaiana</italic>, <italic>Populus adenopoda</italic>, belonging to the Rosaceae, Theaceae, Salicaceae, as well as herbaceous plants. However, clearcutting weakened interspecies competition by reducing canopy density (<xref ref-type="bibr" rid="B16">Jones et&#xa0;al., 2019</xref>). Consequently, a large number of herbaceous plants, and species belonging to Rosaceae, Theaceae, and Salicaceae migrated; thus, D<sub>3yr</sub> had the highest number of species. Subsequently, <italic>Rhododendron</italic> shrubs gradually occupied the upper space, the canopy density increased, and herbaceous plants gradually disappeared. Therefore, the number of species in D<sub>6yr</sub> was lower compared with that in D<sub>3yr</sub>.</p>
<p>The height and coverage of plants in communities are important indicators for evaluating the plant growth status (<xref ref-type="bibr" rid="B5">Chen et&#xa0;al., 2014</xref>). In this study, the height and coverage of plants in communities after clearcutting were significantly higher than those of plants in the seed renewal communities (<xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>), primarily because the large number of sprouting branches produced by <italic>Rhododendron</italic> shrubs could occupy more space by utilizing the resources, hence promoting rapid formation of community hierarchy. Post clearcutting, the sprouting coverage of <italic>Rhododendron</italic> shrubs in D<sub>6yr</sub> was higher than that of shrubs in CK, primarily because clearcutting promoted shrub tillering and expanded individual canopy width. After clearcutting, the relative height of <italic>Rhododendron</italic> shrubs in D<sub>3yr</sub> and D<sub>6yr</sub> also increased significantly, probably because <italic>Rhododendron</italic> shrubs occupied the upper space of the whole shrub community after clearcutting, playing a dominant role in light absorption and utilization. The absence of other tall trees and shrub vegetation enabled <italic>Rhododendron</italic> plants roots to utilize soil nutrients more effectively. Therefore, clearcutting resulted in the domination of <italic>Rhododendron</italic> plants in the community, with vigorous population renewal and a stronger self-sustaining ability of the population.</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Effects of clearcutting on the diversity of <italic>Rhododendron</italic> shrub communities</title>
<p>As a basic feature of a community, biodiversity is a key factor driving the dynamics and processes of an ecosystem and can indicate the habitat status, composition structure, and distribution pattern of a community (<xref ref-type="bibr" rid="B23">Liu and Bra, 1998</xref>; <xref ref-type="bibr" rid="B38">Tilman et&#xa0;al., 2012</xref>). In this study, the evenness index of the communities changed after clearcutting (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref>). The index value was relatively large, indicating that the heterogeneity of the study area was poor. This was also determined by the characteristics of the sprouting branches of <italic>Rhododendron</italic> shrubs. Although clearcutting increased the number of species, the &#x3b1;-diversity was still lower than that in CK before clearcutting. This may be because <italic>Rhododendron</italic> plants has a strong sprouting ability. After clearcutting, <italic>Rhododendron</italic> plants is likely to occupy space in the form of sprouting branches to inhibit the migration and colonisation of foreign species (<xref ref-type="bibr" rid="B13">Grant and Loneragan, 1999</xref>; <xref ref-type="bibr" rid="B19">Kruger and Midgley, 2001</xref>). Therefore, <italic>Rhododendron</italic> plants with its strong sprouting abilities can reduce species diversity and species turnover in the community to a certain extent.</p>
<p>&#x3b2;-Diversity is usually expressed as the turnover rate of biological species in different habitats. A high &#x3b2;-diversity index indicates a low level of species similarity between different habitats or ecosystems (<xref ref-type="bibr" rid="B42">Yang et&#xa0;al., 2020b</xref>). The results showed that the similarity between CK and D<sub>3yr</sub> was low, that between CK and D<sub>6yr</sub> was moderate, and that between D<sub>3yr</sub> and D<sub>6yr</sub> was high (<xref ref-type="table" rid="T3"><bold>Table&#xa0;3</bold></xref>). The Cody and Whittaker indices were higher before and after clearcutting, whereas the Jaccard and Sorenson indices were lower, indicating that clearcutting changed the species composition and community structure of <italic>Rhododendron</italic>. The similarity between CK and D<sub>3yr</sub> was low but that between CK and D<sub>6yr</sub> was moderate, indicating that with the progression of CK&#x2192;D<sub>3yr</sub>&#x2192;D<sub>6yr</sub>, the number of common plant species first decreased and then increased. Subsequently, the similarity among species composition also first decreased and then increased, indicating that <italic>Rhododendron</italic> plants could recover to the pre-clearcutting community level in a short time through clearcutting&#x2013;sprouting renewal.</p>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Effects of clearcutting on the renewal of <italic>Rhododendron</italic> shrub communities</title>
<p>In <italic>Rhododendron</italic> shrub communities, both seedlings and sprouting seedlings can undergo population renewal. Nevertheless, sprouting seedlings are an important source of population dynamics (<xref ref-type="bibr" rid="B39">Vesk and Westoby, 2004</xref>). Due to the strong sprouting ability of <italic>Rhododendron</italic> plants, deforested communities rapidly self-substitute, which not only affects the community structure but also has an important impact on population dynamics. Clearcutting&#x2013;sprouting renewal avoids the problems of very low sprouting and survival rates as well as weak seedling competitiveness. The results of this study similarly showed that the dominance of the <italic>Rhododendron</italic> population in D<sub>3yr</sub> and D<sub>6yr</sub> after sprouting was greater than that in CK before clearcutting. The growth recovery of <italic>Rhododendron</italic> plants after clearcutting is completely dependent on the amount of stubble, root tiller, and rhizome, whereas the number of sprouts is a common index to characterize the strength of sprouting (<xref ref-type="bibr" rid="B8">Chi et&#xa0;al., 2019</xref>). Although the sprouting ability of <italic>Rhododendron</italic> plants was strong in this study, the recovery ability first increased and then decreased with the age of the <italic>Rhododendron</italic> shrubs. The sprouting ability of <italic>Rhododendron</italic> shrubs of age 10&#x2013;12 years was the strongest (<xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4</bold></xref>). This may be associated with the aging of plants, an excessively large root system, and a lack of rhizome growth, which led to partial necrosis of the underground root system.</p>
<p>The community succession of <italic>Rhododendron</italic> plants does not deviate far from the original direction because of its strong sprouting ability and the effect of the ecological niche after clearcutting (<xref ref-type="bibr" rid="B1">Ahmad et&#xa0;al., 2021</xref>). However, the communities in this study had a high number and proportion of <italic>Rhododendron</italic> shrubs initially, which were <italic>in situ</italic> substituted with a similarly high number, accelerating the recovery to the original structure and succession rate of the community. Therefore, the shrub layer of <italic>Rhododendron</italic> communities after clearcutting was primarily composed of sprouting branches of <italic>Rhododendron</italic>, and the final direction of succession was subtropical evergreen broad-leaved forest. Compared with seed renewal and succession, the clearcutting&#x2013;sprouting succession of <italic>Rhododendron</italic> plants takes a shorter time to achieve maturity (<xref ref-type="bibr" rid="B26">Loucks, 1970</xref>; <xref ref-type="bibr" rid="B7">Chen et&#xa0;al., 2019</xref>).</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusions</title>
<p>In total, 26 species from 22 genera and 14 families were found in the three communities. Clearcutting promoted the migration and colonisation of the families Rosaceae, Theaceae, and Salicaceae and other herbaceous plants, whereas the relative height and coverage of shrubs in the communities increased significantly. After clearcutting, there was a slight change in community evenness and a decrease in the &#x3b1;-diversity. The &#x3b2;-diversity showed that clearcutting improved the dominance of <italic>Rhododendron</italic> plants in the community and promoted sprouting renewal of <italic>Rhododendron</italic> populations. Moreover, the sprouting ability of <italic>Rhododendron</italic> shrubs of age 10&#x2013;12 years was the strongest. Clearcutting did not affect the direction of community succession, but could accelerate the succession rate. Owing to the short observation period of clearcutting and sprouting of <italic>Rhododendron</italic> communities in this study, it was not possible to accurately predict the rate of population renewal and community succession of <italic>Rhododendron</italic> shrubs. In subsequent studies, multiple clearcutting modes and <italic>Rhododendron</italic> population succession stages will be established in combination with the competitive ability of the <italic>Rhododendron</italic> population.</p>
</sec>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>YZ, XZ, and LW contributed to the conception and design of the study. HS confirmed and guided the study. ZW, BJ, and HS carried out the experiments. YW performed the statistical analysis and wrote the first draft of the manuscript. YZ, XZ, and CC guided writing sections of the manuscript. All authors contributed to manuscript revision, read, and approved the submitted version.</p>
</sec>
</body>
<back>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>This research was funded by the Science and Technology Department of Guizhou Province (Qian Ke He Zhicheng [2020]1Y076 and [2021]Yiban503, QKHPTRC-CXTD[2022]011).</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We acknowledge the Baili Rhododendron National Nature Reserve and laboratory of the Department of Grassland Science, Guizhou University. We thank Xin Liu, Guiying Liu, Yini Wang, and Xiaolong Tian for their valuable suggestions and help with the laboratory analysis.</p>
</ack>
<sec id="s9" 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="s10" 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>
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