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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">3</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:F7FCE910-8E78-573F-9C77-7788555F8AAD</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:66878968-5FA3-48E4-A0CA-743FBBB0315D</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">PhytoKeys</journal-title>
        <abbrev-journal-title xml:lang="en">PK</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1314-2011</issn>
      <issn pub-type="epub">1314-2003</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/phytokeys.279.206429</article-id>
      <article-id pub-id-type="publisher-id">206429</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Salicaceae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Cenozoic</subject>
          <subject>Mesozoic</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Asia</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Salicaceae">Salicaceae</tp:taxon-name-part></tp:taxon-name>), a new species from Guangxi, China</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Xu</surname>
            <given-names>Ting</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-9598-0387</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/visualization/">Visualization</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Ouyang</surname>
            <given-names>Zilong</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0005-0787-2615</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Su</surname>
            <given-names>Zhiyong</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0001-3180-4496</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Yang</surname>
            <given-names>Mei</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-6821-166X</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Jia</surname>
            <given-names>Xianglu</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0006-9143-8885</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Gong</surname>
            <given-names>Li</given-names>
          </name>
          <email xlink:type="simple">2007gongli@163.com</email>
          <uri content-type="orcid">https://orcid.org/0009-0009-3886-561X</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Nanning Botanical Garden, Nanning 530002, China</addr-line>
        <institution>Nanning Botanical Garden</institution>
        <addr-line content-type="city">Nanning</addr-line>
        <country>China</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Nanning Qingxiu Mountain Scenic Area Tourism Development Co., Ltd., Nanning 530004, China</addr-line>
        <institution>Nanning Qingxiu Mountain Scenic Area Tourism Development Co., Ltd.</institution>
        <addr-line content-type="city">Nanning</addr-line>
        <country>China</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Nanning Horticultural Expo Garden Administration Center, Nanning 530021, China</addr-line>
        <institution>Nanning Horticultural Expo Garden Administration Center</institution>
        <addr-line content-type="city">Nanning</addr-line>
        <country>China</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Li Gong (<email xlink:type="simple">2007gongli@163.com</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Hanno Schaefer</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>30</day>
        <month>09</month>
        <year>2026</year>
      </pub-date>
      <volume>279</volume>
      <fpage>267</fpage>
      <lpage>272</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/37534E3D-37DC-5154-8560-C60A69F780B2">37534E3D-37DC-5154-8560-C60A69F780B2</uri>
      <history>
        <date date-type="received">
          <day>06</day>
          <month>07</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>04</day>
          <month>09</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Ting Xu, Zilong Ouyang, Zhiyong Su, Mei Yang, Xianglu Jia, Li Gong</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <abstract>
        <label>Abstract</label>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. a new species from Guangxi, Southern China, is described and illustrated. The new species exhibits several distinguishing features: capsule 2.2–3 cm, spherical, leathery, orange-red at maturity, 4- or 5-valved, valves fleshy, 0.5 cm thick. Seeds up to 15–16 per fruit. The relationships among <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="membranacea">membranacea</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name></italic> are shortly discussed. The new species is endemic to limestone mountain forests of Western Guangxi, with its known distribution largely confined to the Guangxi Laohutiao Nature Reserve. On the basis of the IUCN criteria, it is assessed as Critically Endangered.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>
          <italic>
            <tp:taxon-name>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part>
            </tp:taxon-name>
          </italic>
        </kwd>
        <kwd>Guangxi</kwd>
        <kwd>morphology</kwd>
        <kwd>new species</kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="family" reg="Salicaceae">Salicaceae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>2024 Central Financial Forestry and Grassland Ecological Protection and Restoration Fund Project: In-situ Conservation and Propagation of Vatica guangxiensis (Notice No 72 issued by the Forestry Bureau of the Guangxi Zhuang Autonomous Region regarding forest protection in 2024) and the 2025 Central Financial Forestry and Grassland Ecological Protection and Restoration Fund Project: Collection and Preservation of Germplasm Resources of Nationally Key Protected Wild Plants in Guangxi (Notice No 2 issued by the Forestry Bureau of the Guangxi Zhuang Autonomous Region regarding forest protection in 2025</funding-statement>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec1">
      <title>Introduction</title>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic> Jacq. is a pantropical woody genus comprising approximately 200 species, with its greatest diversity occurring in the Neotropics (<xref ref-type="bibr" rid="B6">de Mestier et al. 2023</xref>). It is generally placed in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Salicaceae">Salicaceae</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B5">Chase et al. 2002</xref>; <xref ref-type="bibr" rid="B2">APG IV 2016</xref>), though <xref ref-type="bibr" rid="B1">Alford (2005)</xref> proposed a plausible classification that would segregate <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic> and about a dozen other genera of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Salicaceae">Salicaceae</tp:taxon-name-part></tp:taxon-name> into <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Samydaceae">Samydaceae</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B12">Philpott and Applequist 2023</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic> comprises seven species in China, these seven species are characterized by: flowers arranged in axillary glomerules; disk lobes in same row as stamens; style entire; capsule fleshy (<xref ref-type="bibr" rid="B8">Flora of China Editorial Committee 2007</xref>).</p>
      <p>Recent discoveries of new <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic> species have been largely concentrated in Southeast Africa (<xref ref-type="bibr" rid="B12">Philpott and Applequist 2023</xref>), Central America (<xref ref-type="bibr" rid="B6">de Mestier et al. 2023</xref>), South America (<xref ref-type="bibr" rid="B11">Marquete and Torres 2022</xref>) and West Africa (<xref ref-type="bibr" rid="B4">Breteler and Baldé 2024</xref>). Since the publication of the Flora of China, the taxonomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic> in China has remained static, with no new species having been published in the past 20 years. In this paper, we describe a new species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic> endemic to Southern China, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic> T. Xu, L. Gong, Z. L. Ouyang &amp; Z. Y. Su.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec2">
      <title>Materials and methods</title>
      <p>From 2024 to 2026, the research team conducted a total of 13 field surveys at the type locality. Morphological observations and measurements of the new species were carried out on both living plants in the field and herbarium specimens. Comparisons among the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic> species distributed in China were made using relevant literature and digital images of specimens housed at institutions such as AU, BJFC, CSFI, FJSI, GXMG, HITBC, HUH, IBK, IBSC, IMDY, KUN, NAS, P, PE, SWFC, SZG and WUK. In addition, type specimen images were consulted via JSTOR Global Plants. [<ext-link xlink:href="https://plants.jstor.org" ext-link-type="uri">https://plants.jstor.org</ext-link>] and Chinese Virtual Herbarium Website [<ext-link xlink:href="http://www.cvh.ac.cn/" ext-link-type="uri">http://www.cvh.ac.cn/</ext-link>]. The voucher specimens are deposited in the Herbarium of Nanning Botanical Garden (<bold><abbrev content-type="institution" xlink:title="Herbarium of Nanning Botanical Garden">NNBG</abbrev></bold>), the Herbarium of Chenshan Botanical Garden (<bold><named-content content-type="dwc:institutional_code" xlink:title="Shanghai Chenshan Botanical Garden" xlink:href="https://scientific-collections.gbif.org/institution/9c4e9ee7-0a1d-459d-bf05-cd4753142206">CSH</named-content></bold>) and the Herbarium of Guangxi Institute of Chinese Medicine &amp; Pharmaceutical Sciences (<bold><named-content content-type="dwc:institutional_code" xlink:title="Guangxi Institute of Traditional Medical and Pharmaceutical Sciences" xlink:href="https://scientific-collections.gbif.org/institution/26d59ced-676b-494e-b512-bd4537ff3276">GXMI</named-content></bold>). The Nanning Botanical Garden collected seeds and successfully raised seedlings from them for ex-situ conservation.</p>
    </sec>
    <sec sec-type="Taxonomic treatment" id="sec3">
      <title>Taxonomic treatment</title>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="kingdom" reg="Plantae">Plantae</tp:taxon-name-part>
                </tp:taxon-name>
              </named-content>
            </kwd>
            <kwd>
              <named-content content-type="order">Malpighiales</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family">Salicaceae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <tp:taxon-name><object-id content-type="arpha">94396E47-1B4E-5FCE-ADC4-8AB18324BFCF</object-id>
                		<tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part>
                	
                		<object-id content-type="ipni" xlink:type="simple">urn:lsid:ipni.org:names:77399863-1</object-id>
                	</tp:taxon-name>
          <tp:taxon-authority>T.Xu, L.Gong, Z.L.Ouyang &amp; Z.Y.Su</tp:taxon-authority>
          <tp:taxon-status>sp. nov.</tp:taxon-status>
          <xref ref-type="fig" rid="F1">Figs 1</xref>
          <xref ref-type="fig" rid="F2">, 2</xref>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="Type">
          <title>Type.</title>
          <p>China • Guangxi Zhuang Autonomous Region: Baise City, Napo County, Pingmeng Town, Nianjing Village (Guangxi Laohutiao Nature Reserve), <named-content content-type="dwc:verbatimCoordinates">22°58.97'N, 105°50.60'E</named-content>, 1200 m, a.s.l., 30 August 2024, <italic>XT et al. 20240830-01</italic>, • the same location, 12 May 2026, <italic>XT et al. 260512-07</italic> (holotype, <abbrev content-type="institution" xlink:title="Herbarium of Nanning Botanical Garden">NNBG</abbrev>!; isotype, <named-content content-type="dwc:institutional_code" xlink:title="Shanghai Chenshan Botanical Garden" xlink:href="https://scientific-collections.gbif.org/institution/9c4e9ee7-0a1d-459d-bf05-cd4753142206">CSH</named-content>!, <named-content content-type="dwc:institutional_code" xlink:title="Guangxi Institute of Traditional Medical and Pharmaceutical Sciences" xlink:href="https://scientific-collections.gbif.org/institution/26d59ced-676b-494e-b512-bd4537ff3276">GXMI</named-content>!, <abbrev content-type="institution" xlink:title="Herbarium of Nanning Botanical Garden">NNBG</abbrev>!).</p>
          <fig id="F1">
            <object-id content-type="doi">10.3897/phytokeys.279.206429.figure1</object-id>
            <object-id content-type="arpha">4BAB3514-D968-55DF-AB5E-899BCF029A83</object-id>
            <label>Figure 1.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic>. <bold>A</bold>. Habitat; <bold>B</bold>. Fruiting plant in habitat; <bold>C</bold>. Bark; <bold>D</bold>. Twig and stipules; <bold>E</bold>. Front side of the leaf; <bold>F</bold>. Back side of the leaf; <bold>G</bold>. Inflorescence; <bold>H</bold>. Flower, views from side; <bold>I</bold>, <bold>J</bold>. Flower, front view. Photo (<bold>G</bold>) by L. Gong; Photos (<bold>A–F</bold>, <bold>H–J</bold>) by T. Xu, photo correction and design by T. Xu. Scale bars: 5.0 cm (<bold>E, F</bold>); 5.0 mm (<bold>H, I, J</bold>).</p>
            </caption>
            <graphic xlink:href="phytokeys-279-267_article-206429__-g001.jpg" id="oo_1794070.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1794070</uri>
            </graphic>
          </fig>
          <fig id="F2">
            <object-id content-type="doi">10.3897/phytokeys.279.206429.figure2</object-id>
            <object-id content-type="arpha">26DA1B44-75EB-59C0-AFF8-A9318C6AA7A1</object-id>
            <label>Figure 2.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic>. <bold>A, B</bold>. Fruiting branch; <bold>C, D</bold>. Ripe fruit; <bold>E</bold>. Dehiscent capsule; <bold>F</bold>. Seeds. Photos (<bold>A</bold>, <bold>C</bold>, <bold>F</bold>) by L. Gong; Photos (<bold>B</bold>, <bold>D</bold>) by Z.L.Ouyang; Photo <bold>E</bold> by T. Xu, photo correction and design by T. Xu. Scale bars: 5.0 cm (<bold>E</bold>); 1.0 cm (<bold>C, F</bold>).</p>
            </caption>
            <graphic xlink:href="phytokeys-279-267_article-206429__-g002.jpg" id="oo_1794071.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1794071</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Diagnosis">
          <title>Diagnosis.</title>
          <p>Similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="membranacea">membranacea</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name></italic>, but differs mainly in the stamens 6–9, ca. 2–3 mm, filaments sparsely pubescent. Capsule 2.2–3 cm, spherical, leathery, orange-red at maturity, 4- or 5-valved, valves fleshy, 0.5 cm thick. Seeds up to 15–16 per fruit.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Vernacular name">
          <title>Vernacular name.</title>
          <p>Chinese: 那坡脚骨脆 (nà pō jiăo gŬ cuì).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Description">
          <title>Description.</title>
          <p>Trees or shrubs, 4–8 m tall; trunk straight; bark grayish white. <bold><italic>Young branches</italic></bold> longitudinally ridged, exfoliating in flakes, grayish-white, glabrous. <bold><italic>Stipules</italic></bold> narrowly triangular, ca. 3 mm, entire, caducous; leaves alternate, thinly leathery, obovate-oblong, 11–15(–16) × 3.5–5.6(–6) cm, apex acuminate, acumen ca. 1 cm, base cuneate, margin entire, occasionally with sparse small glandular teeth on the upper portion, adaxially bright green, abaxially pale green, drying blackish-brown, both surfaces glabrous; lateral veins 8–10 pairs, arching upward, prominent abaxially; petiole 0.7–1.5 cm, glabrous. <bold><italic>Flowers</italic></bold> small, bisexual, green, 3–4 clustered in leaf axils, without peduncles, with scale-like bracts at the base; flower diameter ca. 4 mm; <bold><italic>pedicels</italic></bold> slender, ca. 3 mm long, glabrous; <bold><italic>sepals</italic></bold> 5, ovate, ca. 4 mm long, 3 mm wide, margin membranous and ciliate; <bold><italic>petals</italic></bold> absent; stamens 6–9; <bold><italic>filaments</italic></bold> sparsely pubescent, ca. 2–3 mm; anthers oblong-ellipsoid; disk lobes oblong, flattened, densely pubescent. <bold><italic>Capsule</italic></bold> 2.2–3 cm, spherical, leathery, surface smooth, orange-red when ripe, 4- or 5-valved, valves fleshy, 0.5 cm thick, dried fruit severely shrunken, pericarp thin, dark brown, with 5 longitudinal ridges; <bold><italic>seeds</italic></bold> 15–16, white, pale brown when dry, droplet-shaped, 5–7 mm. Fl. Apr–Jun, fr. Aug–Nov.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Etymology">
          <title>Etymology.</title>
          <p>The specific epithet is derived from the county name of the type locality (Napo).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Distribution and habitat">
          <title>Distribution and habitat.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic> is only known from the type location, Guangxi, Southern China. Growing in the limestone mountainous area at an altitude of 1200 meters.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Conservation status">
          <title>Conservation status.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic> is endemic to the Guangxi Laohutiao Nature Reserve, with a restricted distribution. From May 2024 to May 2026, we conducted 13 field surveys in this area but did not discover any additional localities for the species, suggesting that its distribution is extremely limited. At present, seventeen individuals of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic> have been found, including two mature ones. According to the Guidelines for Using the IUCN Red List Categories and Criteria (<xref ref-type="bibr" rid="B9">IUCN Standards and Petitions Committee 2024</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic> should be assessed as Critically Endangered (<abbrev xlink:title="Critically Endangered">CR</abbrev>) based on criteria D1.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
    <sec sec-type="Discussion" id="sec4">
      <title>Discussion</title>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic> is distinguished by capsule 2.2–3 cm, spherical, leathery, orange-red at maturity, 4- or 5-valved. Among the recorded species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic>, this new species is similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="membranacea">membranacea</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name></italic>. But both species have 3-valved capsules, which distinguishes them from the new species. The main differences among the three species are summarized in Table <xref ref-type="table" rid="T1">1</xref>. In the protologue of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name></italic>, the description states: Ovarium conicum, 2 mm altum. Fructus carnosus ellipsoidalis, 3.5 × 2.5 mm, plus minusve verrucosus (<xref ref-type="bibr" rid="B10">Lescot and Sleumer 1970</xref>). This may be a typographical error; the fruit size should probably be 3.5 × 2.5 cm. The two species differ in whether the pericarp is verrucosus, allowing easy distinction from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic>. <xref ref-type="bibr" rid="B3">Bao (1983)</xref> published <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="calciphila">calciphila</tp:taxon-name-part></tp:taxon-name></italic> as a new species. This taxon was later synonymized under <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name></italic> in Flora of China (<xref ref-type="bibr" rid="B8">Flora of China Editorial Committee 2007</xref>); however, material of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name></italic> was not seen by the authors of that flora. According to the <xref ref-type="bibr" rid="B7">Editorial Committee of Flora Reipublicae Popularis Sinicae (1999)</xref>, the capsule of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="calciphila">calciphila</tp:taxon-name-part></tp:taxon-name></italic> is described as ellipsoid, ca. 1.2 cm long, which also serves to differentiate it from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic>.</p>
      <table-wrap id="T1" position="float" orientation="portrait">
        <label>Table 1.</label>
        <caption>
          <p>Morphological comparisons among <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="membranacea">membranacea</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name></italic>.</p>
        </caption>
        <table>
          <tbody>
            <tr>
              <th rowspan="1" colspan="1">
                <bold>Characters</bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="membranacea">membranacea</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </bold>
              </th>
              <th rowspan="1" colspan="1">
                <bold>
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </bold>
              </th>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Branches</td>
              <td rowspan="1" colspan="1">Young branches ridged, grayish-white, exfoliating in flakes,</td>
              <td rowspan="1" colspan="1">Young branches ridged, green</td>
              <td rowspan="1" colspan="1">Young branches wrinkled, brownish-gray, older ones flaky</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Stipules</td>
              <td rowspan="1" colspan="1">Narrowly triangular, ca. 3 mm</td>
              <td rowspan="1" colspan="1">Ovate, ca. 1 mm</td>
              <td rowspan="1" colspan="1">Triangular, ca. 2 mm</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Leaf blade</td>
              <td rowspan="1" colspan="1">Thinly leathery</td>
              <td rowspan="1" colspan="1">Papery, sometimes thinly so, or slightly leathery</td>
              <td rowspan="1" colspan="1">Leathery to thickly leathery</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Lateral veins</td>
              <td rowspan="1" colspan="1">8–10 pairs</td>
              <td rowspan="1" colspan="1">5–7(–10) pairs</td>
              <td rowspan="1" colspan="1">6–8 pairs</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Sepals</td>
              <td rowspan="1" colspan="1">5, ovate, 3 mm, glabrous</td>
              <td rowspan="1" colspan="1">5, broadly elliptic, obovate, or obovate-oblong, ca. 2.5 mm, outside rarely sparsely puberulous</td>
              <td rowspan="1" colspan="1">5, ovate, 3.5–4.5 mm, glabrous except for sparsely ciliate margin</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Stamens</td>
              <td rowspan="1" colspan="1">6–9; ca. 2–3 mm; filaments sparsely pubescent</td>
              <td rowspan="1" colspan="1">8–10; 1–2 mm; filaments usually glabrous, rarely puberulous</td>
              <td rowspan="1" colspan="1">8; ca. 2 mm; filaments sparsely pubescent, glabrescent or glabrous</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Disk</td>
              <td rowspan="1" colspan="1">Lobes oblong, flattened, densely pubescent</td>
              <td rowspan="1" colspan="1">Lobes oblong or triangular, adaxially glabrous, apex densely hairy, hairs white or reddish, long</td>
              <td rowspan="1" colspan="1">Lobes oblong, hairy at apex</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Capsule</td>
              <td rowspan="1" colspan="1">2.5–3 cm, spherical, leathery, pericarp smooth, 4- or 5-valved</td>
              <td rowspan="1" colspan="1">1–1.5(–4) cm, ellipsoid to oblong, leathery, 3-valved</td>
              <td rowspan="1" colspan="1">3.5 × 2.5 mm (may be a typographical error, correct: 3.5 × 2.5 cm), ellipsoid, fleshy, verrucose, 3-valved</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">Seeds</td>
              <td rowspan="1" colspan="1">15–16, 5–7 mm, red aril.</td>
              <td rowspan="1" colspan="1">Several, 4–5 mm, pale yellowish aril.</td>
              <td rowspan="1" colspan="1">Not documented</td>
            </tr>
          </tbody>
        </table>
      </table-wrap>
      <sec sec-type="Key to the Casearia species of China" id="sec5">
        <title>Key to the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">Casearia</tp:taxon-name-part></tp:taxon-name></italic> species of China</title>
        <table-wrap content-type="key" position="anchor" orientation="portrait">
          <table>
            <tbody>
              <tr>
                <td>1</td>
                <td>Stipules narrowly lanceolate or linear-lanceolate, 1.5–10 mm</td>
                <td>
                  <bold>2</bold>
                </td>
              </tr>
              <tr>
                <td>–</td>
                <td>Stipules broadly triangular or broadly ovate, minute, 1–2 mm</td>
                <td>
                  <bold>3</bold>
                </td>
              </tr>
              <tr>
                <td>2</td>
                <td>Stipules narrowly lanceolate, 5–10 mm, early caducous, on young growth leaving a large conspicuous pale brown scar; leaves with 10–14 pairs of lateral veins; in dried material pellucid dots and streaks throughout leaf blade usually clearly visible at × 10</td>
                <td>
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="graveolens">graveolens</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td>–</td>
                <td>Stipules linear-lanceolate, 1.5–3 mm, persistent for some time; leaves with 5–8 pairs of lateral veins; in dried material pellucid dots and streaks throughout leaf blade not clearly visible at × 10 mag. without holding leaf up to light, nor reddish brown</td>
                <td>
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="flexuosa">flexuosa</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td>3</td>
                <td>Abaxial surfaces of mature leaves pubescent, at least along midvein and lateral veins</td>
                <td>
                  <bold>4</bold>
                </td>
              </tr>
              <tr>
                <td>–</td>
                <td>Abaxial surfaces of mature leaves glabrous</td>
                <td>
                  <bold>5</bold>
                </td>
              </tr>
              <tr>
                <td>4</td>
                <td>Pedicels 2–4 mm in flower, 5–6 mm in fruit</td>
                <td>
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="velutina">velutina</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td>–</td>
                <td>Pedicels 5–8 mm in flower, ca. 10 mm in fruit</td>
                <td>
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kurzii">kurzii</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td>5</td>
                <td>Leaves leathery</td>
                <td>
                  <bold>6</bold>
                </td>
              </tr>
              <tr>
                <td>–</td>
                <td>Leaves papery to membranous</td>
                <td>
                  <bold>7</bold>
                </td>
              </tr>
              <tr>
                <td>6</td>
                <td>Capsule 2- or 3-valved</td>
                <td>
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tardieuae">tardieuae</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td>–</td>
                <td>Capsule 4- or 5-valved</td>
                <td><bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="napoensis">napoensis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov</bold>.</td>
              </tr>
              <tr>
                <td>7</td>
                <td>Terminal bud, pedicel, and calyx hairy, stamen filaments usually hairy (rarely nearly glabrous); capsule slightly to not at all ridged, pericarp veined, vesicled, vesicles black and shiny in cross-section</td>
                <td>
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="glomerata">glomerata</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td>–</td>
                <td>Terminal bud hairy or glabrous, pedicel, calyx, and stamen filaments usually glabrous (rarely puberulous); capsule usually strongly ridged, pericarp veined but not conspicuously vesicled</td>
                <td>
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Casearia">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="membranacea">membranacea</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>We are grateful to Dr. Dan Liang (Guangxi Institute of Botany) for providing critical photographic material and to the Herbarium of Guangxi Institute of Chinese Medicine &amp; Pharmaceutical Sciences (<named-content content-type="dwc:institutional_code" xlink:title="Guangxi Institute of Traditional Medical and Pharmaceutical Sciences" xlink:href="https://scientific-collections.gbif.org/institution/26d59ced-676b-494e-b512-bd4537ff3276">GXMI</named-content>) for granting access to specimen collections for examination in June 2026.</p>
    </ack>
    <ref-list>
      <title>References</title>
      <ref id="B1">
        <mixed-citation>Alford MH (2005) Systematic studies in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Flacourtiaceae</tp:taxon-name-part></tp:taxon-name>. PhD Thesis, Cornell University, Ithaca.</mixed-citation>
      </ref>
      <ref id="B2">
        <mixed-citation>APG IV [Angiosperm Phylogeny Group] (2016) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1–20. <ext-link xlink:href="10.1111/boj.12385" ext-link-type="doi">https://doi.org/10.1111/boj.12385</ext-link></mixed-citation>
      </ref>
      <ref id="B3">
        <mixed-citation>Bao SY (1983) Some new plants of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Casearia</tp:taxon-name-part></tp:taxon-name></italic> from Yunnan. Kunming Institute of Botany, Academia Sinica (04): 375–378.</mixed-citation>
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    <sec sec-type="Additional information" id="sec6">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec7">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec8">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec9">
        <title>Artificial Intelligence (AI) use</title>
        <p>The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.</p>
        <p>No AI tools were used in the preparation of this manuscript.</p>
      </sec>
      <sec sec-type="Funding" id="sec10">
        <title>Funding</title>
        <p>2024 Central Financial Forestry and Grassland Ecological Protection and Restoration Fund Project: In-situ Conservation and Propagation of Vatica guangxiensis (Notice No 72 issued by the Forestry Bureau of the Guangxi Zhuang Autonomous Region regarding forest protection in 2024) and the 2025 Central Financial Forestry and Grassland Ecological Protection and Restoration Fund Project: Collection and Preservation of Germplasm Resources of Nationally Key Protected Wild Plants in Guangxi (Notice No 2 issued by the Forestry Bureau of the Guangxi Zhuang Autonomous Region regarding forest protection in 2025.</p>
      </sec>
      <sec sec-type="Author contributions" id="sec11">
        <title>Author contributions</title>
        <p>Conceptualization: LG, TX. Data curation: TX, LG, ZS. Formal analysis: TX. Funding acquisition: OZ, LG. Investigation: TX, LG, ZO, ZS, MY, XJ. Methodology: LG, TX, OZ, MY. Project administration: OZ, LG. Resources: OZ, LG. Supervision: OZ, MY. Visualization: TX. Writing – original draft: TX. Writing – review and editing: LG, MY, ZO.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec12">
        <title>Author ORCIDs</title>
        <p>T. Xu <ext-link xlink:href="https://orcid.org/0000-0001-9598-0387" ext-link-type="uri">https://orcid.org/0000-0001-9598-0387</ext-link></p>
        <p>Z. Ouyang <ext-link xlink:href="https://orcid.org/0009-0005-0787-2615" ext-link-type="uri">https://orcid.org/0009-0005-0787-2615</ext-link></p>
        <p>Z. Su <ext-link xlink:href="https://orcid.org/0009-0001-3180-4496" ext-link-type="uri">https://orcid.org/0009-0001-3180-4496</ext-link></p>
        <p>M. Yang <ext-link xlink:href="https://orcid.org/0000-0002-6821-166X" ext-link-type="uri">https://orcid.org/0000-0002-6821-166X</ext-link></p>
        <p>X. Jia <ext-link xlink:href="https://orcid.org/0009-0006-9143-8885" ext-link-type="uri">https://orcid.org/0009-0006-9143-8885</ext-link></p>
        <p>L. Gong <ext-link xlink:href="https://orcid.org/0009-0009-3886-561X" ext-link-type="uri">https://orcid.org/0009-0009-3886-561X</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec13">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text.</p>
      </sec>
    </sec>
  </back>
</article>
