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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">2</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">ZooKeys</journal-title>
        <abbrev-journal-title xml:lang="en">ZK</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1313-2989</issn>
      <issn pub-type="epub">1313-2970</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/zookeys.1260.161209</article-id>
      <article-id pub-id-type="publisher-id">161209</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Lycosidae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biodiversity &amp; Conservation</subject>
          <subject>Conservation Biology</subject>
          <subject>Systematics</subject>
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Cenozoic</subject>
          <subject>Neogene</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Asia</subject>
          <subject>Asian Russia</subject>
          <subject>Central Asia</subject>
          <subject>China</subject>
          <subject>Far East</subject>
          <subject>Japan</subject>
          <subject>Korea</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>﻿ <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part></tp:taxon-name></italic> gen. nov. (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Araneae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part></tp:taxon-name>), a new genus of wolf spiders from East Asia</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Tao</surname>
            <given-names>Ze-Hong</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0006-5364-4127</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Fu</surname>
            <given-names>Dan</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-9092-4180</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Wu</surname>
            <given-names>Chang-Jun</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0007-3557-1240</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Wang</surname>
            <given-names>Yang</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0004-6748-5548</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Liu</surname>
            <given-names>Li-Juan</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0009-0410-5858</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Luo</surname>
            <given-names>Yu-Fa</given-names>
          </name>
          <email xlink:type="simple">lyf223@126.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-0645-8712</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Key Laboratory of Wetland Biodiversity of the Jianhu Basin of Shaoxing, School of Life and Environmental Sciences, Shaoxing University, Shaoxing 312000, China</addr-line>
        <institution>Shaoxing University</institution>
        <addr-line content-type="city">Shaoxing</addr-line>
        <country>China</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Yu-Fa Luo (<email xlink:type="simple">lyf223@126.com</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Ingi Agnarsson</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2025</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>13</day>
        <month>11</month>
        <year>2025</year>
      </pub-date>
      <volume>1260</volume>
      <fpage>1</fpage>
      <lpage>13</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/4138C916-BA95-5590-B1DF-C30742121ED2">4138C916-BA95-5590-B1DF-C30742121ED2</uri>
      <uri content-type="zenodo_dep_id" xlink:href="https://zenodo.org/record/17615645">17615645</uri>
      <history>
        <date date-type="received">
          <day>05</day>
          <month>06</month>
          <year>2025</year>
        </date>
        <date date-type="accepted">
          <day>13</day>
          <month>10</month>
          <year>2025</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Ze-Hong Tao, Dan Fu, Chang-Jun Wu, Yang Wang, Li-Juan Liu, Yu-Fa Luo</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <abstract>
        <label>﻿Abstract</label>
        <p>The new genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov.</bold> (type species: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963) is described and the new combination <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963), <bold>comb. nov.</bold> is made. Phylogenetic analyses of mitochondrial loci do not place <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> into <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part></tp:taxon-name></italic> or the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lycosinae</tp:taxon-name-part></tp:taxon-name> but rather suggest it is sister to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Wadicosa">Wadicosa</tp:taxon-name-part></tp:taxon-name></italic> Zyuzin, 1985 + <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosa">Pardosa</tp:taxon-name-part></tp:taxon-name></italic> C. L. Koch, 1847. Descriptions of the new genus and known species, supplemented by photographs, a distribution map, and phylogenetic evidence, are provided.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="order">Araneae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>mitochondrial genes</kwd>
        <kwd>morphology</kwd>
        <kwd>new combination</kwd>
        <kwd>new genus</kwd>
        <kwd>phylogenetics</kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>Shaoxing University</funding-statement>
      </funding-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="SECID0E4H">
        <title>Citation</title>
        <p>Tao Z-H, Fu D, Wu C-J, Wang Y, Liu L-J, Luo Y-F (2025) Chicosa gen. nov. (Araneae, Lycosidae), a new genus of wolf spiders from East Asia. ZooKeys 1260: 1–13. <ext-link xlink:type="simple" ext-link-type="doi" xlink:href="10.3897/zookeys.1260.161209">https://doi.org/10.3897/zookeys.1260.161209</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="﻿Introduction" id="SECID0EZAAC">
      <title>﻿Introduction</title>
      <p>The family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part></tp:taxon-name> (wolf spiders) represents one of the most species-rich groups within the order <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Araneae</tp:taxon-name-part></tp:taxon-name>, and its taxonomic classification has long been a focal yet challenging subject in arachnological research. The genus-level classification of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part></tp:taxon-name> has long relied on integrated morphological characters, including the male palpal organ, body striation patterns, and leg spine arrangements. However, certain species in East Asia may exhibit generic misplacements due to historically broad classification criteria or incomplete information or a lack of redescriptions of type specimens (<xref ref-type="bibr" rid="B35">Xu 2010</xref>; <xref ref-type="bibr" rid="B12">Liu et al. 2025</xref>).</p>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part></tp:taxon-name></italic> Simon, 1885, a species-rich lineage within <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part></tp:taxon-name>, currently comprises 172 recognized species, ranking as the third largest genus in the family (<xref ref-type="bibr" rid="B33">World Spider Catalog 2025</xref>). Nevertheless, the genus has long been taxonomically contentious, exhibiting significant regional discrepancies in species composition, and is widely regarded as a wastebasket taxon (<xref ref-type="bibr" rid="B32">Wang et al. 2025</xref>). <xref ref-type="bibr" rid="B13">Marusik and Kovblyuk (2011)</xref> noted that the morphological disparities between some <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part></tp:taxon-name></italic> species and the type species exceeded generic-level variation, suggesting that the genus is likely polyphyletic and requires revision and division into multiple independent genera. Despite over a century of research, regional limitations in early studies and a lack of international collaboration have perpetuated numerous misidentifications and synonymies. More critically, the current classification system often accommodates species with morphological traits that deviate from established generic diagnoses, a practice that has compromised the accuracy of phylogenetic reconstructions (<xref ref-type="bibr" rid="B6">Framenau 2015</xref>; <xref ref-type="bibr" rid="B4">Esyunin and Ponomarev 2018</xref>; <xref ref-type="bibr" rid="B18">Naumova and Deltshev 2023</xref>).</p>
      <p>Examination of specimens from several provinces of China (Hebei, Inner Mongolia, Liaoning, and Shandong) revealed significant taxonomic discrepancies between both sexes of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963) and the type species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="fabrilis">fabrilis</tp:taxon-name-part></tp:taxon-name></italic> (Clerck, 1757). Molecular phylogeny also suggested the significant divergence of this species from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part></tp:taxon-name></italic>. To resolve these inconsistencies, we establish a new genus, thereby refining the classification of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part></tp:taxon-name>. Comprehensive morphological descriptions and illustrations are provided for the new taxon.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0EHEAC">
      <title>﻿Material and methods</title>
      <sec sec-type="﻿Morphological treatment" id="SECID0ELEAC">
        <title>﻿Morphological treatment</title>
        <p>All specimens were preserved in 95% ethanol. The specimens were examined, expounded, photographed, and measured using a Phenix stereomicroscope. Palps and epigynes were examined and described after dissection using a Sony digital Camera and Wemacro software. Epigynes were cleared by immersing them in pancreatin (<xref ref-type="bibr" rid="B1">Álvarez-Padilla and Hormiga 2007</xref>). Images were focus-stacked using the software Helicon Focus ver. 3.10. Scanning electron microscope (<abbrev xlink:title="Scanning electron microscope" id="ABBRID0EVEAC">SEM</abbrev>) microphotographs were obtained with a Zeiss SIGMA300 <abbrev xlink:title="Scanning electron microscope" id="ABBRID0EZEAC">SEM</abbrev> (preparation procedures as in <xref ref-type="bibr" rid="B34">Xiao and Li 2012</xref>). Eye sizes were defined as the maximum dorsal diameter. Leg measurements are given as: total length (femur, patella and tibia, metatarsus, tarsus). All measurements are reported in millimeters (mm).</p>
        <p>The following abbreviations are used in the text and figures: ALE-anterior lateral eye; AME-anterior median eye; PLE-posterior lateral eye; PME-posterior median eye.</p>
        <p>Specimens examined here are deposited in the Key Laboratory of Wetland Biodiversity of the Jianhu Basin of Shaoxing, School of Life and Environmental Sciences, Shaoxing University, Shaoxing, China.</p>
      </sec>
      <sec sec-type="﻿DNA extraction, amplification and sequencing" id="SECID0EDFAC">
        <title>﻿DNA extraction, amplification and sequencing</title>
        <p>Genomic DNA was extracted from two legs of the paratype (LCG_11) with the TIANamp Genomic DNA Kit (Tiangen, Beijing, China). Partial sequences of the mitochondrial <italic>COI</italic>, 12S rRNA, and 16S rRNA genes were amplified using the primer pairs: LCO1490/HCO2198 for <italic>COI</italic> (<xref ref-type="bibr" rid="B5">Folmer et al. 1994</xref>); 12Sai/12Sbi for 12S rRNA (<xref ref-type="bibr" rid="B10">Kocher et al. 1989</xref>); and 16Sar/16SB2 for 16S rRNA (<xref ref-type="bibr" rid="B27">Tan et al. 1999</xref>). PCR reactions (25 μl volume) contained: 12.5 μl MyFi Mix (Bioline, USA), 1 μl each primer (10 μM), 2 μl DNA template, and 8.5 μl ddH<sub>2</sub>O. For <italic>COI</italic> amplification, the PCR protocol was as follows: initial denaturation at 94 °C for 5 min; 40 cycles of 94 °C denaturation (30 s), 45 °C annealing (90 s), 72 °C extension (1 min); and final extension at 72 °C for 5 min. For 12S rRNA amplification, the PCR protocol was as follows: an initial denaturation at 94 °C for 1 min; 35 cycles of 94 °C denaturation (25 s), 49 °C annealing (30 s), 72 °C extension (20 s); and a final extension at 72 °C for 6 min. For 16S rRNA amplification, the PCR protocol was as follows: initial denaturation at 94 °C for 1 min; 35 cycles of 94 °C denaturation (30 s), 49 °C annealing (45 s), 72 °C extension (1 min); and a final extension at 72 °C for 5 min. All PCR products were purified and directly sequenced via primer walking using BigDye technology on an ABI 3730 automated sequencer (Applied Biosystems, Foster City, CA, USA) at Tsingke Biotechnology Co., Ltd., Hangzhou, China. New sequences were submitted to GenBank (Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>).</p>
      </sec>
      <sec sec-type="﻿Phylogenetic reconstruction" id="SECID0EBGAC">
        <title>﻿Phylogenetic reconstruction</title>
        <p>Each gene was aligned using MAFFT ver. 7.0, applying the G‐INS‐i algorithm for highly conserved sequences (<italic>COI</italic>) and Q‐INS‐i for sequences with more variable regions (12S rRNA and 16S rRNA). Ambiguously aligned sites/regions were eliminated using Gblocks (<xref ref-type="bibr" rid="B2">Castresana 2000</xref>). We first used the concatenated <italic>COI</italic>, 12S rRNA and 16S rRNA dataset (Suppl. material <xref ref-type="supplementary-material" rid="S2">2</xref>) of 329 wolf spider species, including the sequences of 328 species downloaded from GenBank and BOLDSYSTEM (Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>), to construct the maximum likelihood (<abbrev xlink:title="maximum likelihood" id="ABBRID0EXGAC">ML</abbrev>) tree in W-IQ-TREE (Trifinopoulos et al. 2016). The FreeRate heterogeneity and Bayesian Information Criterion identified the best partition-specific substitution model for each gene partition (GTR+F+R6 for <italic>COI</italic>; GTR+F+I+G4 for 12S rRNA; TIM2+F+I+G4 for 16S rRNA). Perturbation strength (p) and number of iterations since the last optimal tree discovery (c) were specified as 0.5 and 1000, respectively. Branch support was assessed via ultrafast bootstrap (<abbrev xlink:title="ultrafast bootstrap" id="ABBRID0E4GAC">UFBoot</abbrev>; <xref ref-type="bibr" rid="B15">Minh et al. 2013</xref>) and SH-aLRT (<xref ref-type="bibr" rid="B7">Guindon et al. 2010</xref>) with 1000 maximum replicates and a minimum correlation coefficient threshold of 0.99. The outgroups include <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pisaura">Pisaura</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ancora">ancora</tp:taxon-name-part></tp:taxon-name></italic> Paik, 1969, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pisaura">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lama">lama</tp:taxon-name-part></tp:taxon-name></italic> Bösenberg &amp; Strand, 1906 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pisaura">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mirabilis">mirabilis</tp:taxon-name-part></tp:taxon-name></italic> Clerck, 1757. Second, we reconstructed the <abbrev xlink:title="maximum likelihood" id="ABBRID0EKIAC">ML</abbrev> tree of 85 species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lycosinae</tp:taxon-name-part></tp:taxon-name> + <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pardosinae</tp:taxon-name-part></tp:taxon-name> using concatenated <italic>COI</italic>, 12S rRNA and 16S rRNA sequences (Suppl. material <xref ref-type="supplementary-material" rid="S3">3</xref>) with the outgroups <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hippasa">Hippasa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pantherine">pantherine</tp:taxon-name-part></tp:taxon-name></italic> Pocock, 1899, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hippasa">H.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="deserticola">deserticola</tp:taxon-name-part></tp:taxon-name></italic> Simon, 1889 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hippasa">H.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="madraspatana">madraspatana</tp:taxon-name-part></tp:taxon-name></italic> Gravely, 1924. This analysis was also performed in W-IQ-TREE with the optimal substitution models (GTR+F+I+G4 for <italic>COI</italic>; TIM2+F+I+G4 for 12S rRNA; and TIM2+F+R3 for 16S rRNA) using the same sets as above.</p>
      </sec>
    </sec>
    <sec sec-type="﻿Taxonomy" id="SECID0EBKAC">
      <title>﻿Taxonomy</title>
      <sec sec-type="﻿Family Lycosidae Sundevall, 1833" id="SECID0EFKAC">
        <title>﻿Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part></tp:taxon-name> Sundevall, 1833</title>
        <p>Common name 狼蛛科</p>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Araneae</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Lycosidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">9E712250-A451-50FB-B15B-3F9C2B1ED9F8</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part>
              <object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/8E68A01E-D4F4-41D1-9588-2F206D3E2562</object-id>
            </tp:taxon-name>
            <tp:taxon-status>gen. nov.</tp:taxon-status>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="type species" id="SECID0EVLAC">
            <title>Type species.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> Schenkel, 1963.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="etymology" id="SECID0EGMAC">
            <title>Etymology.</title>
            <p>The generic name <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part></tp:taxon-name></italic> is a combination of the Mandarin Pinyin chi (from 螭, chī, a hornless dragon in Chinese mythology) and the common lycosid suffix -cosa (from Greek kósos, meaning ‘creature’). The gender is feminine.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="diagnosis" id="SECID0ETMAC">
            <title>Diagnosis.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part></tp:taxon-name></italic> gen. nov. is distinguished by the following combination of characters. Terminal apophysis (<abbrev xlink:title="terminal apophysis" id="ABBRID0EANAC">TA</abbrev>) well-developed, hook-shaped (Figs <xref ref-type="fig" rid="F1">1A, E</xref>, <xref ref-type="fig" rid="F2">2A, E</xref>); embolus (<abbrev xlink:title="embolus" id="ABBRID0EMNAC">EM</abbrev>) originates low, extends transversely, then twists retrolaterally before curving to anterior part of bulb (Figs <xref ref-type="fig" rid="F1">1C, D, E</xref>, <xref ref-type="fig" rid="F2">2A, B, D</xref>); median apophysis (<abbrev xlink:title="median apophysis" id="ABBRID0EYNAC">MA</abbrev>) undivided, cymbiform (Figs <xref ref-type="fig" rid="F1">1C, E, F</xref>, <xref ref-type="fig" rid="F2">2A, B, E</xref>). Median septum (<abbrev xlink:title="Median septum" id="ABBRID0EEOAC">MS</abbrev>) inverted T-shaped (Figs <xref ref-type="fig" rid="F1">1H</xref>, <xref ref-type="fig" rid="F3">3A</xref>); copulatory ducts (<abbrev xlink:title="copulatory ducts" id="ABBRID0EQOAC">CD</abbrev>) extremely long, spirally coiled (Figs <xref ref-type="fig" rid="F1">1G</xref>, <xref ref-type="fig" rid="F3">3B, C</xref>). Differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part></tp:taxon-name></italic> in: <abbrev xlink:title="median apophysis" id="ABBRID0EDPAC">MA</abbrev> cymbiform and entire (vs. typically bipartite), <abbrev xlink:title="terminal apophysis" id="ABBRID0EHPAC">TA</abbrev> hook-shaped (vs. often dentiform), <abbrev xlink:title="embolus" id="ABBRID0ELPAC">EM</abbrev> basally originating (vs. non-basal), and <abbrev xlink:title="copulatory ducts" id="ABBRID0EPPAC">CD</abbrev> extremely long and spiral (vs. shorter and simpler). Differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosa">Pardosa</tp:taxon-name-part></tp:taxon-name></italic> C. L. Koch, 1847 in: <abbrev xlink:title="median apophysis" id="ABBRID0E1PAC">MA</abbrev> cymbiform and entire (vs. usually bipartite), <abbrev xlink:title="terminal apophysis" id="ABBRID0E5PAC">TA</abbrev> hook-shaped (vs. dentiform), and <abbrev xlink:title="embolus" id="ABBRID0EDAAE">EM</abbrev> basally originating (vs. non-basal). Although sharing two retromarginal cheliceral teeth with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosops">Pardosops</tp:taxon-name-part></tp:taxon-name></italic> Roewer, 1955, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part></tp:taxon-name></italic> exhibits a more complex palpal organ: <abbrev xlink:title="median apophysis" id="ABBRID0EVAAE">MA</abbrev> cymbiform (vs. short and robust), <abbrev xlink:title="terminal apophysis" id="ABBRID0EZAAE">TA</abbrev> elongate and hook-shaped (vs. two spinose processes). Differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Acroniops">Acroniops</tp:taxon-name-part></tp:taxon-name></italic> Simon, 1898 in possessing four eyes (vs. two) in the anterior row. Distinguished from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorilycosa">Chorilycosa</tp:taxon-name-part></tp:taxon-name></italic> Roewer, 1960 by the straight (vs. recurved) anterior eye row and hook-shaped <abbrev xlink:title="terminal apophysis" id="ABBRID0ELBAE">TA</abbrev> (vs. two distinct spines). Differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leimonia">Leimonia</tp:taxon-name-part></tp:taxon-name></italic> C. L. Koch, 1847 in the straight anterior eye row with subequal width to second row (vs. strongly recurved and wider). Differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Draposa">Draposa</tp:taxon-name-part></tp:taxon-name></italic> Kronestedt, 2010 by the undivided, cymbiform <abbrev xlink:title="median apophysis" id="ABBRID0E4BAE">MA</abbrev> with a single acute apex bent ventrolaterally (vs. <abbrev xlink:title="median apophysis" id="ABBRID0EBCAE">MA</abbrev> transversely extended, with broad base bearing processes and narrow distal part bearing a small subapical projection), and the complete inverted T-shaped <abbrev xlink:title="Median septum" id="ABBRID0EFCAE">MS</abbrev> (vs. epigynal atrium only partially divided by a tongue-like septum). Differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Wadicosa">Wadicosa</tp:taxon-name-part></tp:taxon-name></italic> Zyuzin, 1985 in: retromargin with two teeth (vs. three), subtegulum small and positioned directly below tegulum (vs. large and anteroventrally shifted), <abbrev xlink:title="terminal apophysis" id="ABBRID0EQCAE">TA</abbrev> without spiral torsion (vs. strongly twisted), and <abbrev xlink:title="embolus" id="ABBRID0EUCAE">EM</abbrev> originating low and extending transversely (vs. originating antero-apically); female with distinct inverted T-shaped <abbrev xlink:title="Median septum" id="ABBRID0EYCAE">MS</abbrev> (vs. absent).</p>
            <fig id="F1" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1260.161209.figure1</object-id>
              <object-id content-type="zenodo_dep_id">17615700</object-id>
              <object-id content-type="arpha">0C20C5D1-4A8F-576C-8359-07EF4186803D</object-id>
              <label>Figure 1.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963), comb. nov. A. Male habitus, dorsal view; B. Female habitus, dorsal view; C. Left male pedipalp bulb, ventral view; D. Same, retrolateral view; E. Left male palp, ventral view; F. Same, retrolateral view; G. Epigyne, dorsal view; H. Same, ventral view. Scale bars: 1 mm (A, B); 0.4 mm (E, F); 0.2 mm (C, D, G, H). Abbreviations: <abbrev xlink:title="copulatory ducts" id="ABBRID0EPDAE">CD</abbrev> = copulatory duct; CO = copulatory opening; <abbrev xlink:title="embolus" id="ABBRID0ETDAE">EM</abbrev> = embolus; FD = fertilization duct; <abbrev xlink:title="median apophysis" id="ABBRID0EXDAE">MA</abbrev> = median apophysis; <abbrev xlink:title="Median septum" id="ABBRID0E2DAE">MS</abbrev> = median septum; S = spermatheca; ST = subtegulum; <abbrev xlink:title="terminal apophysis" id="ABBRID0E6DAE">TA</abbrev> = terminal apophysis; and TE = tegulum.</p>
              </caption>
              <graphic xlink:href="zookeys-1260-001_article-161209__-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1463990.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1463990</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Phylogeny" id="SECID0EIEAE">
            <title>Phylogeny.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> comb. nov. clusters together with the species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pardosinae</tp:taxon-name-part></tp:taxon-name>, and it is sister to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Wadicosa">Wadicosa</tp:taxon-name-part></tp:taxon-name></italic> + <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosa">Pardosa</tp:taxon-name-part></tp:taxon-name></italic> (Fig. <xref ref-type="fig" rid="F4">4</xref>).</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EQFAE">
            <title>Description.</title>
            <p>See species description.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Composition" id="SECID0EVFAE">
            <title>Composition.</title>
            <p>Only the type species.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="distribution" id="SECID0E1FAE">
            <title>Distribution.</title>
            <p><bold>China</bold> (Anhui, Beijing, Hebei, Henan, Hunan, Inner Mongolia, Jilin, Shanxi, Shandong, Xinjiang and Zhejiang provinces), Japan, Kazakhstan, North Korea and Russia (Fig. <xref ref-type="fig" rid="F5">5</xref>).</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Araneae</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Lycosidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">90338645-3497-56DD-87F5-157A8596D6CE</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>(Schenkel, 1963)</tp:taxon-authority>
            <tp:taxon-status>comb. nov.</tp:taxon-status>
            <xref ref-type="fig" rid="F1">Figs 1</xref>
            <xref ref-type="fig" rid="F2">, 2</xref>
            <xref ref-type="fig" rid="F3">, 3</xref>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Tarentula">Tarentula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name>
                <comment>Schenkel, 1963: 333, fig. 192 (♀); Hu 1984: 246, fig. 258. 1–2 (♀♂).</comment>
              </tp:nomenclature-citation>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosa">Pardosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lusisi">lusisi</tp:taxon-name-part></tp:taxon-name>
                <comment>: <xref ref-type="bibr" rid="B26">Šternbergs 1981</xref>: 60, fig. 1 (♀♂).</comment>
              </tp:nomenclature-citation>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name>
                <comment>: <xref ref-type="bibr" rid="B11">Li and Chen 1982</xref>: 66, figs 1–3 (♀♂); <xref ref-type="bibr" rid="B21">Song 1982</xref>: 76, figs 3–4 (♂); <xref ref-type="bibr" rid="B40">Zhu and Shi 1985</xref>: 129, fig. 113a–c (♀); <xref ref-type="bibr" rid="B22">Song 1986</xref>: 78, figs 19–22 (♀); <xref ref-type="bibr" rid="B23">Song 1987</xref>: 215, fig. 174 (♀♂); <xref ref-type="bibr" rid="B28">Tanaka 1987</xref>: 17, figs 1–4 (♀♂); <xref ref-type="bibr" rid="B38">Zhang 1987</xref>: 144, fig. 119.1–3 (♀♂); <xref ref-type="bibr" rid="B19">Paik 1988</xref>: 92, figs 25–34 (♀♂); <xref ref-type="bibr" rid="B8">Hu and Wu 1989</xref>: 186, fig. 155.1–5 (♀♂); <xref ref-type="bibr" rid="B3">Chen and Zhang 1991</xref>: 216, fig. 221.1–4 (♀♂); <xref ref-type="bibr" rid="B29">Tanaka 1992</xref>: 326, figs 13–16 (♀♂); <xref ref-type="bibr" rid="B36">Yin et al. 1997</xref>: 60, fig. 25a–f (♀♂); <xref ref-type="bibr" rid="B24">Song et al. 1999</xref>: 316, figs 186K, 187B (♀♂); <xref ref-type="bibr" rid="B14">Marusik et al. 2000</xref>: 77; <xref ref-type="bibr" rid="B25">Song et al. 2001</xref>: 231, fig. 141A–D (♀♂); <xref ref-type="bibr" rid="B16">Namkung 2002</xref>: 313, fig. 20.7a–b (♀♂); <xref ref-type="bibr" rid="B17">Namkung 2003</xref>: 315, fig. 20.7a–b (♀♂); <xref ref-type="bibr" rid="B30">Tanaka 2009</xref>: 238, figs 93–94 (♀♂); <xref ref-type="bibr" rid="B41">Zhu and Zhang 2011</xref>: 253, fig. 182A–D (♀♂); <xref ref-type="bibr" rid="B37">Yin et al. 2012</xref>: 791, fig. 395a–f (♀♂); <xref ref-type="bibr" rid="B39">Zhang et al. 2022</xref>: 123, fig. 87A–H (♀♂).</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="material" id="SECID0EYLAE">
            <title>Material examined.</title>
            <p><bold>China, Shandong</bold>: • 2♂ 3♀, Taian City, Taishan District, Xujiazhuang village, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[117.254722,36.172222]}" id="NCID0EDMAE">36°10'20"N, 117°15'17"E</named-content></named-content>, elev. 135.4 m, 3 August 2023, L.J. Liu and Y. Cheng leg. • 1♂ 2♀, Taian City, Daiyue District, Yunbeishan village, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[117.345000,36.208889]}" id="NCID0ELMAE">36°12'32"N, 117°20'42"E</named-content></named-content>, elev. 141.5 m, 4 August 2023, L.J. Liu and Y. Cheng leg. <bold>Liaoning</bold>: • 22♀, Tieling City, Changtu County, Hongshan village, Hongshan Reservoir, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[124.123333,42.881944]}" id="NCID0EVMAE">42°52'55"N, 124°07'24"E</named-content></named-content>, elev. 150.7 m, 1 August 2023, L.J. Liu and Y. Cheng leg. <bold>Hebei</bold>: • 1♂ 1♀, Shijiazhuang City, Yuanshi County, Panlong Lake, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[114.362222,37.796389]}" id="NCID0E6MAE">37°47'47"N, 114°21'44"E</named-content></named-content>, elev. 112.8 m, 16 July 2023, L.J. Liu and Y. Cheng leg. <bold>Inner Mongolia</bold>: • 6♀, Bayannur City, Urad Front Banner, Shunda Petroleum Gas Station, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[108.684444,40.725556]}" id="NCID0EJNAE">40°43'32"N, 108°41'4"E</named-content></named-content>, elev. 1054.2 m, 13 August 2024, Z.H. Tao and H.F. Shi leg. • 2♀, Bayannur City, Hanggin Rear Banner Railway Station, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[107.114722,40.696111]}" id="NCID0ERNAE">40°41'46"N, 107°6'53"E</named-content></named-content>, elev. 1035.5 m, 20 August 2024, Z.H. Tao and H.F. Shi leg.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="diagnosis" id="SECID0EWNAE">
            <title>Diagnosis.</title>
            <p>See generic diagnosis.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0E2NAE">
            <title>Description.</title>
            <p><bold>Male</bold> (Fig. <xref ref-type="fig" rid="F1">1A</xref>): Total length 7.15. Carapace 3.80 long, 3.15 wide; opisthosoma 3.28 long, 1.87 wide. Carapace yellow-green; black longitudinal median band present; cervical and radial furrows faint. Ocular area black, with white, orange and black setae. Eyes: AME 0.20, ALE 0.14, PME 0.32, PLE 0.30; AME–AME 0.14, AME–ALE 0.04, PME–PME 0.31, PME–PLE 0.35. Clypeus height 0.25. Chelicerae pale green; promargin with 3 teeth, retromargin with 2 teeth. Maxillae and labium faint yellow. Sternum pale yellow with black markings. Leg measurements: I 14.64 (3.90, 5.02, 3.62, 2.10); II 13.92 (3.73, 4.78, 3.44, 1.97); III 12.23 (3.41, 4.04, 3.07, 1.71); IV 16.85 (4.40, 5.22, 4.79, 2.44). Leg formula: 4-1-2-3. Opisthosoma long-oval; dorsum grayish-black with light brown maculations; cardiac mark elongated, paired lateral stripes distinct; venter center grayish-yellowish brown, with scattered grayish-black patches laterally.</p>
            <p><bold><italic>Palp</italic></bold> (Figs <xref ref-type="fig" rid="F1">1C–F</xref>, <xref ref-type="fig" rid="F2">2</xref>). Terminal apophysis twisted distally, ending in elongated digitiform process curved retrolaterally and ventrally; median apophysis cymbiform, with acuminate apex bent ventrolaterally; embolus filiform, originating from median apophysis base, initially curved dorsally then descending sharply.</p>
            <fig id="F2" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1260.161209.figure2</object-id>
              <object-id content-type="zenodo_dep_id">17615702</object-id>
              <object-id content-type="arpha">2E2BA64C-5635-54D5-B670-D67C1D7459FA</object-id>
              <label>Figure 2.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963), comb. nov. A. Left male pedipalp bulb, ventral view; B. Median apophysis, ventral view; C. Embolus, retrolateral view; D. Embolus, ventral view; E. Terminal apophysis and median apophysis, ventral view. Scale bars: 0.1 mm (A−E). Abbreviations: <abbrev xlink:title="embolus" id="ABBRID0EJPAE">EM</abbrev> = embolus; <abbrev xlink:title="median apophysis" id="ABBRID0ENPAE">MA</abbrev> = median apophysis; <abbrev xlink:title="terminal apophysis" id="ABBRID0ERPAE">TA</abbrev> = terminal apophysis.</p>
              </caption>
              <graphic xlink:href="zookeys-1260-001_article-161209__-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1463991.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1463991</uri>
              </graphic>
            </fig>
            <p><bold>Female</bold> (Fig. <xref ref-type="fig" rid="F1">1B</xref>): Total length 8.53. Carapace 4.22 long, 3.67 wide; opisthosoma 4.43 long, 2.85 wide. Carapace tawny; black longitudinal median band present; cervical and radial furrows faint. Ocular area as in male. Eyes: AME 0.22, ALE 0.15, PME 0.35, PLE 0.34; AME–AME 0.15, AME–ALE 0.06, PME–PME 0.35, PME–PLE 0.43. Chelicerae brownish-yellow with black stripes; promargin with 3 teeth, retromargin with 2 teeth. Maxillae faint yellow, labium yellow. Sternum yellow with black markings. Leg measurements: I 12.19 (3.43, 4.27, 2.78, 1.71); II 11.52 (3.35, 3.99, 2.55, 1.63); III 10.79 (3.12, 3.61, 2.54, 1.52); IV 15.99 (4.35, 4.99, 4.47, 2.18). Leg formula: 4-1-2-3.</p>
            <p><bold><italic>Epigyne</italic></bold> (Figs <xref ref-type="fig" rid="F1">1G, H</xref>, <xref ref-type="fig" rid="F3">3</xref>). Median septum inverted T-shaped; depression present near insertion pore. Copulatory duct extremely long, spirally coiled; spermatheca rod-shaped; fertilization duct slender, curved.</p>
            <fig id="F3" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1260.161209.figure3</object-id>
              <object-id content-type="zenodo_dep_id">17615704</object-id>
              <object-id content-type="arpha">63686492-11CD-5ED7-8AF3-383B2FEC854E</object-id>
              <label>Figure 3.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963), comb. nov. A. Epigyne, ventral view; B. Epigyne, dorsal view; C. Copulatory duct, dorsal view. Scale bars: 0.1 mm (A–C).</p>
              </caption>
              <graphic xlink:href="zookeys-1260-001_article-161209__-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1463992.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1463992</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
      <sec sec-type="﻿Molecular phylogeny" id="SECID0EJRAE">
        <title>﻿Molecular phylogeny</title>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> comb. nov. and all analyzed species of the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pardosinae</tp:taxon-name-part></tp:taxon-name> cluster into a clade, and it is sister to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosa">Pardosa</tp:taxon-name-part></tp:taxon-name></italic> + <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Wadicosa">Wadicosa</tp:taxon-name-part></tp:taxon-name></italic> (Fig. <xref ref-type="fig" rid="F4">4</xref>).</p>
        <fig id="F4" position="float" orientation="portrait">
          <object-id content-type="doi">10.3897/zookeys.1260.161209.figure4</object-id>
          <object-id content-type="zenodo_dep_id">17615706</object-id>
          <object-id content-type="arpha">60B4237F-2A5C-5418-A1DA-1E2FDCBE98A2</object-id>
          <label>Figure 4.</label>
          <caption>
            <p>Phylogenetic trees reconstructed with the <abbrev xlink:title="maximum likelihood" id="ABBRID0EZSAE">ML</abbrev> method. A. <abbrev xlink:title="maximum likelihood" id="ABBRID0E4SAE">ML</abbrev> tree constructed using the concatenated <italic>COI</italic>, 12S rRNA and 16S rRNA dataset of 329 species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part></tp:taxon-name>; B. <abbrev xlink:title="maximum likelihood" id="ABBRID0EITAE">ML</abbrev> tree reconstructed using the concatenated <italic>COI</italic>, 12S rRNA and 16S rRNA sequences of 85 species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lycosinae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pardosinae</tp:taxon-name-part></tp:taxon-name>. The numbers at the nodes represent bootstrap support values from the <abbrev xlink:title="maximum likelihood" id="ABBRID0EYTAE">ML</abbrev> analyses. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> comb. nov. is shown in red.</p>
          </caption>
          <graphic xlink:href="zookeys-1260-001_article-161209__-g004.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1463993.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1463993</uri>
          </graphic>
        </fig>
        <fig id="F5" position="float" orientation="portrait">
          <object-id content-type="doi">10.3897/zookeys.1260.161209.figure5</object-id>
          <object-id content-type="zenodo_dep_id">17615708</object-id>
          <object-id content-type="arpha">DB134C6A-65DA-56AC-AA22-6092437B8206</object-id>
          <label>Figure 5.</label>
          <caption>
            <p>Geographic distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963).</p>
          </caption>
          <graphic xlink:href="zookeys-1260-001_article-161209__-g005.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1463994.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1463994</uri>
          </graphic>
        </fig>
      </sec>
    </sec>
    <sec sec-type="﻿Discussion" id="SECID0EEVAE">
      <title>﻿Discussion</title>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> was originally placed in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tarentula">Tarentula</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963), with subsequent revisions by <xref ref-type="bibr" rid="B26">Šternbergs (1981)</xref> and <xref ref-type="bibr" rid="B11">Li and Chen (1982)</xref>, until its monospecific status was confirmed by <xref ref-type="bibr" rid="B14">Marusik et al. (2000)</xref>. Based on autapomorphies in reproductive structures and phylogenomic evidence, we hereby establish the new genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part></tp:taxon-name></italic> gen. nov.</p>
      <p>The male palp exhibits a well-developed, hook-shaped terminal apophysis (<abbrev xlink:title="terminal apophysis" id="ABBRID0ERWAE">TA</abbrev>); a slender, sinuous, basally originating embolus (<abbrev xlink:title="embolus" id="ABBRID0EVWAE">EM</abbrev>); and an undivided, cymbiform median apophysis (<abbrev xlink:title="median apophysis" id="ABBRID0EZWAE">MA</abbrev>). The female epigyne possesses an inverted T-shaped median septum (<abbrev xlink:title="Median septum" id="ABBRID0E4WAE">MS</abbrev>) and extremely long, spirally coiled copulatory ducts (<abbrev xlink:title="copulatory ducts" id="ABBRID0EBXAE">CD</abbrev>). These traits differ markedly from all known <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Alopecosa">Alopecosa</tp:taxon-name-part></tp:taxon-name></italic> species, particularly the unique configuration of the <abbrev xlink:title="copulatory ducts" id="ABBRID0EMXAE">CD</abbrev>.</p>
      <p>Although <xref ref-type="bibr" rid="B36">Yin et al. (1997</xref>, <xref ref-type="bibr" rid="B37">2012</xref>) noted convergences in cymbial morphology with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosa">Pardosa</tp:taxon-name-part></tp:taxon-name></italic>, the new genus is readily distinguished by its basally originating <abbrev xlink:title="embolus" id="ABBRID0EBYAE">EM</abbrev> (vs. non-basal), hook-shaped <abbrev xlink:title="terminal apophysis" id="ABBRID0EFYAE">TA</abbrev> (vs. dentiform), and spirally coiled <abbrev xlink:title="copulatory ducts" id="ABBRID0EJYAE">CD</abbrev> (vs. short and straight). Furthermore, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part></tp:taxon-name></italic> differs fundamentally from other available genera: from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pardosops">Pardosops</tp:taxon-name-part></tp:taxon-name></italic> Roewer, 1955 (short and robust <abbrev xlink:title="median apophysis" id="ABBRID0E2YAE">MA</abbrev>; two spinose TAs); <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Acroniops">Acroniops</tp:taxon-name-part></tp:taxon-name></italic> Simon, 1898 (two-eyed anterior row); <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chorilycosa">Chorilycosa</tp:taxon-name-part></tp:taxon-name></italic> Roewer, 1960 (recurved anterior eye row; two spines on <abbrev xlink:title="terminal apophysis" id="ABBRID0ENZAE">TA</abbrev>); and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leimonia">Leimonia</tp:taxon-name-part></tp:taxon-name></italic> C. L. Koch, 1847 (strongly recurved and wider anterior eye row).</p>
      <p>The unique combination of genitalic and ocular characters precludes its assignment to any known lycosid genus, supporting the establishment of a new genus to reflect its evolutionary distinctness.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>﻿Acknowledgements</title>
      <p>Great thanks are given to Yuri M. Marusik (Magadan, Russia) and Kirill G. Mikhailov (Moscow, Russia) for providing specific distribution information on <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chicosa">Chicosa</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cinnameopilosa">cinnameopilosa</tp:taxon-name-part></tp:taxon-name></italic> (Schenkel, 1963) in Kazakhstan and Russia.</p>
    </ack>
    <sec sec-type="﻿Additional information" id="SECID0EJ1AE">
      <title>﻿Additional information</title>
      <sec sec-type="Conflict of interest" id="SECID0EN1AE">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="SECID0ES1AE">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Use of AI" id="SECID0EX1AE">
        <title>Use of AI</title>
        <p>No use of AI was reported.</p>
      </sec>
      <sec sec-type="Funding" id="SECID0E31AE">
        <title>Funding</title>
        <p>This research was supported by the National Natural Science Foundation of China (Grant No. 32170463, 31860602, 31660611), and the Zhejiang Provincial Natural Science Founda-tion of China (Grant No. LTGN24C140006).</p>
      </sec>
      <sec sec-type="Author contributions" id="SECID0EB2AE">
        <title>Author contributions</title>
        <p>All authors have contributed equally.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="SECID0EG2AE">
        <title>Author ORCIDs</title>
        <p>Ze-Hong Tao <ext-link xlink:href="https://orcid.org/0009-0006-5364-4127" ext-link-type="uri" xlink:type="simple">https://orcid.org/0009-0006-5364-4127</ext-link></p>
        <p>Dan Fu <ext-link xlink:href="https://orcid.org/0000-0002-9092-4180" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-9092-4180</ext-link></p>
        <p>Chang-Jun Wu <ext-link xlink:href="https://orcid.org/0009-0007-3557-1240" ext-link-type="uri" xlink:type="simple">https://orcid.org/0009-0007-3557-1240</ext-link></p>
        <p>Yang Wang <ext-link xlink:href="https://orcid.org/0009-0004-6748-5548" ext-link-type="uri" xlink:type="simple">https://orcid.org/0009-0004-6748-5548</ext-link></p>
        <p>Li-Juan Liu <ext-link xlink:href="https://orcid.org/0009-0009-0410-5858" ext-link-type="uri" xlink:type="simple">https://orcid.org/0009-0009-0410-5858</ext-link></p>
        <p>Yu-Fa Luo <ext-link xlink:href="https://orcid.org/0000-0002-0645-8712" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-0645-8712</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="SECID0EO3AE">
        <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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    <sec sec-type="supplementary-material">
      <title>Supplementary materials</title>
      <supplementary-material id="S1" position="float" orientation="portrait" xlink:type="simple">
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        <label>Supplementary material 1</label>
        <caption>
          <p>Samples used in this study</p>
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            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Ze-Hong Tao, Dan Fu, Chang-Jun Wu, Yang Wang, Li-Juan Liu, Yu-Fa Luo</attrib>
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        <object-id content-type="doi">10.3897/zookeys.1260.161209.suppl2</object-id>
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        <label>Supplementary material 2</label>
        <caption>
          <p>The concatenated <italic>COI</italic>, 12S and 16S dataset of 329 <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lycosidae</tp:taxon-name-part></tp:taxon-name> species</p>
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          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Ze-Hong Tao, Dan Fu, Chang-Jun Wu, Yang Wang, Li-Juan Liu, Yu-Fa Luo</attrib>
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      <supplementary-material id="S3" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/zookeys.1260.161209.suppl3</object-id>
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        <label>Supplementary material 3</label>
        <caption>
          <p>The concatenated <italic>COI</italic>, 12S and 16S dataset of 85 <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lycosinae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pardosinae</tp:taxon-name-part></tp:taxon-name> species</p>
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        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Ze-Hong Tao, Dan Fu, Chang-Jun Wu, Yang Wang, Li-Juan Liu, Yu-Fa Luo</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
