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  <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.1266.169869</article-id>
      <article-id pub-id-type="publisher-id">169869</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Staphylinidae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Ecuador</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>﻿First country record of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> Raffray, 1908 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Staphylinidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Pselaphinae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Metopiasini</tp:taxon-name-part></tp:taxon-name>) from Ecuador, with description of two new species</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Tapuy-Avilés</surname>
            <given-names>Yarina</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0001-2357-7916</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Díaz-Guevara</surname>
            <given-names>David R.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0001-4632-4460</uri>
          <xref ref-type="aff" rid="A2">2</xref>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Caterino</surname>
            <given-names>Michael S.</given-names>
          </name>
          <email xlink:type="simple">mcateri@clemson.edu</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-2597-5707</uri>
          <xref ref-type="aff" rid="A3">3</xref>
          <xref ref-type="aff" rid="A4">4</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Pontificia Universidad Católica de Ecuador, Quito, Ecuador</addr-line>
        <institution>Fundación Uru</institution>
        <addr-line content-type="city">Quito</addr-line>
        <country>Ecuador</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Fundación Uru, Quito, Ecuador</addr-line>
        <institution>Pontificia Universidad Católica de Ecuador</institution>
        <addr-line content-type="city">Quito</addr-line>
        <country>Ecuador</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Instituto Nacional de Biodiversidad, Quito, Ecuador</addr-line>
        <institution>Instituto Nacional de Biodiversidad</institution>
        <addr-line content-type="city">Quito</addr-line>
        <country>Ecuador</country>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">Clemson University, Clemson, South Carolina, USA</addr-line>
        <institution>Clemson University</institution>
        <addr-line content-type="city">Clemson</addr-line>
        <country>United States of America</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Michael S. Caterino (<email xlink:type="simple">mcateri@clemson.edu</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Zi-Wei Yin</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>08</day>
        <month>01</month>
        <year>2026</year>
      </pub-date>
      <volume>1266</volume>
      <fpage>127</fpage>
      <lpage>135</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/5CAE7197-F543-5E41-A728-18CA0A7F0215">5CAE7197-F543-5E41-A728-18CA0A7F0215</uri>
      <uri content-type="zenodo_dep_id" xlink:href="https://zenodo.org/record/0">0</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/F0CA51CB-04EB-4454-8B02-6E6400DBF2C6">F0CA51CB-04EB-4454-8B02-6E6400DBF2C6</uri>
      <history>
        <date date-type="received">
          <day>26</day>
          <month>08</month>
          <year>2025</year>
        </date>
        <date date-type="accepted">
          <day>11</day>
          <month>12</month>
          <year>2025</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Yarina Tapuy-Avilés, David R. Díaz-Guevara, Michael S. Caterino</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/F0CA51CB-04EB-4454-8B02-6E6400DBF2C6</self-uri>
      <abstract>
        <label>﻿Abstract</label>
        <p>Two new species from the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> Raffray, 1908 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Staphylinidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Pselaphinae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Metopiasini</tp:taxon-name-part></tp:taxon-name>) are described from both sides of the Ecuadorian Andes: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov.</bold> from the Amazon in eastern Ecuador and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov.</bold> from montane cloud forest in western Ecuador. These findings represent the first country record of the genus.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>Ant-like litter beetles</kwd>
        <kwd>identification key</kwd>
        <kwd>myrmecophily</kwd>
        <kwd>new record</kwd>
        <kwd>pselaphine beetles</kwd>
        <kwd>taxonomy</kwd>
        <kwd>tropical biodiversity</kwd>
      </kwd-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="SECID0ETG">
        <title>Citation</title>
        <p>Tapuy-Avilés Y, Díaz-Guevara DR, Caterino MS (2026) First country record of <italic>Metopiellus</italic> Raffray, 1908 (Staphylinidae, Pselaphinae, Metopiasini) from Ecuador, with description of two new species. ZooKeys 1266: 127–135. <ext-link xlink:href="10.3897/zookeys.1266.169869" ext-link-type="doi" xlink:type="simple">https://doi.org/10.3897/zookeys.1266.169869</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="﻿Introduction" id="SECID0EUH">
      <title>﻿Introduction</title>
      <p><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Metopiasini</tp:taxon-name-part></tp:taxon-name> is a tribe of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Pselaphinae</tp:taxon-name-part></tp:taxon-name> rove beetles suspected to be myrmecophiles (<xref ref-type="bibr" rid="B5">Parker 2016</xref>). This tribe comprises nine genera, eight of which are restricted to the Neotropical region (<xref ref-type="bibr" rid="B4">Mário Chaul and Lopes-Andrade 2024</xref>). One of these Neotropical genera is <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> Raffray, 1908, which is distributed from Colombia to Argentina (<xref ref-type="bibr" rid="B3">Fiorentino et al. 2022</xref>; <xref ref-type="bibr" rid="B4">Mário Chaul and Lopes-Andrade 2024</xref>). This genus is composed of seven described species, five of them (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aglenus">aglenus</tp:taxon-name-part></tp:taxon-name></italic> Reitter, 1885, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hirtus">hirtus</tp:taxon-name-part></tp:taxon-name></italic> Reitter, 1885, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="painensis">painensis</tp:taxon-name-part></tp:taxon-name></italic><xref ref-type="bibr" rid="B1">Asenjo et al., 2017</xref>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="crypticus">crypticus</tp:taxon-name-part></tp:taxon-name></italic><xref ref-type="bibr" rid="B2">Asenjo et al., 2023</xref>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="emavieirae">emavieirae</tp:taxon-name-part></tp:taxon-name></italic> Mário Chaul &amp; Lopes-Andrade, 2024) from Brazil, one from Colombia (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic> Fiorentino, Tocora &amp; Ramirez, 2022), and a final one from Argentina (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="silvaticus">silvaticus</tp:taxon-name-part></tp:taxon-name></italic> Bruch, 1933). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> is distinguished by a second antennomere much longer than the third, protibia carinated anteromedially, metacoxae contiguous or nearly so, and pronotum without sharp spines on the lateral edge, at most having thickened spinose protuberances mediolaterally on the pronotal disc (<xref ref-type="bibr" rid="B4">Mário Chaul and Lopes-Andrade 2024</xref>).</p>
      <p>Until now, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> has not been reported from Ecuador. Herein, we officially describe two species of this genus, from both sides of the Andes: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. from an Amazon ecosystem in eastern Ecuador and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. from a montane cloud forest in western Ecuador.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0E2EAC">
      <title>﻿Material and methods</title>
      <p>Some specimens of the new species were obtained by sifting forest-floor leaf litter from the Minga Reserve in Napo Province, Ecuador (<named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-77.268330,-0.888530]}" id="NCID0EEFAC">0.88853°S, 77.26833°W</named-content></named-content>) and from the Otongachi Reserve in Pichincha Province, Ecuador (<named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-78.934420,-0.330510]}" id="NCID0EMFAC">0.330510°S, 78.934420°W</named-content></named-content>). One was hand-collected at Tiputini Biodiversity Station (<named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-76.149900,-0.637600]}" id="NCID0EUFAC">0.6376°S, 76.1499°W</named-content></named-content>) under permit FAUNA (X) 0020-MA-DPO-PNY (Orellana Province). Sifted litter was processed using Winkler extractors, with specimens collected, stored in 95% EtOH, and finally deposited in the National Biodiversity Institute collection (<abbrev content-type="institution" xlink:title="National Biodiversity Institute collection" id="ABBRID0EZFAC">MECN</abbrev>) and the <named-content xlink:type="simple" content-type="institution" xlink:href="http://grbio.org/institution/museo-de-zoologia-pontificia-universidad-catolica-del-ecuador">Pontificia Universidad Católica del Ecuador</named-content> (<named-content content-type="dwc:institutional_code" xlink:title="Pontificia Universidad Católica del Ecuador" xlink:href="http://grbio.org/institution/museo-de-zoologia-pontificia-universidad-catolica-del-ecuador">QCAZ</named-content>). Genitalia were cleared using 10% KOH, rinsed in water, and stored in glycerin in microvials pinned beneath the specimens. Photos were taken with a Macropod Pro photographic system (Macroscopic Solutions LLC), using a Canon MP-E 1–5X macro lens, combining the resulting stacks with Zerene Stacker (Richland, WA, USA).</p>
      <p>Species recognition was based on unique combinations of external and male genitalic characters. The abbreviations used in the text and figures are: <bold><abbrev xlink:title="abdomen length" id="ABBRID0EGGAC">AL</abbrev></bold>—abdomen length, <bold><abbrev xlink:title="body length" id="ABBRID0ELGAC">BL</abbrev></bold>—body length (from the anterior margin of the prolongation of the head to the posterior margin of tergite VIII), <bold><abbrev xlink:title="body width" id="ABBRID0EQGAC">BW</abbrev></bold>—body width (maximum width of elytra in dorsal view), <bold><abbrev xlink:title="elytral length" id="ABBRID0EVGAC">EL</abbrev></bold>—elytral length (maximum in dorsal view), <bold><abbrev xlink:title="elytral width" id="ABBRID0E1GAC">EW</abbrev></bold>—elytral width (maximum in dorsal view), <bold><abbrev xlink:title="head length" id="ABBRID0E6GAC">HL</abbrev></bold>—head length (from the anterior margin of the prolongation of the head to the posterior margin of the head disc in dorsal view), <bold><abbrev xlink:title="head width" id="ABBRID0EEHAC">HW</abbrev></bold>—head width (maximum, including eyes, in dorsal view), <bold><abbrev xlink:title="neck width" id="ABBRID0EJHAC">NW</abbrev></bold>—neck width (minimum in dorsal view), <bold><abbrev xlink:title="pronotum length" id="ABBRID0EOHAC">PL</abbrev></bold>—pronotum length (maximum in dorsal view), <bold><abbrev xlink:title="pronotum width" id="ABBRID0ETHAC">PW</abbrev></bold>—pronotum width (maximum in dorsal view).</p>
    </sec>
    <sec sec-type="﻿Results" id="SECID0EXHAC">
      <title>﻿Results</title>
      <sec sec-type="﻿Key to species of Metopiellus" id="SECID0E2HAC">
        <title>﻿Key to species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic></title>
        <p>Note: Adapted from <xref ref-type="bibr" rid="B1">Asenjo et al. (2017)</xref>, <xref ref-type="bibr" rid="B3">Fiorentino et al. (2022)</xref>, and <xref ref-type="bibr" rid="B4">Mário Chaul and Lopes-Andrade (2024)</xref>. See those references for additional images and illustrations.</p>
        <table-wrap content-type="key" position="anchor" orientation="portrait">
          <table id="TID0ENDAC" rules="all">
            <tbody>
              <tr>
                <td rowspan="1" colspan="1">1</td>
                <td rowspan="1" colspan="1">Vertex with projections; pronotum with two pairs of mediolateral thick, spinose projections, either simply convex or with a series of well-defined tumuli</td>
                <td rowspan="1" colspan="1">
                  <bold>2</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Vertex without projections</td>
                <td rowspan="1" colspan="1">
                  <bold>5</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">2</td>
                <td rowspan="1" colspan="1">Vertex with one horn-like projection</td>
                <td rowspan="1" colspan="1">
                  <bold>3</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Vertex with a pair of horn-like projections</td>
                <td rowspan="1" colspan="1">
                  <bold>4</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">3</td>
                <td rowspan="1" colspan="1">Pronotum with one pair of mediolateral, thick, spinose projections (Fig. <xref ref-type="fig" rid="F1">1A</xref>); each elytron with two basal foveae (Fig. <xref ref-type="fig" rid="F1">1A</xref>)</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Pronotum with two pairs of mediolateral, thick, spinose projections; basal elytral fovea absent</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic> Fiorentino et al., 2022</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">4</td>
                <td rowspan="1" colspan="1">Vertexal projections horn-like, apically acute; pronotum with two pairs of mediolateral, thick, spinose projections and one lateral process on each side; sternite VI with a pair of mediolateral setose processes and tergite VIII with a medial subquadrate thick process in the male</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="emavieirae">emavieirae</tp:taxon-name-part></tp:taxon-name></italic> Mário Chaul &amp; Lopes-Andrade, 2024</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Vertexal projections forming transverse pair of ridge-like projections (Fig. <xref ref-type="fig" rid="F1">1F</xref>); pronotum with single pair of obliquely transverse mediolateral ridge-like processes, the inner corner of each with acute point in the lateral pronotal one (Fig. <xref ref-type="fig" rid="F1">1E</xref>), in addition to acute lateral process on each side; male sternite VI and tergite VIII lacking secondary modifications</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">5</td>
                <td rowspan="1" colspan="1">Head similar in width to pronotum; eyes absent</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aglenus">aglenus</tp:taxon-name-part></tp:taxon-name></italic> (Reitter, 1885)</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Head narrower than pronotum; eyes small, but conspicuous or very reduced, inconspicuous</td>
                <td rowspan="1" colspan="1">
                  <bold>6</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">6</td>
                <td rowspan="1" colspan="1">Antennomere II about half as long as scape; antennomere V almost as long as antennomeres III and IV together</td>
                <td rowspan="1" colspan="1">
                  <bold>7</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Antennomere II less than half as long as scape; antennomere V shorter than the length of antennomeres III and IV together</td>
                <td rowspan="1" colspan="1">
                  <bold>8</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">7</td>
                <td rowspan="1" colspan="1">Antennomere VII rounded; dorsolateral margins strongly constricted anterad the eyes; paramere with apex bifurcated; median lobe curved and edge with long line of small teeth</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="crypticus">crypticus</tp:taxon-name-part></tp:taxon-name></italic> Asenjo et al., 2023</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Antennomere VII rectangular; dorsolateral margins of head gradually converging anterad the eyes; paramere not bifurcated apically; median lobe almost straight, with the apex curved, without small teeth along its length</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="painensis">painensis</tp:taxon-name-part></tp:taxon-name></italic> Asenjo et al., 2017</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">8</td>
                <td rowspan="1" colspan="1">Antennomere VIII transverse; eyes small, but conspicuous</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hirtus">hirtus</tp:taxon-name-part></tp:taxon-name></italic> (Reitter, 1885)</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Antennomere VIII obconical; eyes very reduced, inconspicuous</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="silvaticus">silvaticus</tp:taxon-name-part></tp:taxon-name></italic> Bruch, 1933</bold>
                </td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
      <sec sec-type="﻿Taxonomy" id="SECID0EUDAE">
        <title>﻿Taxonomy</title>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Coleoptera</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Staphylinidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">ECADB036-E18D-5186-866F-C8C250313EE4</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part>
              <object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/F67D4393-8C7F-4839-B576-FFC41B318BDD</object-id>
            </tp:taxon-name>
            <tp:taxon-status>sp. nov.</tp:taxon-status>
            <xref ref-type="fig" rid="F1">Figs 1A–C</xref>
            <xref ref-type="fig" rid="F2">, 2A–C</xref>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="type material" id="SECID0EJFAE">
            <title>Type material.</title>
            <p><bold><italic>Holotype</italic></bold> ♂ (<abbrev content-type="institution" xlink:title="National Biodiversity Institute collection" id="ABBRID0EUFAE">MECN</abbrev>-EN 23780): “Ecuador. Napo: Chontapunta, R. Minga, -0.88692, -77.27049, 300 m. Winkler. 07-feb-2023, Díaz-Guevara &amp; N. Berrazueta” / “<abbrev content-type="institution" xlink:title="National Biodiversity Institute collection" id="ABBRID0EZFAE">MECN</abbrev>-EN 23780”; deposited in <abbrev content-type="institution" xlink:title="National Biodiversity Institute collection" id="ABBRID0E5FAE">MECN</abbrev>.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="type material" id="SECID0EDGAE">
            <title>Additional, non-type material.</title>
            <p>Ecuador • 1 ♀; Orellana; <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-76.149900,-0.637600]}" id="NCID0EMGAE">0.6376°S, 76.1499°W</named-content></named-content>; Tiputini Biodiv. Sta.; 2-9.vi.2011; general hand collecting, AT1341, M. Caterino, A. Tishechkin; Caterino DNA voucher, Ext. MSC-12663; ZSFQ-i16787 • 1 ♀; Panayacu, Yasuní, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-76.671028,-0.704889]}" id="NCID0EUGAE">0°42'17.6"S, 76°40'15.7"W</named-content></named-content>; 340 m; Winkler; 17 Jul. 2021; Pazmiño-Palomino; <abbrev content-type="institution" xlink:title="National Biodiversity Institute collection" id="ABBRID0EZGAE">MECN</abbrev>-EN 48730.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Comparative diagnosis" id="SECID0E5GAE">
            <title>Comparative diagnosis.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. strongly resembles <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic>, having a vertex with a single horn-like projection. However, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> can be easily differentiated by its single pair of mediolateral, thick, spinose pronotal projections (vs. two pairs in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic>), the presence of basal elytral foveae (absent from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic>), and by aedeagal characters including a more elongate basal bulb, and differently shaped articulated armature (Fig. <xref ref-type="fig" rid="F2">2</xref>), bifid at the tip with subapical hook bent at ~90° (vs. aedeagal armature simply tapered, slightly sinuate at apex in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic>; aedeagal armature with four spatulate, curved tips in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="emavieirae">emavieirae</tp:taxon-name-part></tp:taxon-name></italic>).</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EVJAE">
            <title>Description.</title>
            <p>Avg. <abbrev xlink:title="body length" id="ABBRID0E2JAE">BL</abbrev> = 2.7 mm (<italic>N</italic> = 2; see Table <xref ref-type="table" rid="T1">1</xref> for other measurements). Body (Fig. <xref ref-type="fig" rid="F1">1A–C</xref>) densely setose and brown; eyes prominent, comprising ~11 ommatidia; head pyriform with two vertexal foveae near posterior margin with a single median process similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic>; antennae with 11 antennomeres, antennal scape base thick and slightly narrowed before the apex (in dorsal view), antennomere I as long as rest of the antennomeres combined, antennomere II about as long as III–VIII combined, III-IV and VI slightly longer than wide, V distinctly longer, VII globular, VIII small, IX and X slightly transverse, XI almost as long as IX–X combined, tapered in apical half; pronotum trapezoidal, wider anteriorly and narrower posteriorly, with two pairs of mediolateral, thick, spinose projections; each elytron with two basal foveae, and with longitudinal impressions extending from them posteriorly approximately two-thirds elytron length. Aedeagus (Fig. <xref ref-type="fig" rid="F2">2A–C</xref>) with large, elongate basal bulb, slightly asymmetrical, with smoothly rounded basal margin; dorsal diaphragm large, elongate oval; tegmen narrowed toward flat, paddle-shaped apex; elongate armature articulated within basal bulb, apex bifid, with acute, subapical hook bent at ~90°, other tip slightly diverging to left, acute.</p>
            <fig id="F1" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1266.169869.figure1</object-id>
              <object-id content-type="zenodo_dep_id">0</object-id>
              <object-id content-type="arpha">3BE3100E-BE6D-5B81-A6C9-ABC9C4CCA709</object-id>
              <label>Figure 1.</label>
              <caption>
                <p>Habitus of new <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> species. <bold>A–C.</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. (female dorsal, female lateral, male holotype ventral); <bold>D–F.</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. (male holotype dorsal, female paratype head + pronotum, male non-type head only). Scale bars: 1 mm (unless otherwise indicated).</p>
              </caption>
              <graphic xlink:href="zookeys-1266-127_article-169869__-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1507666.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1507666</uri>
              </graphic>
            </fig>
            <fig id="F2" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1266.169869.figure2</object-id>
              <object-id content-type="zenodo_dep_id">0</object-id>
              <object-id content-type="arpha">B1AF86ED-6D53-5BD9-B285-90DAED1766B7</object-id>
              <label>Figure 2.</label>
              <caption>
                <p>Aedeagus of new <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> species (dorsal, lateral, ventral). <bold>A–C.</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.; <bold>D–F.</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</p>
              </caption>
              <graphic xlink:href="zookeys-1266-127_article-169869__-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1507667.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1507667</uri>
              </graphic>
            </fig>
            <table-wrap id="T1" position="float" orientation="portrait">
              <label>Table 1.</label>
              <caption>
                <p>Measurements of all species, in millimeters (mm).</p>
              </caption>
              <table id="TID0E6RAE" rules="all">
                <tbody>
                  <tr>
                    <th rowspan="1" colspan="1">Species</th>
                    <th rowspan="1" colspan="1">
                      <italic>N</italic>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="abdomen length" id="ABBRID0EOOAE">AL</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="body length" id="ABBRID0EWOAE">BL</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="body width" id="ABBRID0E5OAE">BW</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="elytral length" id="ABBRID0EGPAE">EL</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="elytral width" id="ABBRID0EOPAE">EW</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="head length" id="ABBRID0EWPAE">HL</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="head width" id="ABBRID0E5PAE">HW</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="neck width" id="ABBRID0EGQAE">NW</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="pronotum length" id="ABBRID0EOQAE">PL</abbrev>
                    </th>
                    <th rowspan="1" colspan="1">
                      <abbrev xlink:title="pronotum width" id="ABBRID0EWQAE">PW</abbrev>
                    </th>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> (♂)</td>
                    <td rowspan="1" colspan="1">1</td>
                    <td rowspan="1" colspan="1">1.75</td>
                    <td rowspan="1" colspan="1">2.91</td>
                    <td rowspan="1" colspan="1">1.00</td>
                    <td rowspan="1" colspan="1">0.86</td>
                    <td rowspan="1" colspan="1">1.00</td>
                    <td rowspan="1" colspan="1">0.58</td>
                    <td rowspan="1" colspan="1">0.55</td>
                    <td rowspan="1" colspan="1">0.26</td>
                    <td rowspan="1" colspan="1">0.58</td>
                    <td rowspan="1" colspan="1">0.64</td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> (♀)</td>
                    <td rowspan="1" colspan="1">1</td>
                    <td rowspan="1" colspan="1">1.72</td>
                    <td rowspan="1" colspan="1">2.52</td>
                    <td rowspan="1" colspan="1">0.98</td>
                    <td rowspan="1" colspan="1">0.83</td>
                    <td rowspan="1" colspan="1">0.98</td>
                    <td rowspan="1" colspan="1">0.58</td>
                    <td rowspan="1" colspan="1">0.49</td>
                    <td rowspan="1" colspan="1">0.30</td>
                    <td rowspan="1" colspan="1">0.63</td>
                    <td rowspan="1" colspan="1">0.67</td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> (♂)</td>
                    <td rowspan="1" colspan="1">1</td>
                    <td rowspan="1" colspan="1">2.16</td>
                    <td rowspan="1" colspan="1">3.24</td>
                    <td rowspan="1" colspan="1">1.07</td>
                    <td rowspan="1" colspan="1">0.94</td>
                    <td rowspan="1" colspan="1">1.07</td>
                    <td rowspan="1" colspan="1">0.56</td>
                    <td rowspan="1" colspan="1">0.56</td>
                    <td rowspan="1" colspan="1">0.25</td>
                    <td rowspan="1" colspan="1">0.52</td>
                    <td rowspan="1" colspan="1">0.70</td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1"><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> (♀)</td>
                    <td rowspan="1" colspan="1">1</td>
                    <td rowspan="1" colspan="1">2.02</td>
                    <td rowspan="1" colspan="1">3.24</td>
                    <td rowspan="1" colspan="1">1.03</td>
                    <td rowspan="1" colspan="1">0.96</td>
                    <td rowspan="1" colspan="1">1.03</td>
                    <td rowspan="1" colspan="1">0.64</td>
                    <td rowspan="1" colspan="1">0.56</td>
                    <td rowspan="1" colspan="1">0.24</td>
                    <td rowspan="1" colspan="1">0.58</td>
                    <td rowspan="1" colspan="1">0.67</td>
                  </tr>
                </tbody>
              </table>
            </table-wrap>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="distribution" id="SECID0E5WAE">
            <title>Distribution.</title>
            <p>This species is known from the type locality in Chontapunta, Ecuador (Fig. <xref ref-type="fig" rid="F3">3</xref>), and probably also from the Tiputini region of Orellana (if the associations of the females under ‘other material’ are correct). The sites are separated by about 125 km, but are all on the same (south) side of the Rio Napo.</p>
            <fig id="F3" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1266.169869.figure3</object-id>
              <object-id content-type="zenodo_dep_id">0</object-id>
              <object-id content-type="arpha">B825332A-8F88-58E1-959A-C5447671921E</object-id>
              <label>Figure 3.</label>
              <caption>
                <p>Map of Ecuador showing type localities of the two new species.</p>
              </caption>
              <graphic xlink:href="zookeys-1266-127_article-169869__-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1507668.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1507668</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="etymology" id="SECID0EUXAE">
            <title>Etymology.</title>
            <p>The specific name derives from a Kichwa legend in which Palamaku is the divine source of insects.</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" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Coleoptera</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Staphylinidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">FABB52E7-EA8A-580E-8CCD-D2A5995C0AFA</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part>
              <object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/636C2C0D-06BF-4B70-A4C3-4B1DA9D2F850</object-id>
            </tp:taxon-name>
            <tp:taxon-status>sp. nov.</tp:taxon-status>
            <xref ref-type="fig" rid="F1">Figs 1D–F</xref>
            <xref ref-type="fig" rid="F2">, 2D–F</xref>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="type material" id="SECID0EKZAE">
            <title>Type material.</title>
            <p><bold><italic>Holotype</italic></bold> ♂ (<named-content content-type="dwc:institutional_code" xlink:title="Pontificia Universidad Católica del Ecuador" xlink:href="http://grbio.org/institution/museo-de-zoologia-pontificia-universidad-catolica-del-ecuador">QCAZ</named-content> 280376): “Ecuador. Pichincha: Otongachi Reserve <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-78.934420,-0.330510]}" id="NCID0E4ZAE">-0.330510, -78.934420</named-content></named-content>, 1087m.20-Feb-2020, E. Tapia &amp; N. Dupérré”; deposited in <named-content content-type="dwc:institutional_code" xlink:title="Pontificia Universidad Católica del Ecuador" xlink:href="http://grbio.org/institution/museo-de-zoologia-pontificia-universidad-catolica-del-ecuador">QCAZ</named-content>. <bold><italic>Paratypes</italic></bold> (4 ♀, same general locality as type): <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-78.934420,-0.330510]}" id="NCID0EN1AE">-0.330510, -78.934420</named-content></named-content>, 1087 m (<named-content content-type="dwc:institutional_code" xlink:title="Pontificia Universidad Católica del Ecuador" xlink:href="http://grbio.org/institution/museo-de-zoologia-pontificia-universidad-catolica-del-ecuador">QCAZ</named-content> 280377 to 280380).</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="type material" id="SECID0EX1AE">
            <title>Additional, non-type material.</title>
            <p>Ecuador • 4 ♂, 4 ♀: Zamora Chinchipe; Las Orquídeas; <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-78.659500,-4.248200]}" id="NCID0EA2AE">-4.2482, -78.6595</named-content></named-content>; 877 m; E. Tapia; Sifting litter; 29-Jul-2024; <named-content content-type="dwc:institutional_code" xlink:title="Pontificia Universidad Católica del Ecuador" xlink:href="http://grbio.org/institution/museo-de-zoologia-pontificia-universidad-catolica-del-ecuador">QCAZ</named-content> 280381 to 280388.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Comparative diagnosis" id="SECID0EK2AE">
            <title>Comparative diagnosis.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> resembles <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="emavieirae">emavieirae</tp:taxon-name-part></tp:taxon-name></italic> in the presence of a pair of (rather than a single) vertexal structures and by having two pairs of mediolateral pronotal projections. However, it can be distinguished from it by the vertexal and median pronotal processes being transversely ridge-like, and by the lateral pronotal processes having a posterior acute point (vs. horn-like vertexal and pronotal structures in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="emavieirae">emavieirae</tp:taxon-name-part></tp:taxon-name></italic>). The aedeagi differ strongly, with that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> having a strongly bifurcate dorsal armature, and that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="emavieirae">emavieirae</tp:taxon-name-part></tp:taxon-name></italic> being subdivided subapically into four curved, spatulate tips.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EH4AE">
            <title>Description.</title>
            <p>Avg. <abbrev xlink:title="body length" id="ABBRID0EN4AE">BL</abbrev> = 3.2 mm (<italic>N</italic> = 2; see Table <xref ref-type="table" rid="T1">1</xref> for other measurements). Body (Fig. <xref ref-type="fig" rid="F1">1D–F</xref>) densely setose and brown-reddish; eyes prominent, borne on small elevated base, comprising ~8 ommatidia; head widest at base, rounded, narrowed anterad, with two posterior vertexal foveae (Fig. <xref ref-type="fig" rid="F1">1E</xref>) immediately in front of two low posterior vertexal ridges; antennae with 11 antennomeres, antennal scape about as long as antennomeres II–VIII combined, antennomere II as long as III–VII combined, III–VI elongate, V longer than others, VII globular, VIII smallest, IX–XI forming weak club, IX and X transverse, XI almost as long as IX and X combined, tapered in apical two-thirds; pronotum densely setose, with 4 acute lobes, lateral lobes each with posterior acute point, median ones forming oblique ridges, prominent at their inner corners; each elytron with two basal foveae, and with longitudinal impressions extending from them posteriorly approximately two-thirds elytron length; protibia rather slender. Aedeagus (Fig. <xref ref-type="fig" rid="F2">2D–F</xref>) with basal bulb asymmetrical and globose, with basal rounded margin, elevated distad, dorsal diaphragm elongate oval; tegmen with ventral portion long, slender, curved dorsad and then distad apically; articulated armature robust, deeply bifurcated, one bifurcation is hook-like, longer and evenly tapered, other more robust with small projection oriented towards distal apex of basal bulb.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="distribution" id="SECID0ED5AE">
            <title>Distribution.</title>
            <p>This species probably only occurs in the Otongachi Reserve in northern Ecuador, in the province of Pichincha (Fig. <xref ref-type="fig" rid="F3">3</xref>). We believe the ‘additional material’ cited from Zamora-Chinchipe to be mislabeled, since that locality is over 400 km to the south, and located on the Amazonian side of the Andean crest, an unlikely distribution for a flightless beetle species. The male genitalia is identical between the two alleged localities.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="etymology" id="SECID0EN5AE">
            <title>Etymology.</title>
            <p>The specific epithet “chasqui” refers to the agile messengers of the Inca Empire, evoking the beetle’s slender posterior legs.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
    </sec>
    <sec sec-type="﻿Discussion" id="SECID0E35AE">
      <title>﻿Discussion</title>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> now contains nine known species. These two new species extend the geographic range of the genus into Ecuador. The closest previous record was from Amazonian Colombia, while the bulk of the species are known from eastern and southeastern Brazil. Given the existence of the Colombian species (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic>), a record of the genus from the Ecuadorean Amazon shouldn’t be too surprising. However, the new species from the cloud forests of the western slope of the Andes represents an important extension not only geographically but also ecologically. Clearly, further sampling of leaf-litter habitats will result in continued discovery of new species across the Neotropical region.</p>
      <p><xref ref-type="bibr" rid="B4">Mário Chaul and Lopes-Andrade (2024)</xref> considered their <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="emavieirae">emavieirae</tp:taxon-name-part></tp:taxon-name></italic> to be closely related to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic>, but our <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="palamaku">palamaku</tp:taxon-name-part></tp:taxon-name></italic> is even closer to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guanano">guanano</tp:taxon-name-part></tp:taxon-name></italic>, differing almost exclusively in fairly minor aedeagal characters. So if species groups are to be delineated, the three should be grouped together. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chasqui">chasqui</tp:taxon-name-part></tp:taxon-name></italic> also shares the horn-like projections of the vertex that these three do, and would appear to be closely related as well, although its aedeagus is radically different from any of the others. Of these four, only <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="emavieirae">emavieirae</tp:taxon-name-part></tp:taxon-name></italic> exhibits any significant male secondary sexual characters, with projections of the terminal tergite and sternites.</p>
      <p>We have little to contribute to speculation on the general natural history of species in the genus, with all our specimens coming from fairly generic leaf-litter samples. Early records of some species associated with ants’ colonies have not been repeated, and the collection of one <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">Metopiellus</tp:taxon-name-part></tp:taxon-name></italic> species (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metopiellus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="painensis">painensis</tp:taxon-name-part></tp:taxon-name></italic>; <xref ref-type="bibr" rid="B1">Asenjo et al. 2017</xref>) from caves suggests something rather different: that members of the genus may have subterranean propensities, habits that would be consistent with their generally reduced eyes and elongated appendages. We can only hope that future direct collections take closer note of more significant natural history associations to further illuminate the biology of these impressive pselaphine beetles.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>﻿Acknowledgements</title>
      <p>We express our heartfelt gratitude to Minga Lodge &amp; Reserve, Kamana Pacha, and the Minga Community Development Foundation for their invaluable support. A special acknowledgment to Tamia Camila Torres for her dedication and leadership in promoting taxonomy and conservation initiatives at the Minga Reserve. We are also grateful to Giovanni Onore, David Melo, and Gabriela Simbaña of the Otonga Foundation for their continued collaboration. MSC thanks the staff of the Tiputini Biodiversity Station and the Zoological Museum of the Universidad San Francisco de Quito (ZSFQ), past and present, for their assistance with fieldwork and specimen management. We sincerely thank Alex Pazmiño-Palomino and Ana B. García-Ruilova, researchers at the National Institute of Biodiversity (INABIO), for their essential contributions. We also extend our deep appreciation to Doris Vela, curator of the invertebrate collection at the <named-content content-type="dwc:institutional_code" xlink:title="Pontificia Universidad Católica del Ecuador" xlink:href="http://grbio.org/institution/museo-de-zoologia-pontificia-universidad-catolica-del-ecuador">QCAZ</named-content> Zoology Museum, and Fernanda Salazar, collection manager at the Pontificia Universidad Católica del Ecuador, for their dedicated support and scientific insight. This represents Technical Contribution No. 7488 of the Clemson University Experiment Station.</p>
    </ack>
    <sec sec-type="﻿Additional information" id="SECID0E6CAG">
      <title>﻿Additional information</title>
      <sec sec-type="Conflict of interest" id="SECID0EDDAG">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="SECID0EIDAG">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Use of AI" id="SECID0ENDAG">
        <title>Use of AI</title>
        <p>No use of AI was reported.</p>
      </sec>
      <sec sec-type="Funding" id="SECID0ESDAG">
        <title>Funding</title>
        <p>No funding was reported.</p>
      </sec>
      <sec sec-type="Author contributions" id="SECID0EXDAG">
        <title>Author contributions</title>
        <p>Conceptualization: YTA, MSC, DRDG. Data curation: DRDG, YTA, MSC. Investigation: YTA, MSC, DRDG. Writing – original draft: MSC, DRDG, YTA. Writing – review and editing: YTA, MSC, DRDG.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="SECID0E3DAG">
        <title>Author ORCIDs</title>
        <p>Yarina Tapuy-Avilés <ext-link xlink:href="https://orcid.org/0009-0001-2357-7916" ext-link-type="uri" xlink:type="simple">https://orcid.org/0009-0001-2357-7916</ext-link></p>
        <p>David R. Díaz-Guevara <ext-link xlink:href="https://orcid.org/0009-0001-4632-4460" ext-link-type="uri" xlink:type="simple">https://orcid.org/0009-0001-4632-4460</ext-link></p>
        <p>Michael S. Caterino <ext-link xlink:href="https://orcid.org/0000-0002-2597-5707" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-2597-5707</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="SECID0ESEAG">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text.</p>
      </sec>
    </sec>
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</article>
