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<article xmlns:tp="http://www.plazi.org/taxpub" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" article-type="research-article" xml:lang="en">
  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">101</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:DF2AD546-A4F0-51AC-B38C-88216742E553</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:9B96196F-9505-457E-8F01-0E12D644C289</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Zitteliana</journal-title>
        <abbrev-journal-title xml:lang="en">ZITT</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">2512-5338</issn>
      <issn pub-type="epub">2747-8106</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/zitteliana.100.197055</article-id>
      <article-id pub-id-type="publisher-id">197055</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Psychidae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Palaeontology</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Paleogene</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Russia</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Adult male bagworm moths from Ukrainian and Baltic Eocene amber (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Lepidoptera">Lepidoptera</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>Arnscheid</surname>
            <given-names>Wilfried R.</given-names>
          </name>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Hausmann</surname>
            <given-names>Axel</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-0358-9928</uri>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Gröhn</surname>
            <given-names>Carsten</given-names>
          </name>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Fischer</surname>
            <given-names>Thilo C.</given-names>
          </name>
          <email xlink:type="simple">thilo.c.fischer@gmx.de</email>
          <uri content-type="orcid">https://orcid.org/0000-0001-7371-4053</uri>
          <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>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Staatliches Museum für Naturkunde Karlsruhe, Erbprinzenstr. 13, 76133 Karlsruhe, Germany</addr-line>
        <institution>University of Hamburg</institution>
        <addr-line content-type="city">Hamburg</addr-line>
        <country>Germany</country>
        <uri content-type="ror">https://ror.org/00g30e956</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">SNSB - Bavarian State Collection of Zoology, Münchhausenstraße 21, D-81247 Munich, Germany</addr-line>
        <institution>Staatliches Museum für Naturkunde Karlsruhe</institution>
        <addr-line content-type="city">Karlsruhe</addr-line>
        <country>Germany</country>
        <uri content-type="ror">https://ror.org/035hn3t86</uri>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">University of Hamburg, Mittelweg 177, 20148 Hamburg, Germany</addr-line>
        <institution>SNSB - Bavarian State Collection of Zoology</institution>
        <addr-line content-type="city">Munich</addr-line>
        <country>Germany</country>
        <uri content-type="ror">https://ror.org/04rekk491</uri>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Thilo C. Fischer (<email xlink:type="simple">thilo.c.fischer@gmx.de</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Michael Krings</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>22</day>
        <month>06</month>
        <year>2026</year>
      </pub-date>
      <volume>100</volume>
      <fpage>117</fpage>
      <lpage>127</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/E995C99D-D4A0-5B90-9E20-2B1F3F06B59F">E995C99D-D4A0-5B90-9E20-2B1F3F06B59F</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/0266C759-5734-47A6-8A56-3B2A2C500637">0266C759-5734-47A6-8A56-3B2A2C500637</uri>
      <history>
        <date date-type="received">
          <day>24</day>
          <month>04</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>28</day>
          <month>05</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Wilfried R. Arnscheid, Axel Hausmann, Carsten Gröhn, Thilo C. Fischer</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/0266C759-5734-47A6-8A56-3B2A2C500637</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>Psychid moths from Ukrainian and Baltic amber are described for the first time by male adults. Recognition was possible by their pectinate antennae and reduced mouthparts at first. In combination with further characters, affiliation with the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part></tp:taxon-name> and the extant genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> is established. This affiliation is in accordance with the previous description of a bag belonging to this genus in Baltic amber; three new specimens of such bags with caterpillars are also reported here. Conclusions are drawn on the mode of life of the fossil species by comparison to the extant species.</p>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>New species</kwd>
        <kwd>pectinate antenna</kwd>
        <kwd>
          <italic>
            <tp:taxon-name>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part>
            </tp:taxon-name>
          </italic>
        </kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>taphonomy</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec1">
      <title>Introduction</title>
      <p><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> (bagworm moths) are a family of basal ditrysians within the superfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily" reg="Tineoidea">Tineoidea</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B2">Davis 1964</xref>; <xref ref-type="bibr" rid="B3">Davis and Robinson 1998</xref>). The moths are small to medium sized; males are fully winged, whereas females of most subfamilies are flightless, brachypterous, apterous, or larviform. Females of several species spend their entire lives in their larval bags (<xref ref-type="bibr" rid="B9">Scoble 1995</xref>; <xref ref-type="bibr" rid="B3">Davis and Robinson 1998</xref>; <xref ref-type="bibr" rid="B7">Rhainds et al. 2009</xref>; <xref ref-type="bibr" rid="B1">Arnscheid and Weidlich 2017</xref>). <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> are one of the few families in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Lepidoptera">Lepidoptera</tp:taxon-name-part></tp:taxon-name> in which parthenogenesis occurs; this has been established in several European and Asiatic species (<xref ref-type="bibr" rid="B1">Arnscheid and Weidlich 2017</xref>). The family comprises some thousand species; larval development takes place in bags formed of silk and covered by exogenous particles or faeces (<xref ref-type="bibr" rid="B7">Rhainds et al. 2009</xref>). <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> are polyphagous and caterpillars defoliate a broad spectrum of host plants; however, omnivory occurs in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Oiketicinae">Oiketicinae</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B7">Rhainds et al. 2009</xref>). As males are quite short-lived (one to two days, with females living a little longer, up to one week) and do not feed, their mouthparts (proboscis, maxillary and labial palpi) are rudimentary and useless. Antennae can be bipectinate (<xref ref-type="bibr" rid="B3">Davis and Robinson 1998</xref>), which may serve to facilitate the search for females by their released pheromones.</p>
      <p>Bags of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name>, often with caterpillars in them, are common finds in succinite (Baltic and Bitterfeld amber, Ukrainian amber) and are reported in both scientific as well as popular literature on amber inclusions. Multiple taxa have been described, among them is a single bag attributed to the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B11">Sobczyk and Kobbert 2009</xref>). However, imagos, and especially the winged males of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> in general, were unknown from succinite (<xref ref-type="bibr" rid="B12">Sohn et al. 2012</xref>; <xref ref-type="bibr" rid="B13">Sohn and Lamas 2013</xref>; and later literature on fossil <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Lepidoptera">Lepidoptera</tp:taxon-name-part></tp:taxon-name>). Due to their short lifespan of hours to only one to two days (<xref ref-type="bibr" rid="B7">Rhainds et al. 2009</xref>), they can be expected to be underrepresented in amber compared to long-lived lepidopteran imagos. Furthermore, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name>, as members of superfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily" reg="Tineoidea">Tineoidea</tp:taxon-name-part></tp:taxon-name>, may easily be mistaken, due to their similar habitus, for <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Tineidae">Tineidae</tp:taxon-name-part></tp:taxon-name>, which represent some thirty percent of all adult moths in Baltic amber (<xref ref-type="bibr" rid="B10">Skalski 1976</xref>). Such a bias against <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> moths is also reported for collections of extant moths (<xref ref-type="bibr" rid="B6">Parenti 2000</xref>). However, despite their overall tineoid habitus, psychid male moths can be detected when they show bipectinate antennae and strongly reduced mouthparts.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec2">
      <title>Material and methods</title>
      <p>The two Ukrainian ambers with the described imagos are kept in sealed plastic bags inside a metal box, excluding access of air and light, currently being kept in the collection of the author TF, and to be transferred to the Bavarian State Collection of Natural History upon publication. The Baltic amber specimen is also kept in a sealed plastic bag and stored in the collection of author CG. Authenticity of the specimen was ensured by obtaining them from trusted amber dealers known from long-term relationships. Furthermore, the ambers contain trichomes typical for succinite ambers.</p>
    </sec>
    <sec sec-type="Results" id="sec3">
      <title>Results</title>
      <sec sec-type="Identification of bipectinate moth specimens" id="sec4">
        <title>Identification of bipectinate moth specimens</title>
        <p>The three specimens from Eocene ambers with bipectinate antennae were studied by comparison with many <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Lepidoptera">Lepidoptera</tp:taxon-name-part></tp:taxon-name> families in which such antennae have been described. All families except <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> could be excluded (Table <xref ref-type="table" rid="T1">1</xref>, top part). On this basis, subfamilies of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> were checked for more detailed characters. Special focus was on subfamilies that have been reported from Baltic amber via their bags (<xref ref-type="bibr" rid="B11">Sobczyk and Kobbert 2009</xref>). The SNSB collection was used for comparison with male moths of extant genera, resulting in an affiliation to subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part></tp:taxon-name> (Table <xref ref-type="table" rid="T1">1</xref>, middle part).</p>
        <table-wrap id="T1" position="float" orientation="portrait">
          <label>Table 1.</label>
          <caption>
            <p>Family identification of the new amber records.</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <th rowspan="1" colspan="2">
                  <bold>Family identification</bold>
                </th>
              </tr>
              <tr>
                <td rowspan="1" colspan="2">Bipectinate fam. candidate</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="family" reg="Cossidae">Cossidae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">too large, bushy palpi</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="family" reg="Zygaenidae">Zygaenidae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">too large, long palpi, eyes smaller, antennal branches short</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1"><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Geometridae">Geometridae</tp:taxon-name-part></tp:taxon-name> - Scopula</td>
                <td rowspan="1" colspan="1">too large, proboscis well-developed</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="family" reg="Incurvariidae">Incurvariidae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">larger, bushy palpi, antennal branches clavate</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="family" reg="Prodoxidae">Prodoxidae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">larger, bushy palpi, antennal branches clavate</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="family" reg="Eriocottidae">Eriocottidae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">too large, bushy palpi, antennal branches longer</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">
                  <bold>adult size and antennal pectination similar in some taxa</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="2">
                  <bold><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> subfamily identification</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="2">Subfam. candidate</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1"><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Naryciinae">Naryciinae</tp:taxon-name-part></tp:taxon-name> (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dahlica">Dahlica</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="triq">triq</tp:taxon-name-part></tp:taxon-name></italic>.)</td>
                <td rowspan="1" colspan="1">antennae filiform, not bipectinate</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="subfamily" reg="Typhoniinae">Typhoniinae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">antennae filiform, not bipectinate</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="subfamily" reg="Oiketicinae">Oiketicinae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">frons with bushy scales, palpi tiny, antennal branches longer, imagos large</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="subfamily" reg="Epichnopteriginae">Epichnopteriginae</tp:taxon-name-part>
                  </tp:taxon-name>
                </td>
                <td rowspan="1" colspan="1">antennal pectination similar, but frons and palpi with bushy scales</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1"><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part></tp:taxon-name> (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="betulina">betulina</tp:taxon-name-part></tp:taxon-name></italic>; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Psyche">Psyche</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="casta">casta</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Bacotia">Bacotia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="claustrella">claustrella</tp:taxon-name-part></tp:taxon-name></italic>)</td>
                <td rowspan="1" colspan="1">antennal pectination and other imaginal characters are very similar within these genera; there are only small differences in scalation and morphology of the forelegs (tibial epiphysis present or absent). The classification in the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> should be regarded as provisional. However, the visible characteristics, in combination with the overall visible morphology, make classification within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> of the two specimens dealing with more than likely.</td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
      <sec sec-type="Description of Proutia damzeni" id="sec5">
        <title>Description of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic></title>
        <sec sec-type="Systematic Paleontology" id="sec6">
          <title>Systematic Paleontology</title>
          <tp:taxon-treatment>
            <tp:treatment-meta>
              <kwd-group>
                <label>Taxon classification</label>
                <kwd>
                  <named-content content-type="order">Lepidoptera</named-content>
                </kwd>
                <kwd>
                  <named-content content-type="family">Psychidae</named-content>
                </kwd>
              </kwd-group>
            </tp:treatment-meta>
            <tp:nomenclature>
              <tp:taxon-name><object-id content-type="arpha">0B35C18F-D047-546D-B0D9-70EE2D89DE33</object-id>
                        		<tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part>
                        	</tp:taxon-name>
              <tp:taxon-authority>Boisduval, 1829</tp:taxon-authority>
            </tp:nomenclature>
            <tp:treatment-sec sec-type="Taxon">
              <title>Taxon.</title>
              <p>Genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> Tutt, 1899.</p>
            </tp:treatment-sec>
          </tp:taxon-treatment>
          <tp:taxon-treatment>
            <tp:treatment-meta>
              <kwd-group>
                <label>Taxon classification</label>
                <kwd>
                  <named-content content-type="order">Lepidoptera</named-content>
                </kwd>
                <kwd>
                  <named-content content-type="family">Psychidae</named-content>
                </kwd>
              </kwd-group>
            </tp:treatment-meta>
            <tp:nomenclature>
              <label>Species:</label>
              <tp:taxon-name><object-id content-type="arpha">4B6C6DC6-6E6B-5CD3-A18B-5197358967CC</object-id>
                        		<tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part>
                        	
                        		<object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/2EE7DD41-ED78-44CB-B80E-B0A592D28633</object-id>
                        	</tp:taxon-name>
              <tp:taxon-status>sp. nov.</tp:taxon-status>
            </tp:nomenclature>
            <tp:treatment-sec sec-type="Name">
              <title>Name.</title>
              <p>Etymology: The species is named in honour to Jonas Damzen who found and donated this rare specimen among succinite moths.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Typus">
              <title>Typus.</title>
              <p>Holotypus: Specimen SNSB-ZSM-LEP amb013 (former JDC 13768Rovno), Fig. <xref ref-type="fig" rid="F1">1</xref>.</p>
              <fig id="F1">
                <object-id content-type="doi">10.3897/zitteliana.100.197055.figure1</object-id>
                <object-id content-type="arpha">BC185B67-408F-5219-8D38-56C12C0DEB64</object-id>
                <label>Figure 1.</label>
                <caption>
                  <p>Specimen JDC 13768 from Eocene Ukrainian (Rovno) amber, holotype of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic>. <bold>A</bold>. Right and frontal aspect; <bold>B</bold>. Left and dorsal aspect. Length 5.4 mm.</p>
                </caption>
                <graphic xlink:href="zitteliana-100-117-g001.jpg" id="oo_1687366.jpg">
                  <uri content-type="original_file">https://binary.pensoft.net/fig/1687366</uri>
                </graphic>
              </fig>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Locus typicus">
              <title>Locus typicus.</title>
              <p>Ukrainian amber, Rovno.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Stratum typicum">
              <title>Stratum typicum.</title>
              <p>Eocene sediments with amber, Ukraine.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Repository">
              <title>Repository.</title>
              <p>Bavarian State Collection of Zoology SNSB-ZSM-LEP amb13.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Description of species">
              <title>Description of species.</title>
              <p>Imago 5.4 mm. Head. Rough hairy, frons and vertex covered with broad (classes 4–5) and lanceolate scales. Largest eye-distance 1.25-fold of eye height. Ocelli (accessory “eyes”) absent. Antennae bipectinate, of nearly 2/3 times of costa length, with 29 segments. Pecten between segments 1–15 of 1.5 times of segment length. Segments basally densely covered with short setae. Pecten also distally covered with long setae. Thorax. Forewings stretched, slender, apex pointed, termen oblique. Forewing length 4.4 mm, forewing width 1.3 mm, semi-hyaline. Venation partially visible. Forewings probably with 9 veins. Intercalary cell in the divided media not clearly visible but seemingly present. Forewings covered with broad scales (classes 4–5, after <xref ref-type="bibr" rid="B8">Sauter 1956</xref>). An accessory cell is not visible. Legs partially visible. Forelegs with epiphysis. Midlegs and Hindlegs clearly visible. Midlegs with apical tibial spurs of 260 µm length. Hindlegs with apical and medial tibial spurs of 290 and 460 µm length. The inner spur is slightly shorter in each case. Tarsus with 5 segments. Praetarsus with 2 curved crawls. Legs densely covered with broad scales, and some smaller scales. Abdomen not visible.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Diagnosis">
              <title>Diagnosis.</title>
              <p>Medium sized male imagos, head rough hairy, vertex with broad and lanceolate scales, eye distance larger than eye height. Ocelli absent. Antennae bipectinate, almost two thirds of costa length, with twenty-nine segments, pecten in proximal part of antennae of one-and-a-half times segment length, segments basally with short setae, pecten also distally with long setae. Forewings stretched, slender, apex pointed, termen oblique, covered with broad scales, semihyaline, probably with nine veins, putative intercalary cell in divided media. Forelegs with epiphysis, midlegs with apical tibial spurs, hindlegs with apical and medial tibial spurs, all inner spurs slightly shorter. Tarsus with five segments. Praetarsus with two curved crawls. Legs densely covered with broad and some smaller scales.</p>
            </tp:treatment-sec>
            <tp:treatment-sec sec-type="Differential diagnosis and discussion">
              <title>Differential diagnosis and discussion.</title>
              <p>The new species is characterised by its relatively short antennae, which are just over half the length of the costa. The only relevant extant species in this geographical area, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="betulina">betulina</tp:taxon-name-part></tp:taxon-name></italic>, has significantly longer antennae (2/3 of the costal length). However, the number of antennal segments, including the scapus and pedicellus, is 29, which is significantly higher than the number of antennal segments in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="betulina">betulina</tp:taxon-name-part></tp:taxon-name></italic>, which has an average of only 20–25 segments. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. differs markedly from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="altera">altera</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. (see below) in its narrower cover scales and the smaller number of antenna segments.</p>
            </tp:treatment-sec>
          </tp:taxon-treatment>
        </sec>
      </sec>
      <sec sec-type="Systematic affiliation" id="sec7">
        <title>Systematic affiliation</title>
        <p>For systematic affiliation, extant genera which are morphologically closest to the new records from amber were compared with respect to characters of diagnostic relevance (Table <xref ref-type="table" rid="T2">2</xref>).</p>
        <table-wrap id="T2" position="float" orientation="portrait">
          <label>Table 2.</label>
          <caption>
            <p>Comparison of characters of the new amber records to close extant genera.</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <th rowspan="1" colspan="1">
                  <bold>Character</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Genus <italic>Psyche</italic></bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>Genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic></bold>
                </th>
                <th rowspan="1" colspan="1"><bold><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia"/><tp:taxon-name-part taxon-name-part-type="species">damzeni</tp:taxon-name-part></tp:taxon-name> sp. nov</bold>.</th>
                <th rowspan="1" colspan="1"><bold><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia"/><tp:taxon-name-part taxon-name-part-type="species">altera</tp:taxon-name-part></tp:taxon-name> sp. nov</bold>.</th>
                <th rowspan="1" colspan="1">
                  <bold>Genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Bacotia">Bacotia</tp:taxon-name-part></tp:taxon-name></italic></bold>
                </th>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Cloaking scales (classes after <xref ref-type="bibr" rid="B8">Sauter 1956</xref>)</td>
                <td rowspan="1" colspan="1">classes 5–6</td>
                <td rowspan="1" colspan="1">classes 4–5</td>
                <td rowspan="1" colspan="1">classes 4–5</td>
                <td rowspan="1" colspan="1">classes 5–6</td>
                <td rowspan="1" colspan="1">classes 4–5</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Veins on forewings from discal cell</td>
                <td rowspan="1" colspan="1">9</td>
                <td rowspan="1" colspan="1">9</td>
                <td rowspan="1" colspan="1">probably 9</td>
                <td rowspan="1" colspan="1">invisible</td>
                <td rowspan="1" colspan="1">9</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Veins on hindwings from discal cell</td>
                <td rowspan="1" colspan="1">6</td>
                <td rowspan="1" colspan="1">6</td>
                <td rowspan="1" colspan="1">invisible</td>
                <td rowspan="1" colspan="1">invisible</td>
                <td rowspan="1" colspan="1">6</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Intercalary cell</td>
                <td rowspan="1" colspan="1">absent</td>
                <td rowspan="1" colspan="1">present</td>
                <td rowspan="1" colspan="1">?<bold><sup>1</sup></bold></td>
                <td rowspan="1" colspan="1">?<sup>1</sup></td>
                <td rowspan="1" colspan="1">absent</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Accessory cell</td>
                <td rowspan="1" colspan="1">absent</td>
                <td rowspan="1" colspan="1">absent</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">present</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Tibial epiphysis on foreleg</td>
                <td rowspan="1" colspan="1">present</td>
                <td rowspan="1" colspan="1">present</td>
                <td rowspan="1" colspan="1">invisible</td>
                <td rowspan="1" colspan="1">present</td>
                <td rowspan="1" colspan="1">present</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Antennae length</td>
                <td rowspan="1" colspan="1">half of costa</td>
                <td rowspan="1" colspan="1">2/3 costa length</td>
                <td rowspan="1" colspan="1">more than half costa length</td>
                <td rowspan="1" colspan="1">more than half costa length</td>
                <td rowspan="1" colspan="1">less than half costa length</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Antennal segments</td>
                <td rowspan="1" colspan="1">18–27</td>
                <td rowspan="1" colspan="1">21–27</td>
                <td rowspan="1" colspan="1">29</td>
                <td rowspan="1" colspan="1">app. 32</td>
                <td rowspan="1" colspan="1">25–29</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Tarsus<bold><sup>2</sup></bold></td>
                <td rowspan="1" colspan="1">5 segments</td>
                <td rowspan="1" colspan="1">5 segments</td>
                <td rowspan="1" colspan="1">5 segments</td>
                <td rowspan="1" colspan="1">5 segments</td>
                <td rowspan="1" colspan="1">5 segments</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Larval case</td>
                <td rowspan="1" colspan="1">grass stems</td>
                <td rowspan="1" colspan="1">grass stems</td>
                <td rowspan="1" colspan="1">compare with case inclusions (Fig. <xref ref-type="fig" rid="F2">2</xref>)</td>
                <td rowspan="1" colspan="1">compare with case inclusions (Fig. <xref ref-type="fig" rid="F2">2</xref>)</td>
                <td rowspan="1" colspan="1">bark, lichens</td>
              </tr>
            </tbody>
          </table>
          <table-wrap-foot>
            <fn>
              <p><bold><sup>1</sup></bold> Unfortunately, the intercalary cell in the divided media (a typical feature of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic>) is not clearly visible. However, it is possible to visualize it.</p>
              <p><bold><sup>2</sup></bold> Legs: Characteristic of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part></tp:taxon-name>: Praetarsus with 2 curved claws. Mesotibia terminal with a pair of spurs, metatibia 2 pairs, one terminal and one on the distal third. The inner spur is slightly shorter in each case.</p>
            </fn>
          </table-wrap-foot>
        </table-wrap>
        <p>It is challenging to establish a connection between the fossil preserved in amber and the modern genera of the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name>. It is evident that this specimen belongs to the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part></tp:taxon-name>, a group that is currently distributed worldwide with the exception of Antarctica. The majority of these can be characterized as follows: mostly small moths, dark blackish-brown or brownish-grey, sometimes with distinct reticular pattern. The wings are stretched, and the head and thorax are roughly hairy. Ocelli (accessory eyes) are absent, the labial palp is reduced to one segment, and the antennae are bipectinate with long pecten. On the forewings 9 veins emanate from discal cell, the hindwings have 5 veins from the discal cell. The intercalary cell and accessory cell are either absent or present. Scales are mostly broad (classes 4–5 after <xref ref-type="bibr" rid="B8">Sauter 1956</xref>). On foreleg a tibial epiphysis is present. Midlegs bear apical tibial spurs, hindlegs are with apical and medial tibial spurs. The larval cases are variable, tubular or conical and covered longitudinally with grass stems, bark, lichens or fine grains of sand and soil.</p>
        <p>The above table presents a summary of the most significant characteristics of the genera within the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part></tp:taxon-name> that are relevant in Central Europe. However, it should be noted that some of these characteristics are difficult or impossible to observe in the fossil specimen enclosed in amber.</p>
        <p><bold>Diagnosis</bold>: The classification within the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> remains provisional and speculative, pending further findings. The presence of taxa from the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part></tp:taxon-name> in Baltic amber is a noteworthy occurrence. <xref ref-type="bibr" rid="B11">Sobczyk and Kobbert (2009)</xref> provide sufficient evidence to support this claim, using preserved larval cases as crucial evidence. These are consistent with larval cases of the recent genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> Tutt, 1899, and the classification of the taxon described here as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> is supported by this evidence. It is highly probable that some of the cases found to date can be attributed to the adult specimen described herein, but this cannot be proven.</p>
        <p><bold>Taphonomy</bold>: The inclusion is preserved in a polished piece of Ukrainian amber 27 mm × 12 mm × 4 mm in size (cabochon). Syninclusions: only shed scales from the moth.</p>
      </sec>
      <sec sec-type="Description of Proutia altera" id="sec8">
        <title>Description of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="altera">altera</tp:taxon-name-part></tp:taxon-name></italic></title>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="order">Lepidoptera</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family">Psychidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>Species:</label>
            <tp:taxon-name><object-id content-type="arpha">AEB713E3-BDA3-5626-B609-49B4C9A59893</object-id>
                    		<tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="altera">altera</tp:taxon-name-part>
                    	
                    		<object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/F0DBAA83-359A-4723-8332-4496FB9B2EDD</object-id>
                    	</tp:taxon-name>
            <tp:taxon-status>sp. nov.</tp:taxon-status>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="Name">
            <title>Name.</title>
            <p>Etymology: The species name latin “the other” refers to the first find of the other species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic>.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Typus">
            <title>Typus.</title>
            <p>Holotypus: Specimen SNSB-ZSM-LEP amb014 (former JDC 13954Rovno) Fig. <xref ref-type="fig" rid="F3">3</xref>.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Locus typicus">
            <title>Locus typicus.</title>
            <p>Ukrainian amber, Rovno.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Stratum typicum">
            <title>Stratum typicum.</title>
            <p>Eocene sediments with amber, Ukraine.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Repository">
            <title>Repository.</title>
            <p>Bavarian State Collection of Zoology SNSB-ZSM-LEP amb14.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Description of species">
            <title>Description of species.</title>
            <p>Imago 6.5 mm. Head. Rough hairy, frons and vertex covered with broad (classes 4–5) and lanceolate scales. Largest eye-distance equal to 0.83 of eye height, which is in contrast to that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic> with 1.25. Labial palps thick, short, upwards pointing, and densely covered with scales. No other mouthparts present. Ocelli not observable. Antennae bipectinate, of about 2/3 times of costa length, with 32 segments. Pecten between segments 1–15 of 1.5 times of segment length. Segments without setae. Pecten distally covered with long setae. Tip of antennae noticeable and pronounced: single segments less separated and last five segments with teeth of pecten much shorter in comparison to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic>. Thorax. Forewings stretched, apex pointed, termen oblique. Forewing length some 5 mm, broad, semi-hyaline. Venation only partially visible. Forewings covered with broad scales (classes 4–5, after <xref ref-type="bibr" rid="B8">Sauter 1956</xref>). Legs. All legs covered with scales; femurs covered with very long slender scales strictly directed distally. Forelegs with epiphysis. One midleg with apical tibial spurs visible. Hindlegs with prominent apical and medial tibial spurs. Tarsus with 5 segments. Praetarsus (visible only at right mid leg) with 2 curved crawls. In contrast to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic>, legs of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="altera">altera</tp:taxon-name-part></tp:taxon-name></italic> are densely covered with small stretched scales which are mostly lanceolate. Abdomen. Only distal end visible, this densely covered with scales, clasping organ with valves visible.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Diagnosis">
            <title>Diagnosis.</title>
            <p>Medium sized male imagos, head rough hairy, vertex with broad and lanceolate scales, eye distance shorter than eye height. Labial palps thick, short, upwards pointing, and densely covered with scales. No other mouthparts present. Antennae bipectinate, about two thirds of costa length, about thirty-two segments, pecten in proximal part of antennae of one-and-a-half times of segment length, segments without setae, pecten distally with long setae. Tip of antennae noticeable and pronounced. Forewings stretched, apex pointed, termen oblique, covered with broad scales, broad, semihyaline. Forelegs with epiphysis. Legs with small stretched (almost lanceolate) scales, femurs with very long slender scales. Midlegs with apical tibial spurs, hindlegs with prominent apical and medial tibial spurs. Tarsus with five segments. Praetarsus with two curved crawls. Legs densely covered with small stretched scales, mostly lanceolate.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Differential diagnosis and discussion">
            <title>Differential diagnosis and discussion.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="altera">altera</tp:taxon-name-part></tp:taxon-name></italic> differs from the most closely related species in this geographical region, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="betulina">betulina</tp:taxon-name-part></tp:taxon-name></italic>, in that it has a high number of antenna segments (32), whereas <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="betulina">betulina</tp:taxon-name-part></tp:taxon-name></italic> has an average of 20–25 segments. Tip of antennae with single segments less separated and last five segments with teeth of pecten much shorter in comparison to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic>. In contrast to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic>, legs of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="altera">altera</tp:taxon-name-part></tp:taxon-name></italic> are densely covered with small stretched scales which are mostly lanceolate. Furthermore, the eye distance is significantly shorter than in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="damzeni">damzeni</tp:taxon-name-part></tp:taxon-name></italic> (0.83 versus 1.25). On average, the scales of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="altera">altera</tp:taxon-name-part></tp:taxon-name></italic> are wider than those of the previous species and are also slightly above the average for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="betulina">betulina</tp:taxon-name-part></tp:taxon-name></italic>.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Taphonomy">
            <title>Taphonomy.</title>
            <p>The inclusion is preserved in a polished piece of Ukrainian amber 23 mm × 13 mm × 4 mm in size, broken off at one end. Syninclusions: remains of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Diptera">Diptera</tp:taxon-name-part></tp:taxon-name>, cf. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Tipulidae">Tipulidae</tp:taxon-name-part></tp:taxon-name>; some detritus.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
      <sec sec-type="Specimen of Psychidae from Baltic amberCG # 7168, Fig. 4" id="sec9">
        <title>Specimen of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> from Baltic amberCG # 7168, Fig. 4</title>
        <p><bold>Description</bold>: Imago some 6.5 mm. Head. Hind part covered with small scales, vertex and frons covered with milky emulsion. Largest eye-distance about equal to eye height. Labial palps thin, short, upwards pointing, and densely covered with small scales. No other mouthparts present. Ocelli not observable. Antennae bipectinate, of nearly 2/3 times of costa length, with 29 segments. Pecten between segments 1–15 of 1.5 times of segment length. Segments basally densely covered with short setae. Pecten also distally covered with long setae. Pecten elements forked. Thorax. Forewings stretched, apex pointed, termen oblique. Forewing length some 5 mm, forewing width some 2.2 mm, semi-hyaline. Venation partially visible. Forewings covered with broad scales (classes 4–5, after <xref ref-type="bibr" rid="B8">Sauter 1956</xref>) and with cilia. Forelegs with epiphysis. Tarsus with 5 segments. Praetarsus with 2 curved crawls. Midleg with apical tibial spurs; hindlegs with apical and medial tibial spurs. Abdomen. Only distal end visible, this densely covered with scales, clasping organ with valves visible.</p>
        <p>The specimen can be recognized as a member of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name>, but is too incompletely preserved to allow for a subfamily affiliation.</p>
        <p><bold>Taphonomy</bold>: The inclusion is preserved in a polished piece of Baltic amber 26 mm × 11 mm × 8 mm in size. Syninclusions: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Chironomidae">Chironomidae</tp:taxon-name-part></tp:taxon-name>, isolated Acarus (mite), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Quercus">Quercus</tp:taxon-name-part></tp:taxon-name></italic>-type trichomes, some detritus.</p>
      </sec>
      <sec sec-type="Cases and caterpillars of Proutia in Baltic amber" id="sec10">
        <title>Cases and caterpillars of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> in Baltic amber</title>
        <p>From Baltic amber, three specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic>-type bags with caterpillars have been identified (Fig. <xref ref-type="fig" rid="F2">2</xref>).</p>
        <fig id="F2">
          <object-id content-type="doi">10.3897/zitteliana.100.197055.figure2</object-id>
          <object-id content-type="arpha">0B0013E1-64B7-5F72-B46C-3ECF16970424</object-id>
          <label>Figure 2.</label>
          <caption>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic>-type bags with caterpillars from Baltic amber: <bold>A</bold>. Specimen # TF 4190 (length 8 mm); <bold>B</bold>. # TF 4908 (length 7 mm) dorsal aspect; <bold>C</bold>. Ventral aspect; <bold>D</bold>. # TF 8652 (length 15 mm).</p>
          </caption>
          <graphic xlink:href="zitteliana-100-117-g002.jpg" id="oo_1687367.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1687367</uri>
          </graphic>
        </fig>
        <fig id="F3">
          <object-id content-type="doi">10.3897/zitteliana.100.197055.figure3</object-id>
          <object-id content-type="arpha">477DC260-D7DB-51EC-A3BF-341A05348BE0</object-id>
          <label>Figure 3.</label>
          <caption>
            <p>Specimen JDC 13954 from Eocene Ukrainian (Rovno) amber, holotype of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="altera">altera</tp:taxon-name-part></tp:taxon-name></italic>. Right and frontal aspect, detail with head and antennae. Complete length 6.5 mm.</p>
          </caption>
          <graphic xlink:href="zitteliana-100-117-g003.jpg" id="oo_1687368.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1687368</uri>
          </graphic>
        </fig>
        <fig id="F4">
          <object-id content-type="doi">10.3897/zitteliana.100.197055.figure4</object-id>
          <object-id content-type="arpha">DE4985D9-CB15-5AAF-9CA0-F6B06BB23632</object-id>
          <label>Figure 4.</label>
          <caption>
            <p>Specimen CG # 7168 from Eocene Baltic amber. <bold>A</bold>. Ventral-right aspect; <bold>B</bold>. Dorsal-left aspect. Visible length each 2.5 mm.</p>
          </caption>
          <graphic xlink:href="zitteliana-100-117-g004.jpg" id="oo_1687369.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1687369</uri>
          </graphic>
        </fig>
      </sec>
    </sec>
    <sec sec-type="Discussion" id="sec11">
      <title>Discussion</title>
      <sec sec-type="Identification" id="sec12">
        <title>Identification</title>
        <p>The noticeable character of bipectinate antennae drew attention to these moth inclusions in Ukrainian and Baltic ambers. Screening Lepidopteran families for which bipectinate antennae have been reported confirmed the initial assumption that the specimens are male bagworm moths. With the already described bags affiliated with the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic>, the male adult counterparts to the juveniles have now been found from succinite.</p>
      </sec>
      <sec sec-type="Taphonomy" id="sec13">
        <title>Taphonomy</title>
        <p>Taphonomy of caterpillars and their bags is very different to that of the male imagos. Bags of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Psychidae">Psychidae</tp:taxon-name-part></tp:taxon-name> are often fixed to trees and are durable. The caterpillars are polyphagous and occur on diverse plants. Hence, caterpillars and bags are predisposed for inclusion in resin, and, consequently, frequent in amber. In contrast, males are short living and mobile, with lower likelihood of capture by resin. Furthermore, individuals overwinter as late instars and emerge as adults in late spring (<xref ref-type="bibr" rid="B7">Rhainds et al. 2009</xref>), which may not have been the maximum season in resin production, further diminishing chances of capturing males. Extant males fly during daytime as well as by night, depending on species. Bipectinate antennae suggest the use of pheromones by females for attraction of males, which are equipped with highly efficient chemoreceptors. Seemingly, in Eocene <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> male imagos were also the searching sex, like the extant ones. Capture in resin and preservation in amber may be due to sexual attraction by females positioned with their bags at the bark of resin producing trees, but no syninclusion of female and male is known yet. Only a single case of fossil tineid moths in copula has been described up to now (<xref ref-type="bibr" rid="B4">Fischer and Hörnig 2019</xref>). Also, the female flightless counterparts to the males have not yet been found or are still unrecognized.</p>
      </sec>
      <sec sec-type="Actualism" id="sec14">
        <title>Actualism</title>
        <p>Morphology and, in conclusion, life habits of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> are seemingly conservative, as evidenced from the new findings in comparison to extant species of the genus (Figs <xref ref-type="fig" rid="F5">5</xref>, <xref ref-type="fig" rid="F6">6</xref>), however, fossil females are still unknown. Bags of the psychid genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dahlica">Dahlica</tp:taxon-name-part></tp:taxon-name></italic> from Baltic amber had been found before, and are indistinguishable from those of extant <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dahlica">Dahlica</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="triquetrella">triquetrella</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B11">Sobczyk and Kobbert 2009</xref>). Consequently, and in contrast to common practice with amber inclusions, they were given the name of the extant species. In this work, the finds are affiliated with the extant genus, but as fossil species.</p>
        <fig id="F5">
          <object-id content-type="doi">10.3897/zitteliana.100.197055.figure5</object-id>
          <object-id content-type="arpha">1957B340-BBD5-5B44-8AAB-5461FD269429</object-id>
          <label>Figure 5.</label>
          <caption>
            <p>Extant <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="betulina">betulina</tp:taxon-name-part></tp:taxon-name></italic>. <bold>A</bold>. Male imago (photo Rudolf Bryner, Lepiforum); <bold>B</bold>. Caterpillar with case (photo Julia Bastian, Lepiforum).</p>
          </caption>
          <graphic xlink:href="zitteliana-100-117-g005.jpg" id="oo_1687370.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1687370</uri>
          </graphic>
        </fig>
        <fig id="F6">
          <object-id content-type="doi">10.3897/zitteliana.100.197055.figure6</object-id>
          <object-id content-type="arpha">26DEA26E-53BF-5F82-935C-0DDD6FCE967C</object-id>
          <label>Figure 6.</label>
          <caption>
            <p>Extant <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="comitella">comitella</tp:taxon-name-part></tp:taxon-name></italic>. <bold>A</bold>. Male imago; <bold>B</bold>. Caterpillar with case (photos Michel Billard).</p>
          </caption>
          <graphic xlink:href="zitteliana-100-117-g006.jpg" id="oo_1687371.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1687371</uri>
          </graphic>
        </fig>
      </sec>
    </sec>
    <sec sec-type="Conclusion" id="sec15">
      <title>Conclusion</title>
      <p>After manifold reports on psychid bags in amber (succinite) literature, male psychid counterparts are here described for the first time based on three imago specimens. The taxa show a conservative morphology and could be recognized as members of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Psychinae">Psychinae</tp:taxon-name-part></tp:taxon-name>), which was also known from larval bags. Such males are strongly underrepresented in collections as compared to the bags of the juveniles, which is in agreement with a quite short lifespan. This further indicates that the extreme sexual dimorphism and mode of living might already have been fully evolved in Eocene some 40 million years ago. Future focus should be on recognition of other male psychid imagos among the abundant “tineids” from Baltic and Ukrainian amber, but also finding their female counterparts.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgement</title>
      <p>Julia Bastian and Rudolf Bryner, both active for Lepiforum, are gratefully acknowledged for allowing the use of and providing the photos of extant male and juvenile stages of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="betulina">betulina</tp:taxon-name-part></tp:taxon-name></italic> Zeller, 1839. Michel Billard is acknowledged for the photos of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Proutia">Proutia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="comitella">comitella</tp:taxon-name-part></tp:taxon-name></italic>. We are grateful to Luisa Böttner, Spring/Texas (USA), who kindly provided English and linguistic corrections. Ethan Beaver (Canberra, Australia) is acknowledged for reviewing the work and helping to improve it.</p>
    </ack>
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</article>
