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<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:tp="http://www.plazi.org/taxpub" article-type="research-article">
  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">1</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:F9B2E808-C883-5F47-B276-6D62129E4FF4</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:245B00E9-BFE5-4B4F-B76E-15C30BA74C02</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Biodiversity Data Journal</journal-title>
        <abbrev-journal-title xml:lang="en">BDJ</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1314-2836</issn>
      <issn pub-type="epub">1314-2828</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/BDJ.14.e211811</article-id>
      <article-id pub-id-type="publisher-id">211811</article-id>
      <article-id pub-id-type="manuscript">32017</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Taxonomy &amp; Inventories</subject>
        </subj-group>
        <subj-group subj-group-type="Taxon classification">
          <subject>Ephemeroptera</subject>
        </subj-group>
        <subj-group subj-group-type="Subject classification">
          <subject>Zoology &amp; Animal Biology</subject>
        </subj-group>
        <subj-group subj-group-type="Chronological classification">
          <subject>Cenozoic</subject>
        </subj-group>
        <subj-group subj-group-type="Geographical classification">
          <subject>China</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Reared adults of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 from China (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Ephemeroptera">Ephemeroptera</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Neoephemeridae">Neoephemeridae</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>Gong</surname>
            <given-names>Dewen</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-3740-7125</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Zhou</surname>
            <given-names>Changfa</given-names>
          </name>
          <email xlink:type="simple">zhouchangfa@njnu.edu.cn</email>
          <uri content-type="orcid">https://orcid.org/0000-0001-8785-5228</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Nanjing Normal University, Nanjing, China</addr-line>
        <institution>Nanjing Normal University</institution>
        <addr-line content-type="city">Nanjing</addr-line>
        <country>China</country>
        <uri content-type="ror">https://ror.org/036trcv74</uri>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Changfa Zhou (<email xlink:type="simple">zhouchangfa@njnu.edu.cn</email>).</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Jean-Luc Gattolliat</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>29</day>
        <month>09</month>
        <year>2026</year>
      </pub-date>
      <volume>14</volume>
      <elocation-id>e211811</elocation-id>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/D64BF688-C29F-52F7-A32A-BFBA67EAEA8D">D64BF688-C29F-52F7-A32A-BFBA67EAEA8D</uri>
      <history>
        <date date-type="received">
          <day>14</day>
          <month>08</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>18</day>
          <month>09</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Dewen Gong, Changfa Zhou</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <abstract>
        <label>Abstract</label>
        <sec sec-type="background">
          <title>Background</title>
          <p>The nymphs of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 possess filtering mouthparts and modified fore-legs. However, no adults are known.</p>
        </sec>
        <sec sec-type="new information">
          <title>New information</title>
          <p>In this study, both nymphal and imaginal stages of this species were collected and reared from China. Morphological examination reveals that these specimens differ markedly from all other members of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic> Lestage, 1931. In addition to the filtering nymphal morphology, the males of this species are characterised by more distinctly developed penes, longer tibiae and cerci and smaller compound eyes compared to those of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic>. Significantly, its distinct penes not only validates its independent species status, but also reveals the evolutionary process of the reduction of neoephemerid male genitalia and provides the original clue of the vestigial penes in several other lineages of the order <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Ephemeroptera">Ephemeroptera</tp:taxon-name-part></tp:taxon-name> as well.</p>
        </sec>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>evolution</kwd>
        <kwd>imago</kwd>
        <kwd>mayfly</kwd>
        <kwd>Oriental realm</kwd>
        <kwd>taxonomy</kwd>
      </kwd-group>
      <counts>
        <fig-count count="9"/>
        <table-count count="0"/>
        <ref-count count="19"/>
      </counts>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction">
      <title>Introduction</title>
      <p>The species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> was originally described by <xref ref-type="bibr" rid="B14406931">Nguyen and Bae (2003)</xref>, based on nymphs collected from Vietnam. Since then, no further report on this species has been published.</p>
      <p>Unlike any other member of the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Neoephemeridae">Neoephemeridae</tp:taxon-name-part></tp:taxon-name>, nymphs of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> possess filtering fore-legs, with both fore-tibiae and fore-tarsi bearing long, brush-like setae — a morphological trait typically observed in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Isonychiidae">Isonychiidae</tp:taxon-name-part></tp:taxon-name> or <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Oligoneuriidae">Oligoneuriidae</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B14406858">Qiang and Zhou 2023</xref>, <xref ref-type="bibr" rid="B14406769">Chen et al. 2025</xref>). Their mouthparts also exhibit a filtering configuration, as both maxillary and labial palpi are elongated and equipped with long hair-like setae. By contrast, all other genera and species within the same family are characterised by robust, chewing or cutting mouthparts and their bodies generally lack elongate setae (<xref ref-type="bibr" rid="B14406760">Bae and McCafferty 1998</xref>, <xref ref-type="bibr" rid="B14406831">Ma and Zhou 2021</xref>, <xref ref-type="bibr" rid="B14406822">Li et al. 2026</xref>). In their original description, <xref ref-type="bibr" rid="B14406931">Nguyen and Bae (2003)</xref> noted that these nymphs likely warranted the erection of a new genus, but emphasised that adult specimens were necessary for a definitive taxonomic decision.</p>
      <p>Currently, the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Neoephemeridae">Neoephemeridae</tp:taxon-name-part></tp:taxon-name> comprises five extant genera and 17 described species (<xref ref-type="bibr" rid="B14406760">Bae and McCafferty 1998</xref>, <xref ref-type="bibr" rid="B14406831">Ma and Zhou 2021</xref>, <xref ref-type="bibr" rid="B14406822">Li et al. 2026</xref>). Based on characters of the male imaginal genitalia, nymphal gills and caudal filaments, these genera can be divided into two distinct morphological groups: the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Neoephemera">Neoephemera</tp:taxon-name-part></tp:taxon-name></italic>-like group, characterised by well-developed male forceps, two operculate gills II that are connected or jointed medially and nymphal caudal filaments bearing dense spines; and the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic>-like group, which possesses vestigial male forceps, free gills II and nymphal tails with dense setae and spines. The former group includes four genera: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucorhoenanthus">Leucorhoenanthus</tp:taxon-name-part></tp:taxon-name></italic> Lestage, 1931; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Neoephemera">Neoephemera</tp:taxon-name-part></tp:taxon-name></italic> Mcdunnough, 1925; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ochernova">Ochernova</tp:taxon-name-part></tp:taxon-name></italic> Bae and McCafferty, 1998; and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pulchephemera">Pulchephemera</tp:taxon-name-part></tp:taxon-name></italic> Ma and Zhou, 2021 (<xref ref-type="bibr" rid="B14406949">Lestage 1931</xref>, <xref ref-type="bibr" rid="B14406849">Mcdunnough 1925</xref>, <xref ref-type="bibr" rid="B14406760">Bae and McCafferty 1998</xref>, <xref ref-type="bibr" rid="B14406831">Ma and Zhou 2021</xref>Lestage 1931, ), whereas the latter currently contains only a single genus, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic> Lestage, 1931 (<xref ref-type="bibr" rid="B14406787">Kluge 2004</xref>, <xref ref-type="bibr" rid="B14406831">Ma and Zhou 2021</xref>). Further discoveries within this second clade would not only enhance our understanding of its species diversity, but also provide valuable insights into its evolutionary trajectories.</p>
      <p>In spring 2026, we conducted a collecting expedition in Yunnan Province, south-western China. During this survey, dozens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> nymphs were collected. Through careful indoor rearing, more than ten adults were successfully obtained. These newly-acquired materials provide a valuable opportunity to describe the previously unknown adult stage and to reassess the taxonomic and phylogenetic status of this enigmatic species.</p>
    </sec>
    <sec sec-type="Materials and methods">
      <title>Materials and methods</title>
      <sec sec-type="Specimen collection and storage">
        <title>Specimen collection and storage</title>
        <p>Nymphs were collected using hand nets and adults were obtained both by rearing mature nymphs under laboratory conditions and by light trapping. All specimens were preserved in 80% or higher concentration ethanol and are currently deposited in the Mayfly Collection, College of Life Sciences, Nanjing Normal University (NNU), China.</p>
        <p>For rearing, mature nymphs were placed in trays containing natural water, along with small stones, twigs and leaves. Each tray was covered with a small tent constructed from mosquito netting. Throughout the rearing period, a small aerator was used to continuously supply air to the water. Emerging adults were collected from inside the tent as they emerged.</p>
      </sec>
      <sec sec-type="Illustrations">
        <title>Illustrations</title>
        <p>Morphological observations were conducted using a combination of imaging techniques. Whole-body images of both nymphs and adults were captured with a Sony A7RIV interchangeable-lens digital camera, equipped with a 90 mm macro lens. For intermediate-sized structures, such as male genitalia and hind-wings, a Sony A7RIV camera, fitted with a Laowa FF 25 mm F2.8 Ultra Macro lens, was used. Egg morphology was examined using a Thermo Fisher Scientific Apreo 2S scanning electron microscope (SEM).</p>
        <p>Eggs for SEM analysis were dissected from female imagoes. Together with the hind-legs of female subimagos, they were processed following a standard dehydration protocol: sequential immersion in an acetone gradient (70%, 80%, 90%, 98% and 100%, 10–20 min per step), followed by treatment with 1–1.5 ml of hexamethyldisilazane (HMDS) for 30 min. The specimens were then dried overnight in a vacuum desiccator and subsequently sputter-coated with a gold layer prior to imaging.</p>
      </sec>
    </sec>
    <sec sec-type="Taxon treatments">
      <title>Taxon treatments</title>
      <tp:taxon-treatment>
        <tp:nomenclature>
          <tp:taxon-name><object-id content-type="arpha">749A1C66-C85D-5805-B5E1-CA3C8B9E97E9</object-id><tp:taxon-name-part taxon-name-part-type="genus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">unicutibius</tp:taxon-name-part></tp:taxon-name>
          <tp:taxon-authority>Nguyen &amp; Bae, 2003</tp:taxon-authority>
          <tp:nomenclature-citation-list>
            <tp:nomenclature-citation>
              <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name>
              <comment> Nguyen &amp; Bae, 2003 - <xref ref-type="bibr" rid="B14406931">Nguyen and Bae (2003)</xref>: 233. Types: nymph, from Vietnam.</comment>
            </tp:nomenclature-citation>
            <tp:nomenclature-citation>
              <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name>
              <comment>: <xref ref-type="bibr" rid="B14406831">Ma and Zhou (2021)</xref>: 2 (mentioned); <xref ref-type="bibr" rid="B14406778">Garces and Sartori (2022)</xref>: 49 (mentioned).</comment>
            </tp:nomenclature-citation>
          </tp:nomenclature-citation-list>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="materials">
          <title>Materials</title>
          <list list-type="alpha-lower" list-content="occurrences">
            <list-item>
              <p><bold>Type status:</bold> <named-content content-type="dwc:typeStatus" xlink:type="simple">Other material</named-content>. <bold>Occurrence:</bold> recordedBy: <named-content content-type="dwc:recordedBy" xlink:type="simple">De–Wen Gong, Xu–Hong–Yi Zheng</named-content>; individualCount: <named-content content-type="dwc:individualCount" xlink:type="simple">1</named-content>; sex: <named-content content-type="dwc:sex" xlink:type="simple">female</named-content>; lifeStage: <named-content content-type="dwc:lifeStage" xlink:type="simple">imgao</named-content>; occurrenceID: <named-content content-type="dwc:occurrenceID" xlink:type="simple">81807E76-7F97-50AD-B8A3-D4B2DD7633F0</named-content>; <bold>Taxon:</bold> scientificName: <named-content content-type="dwc:scientificName" xlink:type="simple"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic></named-content>; <bold>Location:</bold> country: <named-content content-type="dwc:country" xlink:type="simple">China</named-content>; stateProvince: <named-content content-type="dwc:stateProvince" xlink:type="simple">Yunnan</named-content>; county: <named-content content-type="dwc:county" xlink:type="simple">Yinjiang</named-content>; locality: <named-content content-type="dwc:locality" xlink:type="simple">Nabang town, Yinjiang county, Yunnan Province, China</named-content>; verbatimElevation: <named-content content-type="dwc:verbatimElevation" xlink:type="simple">1106 m</named-content>; verbatimCoordinates: <named-content content-type="dwc:verbatimCoordinates" xlink:type="simple">24°37.39'N 97°35.02'E</named-content>; decimalLatitude: <named-content content-type="dwc:decimalLatitude" xlink:type="simple">24.623242</named-content>; decimalLongitude: <named-content content-type="dwc:decimalLongitude" xlink:type="simple">97.583675</named-content>; georeferenceProtocol: <named-content content-type="dwc:georeferenceProtocol" xlink:type="simple">label</named-content>; <bold>Event:</bold> eventDate: <named-content content-type="dwc:eventDate" xlink:type="simple">18/19-04-2026</named-content>; <bold>Record Level:</bold> language: <named-content content-type="dwc:language" xlink:type="simple">en</named-content>; collectionCode: <named-content content-type="dwc:collectionCode" xlink:type="simple">Insects</named-content>; basisOfRecord: <named-content content-type="dwc:basisOfRecord" xlink:type="simple">PreservedSpecimen</named-content></p>
            </list-item>
            <list-item>
              <p><bold>Type status:</bold> <named-content content-type="dwc:typeStatus" xlink:type="simple">Other material</named-content>. <bold>Occurrence:</bold> recordedBy: <named-content content-type="dwc:recordedBy" xlink:type="simple">De–Wen Gong, Xu–Hong–Yi Zheng</named-content>; individualCount: <named-content content-type="dwc:individualCount" xlink:type="simple">20</named-content>; lifeStage: <named-content content-type="dwc:lifeStage" xlink:type="simple">nymph</named-content>; occurrenceID: <named-content content-type="dwc:occurrenceID" xlink:type="simple">40F6470E-8283-57F6-87AC-A7DF34A40F12</named-content>; <bold>Taxon:</bold> scientificName: <named-content content-type="dwc:scientificName" xlink:type="simple"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic></named-content>; <bold>Location:</bold> country: <named-content content-type="dwc:country" xlink:type="simple">China</named-content>; stateProvince: <named-content content-type="dwc:stateProvince" xlink:type="simple">Yunnan</named-content>; county: <named-content content-type="dwc:county" xlink:type="simple">Yinjiang</named-content>; locality: <named-content content-type="dwc:locality" xlink:type="simple">Nabang town, Yinjiang county, Yunnan Province, China</named-content>; verbatimElevation: <named-content content-type="dwc:verbatimElevation" xlink:type="simple">1106 m</named-content>; verbatimCoordinates: <named-content content-type="dwc:verbatimCoordinates" xlink:type="simple">24°37.39'N 97°35.02'E</named-content>; decimalLatitude: <named-content content-type="dwc:decimalLatitude" xlink:type="simple">24.623242</named-content>; decimalLongitude: <named-content content-type="dwc:decimalLongitude" xlink:type="simple">97.583675</named-content>; georeferenceProtocol: <named-content content-type="dwc:georeferenceProtocol" xlink:type="simple">label</named-content>; <bold>Event:</bold> eventDate: <named-content content-type="dwc:eventDate" xlink:type="simple">18/19-04-2026</named-content>; <bold>Record Level:</bold> language: <named-content content-type="dwc:language" xlink:type="simple">en</named-content>; collectionCode: <named-content content-type="dwc:collectionCode" xlink:type="simple">Insects</named-content>; basisOfRecord: <named-content content-type="dwc:basisOfRecord" xlink:type="simple">PreservedSpecimen</named-content></p>
            </list-item>
            <list-item>
              <p><bold>Type status:</bold> <named-content content-type="dwc:typeStatus" xlink:type="simple">Other material</named-content>. <bold>Occurrence:</bold> recordedBy: <named-content content-type="dwc:recordedBy" xlink:type="simple">De–Wen Gong, Di Wang</named-content>; individualCount: <named-content content-type="dwc:individualCount" xlink:type="simple">1</named-content>; sex: <named-content content-type="dwc:sex" xlink:type="simple">female</named-content>; lifeStage: <named-content content-type="dwc:lifeStage" xlink:type="simple">subimago</named-content>; occurrenceID: <named-content content-type="dwc:occurrenceID" xlink:type="simple">31F914E4-427F-5D22-9AB2-6492C2AA0CB3</named-content>; <bold>Taxon:</bold> scientificName: <named-content content-type="dwc:scientificName" xlink:type="simple"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic></named-content>; <bold>Location:</bold> country: <named-content content-type="dwc:country" xlink:type="simple">China</named-content>; stateProvince: <named-content content-type="dwc:stateProvince" xlink:type="simple">Yunnan</named-content>; county: <named-content content-type="dwc:county" xlink:type="simple">Yinjiang</named-content>; locality: <named-content content-type="dwc:locality" xlink:type="simple">Nabang town, Yinjiang county, Yunnan Province, China</named-content>; verbatimElevation: <named-content content-type="dwc:verbatimElevation" xlink:type="simple">1106 m</named-content>; verbatimCoordinates: <named-content content-type="dwc:verbatimCoordinates" xlink:type="simple">24°37.39'N 97°35.02'E</named-content>; decimalLatitude: <named-content content-type="dwc:decimalLatitude" xlink:type="simple">24.623242</named-content>; decimalLongitude: <named-content content-type="dwc:decimalLongitude" xlink:type="simple">97.583675</named-content>; georeferenceProtocol: <named-content content-type="dwc:georeferenceProtocol" xlink:type="simple">label</named-content>; <bold>Event:</bold> eventDate: <named-content content-type="dwc:eventDate" xlink:type="simple">8/10-05-2026</named-content>; <bold>Record Level:</bold> language: <named-content content-type="dwc:language" xlink:type="simple">en</named-content>; collectionCode: <named-content content-type="dwc:collectionCode" xlink:type="simple">Insects</named-content>; basisOfRecord: <named-content content-type="dwc:basisOfRecord" xlink:type="simple">PreservedSpecimen</named-content></p>
            </list-item>
            <list-item>
              <p><bold>Type status:</bold> <named-content content-type="dwc:typeStatus" xlink:type="simple">Other material</named-content>. <bold>Occurrence:</bold> recordedBy: <named-content content-type="dwc:recordedBy" xlink:type="simple">De–Wen Gong, Di Wang</named-content>; individualCount: <named-content content-type="dwc:individualCount" xlink:type="simple">1</named-content>; sex: <named-content content-type="dwc:sex" xlink:type="simple">male</named-content>; lifeStage: <named-content content-type="dwc:lifeStage" xlink:type="simple">subimago</named-content>; occurrenceID: <named-content content-type="dwc:occurrenceID" xlink:type="simple">E97D1E5C-769B-5B26-BE08-1AA72F1821A7</named-content>; <bold>Taxon:</bold> scientificName: <named-content content-type="dwc:scientificName" xlink:type="simple"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic></named-content>; <bold>Location:</bold> country: <named-content content-type="dwc:country" xlink:type="simple">China</named-content>; stateProvince: <named-content content-type="dwc:stateProvince" xlink:type="simple">Yunnan</named-content>; county: <named-content content-type="dwc:county" xlink:type="simple">Yinjiang</named-content>; locality: <named-content content-type="dwc:locality" xlink:type="simple">Nabang town, Yinjiang county, Yunnan Province, China</named-content>; verbatimElevation: <named-content content-type="dwc:verbatimElevation" xlink:type="simple">1106 m</named-content>; verbatimCoordinates: <named-content content-type="dwc:verbatimCoordinates" xlink:type="simple">24°37.39'N 97°35.02'E</named-content>; decimalLatitude: <named-content content-type="dwc:decimalLatitude" xlink:type="simple">24.623242</named-content>; decimalLongitude: <named-content content-type="dwc:decimalLongitude" xlink:type="simple">97.583675</named-content>; georeferenceProtocol: <named-content content-type="dwc:georeferenceProtocol" xlink:type="simple">label</named-content>; <bold>Event:</bold> eventDate: <named-content content-type="dwc:eventDate" xlink:type="simple">8/10-05-2026</named-content>; <bold>Record Level:</bold> language: <named-content content-type="dwc:language" xlink:type="simple">en</named-content>; collectionCode: <named-content content-type="dwc:collectionCode" xlink:type="simple">Insects</named-content>; basisOfRecord: <named-content content-type="dwc:basisOfRecord" xlink:type="simple">PreservedSpecimen</named-content></p>
            </list-item>
            <list-item>
              <p><bold>Type status:</bold> <named-content content-type="dwc:typeStatus" xlink:type="simple">Other material</named-content>. <bold>Occurrence:</bold> recordedBy: <named-content content-type="dwc:recordedBy" xlink:type="simple">De–Wen Gong, Di Wang</named-content>; individualCount: <named-content content-type="dwc:individualCount" xlink:type="simple">5</named-content>; sex: <named-content content-type="dwc:sex" xlink:type="simple">male</named-content>; lifeStage: <named-content content-type="dwc:lifeStage" xlink:type="simple">imago</named-content>; occurrenceID: <named-content content-type="dwc:occurrenceID" xlink:type="simple">A96BB679-F6D4-55FD-968C-E24A2EAE7BA5</named-content>; <bold>Taxon:</bold> scientificName: <named-content content-type="dwc:scientificName" xlink:type="simple"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic></named-content>; <bold>Location:</bold> country: <named-content content-type="dwc:country" xlink:type="simple">China</named-content>; stateProvince: <named-content content-type="dwc:stateProvince" xlink:type="simple">Yunnan</named-content>; county: <named-content content-type="dwc:county" xlink:type="simple">Yinjiang</named-content>; locality: <named-content content-type="dwc:locality" xlink:type="simple">Nabang town, Yinjiang county, Yunnan Province, China</named-content>; verbatimElevation: <named-content content-type="dwc:verbatimElevation" xlink:type="simple">1106 m</named-content>; verbatimCoordinates: <named-content content-type="dwc:verbatimCoordinates" xlink:type="simple">24°37.39'N 97°35.02'E</named-content>; decimalLatitude: <named-content content-type="dwc:decimalLatitude" xlink:type="simple">24.623242</named-content>; decimalLongitude: <named-content content-type="dwc:decimalLongitude" xlink:type="simple">97.583675</named-content>; georeferenceProtocol: <named-content content-type="dwc:georeferenceProtocol" xlink:type="simple">label</named-content>; <bold>Event:</bold> eventDate: <named-content content-type="dwc:eventDate" xlink:type="simple">8/10-05-2026</named-content>; <bold>Record Level:</bold> language: <named-content content-type="dwc:language" xlink:type="simple">en</named-content>; collectionCode: <named-content content-type="dwc:collectionCode" xlink:type="simple">Insects</named-content>; basisOfRecord: <named-content content-type="dwc:basisOfRecord" xlink:type="simple">PreservedSpecimen</named-content></p>
            </list-item>
            <list-item>
              <p><bold>Type status:</bold> <named-content content-type="dwc:typeStatus" xlink:type="simple">Other material</named-content>. <bold>Occurrence:</bold> recordedBy: <named-content content-type="dwc:recordedBy" xlink:type="simple">De–Wen Gong, Di Wang</named-content>; individualCount: <named-content content-type="dwc:individualCount" xlink:type="simple">6</named-content>; sex: <named-content content-type="dwc:sex" xlink:type="simple">female</named-content>; lifeStage: <named-content content-type="dwc:lifeStage" xlink:type="simple">imago</named-content>; occurrenceID: <named-content content-type="dwc:occurrenceID" xlink:type="simple">5C0FF554-F6EF-51F3-BA46-0534F832C924</named-content>; <bold>Taxon:</bold> scientificName: <named-content content-type="dwc:scientificName" xlink:type="simple"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic></named-content>; <bold>Location:</bold> country: <named-content content-type="dwc:country" xlink:type="simple">China</named-content>; stateProvince: <named-content content-type="dwc:stateProvince" xlink:type="simple">Yunnan</named-content>; county: <named-content content-type="dwc:county" xlink:type="simple">Yinjiang</named-content>; locality: <named-content content-type="dwc:locality" xlink:type="simple">Nabang town, Yinjiang county, Yunnan Province, China</named-content>; verbatimElevation: <named-content content-type="dwc:verbatimElevation" xlink:type="simple">1106 m</named-content>; verbatimCoordinates: <named-content content-type="dwc:verbatimCoordinates" xlink:type="simple">24°37.39'N 97°35.02'E</named-content>; decimalLatitude: <named-content content-type="dwc:decimalLatitude" xlink:type="simple">24.623242</named-content>; decimalLongitude: <named-content content-type="dwc:decimalLongitude" xlink:type="simple">97.583675</named-content>; georeferenceProtocol: <named-content content-type="dwc:georeferenceProtocol" xlink:type="simple">label</named-content>; <bold>Event:</bold> eventDate: <named-content content-type="dwc:eventDate" xlink:type="simple">8/10-05-2026</named-content>; <bold>Record Level:</bold> language: <named-content content-type="dwc:language" xlink:type="simple">en</named-content>; collectionCode: <named-content content-type="dwc:collectionCode" xlink:type="simple">Insects</named-content>; basisOfRecord: <named-content content-type="dwc:basisOfRecord" xlink:type="simple">PreservedSpecimen</named-content></p>
            </list-item>
            <list-item>
              <p><bold>Type status:</bold> <named-content content-type="dwc:typeStatus" xlink:type="simple">Other material</named-content>. <bold>Occurrence:</bold> recordedBy: <named-content content-type="dwc:recordedBy" xlink:type="simple">De–Wen Gong, Di Wang</named-content>; individualCount: <named-content content-type="dwc:individualCount" xlink:type="simple">15</named-content>; lifeStage: <named-content content-type="dwc:lifeStage" xlink:type="simple">nymph</named-content>; occurrenceID: <named-content content-type="dwc:occurrenceID" xlink:type="simple">D3B76374-BDD2-55C0-9828-078CE2FB9A37</named-content>; <bold>Taxon:</bold> scientificName: <named-content content-type="dwc:scientificName" xlink:type="simple"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic></named-content>; <bold>Location:</bold> country: <named-content content-type="dwc:country" xlink:type="simple">China</named-content>; stateProvince: <named-content content-type="dwc:stateProvince" xlink:type="simple">Yunnan</named-content>; county: <named-content content-type="dwc:county" xlink:type="simple">Yinjiang</named-content>; locality: <named-content content-type="dwc:locality" xlink:type="simple">Nabang town, Yinjiang county, Yunnan Province, China</named-content>; verbatimElevation: <named-content content-type="dwc:verbatimElevation" xlink:type="simple">1106 m</named-content>; verbatimCoordinates: <named-content content-type="dwc:verbatimCoordinates" xlink:type="simple">24°37.39'N 97°35.02'E</named-content>; decimalLatitude: <named-content content-type="dwc:decimalLatitude" xlink:type="simple">24.623242</named-content>; decimalLongitude: <named-content content-type="dwc:decimalLongitude" xlink:type="simple">97.583675</named-content>; georeferenceProtocol: <named-content content-type="dwc:georeferenceProtocol" xlink:type="simple">label</named-content>; <bold>Event:</bold> eventDate: <named-content content-type="dwc:eventDate" xlink:type="simple">8/10-05-2026</named-content>; <bold>Record Level:</bold> language: <named-content content-type="dwc:language" xlink:type="simple">en</named-content>; collectionCode: <named-content content-type="dwc:collectionCode" xlink:type="simple">Insects</named-content>; basisOfRecord: <named-content content-type="dwc:basisOfRecord" xlink:type="simple">PreservedSpecimen</named-content></p>
            </list-item>
          </list>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Description">
          <title>Description</title>
          <p>Male imago (in alcohol): body length 8.0–10.0 mm, fore-legs 11.0–12.0 mm, fore-wings 9.0–9.5 mm, hind-wings 3.0–3.2 mm, cerci 40.0–42.0 mm, terminal filament 1.0–1.1 mm; body reddish-brown (Fig. <xref ref-type="fig" rid="F14405037">1</xref>A and E–F). Antennae brown, but apical half of scape and flagella whitish; antennae narrowed from base to apex (Fig. <xref ref-type="fig" rid="F14405039">2</xref>A); vertex white to grey, but with a brown stripe between two compound eyes (Fig. <xref ref-type="fig" rid="F14405039">2</xref>A). Ocelli cylindrical, with chocolate basal rings; lateral ocelli distinctly larger than median one (Fig. <xref ref-type="fig" rid="F14405039">2</xref>A). Compound eyes divided indistinctly into two parts, shortest distance between two eyes about 0.3x diameter of one eye in dorsal view, just slightly wider than median ocellus, but narrower than lateral ocellus (Fig. <xref ref-type="fig" rid="F14405039">2</xref>A). Posterior margin of occiput almost straight (Fig. <xref ref-type="fig" rid="F14405039">2</xref>A). Pronotum brown, posterior margin concave (Fig. <xref ref-type="fig" rid="F14405037">1</xref>E and Fig. <xref ref-type="fig" rid="F14405039">2</xref>A). Mesonotum with four pairs of deep brown dots laterally and one brown median pit (or called medial ocellus, Fig. <xref ref-type="fig" rid="F14405037">1</xref>E). Prosternum triangular, mesothoracic furcasternum with almost straight inner margins (Fig. <xref ref-type="fig" rid="F14405037">1</xref>F). Fore-wings transparent, but longitudinal veins brown, all crossveins pigmented reddish-brown and surrounding clouds (Fig. <xref ref-type="fig" rid="F14405037">1</xref>A and E–F), especially those near margins; MA forked distally to middle point; MP forked at base (Fig. <xref ref-type="fig" rid="F14405039">2</xref>B and Fig. <xref ref-type="fig" rid="F14405055">8</xref>B). Hind-wings also with various reddish to chocolate dots and markings, length ca. 1.5x width (Fig. <xref ref-type="fig" rid="F14405039">2</xref>C); costal projection blunt; R1 straight and zigzag; MA single, MP forked at middle point almost symmetrically (Fig. <xref ref-type="fig" rid="F14405039">2</xref>D and Fig. <xref ref-type="fig" rid="F14405055">8</xref>C). Femora of fore-legs brown to chocolate, tibiae with brown base and apex, segment I of fore-tarsi dark brown, segments II–V with brown apex, other parts yellow (Fig. <xref ref-type="fig" rid="F14405039">2</xref>E). Length ratio of fore-femora: tibiae: tarsi = 1.0: 2.3: 2.7, length ratio of five segments of fore-tarsi = 1.0: 6.3: 4.7: 3.7: 2.3 (Fig. <xref ref-type="fig" rid="F14405039">2</xref>E); two claws of fore-legs black and blunt (Fig. <xref ref-type="fig" rid="F14405039">2</xref>F). Tibiae of mid-legs with brown base and dark apex, tarsi with dark apex too; length ratio of mid-femora: tibiae: tarsi = 1.0: 0.9: 0.6, length ratio of four movable segments of mid-tarsi = 1.0: 1.0: 0.5: 2.5 (Fig. <xref ref-type="fig" rid="F14405039">2</xref>G); claws black, one blunt one acute (Fig. <xref ref-type="fig" rid="F14405039">2</xref>H). Hind-legs similar to mid-legs, length ratio of mid-femora: tibiae: tarsi = 1.0: 0.9: 0.6, length ratio of four movable segments of mid-tarsi = 1.0: 0.8: 0.5: 2.0 (Fig. <xref ref-type="fig" rid="F14405039">2</xref>I). Mid- and hind-tibiae with patellar-tibial fusion suture (Fig. <xref ref-type="fig" rid="F14405039">2</xref>G and I). Terga of abdomen reddish-brown, washed with indistinct pale stripes (Fig. <xref ref-type="fig" rid="F14405037">1</xref>E); sterna pale, but with brown lateral margins (Fig. <xref ref-type="fig" rid="F14405037">1</xref>F). Tergum IX with distinct posterolateral projections (Fig. <xref ref-type="fig" rid="F14405039">2</xref>J–K and Fig. <xref ref-type="fig" rid="F14405055">8</xref>D–E). Genitalia: subgenital plate with almost straight posterior margin, width about 3.5x length (Fig. <xref ref-type="fig" rid="F14405039">2</xref>J); forceps slightly longer than length of subgenital plate, length ratio of segment I: II = 1.0: 0.6; segment I narrowed slightly from base to apex, apex dark; segment II with slightly expanded apex (Fig. <xref ref-type="fig" rid="F14405039">2</xref>J–K). Penes membranous, ventral surface grey, slightly extended out of subgenital plate; gonopore at posterolateral corner of penis (Fig. <xref ref-type="fig" rid="F14405039">2</xref>L); a dorsal plate between penes and subanal plate, posterolateral corner of this plate slightly extended outwards forming an acute apex (Fig. <xref ref-type="fig" rid="F14405039">2</xref>K). Cerci with two kinds of segments: apical half chocolate segment and basal dark ring segment, two kinds of them arranged alternately (Fig. <xref ref-type="fig" rid="F14405037">1</xref>A and E–F). Terminal filament black, with 8–10 segments, distinctly thinner than cerci (Fig. <xref ref-type="fig" rid="F14405039">2</xref>M).</p>
          <p>Female imago (in alcohol): body length 11.0–12.0 mm, fore-legs 5.0–6.0 mm, fore-wings 11.5–12.0 mm, hind-wings 3.5–4.5 mm, cerci 18.0–21.0, terminal filament 1.8–2.0 mm; body similar to male, but markings of wings slightly paler (Fig. <xref ref-type="fig" rid="F14405037">1</xref>B, G, H and Fig. <xref ref-type="fig" rid="F14405041">3</xref>B–D). Shortest distance between two compound eyes about 1.8x eye diameter (Fig. <xref ref-type="fig" rid="F14405041">3</xref>A). Length ratio of fore-femora: tibiae: tarsi = 1.0: 0.8: 0.8, length ratio of five segments of fore-tarsi = 1.0: 2.5: 2.0: 1.0: 3.5, Fig. <xref ref-type="fig" rid="F14405041">3</xref>E–F); length ratio of mid-femora: tibiae: tarsi = 1.0: 0.9: 0.5, length ratio of four movable segments of mid-tarsi =1.0: 1.0: 0.6: 2.5 (Fig. <xref ref-type="fig" rid="F14405041">3</xref>G); length ratio of hind-femora: tibiae: tarsi = 1.0: 0.7: 0.4, length ratio of four movable segments of mid-tarsi = 1.0: 1.0: 0.8: 3.0 (Fig. <xref ref-type="fig" rid="F14405041">3</xref>H). Two claws of all legs dissimilar, one acute, one blunt (Fig. <xref ref-type="fig" rid="F14405041">3</xref>F). Sternum VII slightly thickened and extended posteriorly, forming a triangular lobe (Fig. <xref ref-type="fig" rid="F14405041">3</xref>I). Posterior margin of sternum IX slightly extended laterally, median portion slightly concave (Fig. <xref ref-type="fig" rid="F14405041">3</xref>I). Terminal filament with 10–12 segments (Fig. <xref ref-type="fig" rid="F14405037">1</xref>G and H).</p>
          <p>Male subimago (Fig. <xref ref-type="fig" rid="F14405037">1</xref>C): body length 9.0 mm, fore-legs 5.0 mm, fore-wings 9.0 mm, hind-wings 3.5 mm, cerci 8.0 mm, terminal filament 0.5 mm. Shortest distance between two compound eyes about 0.67x eye diameter. Length ratio of fore-femora: tibiae: tarsi = 1.0: 1.1: 1.5, length ratio of mid-femora: tibiae: tarsi = 1.0: 0.7: 0.6. Hind-legs similar to mid-legs.</p>
          <p>Female subimago (Fig. <xref ref-type="fig" rid="F14405037">1</xref>D): body length 11.0 mm, fore-legs 4.0 mm, fore-wings 11.0 mm, hind-wings 3.5 mm, cerci 7.0 mm, terminal filament 1.0 mm. Shortest distance between two compound eyes about 2.0x eye diameter. Length ratio of fore-femora: tibiae: tarsi = 1.0: 1.1: 1.5, length ratio of mid-femora: tibiae: tarsi = 1.0: 0.7: 0.7. Mid- and hind-legs similar to fore-legs, but paler.</p>
          <p>Mature nymph (in alcohol): (see <xref ref-type="bibr" rid="B14406931">Nguyen and Bae (2003)</xref> for full description).</p>
          <p>Main characters include: body length 8.0–12.0 mm, caudal filaments 3.0–4.0 mm. Body reddish-brown to chocolate (Fig. <xref ref-type="fig" rid="F14405043">4</xref>). Mouthparts as in Fig. <xref ref-type="fig" rid="F14405049">5</xref>A–F, filtering mouthparts, maxillary and labial palpi elongated and with dense long hair–like setae, especially their apical segments, length ration of three segments of maxillary palpi = 1.0: 1.5: 4 (Fig. <xref ref-type="fig" rid="F14405049">5</xref>E), ratio of labial palpi = 1.0: 0.3: 2.0 (Fig. <xref ref-type="fig" rid="F14405049">5</xref>F); fore-legs with a row of subapical transverse spines (Fig. <xref ref-type="fig" rid="F14405049">5</xref>G and H); fore-tibiae with two rows of long hair-like setae on dorsal surface, one transverse, one at base, one oblique row near middle (Fig. <xref ref-type="fig" rid="F14405049">5</xref>G and H); fore-tarsi with a transverse row of long setae at base and two rows along dorsal and inner margins (Fig. <xref ref-type="fig" rid="F14405049">5</xref>G). Length ratio of fore-femora: tibiae: tarsi = 1.0: 0.9: 1.0 (Fig. <xref ref-type="fig" rid="F14405049">5</xref>G), ratio of mid-leg = 1.0: 0.6: 0.6 (Fig. <xref ref-type="fig" rid="F14405049">5</xref>I), hind-leg = 1.0: 0.6: 0.4 (Fig. <xref ref-type="fig" rid="F14405049">5</xref>J), claws hooked. Terga I–II, VI–VII with a median tubercle respectively, that of tergum II more distinct than others (Fig. <xref ref-type="fig" rid="F14405051">6</xref>A). Gills shown in Fig. <xref ref-type="fig" rid="F14405051">6</xref>B–F, gills I two segmented, with tiny hair–like setae (Fig. <xref ref-type="fig" rid="F14405051">6</xref>B); both dorsal and ventral surface of gill opercula smooth, posterior inner margins with dense setae, ventral lamella with fringes (Fig. <xref ref-type="fig" rid="F14405051">6</xref>C); gills III–V similar, with dorsal and ventral lamellae (Fig. <xref ref-type="fig" rid="F14405051">6</xref>D and E); gills VI single (Fig. <xref ref-type="fig" rid="F14405051">6</xref>F). Caudal filaments with stout spines on articulations, cerci with additional short spines on outer side and long spines on inner side; terminal filament with long spines on both lateral sides; articulations with very sparse tiny hair–like setae, especially apical half (Fig. <xref ref-type="fig" rid="F14405051">6</xref>G and H).</p>
          <p>Egg: surface smooth, micropyle at middle (Fig. <xref ref-type="fig" rid="F14455534">7</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Diagnosis">
          <title>Diagnosis</title>
          <p>The nymph of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> can be recognised by its filtering mouthparts and fore-legs: apical segment of maxillary and labial palpi more than 2 times of segment II in length, with dense hairlike setae (Fig. <xref ref-type="fig" rid="F14405049">5</xref>E<bold>–</bold>F); fore-tibiae with two oblique rows of long filtering setae (Fig. <xref ref-type="fig" rid="F14405049">5</xref>G-H), fore-tarsi with filtering hairs (Fig. <xref ref-type="fig" rid="F14405049">5</xref>G). In addition, their relatively short caudal filaments, smooth body and gill opercula, independent gills II, neither pronotum nor mesonotum with distinct expanded anterolateral corner are useful to their identification (Fig. <xref ref-type="fig" rid="F14405043">4</xref>).</p>
          <p>In imaginal stage, its relatively short body (10.0<bold>–</bold>12.0 mm), heavily stained wings (Fig. <xref ref-type="fig" rid="F14405037">1</xref>A<bold>–</bold>B), long fore-legs and its tibiae (longer than 2.0x fore-femora, Fig. <xref ref-type="fig" rid="F14405039">2</xref>E) and widely opened compound eyes (Fig. <xref ref-type="fig" rid="F14405039">2</xref>A) are good identical characteristics. Its male imago has reduced genitalia, but all components distinct, especially its penes (Fig. <xref ref-type="fig" rid="F14405039">2</xref>J<bold>–</bold>L).</p>
          <p>All other congeners in the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic> have no filtering setae on their fore-leg or mouthparts in nymphal stage. In imaginal stage, all other species have no distinct penes, only forceps and dorsal plate are visible. In addition, the dorsal plates of those species are different: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="amabilis">amabilis</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chinensis">chinensis</tp:taxon-name-part></tp:taxon-name></italic> have deeply divided plates and their forceps are three-segmented. Although <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="caenoides">caenoides</tp:taxon-name-part></tp:taxon-name></italic> has two-segmented forceps, its dorsal plate is two semi-circular (see <xref ref-type="bibr" rid="B14406760">Bae and McCafferty (1998)</xref>). In contrast, P. <italic>unicutibius</italic> has two-segmented forceps and its dorsal plate has two distinct posterolateral processes (Fig. <xref ref-type="fig" rid="F14405039">2</xref>L and Fig. <xref ref-type="fig" rid="F14405055">8</xref>E).</p>
          <p>Egg: surface smooth, without distinct mesh or KCTs (knob-terminated coiled threads), micropyle at equator (Fig. <xref ref-type="fig" rid="F14455534">7</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Distribution">
          <title>Distribution</title>
          <p>Yunnan Prov. (China); Lao Cai Prov. (Vietnam).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Ecology">
          <title>Ecology</title>
          <p>This nymphs were found in a narrow creek with large or small stones, mountain area. The width of the creek is about 1–2 m and the flow velocity is 1 m/s (Fig. <xref ref-type="fig" rid="F14405061">9</xref>). An <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Isonychia">Isonychia</tp:taxon-name-part></tp:taxon-name></italic> sp. was also found in the same water.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
    <sec sec-type="Discussion">
      <title>Discussion</title>
      <p>Reduction or vestigialiation of male genitalia, particularly the penes, has been documented in several mayfly lineages. For instance, species within the taxon <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cryptopenella">Cryptopenella</tp:taxon-name-part></tp:taxon-name></italic> of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Choroterpes">Choroterpes</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Leptophlebiidae">Leptophlebiidae</tp:taxon-name-part></tp:taxon-name>) possess indistinct penes, yet retain well-developed forceps and a distinct ventral penis cover (<xref ref-type="bibr" rid="B14406904">Zhou 2006</xref>). Similarly, most species in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Baetidae">Baetidae</tp:taxon-name-part></tp:taxon-name> have concealed membranous penes, but their ventral penis cover and forceps remain prominent (<xref ref-type="bibr" rid="B14406922">Keffermüller 1960</xref>, <xref ref-type="bibr" rid="B14406895">Ying et al. 2021</xref>, <xref ref-type="bibr" rid="B14406885">Wang et al. 2025</xref>). In contrast, species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Caenidae">Caenidae</tp:taxon-name-part></tp:taxon-name> typically exhibit minute forceps, while retaining recognisable fused penes (<xref ref-type="bibr" rid="B14406876">Sun et al. 2026</xref>). Surprisingly, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> displays yet another pattern of vestigial genitalia: its forceps, penes and penis cover (a component of the subgenital plate) are all reduced, while it possesses a distinct dorsal covering plate (Fig. <xref ref-type="fig" rid="F14405039">2</xref>J–L).</p>
      <p>In most mayflies, the gonopores open at the apex of the penes, implying that the penes are three-dimensional structures. However, based on historical accounts (<xref ref-type="bibr" rid="B14406913">Hsu 1936</xref>, <xref ref-type="bibr" rid="B14406940">Tiunova 1991</xref>, <xref ref-type="bibr" rid="B14406760">Bae and McCafferty 1998</xref>) and our own observations of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic> materials, the penes in this genus are membranous and plate-like. In light of the discovery of male <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic>, we interpret the structure previously regarded as the penes of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic> to be, in fact, the dorsal plate of the penes, as observed in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> (Fig. <xref ref-type="fig" rid="F14405039">2</xref>J–L).</p>
      <p>The genital morphology of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> may reflect a possible evolutionary trend within the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Neoephemeridae">Neoephemeridae</tp:taxon-name-part></tp:taxon-name>: initially, the forceps and penes are relatively well developed, as seen in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ephemera">Ephemera</tp:taxon-name-part></tp:taxon-name></italic> or <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthodes">Potamanthodes</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B14406813">Lei and Zhou 2024</xref>, <xref ref-type="bibr" rid="B14406867">Qu et al. 2026</xref>); later, the penes undergo reduction, as observed in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pulchephemera">Pulchephemera</tp:taxon-name-part></tp:taxon-name></italic> or <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ochernova">Ochernova</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B14406831">Ma and Zhou 2021</xref>, <xref ref-type="bibr" rid="B14406822">Li et al. 2026</xref>); and finally, the forceps become diminished and the penes membranous, as in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic>.</p>
      <p>Within the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Neoephemera">Neoephemera</tp:taxon-name-part></tp:taxon-name></italic>-like clade, at least three nymphal morphotypes or evolutionary trends have been recognised: flattened body form (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pulchephemera">Pulchephemera</tp:taxon-name-part></tp:taxon-name></italic>), stout body form (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucorhoenanthus">Leucorhoenanthus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Neoephemera">Neoephemera</tp:taxon-name-part></tp:taxon-name></italic>) and slender body form (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ochernova">Ochernova</tp:taxon-name-part></tp:taxon-name></italic>) (<xref ref-type="bibr" rid="B14406822">Li et al. 2026</xref>). In contrast, the smooth body surface, operculate gills and swimming-adapted caudal filaments of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic> suggest a certain degree of active locomotion or swimming capability in aquatic environments. Intriguingly, the nymphs of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> appear to represent a further specialisation, as they are theoretically capable of filtering particulate matter from flowing water. The adult morphology of this species may offer additional evidence bearing on its evolutionary position.</p>
      <p>From a biological perspective, based on general nymphal morphology, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> and other <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic> species appear to have evolved in different ecological directions. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> may be adapted to filtering particulate matter in flowing water, whereas other <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part></tp:taxon-name></italic> species likely exhibits greater swimming ability and may feed by chewing on fallen leaves or aquatic macrophytes.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>This work was funded by the National Natural Science Foundation of China (Grant number 31750002, 32070475 and 32470464).</p>
    </ack>
    <sec sec-type="Author contributions">
      <title>Author contributions</title>
      <p>Dewen Gong collected materials of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003.</p>
      <p>Dewen Gong and Changfa Zhou made the illustrations, analysed the materials and wrote the descriptions and manuscript.</p>
    </sec>
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  <floats-group>
    <fig id="F14405037" position="float" orientation="portrait">
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      <label>Figure 1.</label>
      <caption>
        <p>Imaginal habitus of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 <bold>A</bold> Male imago; <bold>B</bold> Female imago; <bold>C</bold> Male subimago; <bold>D</bold> Female subimago; <bold>E</bold> Male imago (dorsal view); <bold>F</bold> Male imago (ventral view); <bold>G</bold> Female imago (dorsal view); <bold>H</bold> Female imago (ventral view).</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g001.jpg" position="float" id="oo_1741731.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1741731</uri>
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    </fig>
    <fig id="F14405039" position="float" orientation="portrait">
      <object-id content-type="arpha">0E5DECED-ACD3-5CDA-8C01-8DE9F595317B</object-id>
      <object-id content-type="doi">10.3897/BDJ.14.e211811.figure3</object-id>
      <label>Figure 2.</label>
      <caption>
        <p>Characters of male imago of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 <bold>A</bold> Head (dorsal view); <bold>B</bold> Fore-wing; <bold>C</bold> Hind-wing; <bold>D</bold> Hind-wing enlarged; <bold>E</bold> Fore-leg; <bold>F</bold> Fore-claw; <bold>G</bold> Mid-leg; <bold>H</bold> Mid-claw; <bold>I</bold> Hind-leg; <bold>J</bold> Genitalia (ventral view); <bold>K</bold> Genitalia (dorsal view); <bold>L</bold> Genitalia (apicodorsal view, showing penes).</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g002.jpg" position="float" id="oo_1741742.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1741742</uri>
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    </fig>
    <fig id="F14405041" position="float" orientation="portrait">
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      <object-id content-type="doi">10.3897/BDJ.14.e211811.figure6</object-id>
      <label>Figure 3.</label>
      <caption>
        <p>Characters of female imago of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 <bold>A</bold> Head (dorsal view); <bold>B</bold> Fore-wing; <bold>C</bold> Hind-wing; <bold>D</bold> Hind-wing enlarged; <bold>E</bold> Fore-leg; <bold>F</bold> Fore-claw; <bold>G</bold> Mid-leg; <bold>H</bold> Hind-leg; <bold>I</bold> Terminal abdoman (ventral view).</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g003.jpg" position="float" id="oo_1741743.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1741743</uri>
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    </fig>
    <fig id="F14405043" position="float" orientation="portrait">
      <object-id content-type="arpha">E230FBA1-930B-5A19-8B78-26C418ADE875</object-id>
      <object-id content-type="doi">10.3897/BDJ.14.e211811.figure4</object-id>
      <label>Figure 4.</label>
      <caption>
        <p>Nymphal habitus of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 <bold>A</bold> Dorsal view; <bold>B</bold> Ventral view.</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g004.jpg" position="float" id="oo_1741745.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1741745</uri>
      </graphic>
    </fig>
    <fig id="F14405049" position="float" orientation="portrait">
      <object-id content-type="arpha">FF79C9A3-406C-55C1-BE41-3E44701F94EA</object-id>
      <object-id content-type="doi">10.3897/BDJ.14.e211811.figure4</object-id>
      <label>Figure 5.</label>
      <caption>
        <p>Mouthparts and legs of nymph of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 <bold>A</bold> Labrum (dorsal view); <bold>B</bold> Left mandible; <bold>C</bold> Right mandible; <bold>D</bold> Hypopharynx (ventral view); <bold>E</bold> Maxilla; <bold>F</bold> Labium (ventral view); <bold>G</bold> Fore-leg; <bold>H</bold> Fore-tibia enlarged (arrows indicating the setaous rows); <bold>I</bold> Mid-leg; <bold>J</bold> Hind-leg</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g005.jpg" position="float" id="oo_1741758.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1741758</uri>
      </graphic>
    </fig>
    <fig id="F14405051" position="float" orientation="portrait">
      <object-id content-type="arpha">99C5398C-4C2B-52BD-980C-668598BE047A</object-id>
      <object-id content-type="doi">10.3897/BDJ.14.e211811.figure4</object-id>
      <label>Figure 6.</label>
      <caption>
        <p>Nymphal characters of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 <bold>A</bold> Abdominal terga (arrows indicating the median tubercle); <bold>B</bold> Gills I; <bold>C</bold> Gills II (dorsal view, arrow indicating the ventral lamella); <bold>D</bold> Gills III (arrow indicating the ventral lamella); <bold>E</bold> Gills IV (arrow indicating the ventral lamella); <bold>F</bold> Gills VI; <bold>G</bold> Caudal filaments; <bold>H</bold> Cerci enlarged (showing the spines and setae).</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g006.jpg" position="float" id="oo_1741764.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1741764</uri>
      </graphic>
    </fig>
    <fig id="F14455534" position="float" orientation="portrait">
      <object-id content-type="arpha">F23C4496-680F-5604-ADFF-B0562FC7F09B</object-id>
      <object-id content-type="doi">10.3897/BDJ.14.e211811.figure4</object-id>
      <label>Figure 7.</label>
      <caption>
        <p>SEM pictures of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 <bold>A</bold> Egg; <bold>B</bold> Micophyle.</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g007.jpg" position="float" id="oo_1768281.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1768281</uri>
      </graphic>
    </fig>
    <fig id="F14405055" position="float" orientation="portrait">
      <object-id content-type="arpha">91FE3926-1906-59A8-86E0-B2521A5525A5</object-id>
      <object-id content-type="doi">10.3897/BDJ.14.e211811.figure4</object-id>
      <label>Figure 8.</label>
      <caption>
        <p>Drawing pictures of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003 <bold>A</bold> Nymphal habitus (dorsal view); <bold>B</bold> Fore-wing of male imago; <bold>C</bold> Hind-wing of male imago; <bold>D</bold> Male genitalia (ventral view, showing the forceps and subgenital plate); <bold>E</bold> Male genitalia (apicodorsal view, showing the penes and dorsal plate).</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g008.jpg" position="float" id="oo_1741767.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1741767</uri>
      </graphic>
    </fig>
    <fig id="F14405061" position="float" orientation="portrait">
      <object-id content-type="arpha">C531E24C-8303-57B8-BA0E-4E9837808796</object-id>
      <object-id content-type="doi">10.3897/BDJ.14.e211811.figure4</object-id>
      <label>Figure 9.</label>
      <caption>
        <p>Nymphal Habitat of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Potamanthellus">Potamanthellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unicutibius">unicutibius</tp:taxon-name-part></tp:taxon-name></italic> Nguyen &amp; Bae, 2003</p>
      </caption>
      <graphic xlink:href="bdj-14-e211811-g009.jpg" position="float" id="oo_1741788.jpg" orientation="portrait" xlink:type="simple">
        <uri content-type="original_file">https://binary.pensoft.net/fig/1741788</uri>
      </graphic>
    </fig>
  </floats-group>
</article>
