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  <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.e189638</article-id>
      <article-id pub-id-type="publisher-id">189638</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Animalia</subject>
          <subject>Arachnida</subject>
          <subject>Araneae</subject>
          <subject>Arthropoda</subject>
          <subject>Invertebrata</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biodiversity &amp; Conservation</subject>
          <subject>Species Inventories</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Spiders in a Growing City: New Country and Regional Records of Spiders (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Araneae">Araneae</tp:taxon-name-part></tp:taxon-name>) from Tbilisi, Georgia</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Seropian</surname>
            <given-names>Armen</given-names>
          </name>
          <email xlink:type="simple">armen.seropiani@iliauni.edu.ge</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-3777-9954</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Bulbulashvili</surname>
            <given-names>Natalia</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-6802-1209</uri>
          <xref ref-type="aff" rid="A3">3</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Egidunashvili</surname>
            <given-names>Vano</given-names>
          </name>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Institute of Zoology, Ilia State University, Tbilisi, Georgia</addr-line>
        <institution>Institute of Zoology, Ilia State University</institution>
        <addr-line content-type="city">Tbilisi</addr-line>
        <country>Georgia</country>
        <uri content-type="ror">https://ror.org/051qn8h41</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Institute of Ecology, Ilia State University, Tbilisi, Georgia</addr-line>
        <institution>Institute of Ecology, Ilia State University</institution>
        <addr-line content-type="city">Tbilisi</addr-line>
        <country>Georgia</country>
        <uri content-type="ror">https://ror.org/051qn8h41</uri>
      </aff>
      <aff id="A3">
        <label>3</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>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Armen Seropian (<email xlink:type="simple">armen.seropiani@iliauni.edu.ge)</email></p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Nils Hein</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>07</day>
        <month>05</month>
        <year>2026</year>
      </pub-date>
      <volume>5</volume>
      <fpage>31</fpage>
      <lpage>39</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/FAFDC577-BDA2-567B-8A0C-2C3476115402">FAFDC577-BDA2-567B-8A0C-2C3476115402</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/93E7A55A-754C-4FB6-A725-474B70B40545">93E7A55A-754C-4FB6-A725-474B70B40545</uri>
      <history>
        <date date-type="received">
          <day>24</day>
          <month>02</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>27</day>
          <month>03</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Armen Seropian, Natalia Bulbulashvili, Vano Egidunashvili</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/93E7A55A-754C-4FB6-A725-474B70B40545</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>The spider fauna of Tbilisi was systematically documented in the early 2000s, when 252 species were recorded from the city and its surroundings. Despite a rapid urban expansion over the past two decades, subsequent studies have increased this number to 371 species. In the present study, we provide the first Georgian records of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">Tolokonniella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stigmatisata">stigmatisata</tp:taxon-name-part></tp:taxon-name></italic> (Menge, 1869) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Lathyidae">Lathyidae</tp:taxon-name-part></tp:taxon-name>) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pilosus">pilosus</tp:taxon-name-part></tp:taxon-name></italic> Simon, 1913 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Linyphiidae">Linyphiidae</tp:taxon-name-part></tp:taxon-name>), and record <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">Drassodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serratichelis">serratichelis</tp:taxon-name-part></tp:taxon-name></italic> (Roewer, 1928) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Gnaphosidae">Gnaphosidae</tp:taxon-name-part></tp:taxon-name>) in the Caucasus region for the first time, all based on specimens collected in Tbilisi.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>
          <italic>
            <tp:taxon-name>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">Drassodes</tp:taxon-name-part>
            </tp:taxon-name>
          </italic>
        </kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="family" reg="Gnaphosidae">Gnaphosidae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="family" reg="Lathyidae">Lathyidae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="family" reg="Linyphiidae">Linyphiidae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>
          <italic>
            <tp:taxon-name>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part>
            </tp:taxon-name>
          </italic>
        </kwd>
        <kwd>South Caucasus</kwd>
        <kwd>
          <italic>
            <tp:taxon-name>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">Tolokonniella</tp:taxon-name-part>
            </tp:taxon-name>
          </italic>
        </kwd>
      </kwd-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="sec1">
        <title>Citation</title>
        <p>Seropian A, Bulbulashvili N, Egidunashvili V (2026) Spiders in a Growing City: New Country and Regional Records of Spiders (Araneae) from Tbilisi, Georgia. Caucasiana 5: 31–39. <ext-link xlink:href="10.3897/caucasiana.5.e189638" ext-link-type="doi">https://doi.org/10.3897/caucasiana.5.e189638</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec2">
      <title>Introduction</title>
      <p>Urbanisation is widely recognised as one of the major drivers of biodiversity change, particularly in arthropod communities, where habitat structure, microclimate, and landscape connectivity play a crucial role (<xref ref-type="bibr" rid="B10">McKinney 2008</xref>). Spiders (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Araneae">Araneae</tp:taxon-name-part></tp:taxon-name>), due to their high diversity, ecological specialisation, and sensitivity to habitat transformation, are commonly used as indicators of environmental change in urban and peri-urban landscapes (<xref ref-type="bibr" rid="B6">Marc et al. 1999</xref>; <xref ref-type="bibr" rid="B21">Shochat et al. 2004</xref>).</p>
      <p>The spider fauna of Tbilisi was comprehensively documented in the early 2000s. Based on previously published data and targeted sampling across urban, suburban, and semi-natural habitats, <xref ref-type="bibr" rid="B15">Pkhakadze (2006)</xref> recorded 252 spider species in Tbilisi and its surroundings, providing a robust historical baseline for subsequent comparisons. Since then, Tbilisi has undergone rapid and spatially extensive urban development. Remote-sensing-based studies and urban sustainability assessments indicated a substantial increase in built-up areas, with the proportion of urbanised land rising by approximately 15% over the last two decades (<xref ref-type="bibr" rid="B22">Tsitsagi et al. 2022</xref>). This expansion has been driven not only by densification of the urban core, but also by lateral growth into formerly semi-natural and agricultural areas, accompanied by fragmentation and loss of open habitats. Such landscape transformations are known to have the potential to disproportionately affect ground-dwelling and habitat-specialist spider taxa, while favouring a limited set of synanthropic and ecologically generalist species (<xref ref-type="bibr" rid="B21">Shochat et al. 2004</xref>; <xref ref-type="bibr" rid="B5">Magura et al. 2010</xref>). At the same time, urban expansion and habitat transformation are expected to influence the composition and functional structure of spider assemblages, potentially masking local declines of habitat specialists through the accumulation of synanthropic or ecologically flexible species (<xref ref-type="bibr" rid="B10">McKinney 2008</xref>; <xref ref-type="bibr" rid="B5">Magura et al. 2010</xref>; Piano et al. 2017). Consequently, changes in species richness alone may be insufficient to capture the ecological consequences of ongoing urbanisation.</p>
      <p>Surveys conducted in Tbilisi and its suburban areas do not indicate a straightforward decline in recorded species richness. On the contrary, despite the rapidly increasing rate of urbanisation, continued spider research in Georgia has resulted in the discovery of additional spider species within the urban and peri-urban landscape of the capital. Specifically, 119 species have been added to the fauna of Tbilisi and its surroundings, of which 63 represented first country records, 22 species were firstly recorded in the Caucasian region, and three species were described as new to science (<xref ref-type="bibr" rid="B9">Marusik et al. 2019</xref>; <xref ref-type="bibr" rid="B16">Seropian et al. 2023a</xref>, <xref ref-type="bibr" rid="B17">b</xref>; <xref ref-type="bibr" rid="B18">Seropian et al. 2024a</xref>, <xref ref-type="bibr" rid="B19">b</xref>; <xref ref-type="bibr" rid="B20">Seropian et al. 2025</xref>; <xref ref-type="bibr" rid="B24">Zamani et al. 2025</xref>). As a result, 371 species are currently recorded from Tbilisi. This increase most likely reflects a combination of intensified sampling effort, improved taxonomic resolution, and the inclusion of previously underexplored habitats, rather than a direct positive effect of urbanisation on spider diversity.</p>
      <p>In the present study, we update the current knowledge of the spider fauna of Tbilisi by reporting the first Georgian records of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">Drassodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serratichelis">serratichelis</tp:taxon-name-part></tp:taxon-name></italic> (Roewer, 1928) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Gnaphosidae">Gnaphosidae</tp:taxon-name-part></tp:taxon-name>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">Tolokonniella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stigmatisata">stigmatisata</tp:taxon-name-part></tp:taxon-name></italic> (Menge, 1869) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Lathyidae">Lathyidae</tp:taxon-name-part></tp:taxon-name>), and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pilosus">pilosus</tp:taxon-name-part></tp:taxon-name></italic> Simon, 1913 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Linyphiidae">Linyphiidae</tp:taxon-name-part></tp:taxon-name>), based on material collected in the capital of Georgia. Notably, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serratichelis">serratichelis</tp:taxon-name-part></tp:taxon-name></italic> is recorded for the first time from the Caucasus, while the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stigmatisata">stigmatisata</tp:taxon-name-part></tp:taxon-name></italic> are newly recorded for Georgia.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec3">
      <title>Materials and methods</title>
      <p>All spiders were collected individually, preserved in 96% ethanol, and stored in the refrigerators of the scientific collections of Ilia State University, Institute of Zoology (Tbilisi, Georgia; hereafter – <abbrev content-type="institution" xlink:title="Ilia State University, Institute of Zoology">ISUIZ</abbrev>). The specimens were examined with a Zeiss Stemi 508 Stereo Microscope with 8:1 Zoom and a Zeiss Apo 1.5× FWD 53 mm front lens attached. A.S. prepared drawings based on microscope photographs taken with a Canon EOS 70D camera mounted on a Zeiss Stemi 508 stereomicroscope with an 8:1 zoom, using a custom-made adapter. The drawings were carried out using a Wacom CTH-690 Intuos Medium Pen and Touch Tablet with the programs Krita (version 2.9.7) and Adobe Photoshop CS6 (version 13.0). Drawings show the epigyne and endogyne; perspective and scale bars are given in the plates and their captions. Epigynes were cleared and cleaned prior to illustrations using a 30% potassium hydroxide (KOH) solution. All measurements are in millimetres. The elevations and GPS coordinates (given in WGS84) were obtained using a Garmin GPS <named-content content-type="dwc:institutional_code" xlink:title="Museo archeologico e paleobotanico di Perfugas" xlink:href="https://scientific-collections.gbif.org/institution/eeaeb325-b878-4627-8faf-017135467da6">MAP</named-content> 64s.</p>
    </sec>
    <sec sec-type="Results" id="sec4">
      <title>Results</title>
      <sec sec-type="List of spider species recorded" id="sec5">
        <title>List of spider species recorded</title>
        <p>
          <bold>Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Gnaphosidae">Gnaphosidae</tp:taxon-name-part></tp:taxon-name> Banks, 1892</bold>
        </p>
        <p>
          <bold>Genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">Drassodes</tp:taxon-name-part></tp:taxon-name></italic> Westring, 1851</bold>
        </p>
        <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">Araneae</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family">Gnaphosidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <tp:taxon-name><object-id content-type="arpha">A96D0AD2-B814-5BCA-830A-2D7E6E68116C</object-id>
                    		<tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">Drassodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serratichelis">serratichelis</tp:taxon-name-part>
                    	</tp:taxon-name>
            <tp:taxon-authority>(Roewer, 1928)</tp:taxon-authority>
            <xref ref-type="fig" rid="F1">Figs 1–2</xref>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">Drassodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serratichelis">serratichelis</tp:taxon-name-part></tp:taxon-name>
                <comment>: <xref ref-type="bibr" rid="B2">Chatzaki et al. 2002</xref>: 618, figs 40–41 (♂)</comment>
              </tp:nomenclature-citation>
              <tp:nomenclature-citation>
                <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">Drassodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serratichelis">serratichelis</tp:taxon-name-part></tp:taxon-name>
                <comment>: <xref ref-type="bibr" rid="B4">Kovblyuk 2008</xref>: 20, figs 6.1–2, 7 (♂)</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="Material examined">
            <title>Material examined.</title>
            <p>GEORGIA • ♂ (<abbrev content-type="institution" xlink:title="Ilia State University, Institute of Zoology">ISUIZ</abbrev>; CaBOL-ID 1039479); Dighomi Massive, Tbilisi; <named-content content-type="dwc:verbatimCoordinates">41.7579, 44.7790</named-content>; 440 m a.s.l.; in apartment; leg. A. Seropian; 16 Jun. 2023.</p>
            <fig id="F1">
              <object-id content-type="doi">10.3897/caucasiana.5.e189638.figures1-2</object-id>
              <object-id content-type="arpha">3C833A54-A656-5982-9D8C-A1781F0088AA</object-id>
              <label>Figures 1–2.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">Drassodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serratichelis">serratichelis</tp:taxon-name-part></tp:taxon-name></italic>, male, left palp (<bold>1</bold> – ventral view; <bold>2</bold> – retrolateral view). Abbreviations: VTA – ventral tibial apophysis. Scale bar: 0.1 mm.</p>
              </caption>
              <graphic xlink:href="caucasiana-05-031-g001.jpg" id="oo_1626836.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1626836</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Comments">
            <title>Comments.</title>
            <p>This species is recorded from the Balearic Islands to Kalmykia; introduced to USA (<xref ref-type="bibr" rid="B12">Nentwig et al. 2026</xref>; <xref ref-type="bibr" rid="B25">WSC 2026</xref>). According to <xref ref-type="bibr" rid="B3">Chatzaki and Van Keer (2019)</xref>, it is closest to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mylonasi">mylonasi</tp:taxon-name-part></tp:taxon-name></italic> Chatzaki, 2019 (co-occuring on Crete), from which the males are distinguished by embolus shorter than tegular apophysis, and lack of small tibial apophysis (VTA). Our single specimen conforms to the published diagnosis and illustrations in all respects except for the presence of a small ventral tibial apophysis on both palps. Nevertheless, we assign it to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Drassodes">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serratichelis">serratichelis</tp:taxon-name-part></tp:taxon-name></italic>, presuming that the presence of a small VTA might represent intraspecific variation in this species. This species is new to the Caucasus.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
      <sec sec-type="Family Lathyidae Cala-Riquelme, Montana, Crews &amp; Esposito, 2025" id="sec6">
        <title>Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Lathyidae">Lathyidae</tp:taxon-name-part></tp:taxon-name> Cala-Riquelme, Montana, Crews &amp; Esposito, 2025</title>
        <p><bold>Remarks</bold>. Recently, based on integrative taxonomy, the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Lathyidae">Lathyidae</tp:taxon-name-part></tp:taxon-name> was established to accommodate several genera previously placed within <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Dictynidae">Dictynidae</tp:taxon-name-part></tp:taxon-name> sensu lato. This family is primarily distinguished from <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Dictynidae">Dictynidae</tp:taxon-name-part></tp:taxon-name><italic>sensu stricto</italic> by the conformation of the epigyne (for the diagnosis, see <xref ref-type="bibr" rid="B11">Montana et al. 2025</xref>).</p>
      </sec>
      <sec sec-type="Genus Tolokonniella Cala-Riquelme, Crews &amp; Esposito, 2025" id="sec7">
        <title>Genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">Tolokonniella</tp:taxon-name-part></tp:taxon-name></italic> Cala-Riquelme, Crews &amp; Esposito, 2025</title>
        <p><bold>Remarks</bold>. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">Tolokonniella</tp:taxon-name-part></tp:taxon-name></italic> was established to include several species formerly assigned to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lathys">Lathys</tp:taxon-name-part></tp:taxon-name></italic> that possess a specific set of genital characters (for the diagnosis, see <xref ref-type="bibr" rid="B11">Montana et al. 2025</xref>).</p>
        <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">Araneae</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family">Lathyidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <tp:taxon-name><object-id content-type="arpha">A83A846F-84CE-558C-93A3-E6DCE235F188</object-id>
                    		<tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">Tolokonniella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stigmatisata">stigmatisata</tp:taxon-name-part>
                    	</tp:taxon-name>
            <tp:taxon-authority>(Menge, 1869)</tp:taxon-authority>
            <xref ref-type="fig" rid="F2">Figs 3–4</xref>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lathys">Lathys</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stigmatisata">stigmatisata</tp:taxon-name-part></tp:taxon-name>
                <comment>: <xref ref-type="bibr" rid="B8">Marusik et al. 2009</xref>: 22, figs 29–32, 44, 51–53 (♂)</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="Material examined">
            <title>Material examined.</title>
            <p>GEORGIA • ♂ (<abbrev content-type="institution" xlink:title="Ilia State University, Institute of Zoology">ISUIZ</abbrev>; CaBOL-ID 1039480); Dighomi Village, Tbilisi; <named-content content-type="dwc:verbatimCoordinates">41.7803, 44.7005</named-content>; 722 m a.s.l.; steppe, under stone; leg. N. Bulbulashvili; 04 May 2023.</p>
            <fig id="F2">
              <object-id content-type="doi">10.3897/caucasiana.5.e189638.figures3-4</object-id>
              <object-id content-type="arpha">157E2E1C-2902-50F6-AD82-5D68A52CAF8D</object-id>
              <label>Figures 3–4.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">Tolokonniella</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stigmatisata">stigmatisata</tp:taxon-name-part></tp:taxon-name></italic>, male, left palp (<bold>3</bold> – ventral view; <bold>4</bold> – retrolateral view). Scale bar: 0.1 mm.</p>
              </caption>
              <graphic xlink:href="caucasiana-05-031-g002.jpg" id="oo_1626837.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1626837</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Comments">
            <title>Comments.</title>
            <p>This species has a European range and is distributed from western Europe (Portugal) to the Ural River (<xref ref-type="bibr" rid="B8">Marusik et al. 2009</xref>; <xref ref-type="bibr" rid="B12">Nentwig et al. 2026</xref>; <xref ref-type="bibr" rid="B25">WSC 2026</xref>). In the Caucasus, it was previously known from North Ossetia-Alania, Russia (Krasnodar Kray), and Azerbaijan (<xref ref-type="bibr" rid="B13">Otto 2022</xref>; <xref ref-type="bibr" rid="B7">Marusik et al. 2004</xref>). This is the first record of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tolokonniella">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stigmatisata">stigmatisata</tp:taxon-name-part></tp:taxon-name></italic> from Georgia.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
      <sec sec-type="Family Linyphiidae Blackwall, 1859" id="sec8">
        <title>Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Linyphiidae">Linyphiidae</tp:taxon-name-part></tp:taxon-name> Blackwall, 1859</title>
        <p>
          <bold>Genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part></tp:taxon-name></italic> Simon, 1913</bold>
        </p>
        <p><bold>Remarks</bold>. This small genus of yellowish-orange Palaearctic linyphiids, generally associated with caves or dark crevices, currently comprises seven recognised species, four of which are restricted to the Canary Islands. A single species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pilosus">pilosus</tp:taxon-name-part></tp:taxon-name></italic> Simon, 1913, is known from continental Eurasia (<xref ref-type="bibr" rid="B25">WSC 2026</xref>).</p>
        <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">Araneae</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family">Linyphiidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <tp:taxon-name><object-id content-type="arpha">4FD5300A-50C0-503A-B429-D021CE8BFB45</object-id>
                    		<tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pilosus">pilosus</tp:taxon-name-part>
                    	</tp:taxon-name>
            <tp:taxon-authority>Simon, 1913</tp:taxon-authority>
            <xref ref-type="fig" rid="F3">Figs 5–8</xref>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pilosus">pilosus</tp:taxon-name-part></tp:taxon-name>
                <comment>: <xref ref-type="bibr" rid="B1">Bosmans 2006</xref>: 149, figs 73–79 (♂♀)</comment>
              </tp:nomenclature-citation>
              <tp:nomenclature-citation>
                <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pilosus">pilosus</tp:taxon-name-part></tp:taxon-name>
                <comment>: <xref ref-type="bibr" rid="B23">Zamani et al. 2023</xref>: 357, figs 12A–C, 13A-B (♂)</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="Material examined">
            <title>Material examined.</title>
            <p>GEORGIA • ♀ (<abbrev content-type="institution" xlink:title="Ilia State University, Institute of Zoology">ISUIZ</abbrev>; CaBOL-ID 1012638); Tbilisi; <named-content content-type="dwc:verbatimCoordinates">41.7701, 44.7673</named-content>; 439 m a.s.l.; steppe, under stone; leg. A. Seropian; 25 Mar. 2021. • ♀ (<abbrev content-type="institution" xlink:title="Ilia State University, Institute of Zoology">ISUIZ</abbrev>; CaBOL-ID 1020246); 41.7706°, 44.7669°; 449 m a.s.l.; steppe, under stone; leg. A. Seropian; 07 Nov. 2021. • 3♀♀1♂ (<abbrev content-type="institution" xlink:title="Ilia State University, Institute of Zoology">ISUIZ</abbrev>; CaBOL-IDs 1020247, 1020249, 1020250, 1020251); Dighomi Village; <named-content content-type="dwc:verbatimCoordinates">41.7811, 44.6965</named-content>; 786 m a.s.l.; steppe, under stones; leg. A. Seropian; 14 Nov. 2021. • ♀ (<abbrev content-type="institution" xlink:title="Ilia State University, Institute of Zoology">ISUIZ</abbrev>; CaBOL-ID 1020829); idem; 27 Nov. 2021; • ♀ (<abbrev content-type="institution" xlink:title="Ilia State University, Institute of Zoology">ISUIZ</abbrev>; CaBOL-ID 1039490); Gori, Kvernaki Ridge; <named-content content-type="dwc:verbatimCoordinates">41.98558, 44.14647</named-content>; 623 m a.s.l.; heathland, under stone; leg. A. Seropian; 07 Mar. 2024.</p>
            <fig id="F3">
              <object-id content-type="doi">10.3897/caucasiana.5.e189638.figures5-8</object-id>
              <object-id content-type="arpha">A2227526-4C55-5047-BAB0-A7DE3F73D18E</object-id>
              <label>Figures 5–8.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">Scotargus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pillosus">pillosus</tp:taxon-name-part></tp:taxon-name></italic> (<bold>5</bold> – male, left palp, ventral view; <bold>6</bold> – ditto, retrolateral view; <bold>7</bold> – female, epigyne, ventral view; 8 – ditto, dorsal view). Scale bars: 0.2 mm.</p>
              </caption>
              <graphic xlink:href="caucasiana-05-031-g003.jpg" id="oo_1626838.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1626838</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Comments">
            <title>Comments.</title>
            <p>This species has a trans-Palaearctic distribution, ranging from the Iberian Peninsula to the Russian Far East (<xref ref-type="bibr" rid="B12">Nentwig et al. 2026</xref>; <xref ref-type="bibr" rid="B25">WSC 2026</xref>). In the Caucasus, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scotargus">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pilosus">pilosus</tp:taxon-name-part></tp:taxon-name></italic> was previously known from Azerbaijan and Dagestan (<xref ref-type="bibr" rid="B13">Otto 2022</xref>). The genus is new to Georgia. In Georgia, this orange medium-sized (micro)cavernicolous linyphiid species is found during early spring and late autumn in arid habitats, where it occupies spaces under stones. In contrast, in Iran, it has only been recorded from caves (<xref ref-type="bibr" rid="B23">Zamani et al. 2023</xref>).</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>We are thankful to Nils Hein (Subject Editor) and three anonymous reviewers for handling the MS and their constructive comments and suggestions, which improved this paper.</p>
    </ack>
    <sec sec-type="Additional information" id="sec9">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec10">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec11">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec12">
        <title>Artificial Intelligence (AI) use</title>
        <p>The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.</p>
        <p>No AI tools were used in the preparation of this manuscript.</p>
      </sec>
      <sec sec-type="Funding" id="sec13">
        <title>Funding</title>
        <p>No funding was reported.</p>
      </sec>
      <sec sec-type="Author contributions" id="sec14">
        <title>Author contributions</title>
        <p>Conceptualisation, original draft writing, visualisation (AS); data analysis, curation (AS, VE); material collection (AS, NB).</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec15">
        <title>Author ORCIDs</title>
        <p>Armen Seropian <ext-link xlink:href="https://orcid.org/0000-0003-3777-9954" ext-link-type="uri">https://orcid.org/0000-0003-3777-9954</ext-link></p>
        <p>Natalia Bulbulashvili <ext-link xlink:href="https://orcid.org/0000-0002-6802-1209" ext-link-type="uri">https://orcid.org/0000-0002-6802-1209</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec16">
        <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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