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<article xmlns:tp="http://www.plazi.org/taxpub" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" article-type="rapid-communication" xml:lang="en">
  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">98</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:3686FA2D-1003-5130-A1A9-6F0A4E63DF4D</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:A3B9B45C-7F69-42DF-85B6-B3189A5B43F2</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Caucasiana</journal-title>
        <abbrev-journal-title xml:lang="en">Caucasiana</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="epub">2667-9809</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/caucasiana.5.e176387</article-id>
      <article-id pub-id-type="publisher-id">176387</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Short Communication</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Animalia</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biodiversity &amp; Conservation</subject>
          <subject>Biogeography</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>New Light on Old Ghosts: First Record of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part></tp:taxon-name></italic> Fabricius, 1775 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Lepidoptera">Lepidoptera</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name>) from Georgia</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Japaridze</surname>
            <given-names>Lasha-Giorgi</given-names>
          </name>
          <email xlink:type="simple">lgjaparidze@gmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0001-7171-5589</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Makharadze</surname>
            <given-names>Giorgi</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0008-4443-5432</uri>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Petrov</surname>
            <given-names>Valeri</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0006-4125-894X</uri>
          <xref ref-type="aff" rid="A3">3</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Grehan</surname>
            <given-names>John R.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-3119-1140</uri>
          <xref ref-type="aff" rid="A4">4</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Agricultural University of Georgia, Tbilisi, Georgia</addr-line>
        <institution>Agricultural University of Georgia</institution>
        <addr-line content-type="city">Tbilisi</addr-line>
        <country>Georgia</country>
        <uri content-type="ror">https://ror.org/01mdqeh78</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Unaffiliated, Tbilisi, Georgia</addr-line>
        <institution>Unaffiliated</institution>
        <addr-line content-type="city">Tbilisi</addr-line>
        <country>Georgia</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Simon Janashia Museum of Georgia, Tbilisi, Georgia</addr-line>
        <institution>Simon Janashia Museum of Georgia</institution>
        <addr-line content-type="city">Tbilisi</addr-line>
        <country>Georgia</country>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">OpenScienceLab NZ, Ngaio, New Zealand</addr-line>
        <institution>OpenScienceLab NZ</institution>
        <addr-line content-type="city">Ngaio</addr-line>
        <country>New Zealand</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Lasha-Giorgi Japaridze (<email xlink:type="simple">lgjaparidze@gmail.com)</email></p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Levan Mumladze</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>03</day>
        <month>04</month>
        <year>2026</year>
      </pub-date>
      <volume>5</volume>
      <fpage>25</fpage>
      <lpage>29</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/63940495-4860-5ED9-8A7D-DC0E2F9E9C56">63940495-4860-5ED9-8A7D-DC0E2F9E9C56</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/96964EA9-2E46-484E-A676-EC999E172F47">96964EA9-2E46-484E-A676-EC999E172F47</uri>
      <history>
        <date date-type="received">
          <day>29</day>
          <month>10</month>
          <year>2025</year>
        </date>
        <date date-type="accepted">
          <day>25</day>
          <month>12</month>
          <year>2025</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Lasha-Giorgi Japaridze, Giorgi Makharadze, Valeri Petrov, John R. Grehan</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/96964EA9-2E46-484E-A676-EC999E172F47</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>This study reviews the diversity of ghost moths (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name>) in Georgia, including the first documentation of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part></tp:taxon-name></italic> Fabricius, 1775 with the recording of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="humuli">humuli</tp:taxon-name-part></tp:taxon-name></italic> (Linnaeus, 1758) based on a two male and a single female specimens collected in 3 regions of Georgia in 1961, 1962 and 1970. The newly recorded genus also represents the first record for the South Caucasus more widely.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>Biodiversity</kwd>
        <kwd>biogeography</kwd>
        <kwd>
          <italic>
            <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="humuli">humuli</tp:taxon-name-part></tp:taxon-name>
          </italic>
        </kwd>
        <kwd>new record</kwd>
      </kwd-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="sec1">
        <title>Citation</title>
        <p>Japaridze L-G, Makharadze G, Petrov V, Grehan JR (2026) New Light on Old Ghosts: First Record of the genus Hepialus Fabricius, 1775 (Lepidoptera, Hepialidae) from Georgia. Caucasiana 5: 25–29. <ext-link xlink:href="10.3897/caucasiana.4.e151922" ext-link-type="doi">https://doi.org/10.3897/caucasiana.5.e176387</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec2">
      <title>Introduction</title>
      <p>The family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name> Stephens, 1828, commonly known as swift moths or ghost moths, represents the most diverse family among the primitive <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Lepidoptera">Lepidoptera</tp:taxon-name-part></tp:taxon-name>. With 725 described species (<xref ref-type="bibr" rid="B4">Grehan et al. 2023</xref>; <xref ref-type="bibr" rid="B6">Mielke et al. 2025</xref>), it is the principal family within the infraorder <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Exoporia">Exoporia</tp:taxon-name-part></tp:taxon-name> and is the sister group to the Heteroneura, together encompassing over 90% of all extant butterfly and moth species (<xref ref-type="bibr" rid="B9">Regier et al. 2015</xref>). Globally, the larvae of most <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name> species are subterranean, feeding in, or on, the roots of woody or herbaceous plants, or consuming fallen foliage. Another major feeding mode involves boring into the stems and branches of trees and shrubs, where larvae feed on callus regrowth at the stem surface, or in some species, on soft woody tissues. A smaller minority of species feed on plant detritus throughout development (<xref ref-type="bibr" rid="B4">Grehan et al. 2023</xref>).</p>
      <p><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name> are distributed across all major landmasses worldwide except for Antarctica, and regional absences in Madagascar, the Caribbean islands, and tropical West Africa (<xref ref-type="bibr" rid="B8">Nielsen et al. 2000</xref>; <xref ref-type="bibr" rid="B4">Grehan et al. 2023</xref>). Eight genera comprising a total of 21 species are currently recorded from northern Eurasia. Most species in this group are endemic or represented in the region between Europe and the Caspian Sea (<xref ref-type="bibr" rid="B3">Grehan and Knyazev 2019</xref>). Their southern distributional boundary is the Mediterranean Basin, and eastwards through Jordan, Türkiye, Georgia, and northern Iran. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name> previously known from Georgia are the regionally endemic or/and relic species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Zenophassus">Zenophassus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="schamyl">schamyl</tp:taxon-name-part></tp:taxon-name></italic> (Christoph, 1888), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triodia">Triodia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="laetus">laetus</tp:taxon-name-part></tp:taxon-name></italic> (Staudinger, 1877), and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triodia">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mlokossevitschi">mlokossevitschi</tp:taxon-name-part></tp:taxon-name></italic> (Romanoff, 1884), along with the more broadly distributed European species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Korscheltellus">Korscheltellus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lupulinus">lupulinus</tp:taxon-name-part></tp:taxon-name></italic> (Linnaeus, 1758) (<xref ref-type="bibr" rid="B3">Grehan and Knyazev 2019</xref>). While the natural history of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triodia">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="laetus">laetus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triodia">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mlokossevitschi">mlokossevitschi</tp:taxon-name-part></tp:taxon-name></italic> remains poorly understood, the specific status of the latter is uncertain, as the type specimen has not yet been dissected and described in accordance with the modern standards.</p>
      <p>The early contributions of <xref ref-type="bibr" rid="B10">Romanoff (1884)</xref> and <xref ref-type="bibr" rid="B1">Christoph (1889)</xref>, and the more recent study by <xref ref-type="bibr" rid="B5">Knyazev (2025)</xref> notwithstanding, the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name> fauna of the Caucasus Mountains region comprising Georgia, Russia, northeastern Türkiye, Armenia, and Azerbaijan, has received limited taxonomic attention. Although the biology of the endemic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triodia">Triodia</tp:taxon-name-part></tp:taxon-name></italic> species is poorly known, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Zenophassus">Z.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="schamyl">schamyl</tp:taxon-name-part></tp:taxon-name></italic> has been studied in detail because of its status as a pest of grapevine (<xref ref-type="bibr" rid="B12">Slatshevskij 1929</xref>; <xref ref-type="bibr" rid="B7">Milyanovskii and Mitrofanov 1952</xref>).</p>
      <p>Georgia, together with other countries bordering or overlapping the Caucasus Mountains, occupies a potentially significant boundary for the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name>. The western Eurasian genera, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gazoryctra">Gazoryctra</tp:taxon-name-part></tp:taxon-name></italic> Hübner, 1821, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part></tp:taxon-name></italic> Fabricius, 1775, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pharmacis">Pharmacis</tp:taxon-name-part></tp:taxon-name></italic> Hübner, 1820, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phymatopus">Phymatopus</tp:taxon-name-part></tp:taxon-name></italic> Wallengren, 1869 have previously published distribution ranges north of the Caucasus while <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Korscheltellus">Korscheltellus</tp:taxon-name-part></tp:taxon-name></italic> Börner, 1920 has a southern distributional limit in Georgia. Only <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triodia">Triodia</tp:taxon-name-part></tp:taxon-name></italic> has a range extending well south of the Caucasus, reaching southern Israel and Jordan, however, as with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triodia">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mlokossevitschi">mlokossevitschi</tp:taxon-name-part></tp:taxon-name></italic>, the taxonomic status of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Triodia">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="froitzhami">froitzhami</tp:taxon-name-part></tp:taxon-name></italic> from Israel and Jordan is also uncertain as the type has not been dissected and there are no clear external diagnostic differences.</p>
    </sec>
    <sec sec-type="Results" id="sec3">
      <title>Results</title>
      <sec sec-type="Genus Hepialus Fabricius, 1775" id="sec4">
        <title>Genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part></tp:taxon-name></italic> Fabricius, 1775</title>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="kingdom" reg="Animalia">Animalia</tp:taxon-name-part>
                  </tp:taxon-name>
                </named-content>
              </kwd>
              <kwd>
                <named-content content-type="order">Lepidoptera</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family">Hepialidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <tp:taxon-name><object-id content-type="arpha">56E2C86C-735D-5E9B-93CD-7CDE02DF976A</object-id>
                    		<tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="humuli">humuli</tp:taxon-name-part>
                    	</tp:taxon-name>
            <tp:taxon-authority>(Linnaeus, 1758)</tp:taxon-authority>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="Material examined">
            <title>Material examined.</title>
            <p>GEORGIA • 1♂; Khevsureti (without exact locality); 1800 m. a.s.l.; 12 July 1970; leg: E. Didmanidze; Simon Janashia Museum of Georgia (SJMG); • 1♂; Martvili; 4 July 1962; leg: E. Didmanidze; SJMG; • 1♀; Bakhmaro; 18 July 1961; leg: E. Didmanidze; SJMG.</p>
            <p>Recent examination of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Lepidoptera">Lepidoptera</tp:taxon-name-part></tp:taxon-name> collection housed in the Simon Janashia Museum of Georgia has yielded several specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part></tp:taxon-name></italic> Fabricius, 1775. The material comprises two males and a single female of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="humuli">humuli</tp:taxon-name-part></tp:taxon-name></italic> (Linnaeus, 1758), from the Georgian regions of Mtskheta-Mtianeti (Khevsureti), Samegrelo-Zemo Svaneti (Martvili), and Guria (Bakhmaro). These records are significant as they suggest that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">H.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="humuli">humuli</tp:taxon-name-part></tp:taxon-name></italic> has been present in Georgia for at least 64 years, although it has rarely been collected, probably owing to its crepuscular lifestyle and occurrence in remote mountainous regions. The species is widespread across Europe and eastwards to central Siberia (<xref ref-type="bibr" rid="B11">Simonsen and Huemer 2014</xref>; <xref ref-type="bibr" rid="B3">Grehan and Knyazev 2019</xref>). In western Eurasia, the southern distributional limit of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">H.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="humuli">humuli</tp:taxon-name-part></tp:taxon-name></italic> is along the Pyrenees and north of the Mediterranean along the European Alps, Balkans, and north of the Black Sea. Its presence in the Caucasus (North Ossetia) was reported by <xref ref-type="bibr" rid="B2">Dobronosov and Komarov (2017)</xref>, suggesting that its occurrence in northern Georgia was to be expected. The Georgian record, long overlooked, raises the question as to whether the species may also occur further south in Central Asia, particularly in Kazakhstan and adjacent countries.</p>
            <fig id="F1">
              <object-id content-type="doi">10.3897/caucasiana.5.e176387.figure1</object-id>
              <object-id content-type="arpha">059416C6-7014-560A-8D65-B6BD7A5492AC</object-id>
              <label>Figure 1.</label>
              <caption>
                <p>General habitus of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hepialus">Hepialus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="humuli">humuli</tp:taxon-name-part></tp:taxon-name></italic> (Linnaeus, 1758). <bold>1</bold>: male, Martvili; <bold>2</bold>: male, Khevsureti; <bold>3</bold>: female, Bakhmaro. Scale bars: 10 mm. Photographs were taken with a Canon EOS 60D camera fitted with a Canon EF 60 mm f/2.8 Macro USM lens. Image stacks were processed using Helicon Focus 3.2 and subsequently refined in Adobe Photoshop CS6.</p>
              </caption>
              <graphic xlink:href="caucasiana-05-025-g001.jpg" id="oo_1583083.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1583083</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
      <sec sec-type="Biogeographic significance" id="sec5">
        <title>Biogeographic significance</title>
        <p>The localised endemism of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name> in the Caucasus is comparable to that observed in other montane regions along the northern margin of the Mediterranean Basin. It has been suggested by <xref ref-type="bibr" rid="B3">Grehan and Knyazev (2019)</xref> that diversification of the northern Eurasian genera was influenced by tectonic history and phases of geological uplift resulting from closure of the western Tethys Sea. The only north African representative of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name> immediately south of the Mediterranean is the monotypic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Neohepialiscus">Neohepialiscus</tp:taxon-name-part></tp:taxon-name></italic> Viette, 1948, which may be more related to oxycanine veined genera in southeastern Asia and Australasia than to European genera (Grehan and c 2023).</p>
      </sec>
    </sec>
    <sec sec-type="Acknowledgements" id="sec6">
      <title>Acknowledgements</title>
      <p>We are grateful to the administration of the Simon Janashia Museum of Georgia for granting permission to study their <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Hepialidae">Hepialidae</tp:taxon-name-part></tp:taxon-name> collection. Armen Seropian (Tbilisi, Georgia) kindly reviewed the initial draft and provided valuable suggestions. Thanks to anonymous reviewers for the valuable comments and suggestions.</p>
    </sec>
  </body>
  <back>
    <sec sec-type="Additional information" id="sec7">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec8">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec9">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Funding" id="sec10">
        <title>Funding</title>
        <p>No funding was reported.</p>
      </sec>
      <sec sec-type="Author contributions" id="sec11">
        <title>Author contributions</title>
        <p>Conceptualization (L-GJ), original draft writing (L-GJ, JRG), collection material (L-GJ, VP), visualization (GM), data curation (L-GJ, VP).</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec12">
        <title>Author ORCIDs</title>
        <p>Lasha-Giorgi Japaridze <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0001-7171-5589">https://orcid.org/0000-0001-7171-5589</ext-link></p>
        <p>Giorgi Makharadze <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0009-0008-4443-5432">https://orcid.org/0009-0008-4443-5432</ext-link></p>
        <p>Valeri Petrov <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0009-0006-4125-894X">https://orcid.org/0009-0006-4125-894X</ext-link></p>
        <p>John R. Grehan <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0002-3119-1140">https://orcid.org/0000-0002-3119-1140</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec13">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text or Supplementary Information.</p>
      </sec>
    </sec>
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</article>
