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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">14</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:69C4EDC7-0CE2-5AB0-81F2-EA3B3CC9D365</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:777322AB-37CC-4A41-B3C2-83A76AAD4F84</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Deutsche Entomologische Zeitschrift</journal-title>
        <abbrev-journal-title xml:lang="en">DEZ</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1435-1951</issn>
      <issn pub-type="epub">1860-1324</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/dez.72.170313</article-id>
      <article-id pub-id-type="publisher-id">170313</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Embiidina</subject>
          <subject>Embioptera</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Faunistics &amp; Distribution</subject>
          <subject>Phylogeny</subject>
          <subject>Systematics</subject>
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Asia</subject>
          <subject>China</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>﻿First record of the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Insecta</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Embioptera</tp:taxon-name-part></tp:taxon-name>) from China, with morphological and molecular characterization of a new species</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Chen</surname>
            <given-names>Zhi-Teng</given-names>
          </name>
          <email xlink:type="simple">chenzhiteng@just.edu.cn</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-6331-8978</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Xiong</surname>
            <given-names>Yuan</given-names>
          </name>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Jiang</surname>
            <given-names>Chao</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-1841-1169</uri>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">School of Grain Science and Technology, Jiangsu University of Science and Technology, Zhenjiang 212004, Jiangsu, China</addr-line>
        <institution>Jiangsu University of Science and Technology</institution>
        <addr-line content-type="city">Zhenjiang</addr-line>
        <country>China</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, National Resource Center for Chinese Materia Medica, China Academy of Chinese Medical Sciences, Beijing 100700, China</addr-line>
        <institution>China Academy of Chinese Medical Sciences</institution>
        <addr-line content-type="city">Beijing</addr-line>
        <country>China</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Zhi-Teng Chen (<email xlink:type="simple">chenzhiteng@just.edu.cn</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Susanne Randolf</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2025</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>03</day>
        <month>12</month>
        <year>2025</year>
      </pub-date>
      <volume>72</volume>
      <issue>2</issue>
      <fpage>569</fpage>
      <lpage>579</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/C37CB6CE-2326-5C00-B68C-8FE598FE39A5">C37CB6CE-2326-5C00-B68C-8FE598FE39A5</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/15E2CDF7-E1D2-4B8B-A974-E91EF38D510A">15E2CDF7-E1D2-4B8B-A974-E91EF38D510A</uri>
      <history>
        <date date-type="received">
          <day>28</day>
          <month>08</month>
          <year>2025</year>
        </date>
        <date date-type="accepted">
          <day>24</day>
          <month>11</month>
          <year>2025</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Zhi-Teng Chen, Yuan Xiong, Chao Jiang</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>
      <self-uri content-type="zoobank" xlink:type="simple">https://zoobank.org/15E2CDF7-E1D2-4B8B-A974-E91EF38D510A</self-uri>
      <abstract>
        <label>﻿Abstract</label>
        <p>We report the first record of the webspinner family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> Miller &amp; Edgerly, 2012, from China, represented by a new species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov.</bold>, discovered in Yunnan Province. Detailed morphological examination, particularly the comparative analysis of male terminalia, clearly distinguishes this species from its congeners. The complete mitochondrial genome of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov.</bold> (15,443 bp) was sequenced, exhibiting the canonical insect gene order with strong A+T bias but showing the absence of two tRNAs (trnV and trnS2). Notably, we detected novel tandem repeats between trnI and trnQ, a feature not reported from other sequenced embiopteran mitogenomes. All protein-coding genes evolve under purifying selection, with <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EEF">COX1</abbrev> showing the highest conservation. DNA barcoding (<abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EIF">COX1</abbrev>) and phylogenetic analyses further support species delimitation in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic>, with interspecific divergences between the new species and other <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> (12.6–17.6%) far exceeding frequently used intraspecific thresholds (~2%). The discovery extends the known distribution range of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> northeastward to southwestern China. Our integrative morphological and molecular framework highlights the previously underestimated species richness in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> and provides the first mitogenomic insights into <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name>.</p>
      </abstract>
      <kwd-group>
        <label>Key Words</label>
        <kwd>mitogenomics</kwd>
        <kwd>
          <italic>
            <tp:taxon-name>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part>
            </tp:taxon-name>
          </italic>
        </kwd>
        <kwd>species delimitation</kwd>
        <kwd>taxonomy</kwd>
        <kwd>webspinners</kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>CACMS Innovation Fund (grant no. CI2024E003)</funding-statement>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="﻿Introduction" id="SECID0E6G">
      <title>﻿Introduction</title>
      <p>The monogeneric webspinner family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> Miller &amp; Edgerly, 2012, was established based on the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> Friederichs, 1923, originally described from Indonesia (type species: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="roepkei">roepkei</tp:taxon-name-part></tp:taxon-name></italic> Friederichs, 1923) (<xref ref-type="bibr" rid="B18">Miller et al. 2012</xref>). Historically, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> was placed in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Notoligotomidae</tp:taxon-name-part></tp:taxon-name> together with the eastern Australian genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Notoligotoma">Notoligotoma</tp:taxon-name-part></tp:taxon-name></italic> Davis, 1936 (<xref ref-type="bibr" rid="B8">Davis 1936</xref>; <xref ref-type="bibr" rid="B26">Ross 1963</xref>), but molecular phylogenetic analysis recovered it as a distinct lineage (<xref ref-type="bibr" rid="B18">Miller et al. 2012</xref>). <xref ref-type="bibr" rid="B18">Miller et al. (2012)</xref> thus erected the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic>. Subsequently, <xref ref-type="bibr" rid="B22">Poolprasert and Edgerly (2014)</xref> provided morphological descriptions and biological and distributional information on four new species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> from Thailand, including <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic> Poolprasert &amp; Edgerly, 2014; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="rossi">rossi</tp:taxon-name-part></tp:taxon-name></italic> Poolprasert &amp; Edgerly, 2014; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senathami">senathami</tp:taxon-name-part></tp:taxon-name></italic> Poolprasert &amp; Edgerly, 2014; and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="thaidina">thaidina</tp:taxon-name-part></tp:taxon-name></italic> Poolprasert &amp; Edgerly, 2014. The males of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name>/<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> are characterized by antennae with long setae and white distal segments, distinct wing venation with the anterior media vein forked, densely setose hind basitarsi with a single papilla, an obliquely cleft tergite X, and elaborated male terminalia (e.g., left tergal process short, medial flap with a hooked prong, and fused left cercomeres) (<xref ref-type="bibr" rid="B18">Miller et al. 2012</xref>; <xref ref-type="bibr" rid="B22">Poolprasert and Edgerly 2014</xref>). Mitochondrial DNA barcoding (<abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0E2DAC">COX1</abbrev>) has also been shown to be effective for distinguishing <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> species (<xref ref-type="bibr" rid="B23">Poolprasert et al. 2017</xref>, <xref ref-type="bibr" rid="B24">2021</xref>).</p>
      <p>To date, all known <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> species are known from Indonesia, Myanmar, and Thailand (<xref ref-type="bibr" rid="B18">Miller et al. 2012</xref>; <xref ref-type="bibr" rid="B22">Poolprasert and Edgerly 2014</xref>). In this study, we report the first record of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> and the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> from China. We describe <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. from Yunnan Province, provide detailed morphological characters, and present its complete mitochondrial genome (mitogenome). In addition, we compare the <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EWFAC">COX1</abbrev> barcode of the new species with published <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> sequences to delimit species boundaries. This integrative approach yields novel data on the taxonomy, phylogeny, and biogeography of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name>.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0ENGAC">
      <title>﻿Material and methods</title>
      <sec sec-type="﻿Sample collection and morphological examination" id="SECID0ERGAC">
        <title>﻿Sample collection and morphological examination</title>
        <p>The specimens were manually collected from the road surface in Yingpan, Ailao Mountain, Yunnan Province, China (<named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[101.547600,23.990100]}" id="NCID0E1GAC">23.9901°N, 101.5476°E</named-content></named-content>; altitude 2000 m) on 23 May 2018. They were preserved in 75% ethanol. A morphological study was conducted under an SZM45 stereomicroscope. Images of habitus and structures were taken with a Canon EOS 5DSR digital camera and an MP-E 65 mm macro lens and processed with Adobe Photoshop v2025. The distribution map was produced using QGIS v3.40.11. Morphological terminology follows <xref ref-type="bibr" rid="B27">Ross (2007)</xref> and <xref ref-type="bibr" rid="B22">Poolprasert and Edgerly (2014)</xref>. The holotype and paratype are deposited in the Insect Collection of Jiangsu University of Science and Technology (<bold><abbrev content-type="institution" xlink:title="Insect Collection of Jiangsu University of Science and Technology" id="ABBRID0EIHAC">ICJUST</abbrev></bold>), Zhenjiang, China.</p>
      </sec>
      <sec sec-type="﻿Mitogenome sequencing and assembly" id="SECID0ENHAC">
        <title>﻿Mitogenome sequencing and assembly</title>
        <p>Genomic DNA was extracted from a single leg of the holotype (genseq-1) using the Wizard SV Genomic DNA Purification System Kit (Promega, USA). A paired-end Illumina library (~400 bp insert) was prepared and sequenced (2×150 bp) on the Illumina HiSeq X platform (Annoroad Gene Technology, China). Raw reads were quality-filtered (removing low-quality and short reads), yielding 1.56 Gb of clean data. The mitogenome was assembled independently using GETORGANELLE v1.7.4 and MITOZ v3.6 pipelines (<xref ref-type="bibr" rid="B17">Meng et al. 2019</xref>; <xref ref-type="bibr" rid="B15">Jin et al. 2020</xref>). The consistency of the two assemblies was confirmed by manual checking. The complete mitogenome sequence was deposited in GenBank with accession number <ext-link ext-link-type="gen" xlink:href="PX232380" xlink:type="simple">PX232380</ext-link>.</p>
      </sec>
      <sec sec-type="﻿Mitogenome annotation and analysis" id="SECID0EAIAC">
        <title>﻿Mitogenome annotation and analysis</title>
        <p>Annotation was performed with MITOS2 (<xref ref-type="bibr" rid="B2">Bernt et al. 2013</xref>) and by comparison to published embiopteran mitogenomes. Boundaries of protein-coding genes (<abbrev xlink:title="protein-coding genes" id="ABBRID0EKIAC">PCGs</abbrev>) were confirmed by ORF FINDER (NCBI). The circular mitogenome map was visualized using PROKSEE (<xref ref-type="bibr" rid="B10">Grant et al. 2023</xref>). We calculated nucleotide composition, codon usage, and relative synonymous codon usage (<abbrev xlink:title="relative synonymous codon usage" id="ABBRID0ESIAC">RSCU</abbrev>) with MEGA11 (<xref ref-type="bibr" rid="B28">Tamura et al. 2021</xref>). Pairwise genetic distances (K2P) for <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0E1IAC">COX1</abbrev> genes and synonymous (Ks) and nonsynonymous (Ka) substitution rates between genes of sequenced webspinners and the outgroup <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phasmotaenia">Phasmotaenia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lanyuhensis">lanyuhensis</tp:taxon-name-part></tp:taxon-name></italic> Huang &amp; Brock, 2001 (<xref ref-type="bibr" rid="B14">Huang and Brock 2001</xref>; GenBank No. <ext-link ext-link-type="gen" xlink:href="PP536087" xlink:type="simple">PP536087</ext-link>) were computed using MEGA11 and KAKS_CALCULATOR v3.0 (<xref ref-type="bibr" rid="B29">Zhang 2022</xref>), respectively. Tandem repeats in the mitogenome were identified using Tandem Repeats Finder (<xref ref-type="bibr" rid="B1">Benson 1999</xref>).</p>
      </sec>
      <sec sec-type="﻿Molecular species delimitation and phylogenetic analysis" id="SECID0E1JAC">
        <title>﻿Molecular species delimitation and phylogenetic analysis</title>
        <p>The <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EAKAC">COX1</abbrev> barcode was extracted from the newly sequenced mitogenome. We downloaded all six available <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic><abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0ELKAC">COX1</abbrev> sequences from GenBank (<xref ref-type="bibr" rid="B18">Miller et al. 2012</xref>; <xref ref-type="bibr" rid="B23">Poolprasert et al. 2017</xref>, <xref ref-type="bibr" rid="B24">2021</xref>) and aligned them with the new sequence. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Aposthonia">Aposthonia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="borneensis">borneensis</tp:taxon-name-part></tp:taxon-name></italic> (Hagen, 1885) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Oligotomidae</tp:taxon-name-part></tp:taxon-name>) was used as an outgroup. Genetic distances (K2P) and a Neighbor-Joining (<abbrev xlink:title="Neighbor-Joining" id="ABBRID0ELLAC">NJ</abbrev>) tree were generated in MEGA11 (<xref ref-type="bibr" rid="B28">Tamura et al. 2021</xref>) using pairwise deletion (gaps removed) and 1000 bootstrap replicates. Maximum likelihood (<abbrev xlink:title="Maximum likelihood" id="ABBRID0ETLAC">ML</abbrev>) and Bayesian inference (<abbrev xlink:title="Bayesian inference" id="ABBRID0EXLAC">BI</abbrev>) phylogenies were estimated based on the <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0E2LAC">COX1</abbrev> alignment. The best-fit nucleotide substitution model was selected by MODELFINDER (<xref ref-type="bibr" rid="B16">Kalyaanamoorthy et al. 2017</xref>). <abbrev xlink:title="Bayesian inference" id="ABBRID0EDMAC">BI</abbrev> analysis was run in MRBAYES v3.2.6 (<xref ref-type="bibr" rid="B25">Ronquist et al. 2012</xref>) under the GTR+G+F model (2 parallel runs, 5000000 generations), in which the initial 25% of sampled data were discarded as burn-in. <abbrev xlink:title="Maximum likelihood" id="ABBRID0ELMAC">ML</abbrev> analysis was performed in IQ-TREE v2.2.0 (<xref ref-type="bibr" rid="B21">Nguyen et al. 2015</xref>) under the TIM2+G4+F model for 1000 ultrafast (<xref ref-type="bibr" rid="B19">Minh et al. 2013</xref>) bootstraps, as well as the Shimodaira–Hasegawa–like approximate likelihood-ratio test (<xref ref-type="bibr" rid="B11">Guindon et al. 2010</xref>). Trees were visualized in FIGTREE v1.4.3.</p>
      </sec>
    </sec>
    <sec sec-type="﻿Results" id="SECID0E2MAC">
      <title>﻿Results</title>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Embioptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Ptilocerembiidae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">0433E825-4D1F-5340-B95D-F713B1621D88</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part>
            <object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/980C319C-B7B7-458C-B7B6-7822FF7D719B</object-id>
          </tp:taxon-name>
          <tp:taxon-status>sp. nov.</tp:taxon-status>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="type locality" id="SECID0EHOAC">
          <title>Type locality.</title>
          <p>China, Yunnan Province: Yuxi City, Ailao Mountain, Yingpan, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[101.547600,23.990100]}" id="NCID0EQOAC">23.9901°N, 101.5476°E</named-content></named-content> (Fig. <xref ref-type="fig" rid="F1">1</xref>), 2000 m, on road surface, 23 May 2018.V.23; L. Qiu leg.</p>
          <fig id="F1" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/dez.72.170313.figure1</object-id>
            <object-id content-type="arpha">A9389A0B-68D2-5E48-8F8D-6396C057EEF5</object-id>
            <label>Figure 1.</label>
            <caption>
              <p>General distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> species. Inset: habitat of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. in Mt. Ailaoshan, Yunnan Province, China. The type locality of the new species is indicated by a red dot, while the distribution of other species is indicated by yellow.</p>
            </caption>
            <graphic xlink:href="dez-72-569-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1483495.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1483495</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Type specimens" id="SECID0EYPAC">
          <title>Type specimens.</title>
          <p><bold><italic>Holotype</italic></bold>: • ♂, in alcohol. Original label: “China, Yunnan, Yuxi, Ailao Mountain, Yingpan, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[101.547600,23.990100]}" id="NCID0EHAAE">23.9901°N, 101.5476°E</named-content></named-content>, 2000 m, 23 May 2018.V.23, L Qiu”, “HOLOTYPE / <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. / Det. Chen” (<abbrev content-type="institution" xlink:title="Insect Collection of Jiangsu University of Science and Technology" id="ABBRID0EXAAE">ICJUST</abbrev>). <bold><italic>Paratype</italic></bold>: • 1♂, with same collection event label as the holotype and additional label “PARATYPE / <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. / Det. Chen” (<abbrev content-type="institution" xlink:title="Insect Collection of Jiangsu University of Science and Technology" id="ABBRID0EKBAE">ICJUST</abbrev>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="diagnosis" id="SECID0EPBAE">
          <title>Diagnosis.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is assigned to the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> of family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> by the antennae with long erect setae and white distal segments, bifurcated MA in all wings, hind basitarsus with dense ventral setae and only one papilla, tergite X obliquely divided into two unequal sclerites with the area between the hemitergites depressed, MF with a hook, elongated HP, and the fused left cercomeres (<xref ref-type="bibr" rid="B18">Miller et al. 2012</xref>; <xref ref-type="bibr" rid="B22">Poolprasert and Edgerly 2014</xref>). Among the five known species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> (see Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>: table S1), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is the most similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic> based on the 10R with a median convex along left margin, MF with a raised nodule near base of EP, and LC1 with a large echinulate nodule (<xref ref-type="bibr" rid="B22">Poolprasert and Edgerly 2014</xref>). However, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. can be distinguished from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic> by the submentum with anterior margin truncate (concave in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic>), MS with broad right half (absent in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic> but similar to that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="rossi">rossi</tp:taxon-name-part></tp:taxon-name></italic>), 10LP and 10 RP broad (narrow in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic> and other species), HP elongated and specialized apically (relatively short and simple in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic> and other species) (<xref ref-type="bibr" rid="B9">Friederichs 1923</xref>; <xref ref-type="bibr" rid="B22">Poolprasert and Edgerly 2014</xref>). The dichotomous key in <xref ref-type="bibr" rid="B22">Poolprasert and Edgerly (2014)</xref> could separate <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic> as well as its similar species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. from other congeners.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="description" id="SECID0EIHAE">
          <title>Description.</title>
          <p><bold>Male</bold>: Body length (from anterior of maxillary palps to cercal apex) ca. 15 mm; color pale to brown (Fig. <xref ref-type="fig" rid="F2">2A, B</xref>).</p>
          <fig id="F2" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/dez.72.170313.figure2</object-id>
            <object-id content-type="arpha">309FB7EC-EFAA-5AD1-8341-F494F70C1AC9</object-id>
            <label>Figure 2.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., male holotype. <bold>A.</bold> Habitus, dorsal view; <bold>B.</bold> Habitus, ventral view; <bold>C.</bold> Distal antennal segments, dorsal view. Scale bars: 1 mm.</p>
            </caption>
            <graphic xlink:href="dez-72-569-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1483496.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1483496</uri>
            </graphic>
          </fig>
          <p>Antennae at least 26-segmented, most segments brown and covered by long erect setae, distal segments pale (Fig. <xref ref-type="fig" rid="F2">2C</xref>). Head capsule dark brown (Fig. <xref ref-type="fig" rid="F3">3A, B</xref>), near 1.5 times longer than wide, with reticular pigmentation on posterior 1/3. Eyes small, sides behind eyes rounded, gradually converging posteriorly. Clypeus broad, pale brown; labrum semicircular, dark medially, pale laterally. Maxillary palpi dark brown, five-segmented; labial palpi brown and short, three-segmented. Submentum dark brown, subquadrate, slightly wider than long, anterior margin truncate, posterior margin slightly concave, lateral margins slightly convex.</p>
          <fig id="F3" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/dez.72.170313.figure3</object-id>
            <object-id content-type="arpha">DE1436F7-A55B-5D3F-8F01-2766002CBC68</object-id>
            <label>Figure 3.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., male holotype. <bold>A.</bold> Head and thorax, dorsal view; <bold>B.</bold> Head and thorax, ventral view; <bold>C.</bold> Right hind tarsus, dorsal view. Abbreviations: pa – papilla; SMT – submentum. Scale bars: 1 mm (<bold>A</bold> and <bold>B</bold>); 0.5 mm (<bold>C</bold>).</p>
            </caption>
            <graphic xlink:href="dez-72-569-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1483497.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1483497</uri>
            </graphic>
          </fig>
          <p>Prothorax brown (Fig. <xref ref-type="fig" rid="F3">3A</xref>), subquadrate, slightly wider posteriorly, much shorter and narrower than head, with subequal length and width, surface with deep transverse and longitudinal grooves. Meso- and metathorax generally brown, width subequal to head. Legs brown; hind basitarsus with single ventral papilla (Fig. <xref ref-type="fig" rid="F3">3C</xref>). Wings pale brown (Fig. <xref ref-type="fig" rid="F4">4</xref>), with longitudinal pale stripes; MA of all wings forked subbasally.</p>
          <fig id="F4" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/dez.72.170313.figure4</object-id>
            <object-id content-type="arpha">472AE012-1A0A-5D20-A7E0-1FE731C6AC3C</object-id>
            <label>Figure 4.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., male holotype. <bold>A.</bold> Left forewing, dorsal view; <bold>B.</bold> Right forewing, dorsal view; <bold>C.</bold> Left hind wing, dorsal view; <bold>D.</bold> Right hind wing, dorsal view. Abbreviation: MA – anterior media. Scale bars: 1 mm.</p>
            </caption>
            <graphic xlink:href="dez-72-569-g004.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1483498.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1483498</uri>
            </graphic>
          </fig>
          <p>Abdomen dark brown (Figs <xref ref-type="fig" rid="F2">2</xref>, <xref ref-type="fig" rid="F5">5</xref>). Terminalia with tenth abdominal tergite completely cleft, hemitergites separated basally (Fig. <xref ref-type="fig" rid="F5">5A–E</xref>). MS transverse, sinuate, near fusiform, with a posterior notch on right half. Hemitergite 10L broad, subtrapezoidal, left margin expanded to ventral side; 10LP constricted basally, right margin rounded, left margin greatly projected leftward medially, apex blunt and curved leftward. 10R broad throughout, inner margin sinuate, with a median convex; MF with a long sickle-shaped hook (EP) and a raised nodule; 10RP broad, subquadrate, apex with left angle pointed and right angle notched. LPPT produced as a subtriangular thin sclerite beneath 10L, sharp LPPT-P pointing backwards. LC1 tubular basally, elongated, inner margin with a large echinulate lobe apically. LC2 shorter than LC1, subconical, basal fusion with LC1 slightly constricted. Right cercus slightly longer than left cercus; RC1 slightly shorter than RC2, both covered with long setae and tapering towards apex. H broad (Fig. <xref ref-type="fig" rid="F5">5F, G</xref>), rounded, lateral margins obliquely convex; a slender curved sclerite present posterolateral to H. HP near as long as H; right margin mostly straight except expanded basally and convex apically; left margin convex at basal half, forming a denticulate ridge at apical half.</p>
          <fig id="F5" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/dez.72.170313.figure5</object-id>
            <object-id content-type="arpha">330B7F05-29E0-56DA-9C7B-17264FDB681D</object-id>
            <label>Figure 5.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., male holotype. <bold>A.</bold> Terminalia, dorsal view; <bold>B.</bold> Drawing of terminalia, dorsal view; <bold>C.</bold> Terminalia, slightly dorsocaudal view; <bold>D.</bold> Terminalia, dorsolateral view; <bold>E.</bold> Terminalia, lateral view; <bold>F.</bold> Terminalia, ventral view; <bold>G.</bold> Drawing of terminalia, ventral view. Abbreviation: 10L/R – left/right hemitergite of abdominal segment 10; 10LP/RP – left/right hemitergal process; EP – epiproct; H – hypandrium (ninth sternite); HP – hypandrium process; LPPT – left paraproct; LC1/2 – first/second segment of left cercus; MF – medial flap; MS – medial sclerite; RC1/2 – first/second segment of right cercus. Scale bars: 0.5 mm.</p>
            </caption>
            <graphic xlink:href="dez-72-569-g005.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1483499.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1483499</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="etymology" id="SECID0EGNAE">
          <title>Etymology.</title>
          <p>The species is named after its collector, Dr. Lu Qiu (Mianyang, Sichuan, China).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="distribution" id="SECID0ELNAE">
          <title>Distribution.</title>
          <p>China: Yunnan Province.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="biology" id="SECID0EQNAE">
          <title>Biology.</title>
          <p>The specimens were collected at a high-elevation site in the Ailao Mountain region, situated at the transitional boundary between the Yungui Plateau and the Hengduan Mountains. This area serves as a climatic divide between the Pacific Southeast Monsoon and the Indian Ocean Southwest Monsoon systems. The region experiences a subtropical montane climate characterized by cool annual temperatures, with mild summer conditions and cold winters. Precipitation is abundant and seasonally concentrated, with most rainfall occurring during an extended rainy season, while humidity remains consistently high throughout much of the year. The vegetation surrounding the habitat is dominated by extensive mid-elevation moist evergreen broad-leaved forests, recognized as one of China’s most significant subtropical montane wet evergreen broad-leaved forest ecosystems. These forests are primarily composed of species from the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Fagaceae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Theaceae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lauraceae</tp:taxon-name-part></tp:taxon-name>, and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Magnoliaceae</tp:taxon-name-part></tp:taxon-name> and feature a rich diversity of epiphytes (including mosses and ferns) and lianas. This structurally complex, humid evergreen forest, combined with persistent mist and steep elevational variation, creates stable micro-habitats such as bark crevices, leaf-litter zones, and humid canopy niches that are favorable for webspinners. Given the pronounced vertical climate stratification of the region, the new species may occupy a narrow elevational range with specific humidity, canopy cover, and litter-substrate conditions. Future field studies targeting mid-montane evergreen forest corridors in the Indochina–Yunnan region may reveal a broader distribution of the genus across similar habitats.</p>
          <sec sec-type="﻿Mitogenome characters" id="SECID0EKOAE">
            <title>﻿Mitogenome characters</title>
            <p>The complete mitogenome of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is 15,443 bp in length (GenBank accession No. <ext-link ext-link-type="gen" xlink:href="PX232380" xlink:type="simple">PX232380</ext-link>), which is slightly smaller than other reported webspinner mitogenomes (typically ~15.7–17.3 kb). It is a circular molecule comprising 13 protein-coding genes (<abbrev xlink:title="protein-coding genes" id="ABBRID0EAPAE">PCGs</abbrev>), 20 tRNA genes, and 2 rRNA genes, plus the A+T-rich control region (Fig. <xref ref-type="fig" rid="F6">6</xref>). Two tRNAs, tRNA-Val (trnV) and tRNA-Ser(UGA) (trnS2), are missing (a feature not seen in other embiopteran mitogenomes). The gene order is the canonical insect arrangement as in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drosophila">Drosophila</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="yakuba">yakuba</tp:taxon-name-part></tp:taxon-name></italic><xref ref-type="bibr" rid="B3">Burla (1954)</xref> (<xref ref-type="bibr" rid="B7">Clary and Wolstenholme 1985</xref>), with no rearrangements compared to the ancestral pattern (see Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>: table S2). Base composition is strongly A+T-biased (A 39.2%, T 25.1%, G 7.8%, C 27.9%).</p>
            <fig id="F6" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/dez.72.170313.figure6</object-id>
              <object-id content-type="arpha">492DC4F8-FCF8-5A4B-B16F-4864A317DB1D</object-id>
              <label>Figure 6.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., mitochondrial map. Genes outside the map are transcribed in a clockwise direction, whereas those inside the map are transcribed counterclockwise. The second circle shows the GC content, and the third shows the GC skew. GC content and GC skew are plotted as the deviation from the average value of the entire sequence.</p>
              </caption>
              <graphic xlink:href="dez-72-569-g006.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1483500.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1483500</uri>
              </graphic>
            </fig>
            <p>A notable feature is the presence of ~5.4 tandem repeat units in the intergenic region between tRNA-Ile and tRNA-Gln. Such repeated sequences have not been reported in other sequenced embiopteran mitogenomes. Tandem Repeats Finder analysis confirmed these repeats (<xref ref-type="bibr" rid="B1">Benson 1999</xref>), whereas the control region itself contains no repeated sequences.</p>
            <p>Most <abbrev xlink:title="protein-coding genes" id="ABBRID0E6QAE">PCGs</abbrev> use standard invertebrate start codons (ATA, ATG, ATT) and terminate with TAA; six genes (<abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EDRAE">COX1</abbrev>, COX2, COX3, ND1, ND4, ND5) end with an incomplete stop codon (single T), which is presumed to be completed by post-transcriptional polyadenylation. Relative synonymous codon usage (<abbrev xlink:title="relative synonymous codon usage" id="ABBRID0EHRAE">RSCU</abbrev>) analysis shows a strong bias toward A/T-ending codons; the most frequent codons encode serine (UCU, UCA) and threonine (ACC) (see Suppl. material <xref ref-type="supplementary-material" rid="S2">2</xref>: fig. S1A).</p>
            <p>Analysis of selective pressure (Ka/Ks) for each PCG (vs. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phasmotaenia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lanyuhensis">lanyuhensis</tp:taxon-name-part></tp:taxon-name></italic>) yielded values &lt; 1.0 for all genes (see Suppl. material <xref ref-type="supplementary-material" rid="S2">2</xref>: fig. S1B), indicating overall purifying selection in the sequenced webspinners. <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EASAE">COX1</abbrev> shows the lowest Ka/Ks (highest conservation), while ATP8 shows a relatively higher rate (although still &lt;1). The pattern that <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EESAE">COX1</abbrev> is the slowest-evolving mitochondrial gene and an ATP synthase gene is the fastest is consistent with other <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Embioptera</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B6">Chen et al. 2017</xref>).</p>
          </sec>
          <sec sec-type="﻿Molecular delimitation and phylogeny" id="SECID0ERSAE">
            <title>﻿Molecular delimitation and phylogeny</title>
            <p>The <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EXSAE">COX1</abbrev> barcode of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. was compared to six available <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> sequences from GenBank (except for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senathami">senathami</tp:taxon-name-part></tp:taxon-name></italic>, for which <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EYTAE">COX1</abbrev> is missing). Pairwise K2P distances between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. and other congeners ranged from 0.1259 to 0.1759 (see Suppl. material <xref ref-type="supplementary-material" rid="S2">2</xref>: fig. S2), far exceeding the widely adopted intraspecific thresholds (~2%) in webspinners and other insects (<xref ref-type="bibr" rid="B13">Hebert et al. 2003</xref>; <xref ref-type="bibr" rid="B4">Chen 2022</xref>, <xref ref-type="bibr" rid="B5">2023</xref>). The smallest distance among any congeners was 0.042 (between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="catherinae">catherinae</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="thaidina">thaidina</tp:taxon-name-part></tp:taxon-name></italic>). In phylogenetic trees (<abbrev xlink:title="Bayesian inference" id="ABBRID0ENVAE">BI</abbrev>, <abbrev xlink:title="Neighbor-Joining" id="ABBRID0ERVAE">NJ</abbrev>, and <abbrev xlink:title="Maximum likelihood" id="ABBRID0EVVAE">ML</abbrev>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. grouped with two unnamed <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> specimens sequenced in <xref ref-type="bibr" rid="B18">Miller et al. (2012)</xref>, rather than with any of the formally described species. The phylogenetic placement of the new species was supported by relatively strong topological values, with a posterior probability of 0.82 in the <abbrev xlink:title="Bayesian inference" id="ABBRID0EPWAE">BI</abbrev> tree (Fig. <xref ref-type="fig" rid="F7">7</xref>), a bootstrap value of 66 in the <abbrev xlink:title="Neighbor-Joining" id="ABBRID0EXWAE">NJ</abbrev> tree (see Suppl. material <xref ref-type="supplementary-material" rid="S2">2</xref>: fig. S3A), and a bootstrap value of 52 in the <abbrev xlink:title="Maximum likelihood" id="ABBRID0E6WAE">ML</abbrev> tree (see Suppl. material <xref ref-type="supplementary-material" rid="S2">2</xref>: fig. S3B). These results indicate that the sequenced individual of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. forms a distinct clade, consistent with its morphological uniqueness. In summary, both morphological and <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0ESXAE">COX1</abbrev> data support the delimitation of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. as a new species, divergent from its congeners.</p>
            <fig id="F7" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/dez.72.170313.figure7</object-id>
              <object-id content-type="arpha">BAA4D26F-51D0-5EB0-B94A-DE599E292D26</object-id>
              <label>Figure 7.</label>
              <caption>
                <p>Bayesian inference tree of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> species based on <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EQYAE">COX1</abbrev> sequences. Numbers at the nodes are posterior probabilities.</p>
              </caption>
              <graphic xlink:href="dez-72-569-g007.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1483501.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1483501</uri>
              </graphic>
            </fig>
          </sec>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
    <sec sec-type="﻿Discussion" id="SECID0EZYAE">
      <title>﻿Discussion</title>
      <p>This study reports the first record of the webspinner family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> from China, expanding the known range of this Southeast Asian lineage. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> was previously known from Indonesia through Thailand and Myanmar. The discovery of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. in Yunnan Province (southwestern China) is consistent with the prediction of <xref ref-type="bibr" rid="B27">Ross (2007)</xref> that the montane regions of northern Myanmar, Laos, Vietnam, and southwestern China harbor undiscovered embiopteran taxa. Our finding extends the boundary of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> and suggests that related species may occur across the border regions in Indochina. Although this record is not the northernmost known occurrence of the genus (since Myanmar already hosted species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic>), it nonetheless highlights the broader biogeographic significance of Yunnan Province as a biodiversity hotspot. The presence of this lineage in Yunnan underscores the incompleteness of current knowledge of the embiopteran fauna in China.</p>
      <p>Morphologically, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is clearly distinct from all described <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> species, especially in the characters of its head and male genitalia. The combination of features (e.g., submentum shape, tergal processes, hypandrium form) distinguishes it from the four Thai species (<xref ref-type="bibr" rid="B22">Poolprasert and Edgerly 2014</xref>) and the Indonesian <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="roepkei">roepkei</tp:taxon-name-part></tp:taxon-name></italic>. In our analysis of molecular data, the <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EJ2AE">COX1</abbrev> barcode of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. was highly divergent from those of all sampled congeners. The substantial interspecific distances (&gt;0.12) far exceed usual DNA barcoding thresholds (<xref ref-type="bibr" rid="B13">Hebert et al. 2003</xref>) and indicate that the new specimens are not conspecific with any previously sequenced taxon. The congruence of morphological and genetic results provides strong evidence that the Chinese specimens represent a new species.</p>
      <p>Notably, the genetic analysis revealed considerable interspecific genetic distances, suggesting that the specimens sequenced but not formally named in <xref ref-type="bibr" rid="B18">Miller et al. (2012)</xref> might also represent new species. The relatively small number of described species (currently six worldwide) combined with our findings implies that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> diversity is seriously underestimated. Many additional species likely remain to be discovered in Asia. <xref ref-type="bibr" rid="B26">Ross (1963)</xref> suspected a complex of many species across the region, and our results confirm that even putative colonies in well-surveyed areas can yield new taxa.</p>
      <p>The complete mitogenome of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. provides a first look at the mitogenomics of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name>. It conforms to the typical insect pattern (13 <abbrev xlink:title="protein-coding genes" id="ABBRID0E63AE">PCGs</abbrev>, A+T-biased, ancestral gene order), but with some distinctive features. In particular, we found tandem repeats in a coding-region intergenic spacer (between trnI and trnQ) that have not been reported in other webspinner mitogenomes. Tandem repeat regions are common in the control region of many insect mitogenomes, but their presence in this non-control region suggests lineage-specific evolution. The functional significance of these repeats is unclear. They could be neutral structural features or potentially involved in replication dynamics. In addition, two tRNA genes (trnV and trnS2) were not detected in our mitogenome assembly. Because missing tRNAs are sometimes the result of annotation or sequencing limitations rather than true gene loss (<xref ref-type="bibr" rid="B20">Morrison 2010</xref>), these absences should be interpreted cautiously until confirmed by additional sequencing data. Further comparative mitogenomic analyses of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> and related taxa will help clarify whether these features represent lineage-specific modifications or methodological artifacts. Our Ka/Ks analysis indicated that all protein genes are evolving under strong purifying selection (Ka/Ks &lt; 1), with <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0EO4AE">COX1</abbrev> being the most conserved and ATP8 the least. The elevated Ka/Ks in ATP8 suggests a relaxed constraint on this gene, whereas <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0ES4AE">COX1</abbrev>’s strong constraint supports its use as a robust barcode marker.</p>
      <p>Our study is based on male specimens from a single locality; no females are yet known for this species. More fieldwork is needed to collect females and additional populations of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., which would allow for the study of potential intraspecific variation and female morphology. From a molecular perspective, our phylogenetic analysis used only <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0ED5AE">COX1</abbrev>, which is useful for species delimitation but limited for deeper relationships. Future work should incorporate additional loci (nuclear markers or transcriptomes) and full mitogenomes of multiple species to build a more comprehensive phylogeny of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Embioptera</tp:taxon-name-part></tp:taxon-name>. In particular, multi-gene phylogenomics could test the monophyly of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> and clarify its position relative to other families. Expanding genomic resources will also enable studies of population genetics and biogeography; for example, the tandem repeats found here could serve as genetic markers if conserved in populations.</p>
      <p>In conclusion, we provide the first Chinese record of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ptilocerembiidae</tp:taxon-name-part></tp:taxon-name> with the description of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. Our integrative approach (morphology + mitogenomics) demonstrates the value of combining data types in embiid taxonomy. This discovery highlights the hidden diversity of webspinners and underscores that even well-studied regions can yield novel taxa. As taxonomic work progresses, it will be important to integrate classical morphology, DNA barcoding, and genomic data to achieve a robust classification of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Embioptera</tp:taxon-name-part></tp:taxon-name> and to document the evolutionary history of this insect order.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>﻿Acknowledgments</title>
      <p>We are grateful to Dr. Lu Qiu (Mianyang, Sichuan, China) for providing the specimens. We also thank Dr. Pisit Poolprasert (Kasetsart University, Thailand) for providing valuable information on Thailand webspinners. This study was supported by the CACMS Innovation Fund (grant no. CI2024E003).</p>
    </ack>
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    <sec sec-type="supplementary-material">
      <title>Supplementary materials</title>
      <supplementary-material id="S1" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="arpha">3DA44D55-4B79-578C-B0E7-44D5A5A0F85C</object-id>
        <label>﻿Supplementary material 1</label>
        <caption>
          <p>﻿Supplementary tables</p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>xlsx</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p><bold>table S1.</bold> Comparison of diagnostic characters in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> species. <bold>table S2.</bold> Mitogenome organization of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</p>
        </statement>
        <media xlink:href="dez-72-569-s001.xlsx" mimetype="application" mime-subtype="vnd.openxmlformats-officedocument.spreadsheetml.sheet" position="float" orientation="portrait" xlink:type="simple" id="oo_1483502.xlsx">
          <uri content-type="original_file">https://binary.pensoft.net/file/1483502</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Zhi-Teng Chen, Yuan Xiong, Chao Jiang</attrib>
      </supplementary-material>
      <supplementary-material id="S2" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="arpha">674F917E-2438-571D-930B-33C4F682779E</object-id>
        <label>﻿Supplementary material 2</label>
        <caption>
          <p>﻿Supplementary figures</p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>pdf</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p><bold>fig. S1.</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="qiului">qiului</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., mitogenomic features. <bold>A.</bold> Relative synonymous codon usage (<abbrev xlink:title="relative synonymous codon usage" id="ABBRID0E6GBG">RSCU</abbrev>) of <abbrev xlink:title="protein-coding genes" id="ABBRID0EDHBG">PCGs</abbrev> in the mitogenome; numbers on bars represent the ratio of each amino acid; <bold>B.</bold> Nonsynonymous/synonymous substitution (Ka/Ks) ratios of each PCG in three sequenced webspinner mitogenomes. <bold>fig. S2.</bold> Genetic distance among the DNA barcodes of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> species. <bold>fig. S3.</bold> Phylogenetic trees of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ptilocerembia">Ptilocerembia</tp:taxon-name-part></tp:taxon-name></italic> species based on <abbrev xlink:title="Mitochondrial DNA barcoding" id="ABBRID0E2HBG">COX1</abbrev> sequences. <bold>A.</bold> Neighborjoining tree; numbers at the nodes are bootstrap values; <bold>B.</bold> Maximum likelihood tree; numbers at the nodes are bootstrap values.</p>
        </statement>
        <media xlink:href="dez-72-569-s002.pdf" mimetype="application" mime-subtype="pdf" position="float" orientation="portrait" xlink:type="simple" id="oo_1483503.pdf">
          <uri content-type="original_file">https://binary.pensoft.net/file/1483503</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Zhi-Teng Chen, Yuan Xiong, Chao Jiang</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
