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<article xmlns:tp="http://www.plazi.org/taxpub" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" article-type="research-article" xml:lang="en">
  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">2</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">ZooKeys</journal-title>
        <abbrev-journal-title xml:lang="en">ZK</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1313-2989</issn>
      <issn pub-type="epub">1313-2970</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/zookeys.1293.204408</article-id>
      <article-id pub-id-type="publisher-id">204408</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Moniligastridae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Molecular systematics</subject>
          <subject>Phylogeny</subject>
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Asia</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Establishment of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> species complex and description of a new species from northeastern China (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subclass" reg="Oligochaeta">Oligochaeta</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Moniligastridae">Moniligastridae</tp:taxon-name-part></tp:taxon-name>)</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Qiao</surname>
            <given-names>Saiyu</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0002-9964-1352</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Li</surname>
            <given-names>Jiangni</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-1642-0328</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Zhang</surname>
            <given-names>Yufeng</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-4066-6355</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Aspe</surname>
            <given-names>Nonillon M.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-0025-6017</uri>
          <xref ref-type="aff" rid="A3">3</xref>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Zhao</surname>
            <given-names>Huifeng</given-names>
          </name>
          <email xlink:type="simple">zhaohf@lfnu.edu.cn</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-4243-9671</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Hebei Key Laboratory of Animal Diversity, Langfang Normal University, Langfang 065000, China</addr-line>
        <institution>Hebei Key Laboratory of Animal Diversity, Langfang Normal University</institution>
        <addr-line content-type="city">Langfang</addr-line>
        <country>China</country>
        <uri content-type="ror">https://ror.org/05kyq2m47</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Hebei Key Laboratory of Zoological Systematics and Application, College of Life Sciences, Hebei University, Baoding, 071002, China</addr-line>
        <institution>Hebei Key Laboratory of Zoological Systematics and Application, College of Life Sciences, Hebei University</institution>
        <addr-line content-type="city">Baoding</addr-line>
        <country>China</country>
        <uri content-type="ror">https://ror.org/01p884a79</uri>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">College of Environment and Life Sciences, Mindanao State University at Naawan, Naawan 9023, Misamis Oriental, Philippines</addr-line>
        <institution>College of Environment and Life Sciences, Mindanao State University at Naawan</institution>
        <addr-line content-type="city">Naawan</addr-line>
        <country>Philippines</country>
        <uri content-type="ror">https://ror.org/01f51gm92</uri>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Huifeng Zhao (<email xlink:type="simple">zhaohf@lfnu.edu.cn</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Samuel James</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>01</day>
        <month>10</month>
        <year>2026</year>
      </pub-date>
      <volume>1293</volume>
      <fpage>327</fpage>
      <lpage>346</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/AD19E178-E838-5A9F-BDC3-191F68FA3F5F">AD19E178-E838-5A9F-BDC3-191F68FA3F5F</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/AE7767BC-E035-4B8E-B4E5-DF34B1EBF83F">AE7767BC-E035-4B8E-B4E5-DF34B1EBF83F</uri>
      <history>
        <date date-type="received">
          <day>17</day>
          <month>06</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>12</day>
          <month>09</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Saiyu Qiao, Jiangni Li, Yufeng Zhang, Nonillon M. Aspe, Huifeng Zhao</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/AE7767BC-E035-4B8E-B4E5-DF34B1EBF83F</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>The family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Moniligastridae">Moniligastridae</tp:taxon-name-part></tp:taxon-name> Claus, 1880 remains taxonomically poorly resolved, and its actual species diversity is likely severely underestimated. Despite its ecological significance as an East Asian endemic species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> Michaelsen, 1931 has been studied almost exclusively from non-taxonomic perspectives. Consequently, its species boundaries remain poorly understood. Here, we formally establish the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> species complex (<abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev>), comprising eight morphologically and geographically related species. We employed an integrative taxonomic approach combining morphology and molecular data to examine specimens of <abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev> from northeastern China, leading to the description of a new species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. The new species is characterized by a colorless body in alcohol, 3–5 gizzards, M-shaped prostates, a very short and uncoiled vas deferens, and thick penes with a swollen base and blunt, rounded apex. Phylogenetic analyses based on 13 protein-coding genes of the mitochondrial genome of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part></tp:taxon-name></italic> employing two molecular species delimitation methods—Assemble Species by Automatic Partitioning (<abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev>), and Bayesian Phylogenetics and Phylogeography (<abbrev xlink:title="Bayesian Phylogenetics and Phylogeography">BPP</abbrev>)—recovered <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. as a monophyletic lineage, and COI genetic distances (19.4–22.0% between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>) confirmed the species status of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. Additionally, we provide an updated redescription of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> based on newly examined specimens from multiple localities. Our findings highlight considerable external morphological plasticity in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>, which may likely be environmentally induced, and underscore the necessity of integrative approaches for accurate species delimitation in this group.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="phylum" reg="Annelida">Annelida</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>mitogenome</kwd>
        <kwd>new species</kwd>
        <kwd>species group</kwd>
      </kwd-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="sec1">
        <title>Citation</title>
        <p>Qiao S, Li J, Zhang Y, Aspe NM, Zhao H (2026) Establishment of the <italic>Drawida gisti</italic> species complex and description of a new species from northeastern China (Oligochaeta, Moniligastridae). ZooKeys 1293: 327–346. <ext-link xlink:href="10.3897/zookeys.1293.204408" ext-link-type="doi">https://doi.org/10.3897/zookeys.1293.204408</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec2">
      <title>Introduction</title>
      <p>The family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Moniligastridae">Moniligastridae</tp:taxon-name-part></tp:taxon-name> Claus, 1880 comprises five genera and 193 species. Among them, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part></tp:taxon-name></italic> Michaelsen, 1900 is the largest genus with 163 species and subspecies (<xref ref-type="bibr" rid="B54">Tiwari et al. 2025</xref>, <xref ref-type="bibr" rid="B55">2026</xref>; <xref ref-type="bibr" rid="B2">Ardra et al. 2026</xref>; <xref ref-type="bibr" rid="B18">Fahis et al. 2026</xref>; <xref ref-type="bibr" rid="B46">Qiao et al. 2026a</xref>, <xref ref-type="bibr" rid="B47">2026b</xref>), distributed across South, Southeast, and East Asia (<xref ref-type="bibr" rid="B22">Gates 1936</xref>, <xref ref-type="bibr" rid="B24">1972</xref>; <xref ref-type="bibr" rid="B16">Easton 1981</xref>; <xref ref-type="bibr" rid="B4">Blakemore 2012</xref>; <xref ref-type="bibr" rid="B7">Blakemore et al. 2014</xref>; <xref ref-type="bibr" rid="B1">Anderson et al. 2017</xref>; <xref ref-type="bibr" rid="B65">Zhang et al. 2020</xref>; <xref ref-type="bibr" rid="B41">Narayanan et al. 2025a</xref>, <xref ref-type="bibr" rid="B42">2025b</xref>; <xref ref-type="bibr" rid="B64">Zhang et al. 2026</xref>). Integrative taxonomic methods have become a standard for species delimitation due to widespread cryptic speciation of earthworms (<xref ref-type="bibr" rid="B8">Chang and James 2011</xref>; <xref ref-type="bibr" rid="B26">Han et al. 2024</xref>, <xref ref-type="bibr" rid="B27">2026</xref>; <xref ref-type="bibr" rid="B48">Qin et al. 2025</xref>; <xref ref-type="bibr" rid="B37">Liu et al. 2026</xref>; <xref ref-type="bibr" rid="B57">Xu et al. 2026</xref>). <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Moniligastridae">Moniligastridae</tp:taxon-name-part></tp:taxon-name> remains taxonomically poorly resolved with likely underreported cryptic diversity (<xref ref-type="bibr" rid="B35">Liu et al. 2025a</xref>, <xref ref-type="bibr" rid="B36">2025b</xref>). Within this family, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> Michaelsen, 1931, an ecologically significant East Asian endemic, has been studied almost exclusively from non-taxonomic perspectives (<xref ref-type="bibr" rid="B56">Wang et al. 2020</xref>; <xref ref-type="bibr" rid="B62">Yin et al. 2022</xref>; <xref ref-type="bibr" rid="B33">Liao et al. 2024</xref>), leaving its taxonomy strikingly understudied and its true species diversity potentially obscured.</p>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> is distributed in eastern China (<xref ref-type="bibr" rid="B23">Gates 1939</xref>; <xref ref-type="bibr" rid="B31">Kobayashi 1940</xref>; <xref ref-type="bibr" rid="B7">Blakemore et al. 2014</xref>; <xref ref-type="bibr" rid="B3">Bi et al. 2025</xref>). Original description of the species is from Jinan (in Shandong Province) and Beijing (<xref ref-type="bibr" rid="B40">Michaelsen 1931</xref>), with type specimens deposited in the Hamburg Museum. It measures 77–151 mm length, 6 mm diameter with 180–190 number of segments and is characterized by a slender vas deferens emerging from the testis sacs, forming a tightly coiled loop within segment IX, entering segment X to produce a similar convolution, and finally leading to the proximal portion of the prostate gland. <xref ref-type="bibr" rid="B10">Chen (1933)</xref> described one subspecies, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="nanchangiana">nanchangiana</tp:taxon-name-part></tp:taxon-name></italic>, based on minor morphological differences in the penial chamber and prostate distinguished it from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="anchingiana">anchingiana</tp:taxon-name-part></tp:taxon-name></italic> was concurrently described but later elevated to species level as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anchingiana">anchingiana</tp:taxon-name-part></tp:taxon-name></italic> by <xref ref-type="bibr" rid="B29">Kobayashi (1936)</xref>. Later redescriptions of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> by Gates (<xref ref-type="bibr" rid="B21">1935</xref>, <xref ref-type="bibr" rid="B23">1939</xref>), <xref ref-type="bibr" rid="B30">Kobayashi (1938)</xref>, and <xref ref-type="bibr" rid="B11">Chen (1959)</xref> were added more details on spermathecae, prostate, and the gizzard, yet these accounts remain incomplete by modern standards and lack molecular validation.</p>
      <p>We examined other morphologically similar species recorded from the distributional range of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>. These taxa were only reviewed based on published literature, as type specimens are presumed lost: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cheni">cheni</tp:taxon-name-part></tp:taxon-name></italic> Gates, 1935 (body length 122–177 mm, diameter 4 mm) is distributed in Jiangsu; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="linhaiensis">linhaiensis</tp:taxon-name-part></tp:taxon-name></italic> Chen, 1933 (37–58 mm, 3–4 mm) in Zhejiang, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinica">sinica</tp:taxon-name-part></tp:taxon-name></italic> Chen, 1933 (50–95 mm, 2.2–3.2 mm) in Jiangxi (<xref ref-type="bibr" rid="B10">Chen 1933</xref>; <xref ref-type="bibr" rid="B21">Gates 1935</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sulcata">sulcata</tp:taxon-name-part></tp:taxon-name></italic> Zhong, 1986 (60–86 mm, 9.0–12.5 mm) in Yunnan, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="syringa">syringa</tp:taxon-name-part></tp:taxon-name></italic> Chen, 1933 (50–100 mm, 2.5–3.0 mm) in Zhejiang. These species all occur in eastern and southwestern China, with their type localities situated in relatively geographic proximity, suggesting potential biogeographic affinity. Also, these species highly share similarities in external and internal morphological characters with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>, including the presence of a penis and penial chamber. We therefore hypothesize that they represent closely allied species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>, and we formally establish the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> species complex (<abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev>), comprising <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anqingiana">anqingiana</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="nanchangiana">nanchangiana</tp:taxon-name-part></tp:taxon-name></italic>, with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cheni">cheni</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="linhaiensis">linhaiensis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="syringa">syringa</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinica">sinica</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sulcata">sulcata</tp:taxon-name-part></tp:taxon-name></italic> (Table <xref ref-type="table" rid="T1">1</xref>).</p>
      <table-wrap id="T1" position="float" orientation="portrait">
        <label>Table 1.</label>
        <caption>
          <p>Key morphological characteristics of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> species complex (NA: not available).</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="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov</bold>.</th>
              <th rowspan="1" colspan="1">
                <bold>
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</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="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anchingiana">anchingiana</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="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="nanchangiana">nanchangiana</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="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cheni">cheni</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="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="linhaiensis">linhaiensis</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="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinica">sinica</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="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sulcata">sulcata</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="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="syringa">syringa</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </bold>
              </th>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Color</bold>
              </td>
              <td rowspan="1" colspan="1">light gray</td>
              <td rowspan="1" colspan="1">Red-brown</td>
              <td rowspan="1" colspan="1">light blue-green</td>
              <td rowspan="1" colspan="1">light gray, without pigment</td>
              <td rowspan="1" colspan="1">light blue</td>
              <td rowspan="1" colspan="1">Light blue-gray</td>
              <td rowspan="1" colspan="1">light green</td>
              <td rowspan="1" colspan="1">brown</td>
              <td rowspan="1" colspan="1">dusky flesh</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Length</bold>
              </td>
              <td rowspan="1" colspan="1">48–87</td>
              <td rowspan="1" colspan="1">48–66</td>
              <td rowspan="1" colspan="1">56–80</td>
              <td rowspan="1" colspan="1">55–98</td>
              <td rowspan="1" colspan="1">122–177</td>
              <td rowspan="1" colspan="1">37–58</td>
              <td rowspan="1" colspan="1">50–95</td>
              <td rowspan="1" colspan="1">60–86</td>
              <td rowspan="1" colspan="1">50–100</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Width</bold>
              </td>
              <td rowspan="1" colspan="1">4.5–7.0</td>
              <td rowspan="1" colspan="1">5.0–7.0</td>
              <td rowspan="1" colspan="1">3.0–5.0</td>
              <td rowspan="1" colspan="1">2.2–3.0</td>
              <td rowspan="1" colspan="1">4.0</td>
              <td rowspan="1" colspan="1">3.0–4.0</td>
              <td rowspan="1" colspan="1">2.2–3.2</td>
              <td rowspan="1" colspan="1">9.0–12.5</td>
              <td rowspan="1" colspan="1">2.5–3.0</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Genital markings</bold>
              </td>
              <td rowspan="1" colspan="1">VII–XI, unpaired, irregular</td>
              <td rowspan="1" colspan="1">absent or present VIII–XII</td>
              <td rowspan="1" colspan="1">VII–XI, unpaired</td>
              <td rowspan="1" colspan="1">VII–XIII, XIV, unpaired, numerous</td>
              <td rowspan="1" colspan="1">7–10, irregular</td>
              <td rowspan="1" colspan="1">8–10, irregular</td>
              <td rowspan="1" colspan="1">8–11</td>
              <td rowspan="1" colspan="1">absent</td>
              <td rowspan="1" colspan="1">absent or present in VII–IX</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Clitellum</bold>
              </td>
              <td rowspan="1" colspan="1">X–XIII; Pale pink</td>
              <td rowspan="1" colspan="1">IX/X–XIII/XIV</td>
              <td rowspan="1" colspan="1">X–XIII</td>
              <td rowspan="1" colspan="1">X–XIII</td>
              <td rowspan="1" colspan="1">½IX–XIV,½XV</td>
              <td rowspan="1" colspan="1">X–XIII, pale</td>
              <td rowspan="1" colspan="1">X–XIII, pale white</td>
              <td rowspan="1" colspan="1">NA</td>
              <td rowspan="1" colspan="1">X–XII</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Penis</bold>
              </td>
              <td rowspan="1" colspan="1">10/11, large, rounded at tip, large at base</td>
              <td rowspan="1" colspan="1">10/11, long, large at base gradually tapering to sharp</td>
              <td rowspan="1" colspan="1">10/11, small</td>
              <td rowspan="1" colspan="1">10/11, small, pointed at tip, large at base</td>
              <td rowspan="1" colspan="1">10/11, broad triangular at base</td>
              <td rowspan="1" colspan="1">10/11, large, transversely ovoid, rounded at tip</td>
              <td rowspan="1" colspan="1">10/11, small, rounded or transversely flat at tip, large at base 0.18 mm</td>
              <td rowspan="1" colspan="1">Conical</td>
              <td rowspan="1" colspan="1">10/11 thick at base tapering at tip</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Form of male pores</bold>
              </td>
              <td rowspan="1" colspan="1">small pore on penis</td>
              <td rowspan="1" colspan="1">longitudinal slit on penis</td>
              <td rowspan="1" colspan="1">small pore on penis</td>
              <td rowspan="1" colspan="1">paired</td>
              <td rowspan="1" colspan="1">at tip of penis</td>
              <td rowspan="1" colspan="1">10/11</td>
              <td rowspan="1" colspan="1">small, at tip or slightly lateral of penis</td>
              <td rowspan="1" colspan="1">10/11, oval or eye-shaped</td>
              <td rowspan="1" colspan="1">NA</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Female pore</bold>
              </td>
              <td rowspan="1" colspan="1">11/12</td>
              <td rowspan="1" colspan="1">11/12</td>
              <td rowspan="1" colspan="1">11/12, transverse slit-like</td>
              <td rowspan="1" colspan="1">paired</td>
              <td rowspan="1" colspan="1">12; in ab line</td>
              <td rowspan="1" colspan="1">11/12; in b line</td>
              <td rowspan="1" colspan="1">11/12; in b line</td>
              <td rowspan="1" colspan="1">11/12, Small crack-shaped</td>
              <td rowspan="1" colspan="1">NA</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Spermathecae ducts</bold>
              </td>
              <td rowspan="1" colspan="1">medium, coiled</td>
              <td rowspan="1" colspan="1">medium, coiled</td>
              <td rowspan="1" colspan="1">medium, less coiled</td>
              <td rowspan="1" colspan="1">coiled</td>
              <td rowspan="1" colspan="1">long, coiled</td>
              <td rowspan="1" colspan="1">slender, coiled</td>
              <td rowspan="1" colspan="1">long, coiled</td>
              <td rowspan="1" colspan="1">long, coiled</td>
              <td rowspan="1" colspan="1">coiled</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Spermathecae Ampulla</bold>
              </td>
              <td rowspan="1" colspan="1">VIII</td>
              <td rowspan="1" colspan="1">VIII</td>
              <td rowspan="1" colspan="1">VII</td>
              <td rowspan="1" colspan="1">VIII</td>
              <td rowspan="1" colspan="1">VIII</td>
              <td rowspan="1" colspan="1">VIII</td>
              <td rowspan="1" colspan="1">VIII</td>
              <td rowspan="1" colspan="1">VIII</td>
              <td rowspan="1" colspan="1">VIII</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Spermathecae Atrium</bold>
              </td>
              <td rowspan="1" colspan="1">large, elliptical in 7/8</td>
              <td rowspan="1" colspan="1">large, comma-shaped in 8</td>
              <td rowspan="1" colspan="1">short, cylindrical in 7/8</td>
              <td rowspan="1" colspan="1">thumb-like and small in 8</td>
              <td rowspan="1" colspan="1">NA</td>
              <td rowspan="1" colspan="1">in 7/8</td>
              <td rowspan="1" colspan="1">zigzag, long tube in 7/8</td>
              <td rowspan="1" colspan="1">cylindrical and slightly flattened</td>
              <td rowspan="1" colspan="1">thick and short, stick-shaped</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Ovaries</bold>
              </td>
              <td rowspan="1" colspan="1">ovary chambers present, ovaries in XI–XII</td>
              <td rowspan="1" colspan="1">ovary chambers present, ovaries in X</td>
              <td rowspan="1" colspan="1">ovary chambers present, ovaries in XI</td>
              <td rowspan="1" colspan="1">ovary chambers present, ovaries in XI</td>
              <td rowspan="1" colspan="1">NA</td>
              <td rowspan="1" colspan="1">ovary chambers present, ovaries in XI</td>
              <td rowspan="1" colspan="1">ovary chambers present, ovaries in XI</td>
              <td rowspan="1" colspan="1">ovary chambers present, ovaries in XI</td>
              <td rowspan="1" colspan="1">NA</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Ovisacs</bold>
              </td>
              <td rowspan="1" colspan="1">XIII–XXI</td>
              <td rowspan="1" colspan="1">X–XIII/XVI/XVIII</td>
              <td rowspan="1" colspan="1">NA</td>
              <td rowspan="1" colspan="1">XII–XIIII</td>
              <td rowspan="1" colspan="1">XII, XIV, or XV</td>
              <td rowspan="1" colspan="1">XVI–XX</td>
              <td rowspan="1" colspan="1">X/XI/XII–XVI or XXII</td>
              <td rowspan="1" colspan="1">NA</td>
              <td rowspan="1" colspan="1">Extend to XXV</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Gizzard segments and position</bold>
              </td>
              <td rowspan="1" colspan="1">3–5; IX–XVI, XII/XIII–XVIII or XI–XVII</td>
              <td rowspan="1" colspan="1">3; IX/X–XV/XVII/XVIII</td>
              <td rowspan="1" colspan="1">2 or 3; XI–XIII</td>
              <td rowspan="1" colspan="1">3; XII–XV</td>
              <td rowspan="1" colspan="1">3; XIII–XV</td>
              <td rowspan="1" colspan="1">3; XIII–XVI</td>
              <td rowspan="1" colspan="1">3 or 4; XI–XIII</td>
              <td rowspan="1" colspan="1">3, XI/XII–XIV</td>
              <td rowspan="1" colspan="1">2–5</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Vas deferens</bold>
              </td>
              <td rowspan="1" colspan="1">short, uncoiled</td>
              <td rowspan="1" colspan="1">extremely long, two coils</td>
              <td rowspan="1" colspan="1">long, coiled</td>
              <td rowspan="1" colspan="1">long, coiled</td>
              <td rowspan="1" colspan="1">short, twisted</td>
              <td rowspan="1" colspan="1">multi-coiled</td>
              <td rowspan="1" colspan="1">extremely long, two coils</td>
              <td rowspan="1" colspan="1">short coiled</td>
              <td rowspan="1" colspan="1">NA</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <bold>Prostate</bold>
              </td>
              <td rowspan="1" colspan="1">M-shaped, white, 4.705–11.614 mm</td>
              <td rowspan="1" colspan="1">Shape</td>
              <td rowspan="1" colspan="1">slender, cylindrical, maximum 2 mm</td>
              <td rowspan="1" colspan="1">Slender, short, club-shaped, 2.0–2.5 mm</td>
              <td rowspan="1" colspan="1">multi-curved, 10–12 mm</td>
              <td rowspan="1" colspan="1">large, long</td>
              <td rowspan="1" colspan="1">long</td>
              <td rowspan="1" colspan="1">smooth, long, 5.0–6.8 mm</td>
              <td rowspan="1" colspan="1">short and broad, with thick glands on exterior</td>
            </tr>
          </tbody>
        </table>
      </table-wrap>
      <p>Here, we provide an updated redescription of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> based on the newly examined specimens. Moreover, we employ an integrative taxonomic approach to examine and analyze earthworm specimens belonging to the <abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev> collected from northeastern China, leading to the delimitation of a new species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. Mitochondrial genomes and three nuclear markers were sequenced and analyzed for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec3">
      <title>Material and methods</title>
      <sec sec-type="Sampling" id="sec4">
        <title>Sampling</title>
        <p>Specimens of the new species were collected from Weiyuan Park, Shanhaiguan District, Qinhuangdao Prefecture, Hebei Province, China. Specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> were collected from Mentougou district in Beijing, Rizhao and Heze prefectures in Shandong Province, and Chengde Prefecture in Hebei Province, China (Table <xref ref-type="table" rid="T2">2</xref>). All specimens were collected using hand-sorting and digging methods. Collected earthworms were immediately fixed and stored in 95% ethanol in the field. In the laboratory, all specimens were preserved at −20 °C in the Hebei Key Laboratory of Animal Diversity, Langfang Normal University, Hebei, China (<bold><abbrev content-type="institution" xlink:title="Hebei Key Laboratory of Animal Diversity, Langfang Normal University, Hebei, China">C-HLU</abbrev></bold>).</p>
        <table-wrap id="T2" position="float" orientation="portrait">
          <label>Table 2.</label>
          <caption>
            <p>List of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part></tp:taxon-name></italic> species, incorporated into analysis (hyphen indicate not available).</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <th rowspan="2" colspan="1">
                  <bold>Specimen ID</bold>
                </th>
                <th rowspan="2" colspan="1">
                  <bold>Species</bold>
                </th>
                <th rowspan="2" colspan="1">
                  <bold>Location</bold>
                </th>
                <th rowspan="1" colspan="4">
                  <bold>Accession Number</bold>
                </th>
                <th rowspan="1" colspan="1"/>
              </tr>
              <tr>
                <th rowspan="1" colspan="1">
                  <bold>COI</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>28S</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>AKAP17</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>FLAD1</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Mitogenome</bold>
                </th>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">DLS_35</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Beijing: Dongling Mountain</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PX369606" ext-link-type="gen">PX369606</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484227" ext-link-type="gen">PZ484227</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495527" ext-link-type="gen">PZ495527</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495535" ext-link-type="gen">PZ495535</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PX369606" ext-link-type="gen">PX369606</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">L668_09_01</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Hebei: Wuling Mountain</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462180" ext-link-type="gen">PZ462180</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484228" ext-link-type="gen">PZ484228</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495528" ext-link-type="gen">PZ495528</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495536" ext-link-type="gen">PZ495536</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462180" ext-link-type="gen">PZ462180</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">SDRZ_03</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Shandong: Rizhao</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462182" ext-link-type="gen">PZ462182</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484230" ext-link-type="gen">PZ484230</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495530" ext-link-type="gen">PZ495530</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495537" ext-link-type="gen">PZ495537</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462182" ext-link-type="gen">PZ462182</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">SDHZ04_01</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Shandong: Heze</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462181" ext-link-type="gen">PZ462181</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484229" ext-link-type="gen">PZ484229</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495529" ext-link-type="gen">PZ495529</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495534" ext-link-type="gen">PZ495534</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462181" ext-link-type="gen">PZ462181</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">BJMTG01_II_3</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Beijing: Mentougou</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462178" ext-link-type="gen">PZ462178</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462178" ext-link-type="gen">PZ462178</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">345R_01</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
                <td rowspan="1" colspan="1">China: Hebei: Qinhuangdao</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484221" ext-link-type="gen">PZ484221</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ168347" ext-link-type="gen">PZ168347</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">345R_33</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
                <td rowspan="1" colspan="1">China: Hebei: Qinhuangdao</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ486062" ext-link-type="gen">PZ486062</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484222" ext-link-type="gen">PZ484222</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">345R_34</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
                <td rowspan="1" colspan="1">China: Hebei: Qinhuangdao</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ486063" ext-link-type="gen">PZ486063</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484223" ext-link-type="gen">PZ484223</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">345R_38</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
                <td rowspan="1" colspan="1">China: Hebei: Qinhuangdao</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484225" ext-link-type="gen">PZ484225</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495524" ext-link-type="gen">PZ495524</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495533" ext-link-type="gen">PZ495533</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462175" ext-link-type="gen">PZ462175</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">345R_39</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
                <td rowspan="1" colspan="1">China: Hebei: Qinhuangdao</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ486064" ext-link-type="gen">PZ486064</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">345R_40</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
                <td rowspan="1" colspan="1">China: Hebei: Qinhuangdao</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484224" ext-link-type="gen">PZ484224</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495525" ext-link-type="gen">PZ495525</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495532" ext-link-type="gen">PZ495532</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462176" ext-link-type="gen">PZ462176</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">345R_42</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
                <td rowspan="1" colspan="1">China: Hebei: Qinhuangdao</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ484226" ext-link-type="gen">PZ484226</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495526" ext-link-type="gen">PZ495526</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ495531" ext-link-type="gen">PZ495531</ext-link>
                </td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ462177" ext-link-type="gen">PZ462177</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">HNLN_GR_I1_02</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="henanensis">henanensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Henan: Luonling</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ288546" ext-link-type="gen">PQ288546</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ278996" ext-link-type="gen">PQ278996</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">HNLN_GR_I2_23</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="henanensis">henanensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Henan: Luonling</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ288549" ext-link-type="gen">PQ288549</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ278997" ext-link-type="gen">PQ278997</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">HNLN_GR_I2_34</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinensis">sinensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Henan: Luonling</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ288559" ext-link-type="gen">PQ288559</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ278998" ext-link-type="gen">PQ278998</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">HNLN_GR_I2_40</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinensis">sinensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Henan: Luonling</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ288561" ext-link-type="gen">PQ288561</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ278999" ext-link-type="gen">PQ278999</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">HNLN_GR_I3_01</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="japonica">japonica</tp:taxon-name-part></tp:taxon-name></italic> s. s.</td>
                <td rowspan="1" colspan="1">China: Henan: Luonling</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ288577" ext-link-type="gen">PQ288577</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PQ279000" ext-link-type="gen">PQ279000</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">360R02</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="japonica">japonica</tp:taxon-name-part></tp:taxon-name></italic> s. s.</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ168348" ext-link-type="gen">PZ168348</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ168348" ext-link-type="gen">PZ168348</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">CQWX11_45</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="wuxiensis">wuxiensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Chongqing: Wuxi</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PX552887" ext-link-type="gen">PX552887</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PX369605" ext-link-type="gen">PX369605</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">CQWX05_37</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="wuxiensis">wuxiensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Chongqing: Wuxi</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PX552881" ext-link-type="gen">PX552881</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ210301" ext-link-type="gen">PZ210301</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">CNWN02_01</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xinjiangensis">xinjiangensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Xinjiang: Shuanghe</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PX978833" ext-link-type="gen">PX978833</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ099866" ext-link-type="gen">PZ099866</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">nxGXJ1_1_1</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xinjiangensis">xinjiangensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">China: Ningxia: Xiji</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PX978843" ext-link-type="gen">PX978843</ext-link>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ210303" ext-link-type="gen">PZ210303</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">MN539609</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="japonica">japonica</tp:taxon-name-part></tp:taxon-name></italic> s. l.</td>
                <td rowspan="1" colspan="1">China: Jiangsu: Nanjing</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="MN539609" ext-link-type="gen">MN539609</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Enchyraeus">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irregularis">irregularis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PZ168348" ext-link-type="gen">PZ168348</ext-link>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesenchytraeus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anisodiverticulus">anisodiverticulus</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">
                  <ext-link xlink:href="PV591013" ext-link-type="gen">PV591013</ext-link>
                </td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
      <sec sec-type="Morphological analysis" id="sec5">
        <title>Morphological analysis</title>
        <p>External and internal characters were examined under a ZEISS stereomicroscope (ZEN v. 3.3 Pro). Measurements (to 0.1 mm) and dissections followed <xref ref-type="bibr" rid="B7">Blakemore et al. (2014)</xref> and <xref ref-type="bibr" rid="B39">Michaelsen (1892)</xref>. Terminology follows <xref ref-type="bibr" rid="B58">Xu and Xiao (2011)</xref>.</p>
      </sec>
      <sec sec-type="Molecular data generation" id="sec6">
        <title>Molecular data generation</title>
        <p>Genomic DNA was extracted from posterior tissues (TIANGEN kit). Sanger sequencing targeted four markers: mitochondrial COI, nuclear 28S ribosomal RNA gene, 28S; A-kinase anchor protein 17A, AKAP17; flavin adenine dinucleotide synthetase 1, FLAD1 primers in Table <xref ref-type="table" rid="T3">3</xref>. Sequences were assembled in MEGA 5 (<xref ref-type="bibr" rid="B53">Tamura et al. 2011</xref>); COI distances were calculated under Kimura two-parameter <abbrev xlink:title="Kimura two-parameter">K2P</abbrev> model (<xref ref-type="bibr" rid="B28">Kimura 1980</xref>). For mitogenome reconstruction, high-throughput sequencing (150 bp paired-end, BGI Genomics) was performed on selected specimens, assembled with MitoZ v. 2.4 (<xref ref-type="bibr" rid="B38">Meng et al. 2019</xref>), and circularization verified by circle_check.py (<xref ref-type="bibr" rid="B38">Meng et al. 2019</xref>). All sequences were deposited in GenBank (Table <xref ref-type="table" rid="T2">2</xref>).</p>
        <table-wrap id="T3" position="float" orientation="portrait">
          <label>Table 3.</label>
          <caption>
            <p>Primers used for PCR and sequencing in this study.</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <th rowspan="1" colspan="1">
                  <bold>Marker</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Primer</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Sequence (5’–3’)</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Round</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Source</bold>
                </th>
              </tr>
              <tr>
                <td rowspan="2" colspan="1">COI</td>
                <td rowspan="1" colspan="1">LCO1490</td>
                <td rowspan="1" colspan="1">GGTCAACAAATCATAAAGATATTGG</td>
                <td rowspan="2" colspan="1"/>
                <td rowspan="2" colspan="1">
                  <xref ref-type="bibr" rid="B20">Folmer et al. (1994)</xref>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">HCO2198</td>
                <td rowspan="1" colspan="1">TAAACTTCAGGGTGACCAAAAAATCA</td>
              </tr>
              <tr>
                <td rowspan="2" colspan="1">28S</td>
                <td rowspan="1" colspan="1">28sF1</td>
                <td rowspan="1" colspan="1">GAGTACGTGAAACCGTCTAG</td>
                <td rowspan="2" colspan="1"/>
                <td rowspan="2" colspan="1">
                  <xref ref-type="bibr" rid="B44">Pérez-Losada et al. (2009)</xref>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">28SR1</td>
                <td rowspan="1" colspan="1">CGTTTCGTCCCCAAGGCCTC</td>
              </tr>
              <tr>
                <td rowspan="4" colspan="1">AKAP17</td>
                <td rowspan="1" colspan="1">AKAP17-F1</td>
                <td rowspan="1" colspan="1">AAYTGGGARGTNATGGARAA</td>
                <td rowspan="2" colspan="1">Round1</td>
                <td rowspan="8" colspan="1">
                  <xref ref-type="bibr" rid="B35">Liu et al. (2025a)</xref>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">AKAP17-R1</td>
                <td rowspan="1" colspan="1">TCYTTRAACATNARYTTCAT</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">AKAP17-F2</td>
                <td rowspan="1" colspan="1">AARATGATHAARCCNGAYCARTT</td>
                <td rowspan="2" colspan="1">Round2</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">AKAP17-R2</td>
                <td rowspan="1" colspan="1">GCYTTNACRAANCCCATRTAYTC</td>
              </tr>
              <tr>
                <td rowspan="4" colspan="1">FLAD1</td>
                <td rowspan="1" colspan="1">FLAD1-F1</td>
                <td rowspan="1" colspan="1">GGNCCNACNCAYGAYGAYAT</td>
                <td rowspan="2" colspan="1">Round1</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">FLAD1-R1</td>
                <td rowspan="1" colspan="1">TTNGGRTGNGTRTTYTCCAT</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">FLAD1-F2</td>
                <td rowspan="1" colspan="1">TGYAARGCNTTYTTYGGNACNGA</td>
                <td rowspan="2" colspan="1">Round2</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">FLAD1-R2</td>
                <td rowspan="1" colspan="1">TTNACNCKCATRAAYTCNGGCCA</td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
      <sec sec-type="Phylogenetic inference" id="sec7">
        <title>Phylogenetic inference</title>
        <p>Phylogenetic analyses were performed on a concatenated dataset of 13 protein-coding genes (<abbrev xlink:title="13 protein-coding genes">PCGs</abbrev>) extracted from mitogenomes by a script of gbseqextractor_v2.py (<xref ref-type="bibr" rid="B38">Meng et al. 2019</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mesenchytraeus">Mesenchytraeus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anisodiverticulus">anisodiverticulus</tp:taxon-name-part></tp:taxon-name></italic> Chen, 2012 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Enchyraeus">Enchyraeus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irregularis">irregularis</tp:taxon-name-part></tp:taxon-name></italic> Vejdovsky, 1879 were used as outgroup. Sequences were aligned using MAFFT (<xref ref-type="bibr" rid="B52">Rozewicki et al. 2019</xref>), and the best-fit nucleotide substitution model was selected with jModelTest-NG (<xref ref-type="bibr" rid="B12">Darriba et al. 2020</xref>) under the Akaike Information Criterion (<abbrev xlink:title="Akaike Information Criterion">AIC</abbrev>), and the best model is GTR + I + G. Maximum-likelihood (<abbrev xlink:title="Maximum-likelihood">ML</abbrev>) and Bayesian inference (<abbrev xlink:title="Bayesian inference">BI</abbrev>) trees were reconstructed using RAxML-NG (<xref ref-type="bibr" rid="B32">Kozlov et al. 2019</xref>) and MrBayes 3.2.6 (<xref ref-type="bibr" rid="B51">Ronquist et al. 2012</xref>), respectively. <abbrev xlink:title="Maximum-likelihood">ML</abbrev> nodal support was assessed by 1,000 rapid bootstrap replicates, with bootstrap values (<abbrev xlink:title="bootstrap values">BV</abbrev>) ≥ 70 considered well supported. <abbrev xlink:title="Bayesian inference">BI</abbrev> comprised two million MCMC generations, with convergence confirmed by split frequencies &lt; 0.01 and check ESS values &gt; 200 in Tracer v. 1.7.5 (<xref ref-type="bibr" rid="B49">Rambaut et al. 2018</xref>). Posterior probabilities (<abbrev xlink:title="Posterior probabilities">PP</abbrev>) ≥ 0.95 were accepted as strong support. The resulting trees were visualized in FigTree v. 1.4.4 (<ext-link xlink:href="https://tree.bio.ed.ac.uk/software/figtree/" ext-link-type="uri">https://tree.bio.ed.ac.uk/software/figtree/</ext-link>).</p>
      </sec>
      <sec sec-type="Species delimitation" id="sec8">
        <title>Species delimitation</title>
        <p>Species boundaries were assessed using Assemble Species by Automatic Partitioning (<abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev>, <xref ref-type="bibr" rid="B45">Puillandre et al. 2021</xref>), the Generalized Mixed Yule Coalescent (<abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev>; <xref ref-type="bibr" rid="B15">Drummond et al. 2012</xref>), and Bayesian Phylogenetics and Phylogeography (<abbrev xlink:title="Bayesian Phylogenetics and Phylogeography">BPP</abbrev>; <xref ref-type="bibr" rid="B61">Yang and Rannala 2010</xref>). Single-locus analyses of COI and 28S were performed with <abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev> (<abbrev xlink:title="Kimura two-parameter">K2P</abbrev> distances, Pmin = 0.001, <italic>P</italic><sub>max</sub> = 0.1, steps = 10, <italic>X</italic> = 1.0, 20 bins) and <abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev> (ultrametric trees inferred in BEAST v. 1.7.5 under Yule prior and strict clock; <abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev> performed using the splits package in R; <xref ref-type="bibr" rid="B17">Ezard et al. 2017</xref>). Multi-locus validation used <abbrev xlink:title="Bayesian Phylogenetics and Phylogeography">BPP</abbrev> v. 2.2 (<xref ref-type="bibr" rid="B61">Yang and Rannala 2010</xref>) on the four-gene dataset under the multispecies coalescent model, with an A00 analysis (estimating θ and τ) and an A10 analysis (species delimitation). Prior settings: θ ~ G (3, 1000); root τ0 ~ G (2, 2000); other τ parameters assigned a uniform Dirichlet distribution (<xref ref-type="bibr" rid="B59">Yang 2002</xref>, <xref ref-type="bibr" rid="B60">2015</xref>; <xref ref-type="bibr" rid="B50">Rannala and Yang 2003</xref>; <xref ref-type="bibr" rid="B19">Flouri et al. 2018</xref>).</p>
      </sec>
    </sec>
    <sec sec-type="Results" id="sec9">
      <title>Results</title>
      <sec sec-type="Taxonomy" id="sec10">
        <title>Taxonomy</title>
        <p>
          <bold>Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Moniligastridae">Moniligastridae</tp:taxon-name-part></tp:taxon-name> Claus, 1880</bold>
        </p>
        <sec sec-type="Genus Drawida Michaelsen, 1900" id="sec11">
          <title>Genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part></tp:taxon-name></italic> Michaelsen, 1900</title>
          <tp:taxon-treatment>
            <tp:treatment-meta>
              <kwd-group>
                <label>Taxon classification</label>
                <kwd>
                  <named-content content-type="kingdom">Animalia</named-content>
                </kwd>
                <kwd>
                  <named-content content-type="order">Moniligastrida</named-content>
                </kwd>
                <kwd>
                  <named-content content-type="family">Moniligastridae</named-content>
                </kwd>
              </kwd-group>
            </tp:treatment-meta>
            <tp:nomenclature>
              <tp:taxon-name><object-id content-type="arpha">A4EE0A76-F11C-5CFE-BDAC-28A992F717F9</object-id>
                        		<tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part>
                        	</tp:taxon-name>
              <tp:taxon-authority>Michaelsen, 1931</tp:taxon-authority>
              <xref ref-type="fig" rid="F1">Figs 1</xref>
              <xref ref-type="fig" rid="F3">, 3</xref>
              <tp:nomenclature-citation-list>
                <tp:nomenclature-citation>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  <comment>: <xref ref-type="bibr" rid="B40">Michaelsen 1931</xref>: 525.</comment>
                </tp:nomenclature-citation>
                <tp:nomenclature-citation>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  <comment> f. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida"/><tp:taxon-name-part taxon-name-part-type="species" reg="gisti"/><tp:taxon-name-part taxon-name-part-type="form" reg="typica">typica</tp:taxon-name-part></tp:taxon-name></italic>: <xref ref-type="bibr" rid="B10">Chen 1933</xref>: 195–200.</comment>
                </tp:nomenclature-citation>
                <tp:nomenclature-citation>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  <comment>: <xref ref-type="bibr" rid="B21">Gates 1935</xref>: 2–3.</comment>
                </tp:nomenclature-citation>
                <tp:nomenclature-citation>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  <comment>: <xref ref-type="bibr" rid="B23">Gates 1939</xref>: 406–408.</comment>
                </tp:nomenclature-citation>
                <tp:nomenclature-citation>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  <comment>: <xref ref-type="bibr" rid="B6">Blakemore et al. 2010</xref>: 1; <xref ref-type="bibr" rid="B5">Blakemore and Kupriyanova 2010</xref>: 8.</comment>
                </tp:nomenclature-citation>
              </tp:nomenclature-citation-list>
            </tp:nomenclature>
            <tp:treatment-sec sec-type="Material examined">
              <title>Material examined.</title>
              <p>• Five clitellates (SDRZ_03, SDRZ_04, L668_09_01, DLS_35, SDHZ4_01). • SDRZ_03, SDRZ_04: Rizhao, Shandong, China, Oct-2025, collected by Jiangni Li; • L668_09_01: Xinglong, Hebei, China, Jul-2025, collected by Huifeng Zhao; • DLS_35: Dongling Mountain Beijing, China, Jul-2025; • SDHZ4_01: Heze, Shandong, China, Oct-2025, collected by Yanmeng Bi. All specimens were preserved in the Hebei Key Laboratory of Animal Diversity, Langfang Normal University, Hebei, China (<abbrev content-type="institution" xlink:title="Hebei Key Laboratory of Animal Diversity, Langfang Normal University, Hebei, China">C-HLU</abbrev>).</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="External characteristics (Fig. 1)">
              <title>External characteristics (Fig. <xref ref-type="fig" rid="F1">1</xref>).</title>
              <p>Length 48–66 mm, diameter 5.0–7.0 mm. Segments 135–194. The setae begin on II and are closely paired; <italic>aa</italic> slightly less than <italic>bc</italic>. Setae clearly visible. Prostomium prolobous. Dorsal pores absent. The clitellar coloration variable, extending across segments IX/X–XIII/XIV. A pair of spermathecal pores are broad transverse slits on 7/8, at the middle of the <italic>cd</italic>, with the anterior and posterior margins swollen so that the pores appear elevated. The female pores are very minute, narrow transverse slits on 11/12, the slits in <italic>ab</italic>, external invisible. On 10/11 there is a larger transverse slit, in which a pair of penes is situated in <italic>bc</italic>, slightly nearer to <italic>b</italic> than to <italic>c</italic>. The true male pore is much smaller and opens at the ventral end of the penis as a longitudinal slit, about half the length of the penis. The penis of the examined specimens is 2.827 mm in length.</p>
              <fig id="F1">
                <object-id content-type="doi">10.3897/zookeys.1293.204408.figure1</object-id>
                <object-id content-type="arpha">C9C59EEA-E1CE-5DA2-8F70-74B786E97576</object-id>
                <label>Figure 1.</label>
                <caption>
                  <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> (SDRZ_03). <bold>A</bold>. Ventral view of the prostomium; <bold>B</bold>. Dorsal view of clitellum region; <bold>C, K</bold>. Ventral view of male pores, spermathecal pores, seta, and genital markings; <bold>D</bold>. Lateral view of the spermathecal pores, penis, and clitellum; <bold>E</bold>. Dorsal view of gizzards, ovisacs, and ovary chamber; <bold>F, J</bold>. Dorsal view of right testis sac, with vas deferens and prostate gland; <bold>G</bold>. Dorsal view of left testis sac and penial chamber; <bold>H, I</bold>. Dorsal view of right spermathecae. Scale bars: 1.0 mm (<bold>A, B</bold>); 0.5 mm (<bold>C–K</bold>).</p>
                </caption>
                <graphic xlink:href="zookeys-1293-327_article-204408__-g001.jpg" id="oo_1795857.jpg">
                  <uri content-type="original_file">https://binary.pensoft.net/fig/1795857</uri>
                </graphic>
              </fig>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Internal characteristics (Fig. 1)">
              <title>Internal characteristics (Fig. <xref ref-type="fig" rid="F1">1</xref>).</title>
              <p>Hearts from V to IX. Septa 5/6, 6/7, 7/8 and 8/9 thickened. Three gizzards in IX/X–XV/XVII/XVIII, occupying seven or nine segments. Testis sacs in two pairs, kidney-shaped, the concave side directed ventrally; in IX–X, or IX, or IX–XI, unconstricted by 9/10, but enveloped by fine membranes and fixed in the ventral body cavity. From the middle of each testis sac near the ventral median line a slender vas deferens emerges, first coiled into a tight bunch of loops on the anterior face of 9/10, then into another tight bunch on the posterior face; the anterior bunch smaller. The vas deferens finally opens on the ventral body wall of segment X. Prostate glands are in two parts: glandular portion and the prostatic duct. The glandular portion is covered with an outer layer of glandular tissue; when the outer layer is removed, a firm, slender, glossy central body is visible (Fig. <xref ref-type="fig" rid="F3">3</xref>), gradually narrowing toward the distal end; near the penial chamber the prostatic duct is visible, narrower than the glandular portion and lacking the outer granular tissue, 1.048 mm in length, finally opening at the dorsal center of the penial chamber. The penial chambers are in 10/11, projecting conspicuously into the body cavity, however, this projection is not present in all individuals (in specimen SDRZ_03 the right penial chamber was not observed). The prostates are 7.638 mm in length (including the glandular portion and the terminal prostatic duct). The middle portion is bent into a sort of C-shape and is placed around the penial chambers. The ovary chambers present, ovaries in X, ovisacs in X–XIII/XVI/XVIII, The spermathecae are in VIII; the ampulla small, 0.849 × 0.716 mm (SDRZ_03); the duct tightly coiled, of moderate length, gradually widening toward the ampulla; the atrium large, comma-shaped, 2.373 × 1.726 mm (SDRZ_03). On the inner side of the atrium there is a characteristic swelling, this swelling is an intrinsic part of the atrium (a localized dilation of its wall), and should not be confused with the accessory gland, although it superficially resembles the latter but is noticeably larger.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Distribution">
              <title>Distribution.</title>
              <p>The eastern China (except northeastern China).</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Remarks">
              <title>Remarks.</title>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anqingiana">anqingiana</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="nanchangiana">nanchangiana</tp:taxon-name-part></tp:taxon-name></italic> share several morphological similarities: all three species possess three gizzards, and their prostates terminate in a slender duct connecting to the penial chamber, and ovarian chambers surrounded by septa, and spermathecal ampulla are relatively small with coiled ducts. The differences among the three are as follows: the prostate of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="nanchangiana">nanchangiana</tp:taxon-name-part></tp:taxon-name></italic> measures 2.0–2.5 mm in length, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> is 7.6 mm, and reaching a maximum of 2.0 mm of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anchingiana">anchingiana</tp:taxon-name-part></tp:taxon-name></italic>; the spermathecal atrium of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> is a larger (2.4 mm in length), that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anchingiana">anchingiana</tp:taxon-name-part></tp:taxon-name></italic> is a smaller, short-cylindrical to globose structure reaching a maximum of 2.0 mm, and that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">g.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="nanchangiana">nanchangiana</tp:taxon-name-part></tp:taxon-name></italic> is thumb-shaped and small. Prostate morphology differs substantially among <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> and the other species of the complex. In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cheni">cheni</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="linhaiensis">linhaiensis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinica">sinica</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sulcata">sulcata</tp:taxon-name-part></tp:taxon-name></italic>, the prostates are cylindrical, relatively long, but curved in various configurations. In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="syringa">syringa</tp:taxon-name-part></tp:taxon-name></italic>, however, the prostates are shorter and possess an additional outer glandular layer; comparative measurements are given in Table <xref ref-type="table" rid="T1">1</xref>.</p>
              <p>Moreover, there is variation in penis length within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>. SDRZ_03 and SDRZ_04 possess long, well-developed penises, which is consistent with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>.The penes of the remaining specimens are relatively small and lamellate, presumably due to morphological variation induced by differences in habitat environment. Additionally, these specimens exhibit some color variation, yet their internal morphology is highly uniform.</p>
            </tp:treatment-sec>
          </tp:taxon-treatment>
          <tp:taxon-treatment>
            <tp:treatment-meta>
              <kwd-group>
                <label>Taxon classification</label>
                <kwd>
                  <named-content content-type="kingdom">Animalia</named-content>
                </kwd>
                <kwd>
                  <named-content content-type="order">Moniligastrida</named-content>
                </kwd>
                <kwd>
                  <named-content content-type="family">Moniligastridae</named-content>
                </kwd>
              </kwd-group>
            </tp:treatment-meta>
            <tp:nomenclature>
              <tp:taxon-name><object-id content-type="arpha">EA8B5612-20BA-5C52-8E01-E9CB3F96DE1E</object-id>
                        		<tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part>
                        	
                        		<object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/BE7F4BB7-F944-44BF-AA5D-F70CD7E358EA</object-id>
                        	</tp:taxon-name>
              <tp:taxon-authority>Qiao &amp; Zhao</tp:taxon-authority>
              <tp:taxon-status>sp. nov.</tp:taxon-status>
              <xref ref-type="fig" rid="F2">Fig. 2</xref>
            </tp:nomenclature>
            <tp:treatment-sec sec-type="Material examined">
              <title>Material examined.</title>
              <p><bold><italic>Holotype</italic></bold>: • One clitellate (mature) specimen (345R_39) (87 mm in total length, dissected), Weiyuan Park, Shanhaiguan District, Qinhuangdao Prefecture, Hebei, China, <named-content content-type="dwc:verbatimCoordinates">40.0139°N, 119.7485°E</named-content>, 21-Aug-2019, collected by Yufeng Zhang. <bold><italic>Paratypes</italic></bold>: • Five clitellates (345R_33, 345R_34, 345R_37, 345R_38, 345R_40), data the same as the holotype. All specimens were preserved in <abbrev content-type="institution" xlink:title="Hebei Key Laboratory of Animal Diversity, Langfang Normal University, Hebei, China">C-HLU</abbrev>.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Diagnosis">
              <title>Diagnosis.</title>
              <p>Length 48–87 mm, diameter 4.5–7.0 mm. Segments 160–181. Body light gray in 95% ethanol. Prostomium prolobous, dorsal pores absent. Clitellum pale pink, on segments X–XIII. Gizzards 3–5 in number. Vas deferens very short and uncoiled, a key character distinguishing it from most congeners in the <abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev>. Prostates distinctly M-shaped, having a very slender terminal prostatic duct. Penes thick, with swollen base and blunt rounded apex; male pores minute, opens on the ventral face of the penis. Ovaries enclosed in ovarian chambers in segments XI–XII. Spermathecae situated in segment VIII, with large ampulla and small elliptical atrium.</p>
              <fig id="F2">
                <object-id content-type="doi">10.3897/zookeys.1293.204408.figure2</object-id>
                <object-id content-type="arpha">9A8587A2-1C34-5048-A2AA-CB448F7E1DF7</object-id>
                <label>Figure 2.</label>
                <caption>
                  <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., holotype (345R_39). <bold>A</bold>. Ventral view of the prostomium; <bold>B</bold>. Dorsal view of clitellum region; <bold>C, K</bold>. Ventral view of male pores, spermathecal pores and genital markings; <bold>D</bold>. Lateral view of the spermathecal pores, penis and clitellum; <bold>E</bold>. Dorsal view of ovisacs, ovaries and ovary chamber; <bold>F</bold>. Dorsal view of gizzards; <bold>G, J</bold>. Dorsal view of right testis sac, with vas deferens and prostate gland; <bold>H, I</bold>. Dorsal view of right spermathecae. Scale bars: 1.0 mm (<bold>A–D, K</bold>); 0.5 mm (<bold>E–J</bold>).</p>
                </caption>
                <graphic xlink:href="zookeys-1293-327_article-204408__-g002.jpg" id="oo_1795858.jpg">
                  <uri content-type="original_file">https://binary.pensoft.net/fig/1795858</uri>
                </graphic>
              </fig>
              <fig id="F3">
                <object-id content-type="doi">10.3897/zookeys.1293.204408.figure3</object-id>
                <object-id content-type="arpha">658EE4DB-737D-5BE0-B44D-0C18D32E7A76</object-id>
                <label>Figure 3.</label>
                <caption>
                  <p>Morphology of the prostatic central body. <bold>A</bold>. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> (SDRZ_04); <bold>B</bold>. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="wuxiensis">wuxiensis</tp:taxon-name-part></tp:taxon-name></italic>; <bold>C</bold>. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xinjiangensis">xinjiangensis</tp:taxon-name-part></tp:taxon-name></italic>. Scale bars: 0.5 mm.</p>
                </caption>
                <graphic xlink:href="zookeys-1293-327_article-204408__-g003.jpg" id="oo_1795859.jpg">
                  <uri content-type="original_file">https://binary.pensoft.net/fig/1795859</uri>
                </graphic>
              </fig>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="External characteristics">
              <title>External characteristics.</title>
              <p>Holotype: length 87 mm, diameter 7.0 mm. Segments 164. The setae begin on II and are closely paired; <italic>ab</italic>:<italic>cd</italic>:<italic>aa</italic>:<italic>bc</italic> = 1:1<italic>ab</italic>:10<italic>ab</italic>:10<italic>ab</italic>. Prostomium prolobous. Dorsal pores absent or invisible. Body color brown, head greyish blue, clitellum pale pink. Clitellum X–XIII. The spermathecal pores are narrow transverse slits on 7/8, at <italic>bc</italic>, close to <italic>c</italic> than <italic>b</italic>. The female pores are very minute, narrow transverse slits on 11/12, external invisible. On 10/11 there is a larger transverse slit, in which a pair of penes are situated in <italic>bc</italic>, slightly nearer to <italic>b</italic> than to <italic>c</italic>. The true male pore is much smaller and opens on the ventral face of the penis but is inconspicuous, no longitudinal or transverse slit observed. The left penis is 0.929 mm in length; the right is 1.066 mm in length. The penes of this new species are rather thick, swollen at the base and gradually tapering and the apex blunt resembling a triangular cone. Genital markings present, irregular and three in number, on segment VIII. The genital markings are mainly concentrated around the spermathecal pores.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Internal characteristics">
              <title>Internal characteristics.</title>
              <p>Holotype: hearts from V to IX. Septa 5/6, 6/7, 7/8 and 8/9 thickened. The gizzards three to five, in IX–XVI, XII/XIII–XVIII or XI–XVII, occupying six, seven or eight segments. The testis sacs two pairs, 2.362 × 3.545 mm in size, elliptical; in IX–X, or IX/X, or IX–XI, occupying one to two segments, occasionally three, unconstricted by 9/10. A very short, uncoiled vas deferens arises from the margin at two-thirds of the testis sac, finally opening on the ventral body wall of segment X, 2.977 mm in length. One pair of prostates are M-shaped, 11.614 mm in length, only the glandular portion enveloped by granulations (external glandular layer), and when this outer layer is removed a smooth central body is visible. The prostate width gradually narrows, 0.828 mm at the apex and 0.302 mm at the distal end. On the inner side of the prostate near the ventral median line there are one or two accessory glands. The penial chambers in 10/11. A very short prostatic duct arises from the apex of the prostate, 0.437 mm in length, lying close to the opening of the vas deferens. The ovaries are on each side of the gizzard, in XI–XII, inverted U-shaped; the septa form ovarian chambers, and when these are removed the white ovaries are visible. The ovisacs appear as brown sacs covering the gizzard, extending to segment XXI. The spermathecae are in VIII; the ampulla large, milky white, 1.497 × 1.694 mm in size; the atrium small, elliptical, pale yellow, 1.853 × 0.859 mm in size, situated on the septum 7/8. The duct tightly coiled, gradually widening from the end of the ampulla and finally ending in the middle of the atrium; the anterior portion 0.129 mm in width, the posterior portion 0.173 mm in width, the duct about 10.0 mm in length.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Variation">
              <title>Variation.</title>
              <p>In the paratypes, the genital markings are irregular, unpaired, and scattered on segments VII–XI. The number and position of gizzards vary as follows: specimen 345R_34 has three gizzards in segments 13–18; 345R_33 has five gizzards in segments 9–16; 345R_38 has four gizzards in segments 9–16; 345R_37 has three gizzards in segments 12–18; and 345R_40 has three gizzards in segments 11–17. Thus, the gizzard number ranges from three to five, and the segmental distribution ranges from nine to 18 among the paratypes. This variation is considered intraspecific, as is commonly observed in the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part></tp:taxon-name></italic>. The testis sacs are two pairs, elliptical, measuring 2.362 × 3.545 mm, and occur in IX–X, or at IX/X, or in IX–XI, occupying one or two segments (occasionally three), and are not constricted by septum 9/10. The vas deferens ranges from 2.208 to 3.113 mm in length. The prostates are one pair, M-shaped, 4.705–11.614 mm long. The spermathecal ampulla is large, milky white, about 1.43 × 1.47 mm; the atrium is small, elliptical, pale yellow, about 1.585 × 0.859 mm. The ovisacs appear as brown sacs covering the gizzard and extend to segments XVI–XX.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Etymology">
              <title>Etymology.</title>
              <p>The specific epithet <italic>xuqini</italic> is a patronymic genitive noun formed, in honor of Mr Qin Xu, the first author of “Terrestrial Earthworms (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Opisthopora">Opisthopora</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subclass" reg="Oligochaeta">Oligochaeta</tp:taxon-name-part></tp:taxon-name>) of China”.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Distribution">
              <title>Distribution.</title>
              <p>Known only from the type locality (Qinhuangdao, Hebei).</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Remarks">
              <title>Remarks.</title>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is readily distinguished from other members of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> species complex by a combination of clear morphological characters. The penis is thick, with a swollen base and a blunt, rounded apex, whereas in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> it is slender, elongated, and tapers to a sharp tip. The new species possesses 3–5 gizzards located in segments 9–18 (with intraspecific variation), while <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> and most other species consistently have three gizzards. The vas deferens is very short and entirely uncoiled, a unique character among the complex (vs long and coiled in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="anchingiana">anchingiana</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">g.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="nanchangiana">nanchangiana</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="linhaiensis">linhaiensis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinica">sinica</tp:taxon-name-part></tp:taxon-name></italic>, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="syringa">syringa</tp:taxon-name-part></tp:taxon-name></italic>; short but coiled in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sulcata">sulcata</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cheni">cheni</tp:taxon-name-part></tp:taxon-name></italic>). The spermathecal atrium is elliptical and large in the new species (comma-shaped in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>), and the prostate is M-shaped without a slender terminal duct. The body color in alcohol is light gray (vs reddish brown in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>).</p>
            </tp:treatment-sec>
          </tp:taxon-treatment>
        </sec>
      </sec>
      <sec sec-type="Phylogenetic analyses" id="sec12">
        <title>Phylogenetic analyses</title>
        <p>The mitogenomes of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. are 14,647–14,648 bp and 14,668–14,750 bp in length, respectively, and all of them contain the typical mitochondrial 37 genes: 13 protein-coding genes (<abbrev xlink:title="13 protein-coding genes">PCGs</abbrev>), two ribosomal RNA genes (<abbrev xlink:title="two ribosomal RNA genes">rRNAs</abbrev>), and 22 transfer RNA genes (<abbrev xlink:title="22 transfer RNA genes">tRNAs</abbrev>). (Fig. <xref ref-type="fig" rid="F4">4A, B</xref>). The control region (<abbrev xlink:title="control region">CR</abbrev>) is absent in both species examined here, unlike its typical presence in earthworm mitogenomes (<xref ref-type="bibr" rid="B66">Zhao et al. 2022</xref>) but similar to some species of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="japonica">japonica</tp:taxon-name-part></tp:taxon-name></italic> species complex (<abbrev xlink:title="Drawida japonica species complex">DjSC</abbrev>) (<xref ref-type="bibr" rid="B63">Zhang et al. 2016</xref>; <xref ref-type="bibr" rid="B46">Qiao et al. 2026a</xref>).</p>
        <fig id="F4">
          <object-id content-type="doi">10.3897/zookeys.1293.204408.figure4</object-id>
          <object-id content-type="arpha">EF73D32F-9093-57AB-AAD8-C1C46524F015</object-id>
          <label>Figure 4.</label>
          <caption>
            <p><bold>A, B</bold>. Mitogenomes of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.; <bold>C</bold>. Phylogenetic relationship of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. and other species in <abbrev xlink:title="Drawida japonica species complex">DjSC</abbrev> and <abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev> using 13 coding genes of mitogenomes. Numbers near branches indicate the maximum likelihood bootstrap support/Bayesian posterior probabilities. Values with a bootstrap value below 50 and a posterior probability lower than 80 have been omitted. Orange tips showed the new species.</p>
          </caption>
          <graphic xlink:href="zookeys-1293-327_article-204408__-g004.jpg" id="oo_1795860.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1795860</uri>
          </graphic>
        </fig>
        <p>Phylogenetic analysis based on the concatenated 13 <abbrev xlink:title="13 protein-coding genes">PCGs</abbrev> (Fig. <xref ref-type="fig" rid="F4">4C</xref>) shows that <abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev> and <abbrev xlink:title="Drawida japonica species complex">DjSC</abbrev> forms two independent well-supported monophyletic groups. However, the lineage of <ext-link xlink:href="MN539609" ext-link-type="gen">MN539609</ext-link> identified as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B34">Liu et al. 2020</xref>) is clustered in <abbrev xlink:title="Drawida japonica species complex">DjSC</abbrev>. Within this, the <abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev>, four tips of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. are clustered as a distinct lineage with robust support in both <abbrev xlink:title="Maximum-likelihood">ML</abbrev> and <abbrev xlink:title="Bayesian inference">BI</abbrev> analyses (<abbrev xlink:title="Maximum-likelihood">ML</abbrev> Bootstrap value = 100, <abbrev xlink:title="Bayesian inference">BI</abbrev> Posterior probability = 1.00). The same situation of five individuals of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>. However, SDRZ_03 shows genetic differences with the other four individuals.</p>
      </sec>
      <sec sec-type="Molecular species delimitation" id="sec13">
        <title>Molecular species delimitation</title>
        <p>Three methods—<abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev>, <abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev>, and <abbrev xlink:title="Bayesian Phylogenetics and Phylogeography">BPP</abbrev>—were employed to delimit molecular operational taxonomic units (<abbrev xlink:title="molecular operational taxonomic units">MOTUs</abbrev>) (Fig. <xref ref-type="fig" rid="F5">5</xref>). <abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev> (<abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev> score 1.50) based on COI split <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> into two <abbrev xlink:title="molecular operational taxonomic units">MOTUs</abbrev>, whereas <abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev> based on the same marker further subdivided <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> into three <abbrev xlink:title="molecular operational taxonomic units">MOTUs</abbrev> and split <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. into two <abbrev xlink:title="molecular operational taxonomic units">MOTUs</abbrev>. For the 28S dataset, <abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev> (<abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev> score 1.00) results are the same as morphological evidence (support <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> and new species are independent), but <abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev> recovered two <abbrev xlink:title="molecular operational taxonomic units">MOTUs</abbrev> within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</p>
        <fig id="F5">
          <object-id content-type="doi">10.3897/zookeys.1293.204408.figure5</object-id>
          <object-id content-type="arpha">C2599807-F510-5877-A1F7-9B5D0A5C95A7</object-id>
          <label>Figure 5.</label>
          <caption>
            <p>Species delimitation of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> species complex. The black vertical bar shows the number of species delimited by morphological characteristics and molecular data. The upper left shows field photographs of the new species.</p>
          </caption>
          <graphic xlink:href="zookeys-1293-327_article-204408__-g005.jpg" id="oo_1795861.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1795861</uri>
          </graphic>
        </fig>
        <p><abbrev xlink:title="Bayesian Phylogenetics and Phylogeography">BPP</abbrev> analysis of the four-locus dataset consistently distinguished <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. as two distinct species congruent with morphological evidence. The <abbrev xlink:title="Kimura two-parameter">K2P</abbrev> distances based on COI further supported the genetic integrity of the new species: intraspecific divergence ranged from 0 to 0.5%, whereas distances between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> fell between 19.4% and 22.0% (Table <xref ref-type="table" rid="T4">4</xref>), well above the 15% interspecific threshold routinely applied in earthworm taxonomy (<xref ref-type="bibr" rid="B14">Dong et al. 2019</xref>; <xref ref-type="bibr" rid="B9">Chang et al. 2024</xref>; <xref ref-type="bibr" rid="B67">Zhao et al. 2025</xref>). The subdivision of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> by single-locus methods suggests potential cryptic diversity within this nominal species; however, expanded molecular sampling across the entire <abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev> is required to test this hypothesis and clarify species boundaries. For detailed results of molecular species delimitation, see the Suppl. materials <xref ref-type="supplementary-material" rid="S1">1</xref>–<xref ref-type="supplementary-material" rid="S3">3</xref>.</p>
        <table-wrap id="T4" position="float" orientation="portrait">
          <label>Table 4.</label>
          <caption>
            <p>The COI genetic distance of <abbrev xlink:title="Kimura two-parameter">K2P</abbrev> of groups of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part></tp:taxon-name></italic> (values are percentages; intraspecific genetic distances are given in parentheses).</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <th rowspan="1" colspan="1"/>
                <th rowspan="1" colspan="1">
                  <bold>Species</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>1</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>2</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>3</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>4</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>5</bold>
                </th>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">1</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
                <td rowspan="1" colspan="1">(0–0.5)</td>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1"/>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">2</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">19.4–22.0</td>
                <td rowspan="1" colspan="1">(0.3–5.7)</td>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1"/>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">3</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="henanensis">henanensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">19.9–23.5</td>
                <td rowspan="1" colspan="1">22.6–27.1</td>
                <td rowspan="1" colspan="1">(0.–5.7)</td>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1"/>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">4</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="japonica">japonica</tp:taxon-name-part></tp:taxon-name></italic> s. s.</td>
                <td rowspan="1" colspan="1">23.3–24.3</td>
                <td rowspan="1" colspan="1">22.3–23.4</td>
                <td rowspan="1" colspan="1">19.2–22.8</td>
                <td rowspan="1" colspan="1">(0–0.2)</td>
                <td rowspan="1" colspan="1"/>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">5</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinensis">sinensis</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
                <td rowspan="1" colspan="1">22.3–23.8</td>
                <td rowspan="1" colspan="1">20.8–22.0</td>
                <td rowspan="1" colspan="1">20.2–22.5</td>
                <td rowspan="1" colspan="1">17.6–22.2</td>
                <td rowspan="1" colspan="1">(13.3)</td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
    </sec>
    <sec sec-type="Discussion" id="sec14">
      <title>Discussion</title>
      <p>Morphological examinations reveal considerable external plasticity in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic>, with body color and size varying markedly among specimens from different localities. This external plasticity contrasts with the relatively conserved internal characters, which show only minor variations across specimens. Such disparity complicates species delimitation in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part></tp:taxon-name></italic>, underscoring the necessity of integrative taxonomic approaches (<xref ref-type="bibr" rid="B13">Dayrat 2005</xref>; <xref ref-type="bibr" rid="B43">Padial et al. 2010</xref>; <xref ref-type="bibr" rid="B35">Liu et al. 2025a</xref>, <xref ref-type="bibr" rid="B36">2025b</xref>). For example, mitogenome <ext-link xlink:href="MN539609" ext-link-type="gen">MN539609</ext-link> was previously misidentified as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> by <xref ref-type="bibr" rid="B34">Liu et al. (2020)</xref>, likely due to insufficient morphological expertise and the lack of authentic DNA sequences for this species.</p>
      <p>Notably, specimens with divergent penial morphology from Rizhao populations showed <abbrev xlink:title="Kimura two-parameter">K2P</abbrev> genetic distances of 4.8–5.7% from typical <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> populations, and species-delimitation analyses using <abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev> and <abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev> (based on COI) all recovered them as distinct evolutionary lineages. However, it should be noted that <abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev> tends to separate biological species into several OTUs due to excessive lineage splitting (<xref ref-type="bibr" rid="B25">Goulpeau et al. 2022</xref>), and its application to mitochondrial COI data often yields results incongruent with other delimitation methods (<xref ref-type="bibr" rid="B36">Liu et al. 2025b</xref>). Among the northern specimens examined, only SDRZ_03 and SDRZ_04 exhibited penial morphology consistent with the historical description of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B40">Michaelsen 1931</xref>); the remaining specimens matched <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> in most other internal and external features except for the penis. This suggests that the northern populations likely do not represent simple morphological variants of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> from northward dispersal but rather may constitute cryptic species or novel taxa within the complex awaiting further validation through expanded sampling.</p>
      <p>The discovery of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. at Shanhaiguan in northeastern China hints at an expansion of distribution in <abbrev xlink:title="Drawida gisti species complex">DiSC</abbrev>. The Shanhaiguan population may represent a peripheral isolate that diverged through geographic isolation and local adaptation under different climatic regimes. Conversely, given the wide distribution of allied taxa across eastern China, additional cryptic species may await discovery in the southern portion of the range. Both hypotheses remain to be tested through denser sampling across the northern and southern provinces of China.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>We thanked the anonymous referee for his improvement of the manuscript, and International Joint Research Center for Invertebrate Biology in Hebei Province.</p>
    </ack>
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    <sec sec-type="Additional information" id="sec15">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec16">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec17">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec18">
        <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="sec19">
        <title>Funding</title>
        <p>This study is funded by Science &amp; Technology Fundamental Resources Investigation Program (Grant No. 2025FY100800 and 2024FY100400) and the Hebei Natural Science Foundation (Grant No. C2025408018).</p>
      </sec>
      <sec sec-type="Author contributions" id="sec20">
        <title>Author contributions</title>
        <p>Conceptualization, Qiao S., Li J., Zhang Y. and Zhao H.; specimen identification, Qiao S. and Zhao H.; methodology and experiments, Qiao S. and Zhao H.; data analysis, Qiao S. and Zhao H.; writing-original draft prepara-tion, Qiao S. and Zhao H.; writing-review and editing, Qiao S., Li J., Zhang Y. and Zhao H.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec21">
        <title>Author ORCIDs</title>
        <p>S. Qiao <ext-link xlink:href="https://orcid.org/0009-0002-9964-1352" ext-link-type="uri">https://orcid.org/0009-0002-9964-1352</ext-link></p>
        <p>J. Li <ext-link xlink:href="https://orcid.org/0000-0003-1642-0328" ext-link-type="uri">https://orcid.org/0000-0003-1642-0328</ext-link></p>
        <p>Y. Zhang <ext-link xlink:href="https://orcid.org/0000-0002-4066-6355" ext-link-type="uri">https://orcid.org/0000-0002-4066-6355</ext-link></p>
        <p>N.M. Aspe <ext-link xlink:href="https://orcid.org/0000-0003-0025-6017" ext-link-type="uri">https://orcid.org/0000-0003-0025-6017</ext-link></p>
        <p>H. Zhao <ext-link xlink:href="https://orcid.org/0000-0003-4243-9671" ext-link-type="uri">https://orcid.org/0000-0003-4243-9671</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec22">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text or Supplementary Information.</p>
      </sec>
    </sec>
    <fn-group>
      <fn>
        <p>Saiyu Qiao and Jiangni Li contributed equally to this work.</p>
      </fn>
    </fn-group>
    <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">33B910B8-3824-5BFE-9456-DDE90CDE0864</object-id>
        <label>Supplementary material 1</label>
        <caption>
          <p><abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev> species delimition results</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>. <abbrev xlink:title="Generalized Mixed Yule Coalescent">GMYC</abbrev> species delimition results based on COI (<bold>A</bold>) and 28S (<bold>B</bold>).</p>
        </statement>
        <media xlink:href="zookeys-1293-327_article-204408__-s001.pdf" mimetype="application" mime-subtype="pdf" position="float" orientation="portrait" id="oo_1795862.pdf">
          <uri content-type="original_file">https://binary.pensoft.net/file/1795862</uri>
        </media>
        <permissions>
          <license>
            <license-p>This dataset is made available under the Open Database License (<ext-link ext-link-type="uri" xlink:href="http://opendatacommons.org/licenses/odbl/1.0/">http://opendatacommons.org/licenses/odbl/1.0/</ext-link>). 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"> Saiyu Qiao, Jiangni Li, Yufeng Zhang, Nonillon M. Aspe, Huifeng Zhao</attrib>
      </supplementary-material>
      <supplementary-material id="S2" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="arpha">945D63F3-5C10-5706-BEF5-7AF4719DB434</object-id>
        <label>Supplementary material 2</label>
        <caption>
          <p>The result of molecular species delimitaiton</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 SS1</bold>. The result of molecular species delimitaiton of <abbrev xlink:title="Assemble Species by Automatic Partitioning">ASAP</abbrev> on COI and 28S in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">Drawida</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gisti">gisti</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="gisti">gisti</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drawida">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="xuqini">xuqini</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</p>
        </statement>
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          <uri content-type="original_file">https://binary.pensoft.net/file/1795863</uri>
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        <permissions>
          <license>
            <license-p>This dataset is made available under the Open Database License (<ext-link ext-link-type="uri" xlink:href="http://opendatacommons.org/licenses/odbl/1.0/">http://opendatacommons.org/licenses/odbl/1.0/</ext-link>). 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"> Saiyu Qiao, Jiangni Li, Yufeng Zhang, Nonillon M. Aspe, Huifeng Zhao</attrib>
      </supplementary-material>
      <supplementary-material id="S3" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="arpha">3586AC6E-B9A4-576E-A15D-A4A9E2891CC9</object-id>
        <label>Supplementary material 3</label>
        <caption>
          <p><abbrev xlink:title="Bayesian Phylogenetics and Phylogeography">BPP</abbrev> related files</p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>rar</p>
        </statement>
        <media xlink:href="zookeys-1293-327_article-204408__-s003.rar" mimetype="application" mime-subtype="x-rar-compressed" position="float" orientation="portrait" id="oo_1795864.rar">
          <uri content-type="original_file">https://binary.pensoft.net/file/1795864</uri>
        </media>
        <permissions>
          <license>
            <license-p>This dataset is made available under the Open Database License (<ext-link ext-link-type="uri" xlink:href="http://opendatacommons.org/licenses/odbl/1.0/">http://opendatacommons.org/licenses/odbl/1.0/</ext-link>). 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"> Saiyu Qiao, Jiangni Li, Yufeng Zhang, Nonillon M. Aspe, Huifeng Zhao</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
