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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">104</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:f2cd1fff-21e4-581f-a7fa-850997197b7f</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:B1C81912-2D17-4CD8-8D2C-EFEAAAB2EF75</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Vertebrate Zoology</journal-title>
        <abbrev-journal-title xml:lang="en">VZ</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1864-5755</issn>
      <issn pub-type="epub">2625-8498</issn>
      <publisher>
        <publisher-name>Senckenberg Gesellschaft für Naturforschung</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/vz.76.e176241</article-id>
      <article-id pub-id-type="publisher-id">176241</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Reptilia</subject>
          <subject>Sauria</subject>
          <subject>Scincidae</subject>
          <subject>Squamata</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biogeography</subject>
          <subject>Molecular systematics</subject>
          <subject>Morphology</subject>
          <subject>Systematics</subject>
          <subject>Taxonomy</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Two new highly divergent and isolated <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> species from Nosy Be and the Tsingy de Namoroka, Madagascar (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Squamata">Squamata</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scincidae">Scincidae</tp:taxon-name-part></tp:taxon-name>)</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Miralles</surname>
            <given-names>Aurélien</given-names>
          </name>
          <email xlink:type="simple">miralles.skink@gmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-2538-7710</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
          <role content-type="http://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
          <role content-type="http://credit.niso.org/contributor-roles/visualization/">Visualization</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Scherz</surname>
            <given-names>Mark D.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-4613-7761</uri>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Hyde Roberts</surname>
            <given-names>Sam</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-9757-9121</uri>
          <xref ref-type="aff" rid="A3">3</xref>
          <xref ref-type="aff" rid="A4">4</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Rakotoarison</surname>
            <given-names>Andolalao</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-2620-440X</uri>
          <xref ref-type="aff" rid="A5">5</xref>
          <xref ref-type="aff" rid="A6">6</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Glaw</surname>
            <given-names>Frank</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-4072-8111</uri>
          <xref ref-type="aff" rid="A7">7</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Vences</surname>
            <given-names>Miguel</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-0747-0817</uri>
          <xref ref-type="aff" rid="A8">8</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</role>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Institut de Systématique, Évolution, Biodiversité (ISYEB), Muséum national d’Histoire Naturelle, CNRS, Sorbonne Université, EPHE, Université des Antilles, 57 rue Cuvier, 75231 Paris cedex, France</addr-line>
        <institution>Duke University</institution>
        <addr-line content-type="city">Durham</addr-line>
        <country>United States of America</country>
        <uri content-type="ror">https://ror.org/00py81415</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Natural History Museum Denmark, University of Copenhagen, Universitetsparken 15, 2100, Copenhagen Ø, Denmark</addr-line>
        <institution>Technische Universität Braunschweig</institution>
        <addr-line content-type="city">Braunschweig</addr-line>
        <country>Germany</country>
        <uri content-type="ror">https://ror.org/010nsgg66</uri>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Duke University, Biological Sciences, Science Drive, Durham, North Carolina 27708, USA</addr-line>
        <institution>University of Copenhagen</institution>
        <addr-line content-type="city">Copenhagen</addr-line>
        <country>Denmark</country>
        <uri content-type="ror">https://ror.org/035b05819</uri>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">SEED Madagascar, 7 Bell Yard, London, WC2A 2JR, United Kingdom</addr-line>
        <institution>Museum national d'Histoire naturelle</institution>
        <addr-line content-type="city">Paris</addr-line>
        <country>France</country>
        <uri content-type="ror">https://ror.org/03wkt5x30</uri>
      </aff>
      <aff id="A5">
        <label>5</label>
        <addr-line content-type="verbatim">Mention Environnement, Universite de l’Itasy, Faliarivo Ambohidanerana, 118 Soavinandriana Itasy, Madagascar</addr-line>
        <institution>Zoologische Staatssammlung München (ZSM-SNSB)</institution>
        <addr-line content-type="city">München</addr-line>
        <country>Germany</country>
        <uri content-type="ror">https://ror.org/04rekk491</uri>
      </aff>
      <aff id="A6">
        <label>6</label>
        <addr-line content-type="verbatim">School for International Training, VN 41A Bis Ankazolava Ambohitsoa, Antananarivo, 101 Madagascar</addr-line>
        <institution>SEED Madagascar</institution>
        <addr-line content-type="city">London</addr-line>
        <country>United Kingdom</country>
      </aff>
      <aff id="A7">
        <label>7</label>
        <addr-line content-type="verbatim">Zoologische Staatssammlung München (ZSM-SNSB), Münchhausenstr. 21, 81247 München, Germany</addr-line>
        <institution>Universite de l’Itasy</institution>
        <addr-line content-type="city">Soavinandriana</addr-line>
        <country>Madagascar</country>
      </aff>
      <aff id="A8">
        <label>8</label>
        <addr-line content-type="verbatim">Division of Evolutionary Biology, Zoological Institute, Braunschweig University of Technology, Mendelssohnstr. 4, 38106 Braunschweig, Germany</addr-line>
        <institution>School for International Training</institution>
        <addr-line content-type="city">Antananarivo</addr-line>
        <country>Madagascar</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Aurélien Miralles (<email xlink:type="simple">miralles.skink@gmail.com</email>)</p>
        </fn>
        <fn>
          <p><bold>Academic editor</bold> Uwe Fritz</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>06</day>
        <month>03</month>
        <year>2026</year>
      </pub-date>
      <volume>76</volume>
      <fpage>135</fpage>
      <lpage>156</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/56831F72-CE15-58DD-A116-716F03CC9B93">56831F72-CE15-58DD-A116-716F03CC9B93</uri>
      <uri content-type="zenodo_dep_id" xlink:href="https://zenodo.org/record/0">0</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/21F8A4B3-D34D-4DA6-8A7F-84A739352019">21F8A4B3-D34D-4DA6-8A7F-84A739352019</uri>
      <history>
        <date date-type="received">
          <day>27</day>
          <month>10</month>
          <year>2025</year>
        </date>
        <date date-type="accepted">
          <day>09</day>
          <month>02</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Aurélien Miralles, Mark D. Scherz, Sam Hyde Roberts, Andolalao Rakotoarison, Frank Glaw, Miguel Vences</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/21F8A4B3-D34D-4DA6-8A7F-84A739352019</self-uri>
      <abstract>
        <p>
          <bold>Abstract</bold>
        </p>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> is a genus of quadrupedal skinks endemic to Madagascar, with 12 recognized species described between the 19<sup>th</sup> and 21<sup>st</sup> centuries, occupying diverse habitats from humid forests to arid southern regions and even high-altitude areas. Recent field expeditions uncovered two morphologically distinct forms that did not match any known species, prompting integrative taxonomic analyses that combine multilocus phylogenetics and morphology. Results revealed that each of these forms represents a highly divergent and likely ancient lineage, as evidenced by the substantial branch lengths in both mitochondrial and nuclear phylogenetic trees. These investigations led to the formal description of two new species with restricted distribution ranges: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., likely endemic to the island of Nosy Be, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., apparently restricted to the Tsingy de Namoroka karst system.</p>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>Biogeography</kwd>
        <kwd>herpetofauna</kwd>
        <kwd>integrative taxonomy</kwd>
        <kwd>phylogeny</kwd>
        <kwd>reptile</kwd>
        <kwd>skink</kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>Deutsche Forschungsgemeinschaft (grant MI 2748/1-1)
Agence Nationale de la Recherche (ANR-24-CPJ1-0129-01)</funding-statement>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec1">
      <title>Introduction</title>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> is a genus of fully quadrupedal skinks that belong to the endemic scincine clade that colonized Madagascar during the Paleogene period and successfully diversified on the island (<xref ref-type="bibr" rid="B37">Schmitz et al. 2005</xref>; <xref ref-type="bibr" rid="B10">Crottini et al. 2012</xref>; <xref ref-type="bibr" rid="B22">Miralles et al. 2022</xref>). Based on an integrative taxonomic revision combining multilocus and morphological datasets, <xref ref-type="bibr" rid="B25">Miralles et al. (2016)</xref> recognised a total of 12 described species within the genus, all endemic to Madagascar. Half of them (six) were described during the second half of the 19<sup>th</sup> century: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="polleni">polleni</tp:taxon-name-part></tp:taxon-name></italic> (Grandidier, 1869), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="igneocaudatus">igneocaudatus</tp:taxon-name-part></tp:taxon-name></italic> (Grandidier, 1867), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mouroundavae">mouroundavae</tp:taxon-name-part></tp:taxon-name></italic> (Grandidier, 1872), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="melanopleura">melanopleura</tp:taxon-name-part></tp:taxon-name></italic> (Günther, 1877), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stumpffi">stumpffi</tp:taxon-name-part></tp:taxon-name></italic> (Boettger, 1882) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrolepis">macrolepis</tp:taxon-name-part></tp:taxon-name></italic> (Boulenger, 1888). Two additional species were described during the 20<sup>th</sup> century: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ankodabensis">ankodabensis</tp:taxon-name-part></tp:taxon-name></italic> (Angel, 1930) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minutus">minutus</tp:taxon-name-part></tp:taxon-name></italic> (Raxworthy &amp; Nussbaum, 1993). Finally, four species have been added in the 21<sup>st</sup> century: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nanus">nanus</tp:taxon-name-part></tp:taxon-name></italic> (Andreone &amp; Greer, 2002), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="arenicola">arenicola</tp:taxon-name-part></tp:taxon-name></italic> Miralles, Köhler, Glaw &amp; Vences, 2011, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pyrurus">pyrurus</tp:taxon-name-part></tp:taxon-name></italic> Miralles, Köhler, Glaw &amp; Vences, 2016, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="miafina">miafina</tp:taxon-name-part></tp:taxon-name></italic> Miralles, Köhler, Glaw &amp; Vences, 2016. Widely distributed across Madagascar and occupying a range of different biomes, species of the genus occur from the eastern humid evergreen forests (e.g., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="melanopleura">melanopleura</tp:taxon-name-part></tp:taxon-name></italic>) to the hottest and driest sandy areas of the South (e.g., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="igneocaudatus">igneocaudatus</tp:taxon-name-part></tp:taxon-name></italic>). One species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pyrurus">pyrurus</tp:taxon-name-part></tp:taxon-name></italic>, has even colonized the elevated and cool habitats of Mount Ibity (up to 1700 m a.s.l).</p>
      <p>During two field expeditions carried out in recent years, we had the opportunity to collect two forms of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> that did not match any species previously recorded in the scientific literature morphologically. Their isolated geographical distribution, as well as the singular combination of morphological features they exhibit, prompted us to investigate the taxonomic status of these two forms more thoroughly using an integrative approach, combining multilocus phylogenetic analyses and morphological assessment, and we herein conclude that they represent new species, formally named and described in the taxonomy section below.</p>
      <p>For the sake of simplicity and clarity, we anticipate our taxonomic results and introduce from here on their names. The first species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. was collected on Nosy Be, an island that covers an area of 321 km<sup>2</sup> and is located eight kilometers off the northwestern coast of Madagascar in the Sambirano region. The second species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. was collected in the Tsingy de Namoroka, a labyrinthic karst system extending over ca. 220 km<sup>2</sup> and located within the Mahajanga basin in the North West of Madagascar.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec2">
      <title>Materials and Methods</title>
      <sec sec-type="Sampling, acronyms and definitions" id="sec3">
        <title>Sampling, acronyms and definitions</title>
        <p>In addition to the newly collected material, the comparative morphological dataset is composed of a total of 168 preserved specimens that were genotyped by <xref ref-type="bibr" rid="B29">Miralles and Vences (2013)</xref>, complemented by 40 additional specimens without associated molecular data; as well as data previously published by <xref ref-type="bibr" rid="B2">Andreone and Greer (2002)</xref>, <xref ref-type="bibr" rid="B14">Glaw and Vences (2007)</xref> and Miralles et al. (<xref ref-type="bibr" rid="B24">2011a</xref>, <xref ref-type="bibr" rid="B25">2016</xref>). See File S1 for a complete list of the material examined. All the name-bearing type specimens known for this genus have been examined, except the types of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minutus">minutus</tp:taxon-name-part></tp:taxon-name></italic> Raxworthy &amp; Nussbaum, 1993 (UMMZ 192705) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Sepsina">Sepsina</tp:taxon-name-part></tp:taxon-name> vulsini</italic> Barbour, 1918 (MCZ R-11869), regarded as a junior synonym of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="melanopleura">melanopleura</tp:taxon-name-part></tp:taxon-name></italic> (Günther, 1877), which are both unambiguously members of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="melanopleura">melanopleura</tp:taxon-name-part></tp:taxon-name></italic> clade. The <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nanus">nanus</tp:taxon-name-part></tp:taxon-name></italic> group is herein considered a putative assemblage of miniaturized species within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> requiring formal taxonomic evaluation. It tentatively includes <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nanus">nanus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrolepis">macrolepis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> sp. “baeus”, and a candidate species from Betampona. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrolepis">macrolepis</tp:taxon-name-part></tp:taxon-name></italic> is excluded from most analyses due to its unresolved taxonomic status and the absence of molecular and recent voucher data.</p>
        <p>Specimens are deposited in the Muséum National d’Histoire Naturelle, Paris (<bold><named-content content-type="dwc:institutional_code" xlink:title="MNHN - Museum national d'Histoire naturelle" xlink:href="https://scientific-collections.gbif.org/institution/6a6ac6c5-1b8a-48db-91a2-f8661274ff80">MNHN</named-content></bold>), Museo Regionale di Scienze Naturali, Torino (<bold><named-content content-type="dwc:collectionCode" xlink:title="Collezioni botaniche del Museo regionale di Scienze naturali di Torino" xlink:href="https://scientific-collections.gbif.org/collection/4e693c61-4308-4b54-993d-e157bb5f9eb9">MRSN</named-content></bold>), Natural History Museum, London (<bold><named-content content-type="dwc:institutional_code" xlink:title="Natural History Museum, London" xlink:href="https://scientific-collections.gbif.org/institution/1d808a7c-1f9e-4379-9616-edb749ecf10e">NHMUK</named-content></bold>), Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main (<bold><named-content content-type="dwc:institutional_code" xlink:title="Forschungsinstitut und Naturmuseum Senckenberg" xlink:href="https://scientific-collections.gbif.org/institution/16946ec1-8db3-45b8-b084-7644384cc5f5">SMF</named-content></bold>), Université d’Antananarivo, Département de Biologie Animale (<bold><abbrev content-type="institution" xlink:title="Université d’Antananarivo, Département de Biologie Animale">UADBA</abbrev></bold>), and Zoologische Staatssammlung München (<bold><abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev></bold>). Where applicable, we provide their respective field numbers, belonging to each specimen, using the abbreviations: <bold>FGZC</bold>, <bold>FGMV</bold>, <bold>MV</bold>, and <bold>MgF</bold> referring to Frank Glaw, Miguel Vences and “Frontier, Madagascar” field numbers (cf. Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>). Some specimens deposited at <abbrev content-type="institution" xlink:title="Université d’Antananarivo, Département de Biologie Animale">UADBA</abbrev> have not yet been formally catalogued in that collection, and we refer to these specimens as <abbrev content-type="institution" xlink:title="Université d’Antananarivo, Département de Biologie Animale">UADBA</abbrev> uncatalogued, followed by the respective field number (written on a tag physically attached to the specimens) to allow for unambiguous specimen identification. Abbreviations used for morphological traits are the following: <bold>F</bold>: frontal, <bold><abbrev xlink:title="frontonasal">FN</abbrev></bold>: frontonasal, <bold><abbrev xlink:title="infralabial">IL</abbrev></bold>: infralabial, <bold><abbrev xlink:title="interparietal">IP</abbrev></bold>: interparietal, <bold>L</bold>: loreal, <bold>M</bold>: mental, <bold>N</bold>: enlarged nuchal, <bold>O</bold>: ocular, <bold>P</bold>: parietal, <bold><abbrev xlink:title="postnasal">PN</abbrev></bold>: postnasal, <bold><abbrev xlink:title="preocular">PO</abbrev></bold>: preocular, <bold><abbrev xlink:title="postsubocular">PoS</abbrev></bold>: postsubocular, <bold><abbrev xlink:title="presubocular">PrS</abbrev></bold>: presubocular, <bold><abbrev xlink:title="pretemporal">PT</abbrev></bold>: pretemporal, <bold><abbrev xlink:title="post-supralabial,">PS</abbrev></bold>: post-supralabial, <bold>R</bold>: rostral shield, <bold><abbrev xlink:title="supralabial">SL</abbrev></bold>: supralabial, <bold><abbrev xlink:title="supranasal">SN</abbrev></bold>: supranasal, <bold><abbrev xlink:title="supraocular">SO</abbrev></bold>: supraocular, <bold><abbrev xlink:title="snout–vent length">SVL</abbrev></bold>: snout–vent length, <bold>1</bold> and <bold>2</bold>: primary and secondary temporals. Geographic regions within Madagascar are named according to <xref ref-type="bibr" rid="B7">Boumans et al. (2007)</xref> and <xref ref-type="bibr" rid="B8">Brown et al. (2016)</xref>.</p>
      </sec>
      <sec sec-type="Morphological characters" id="sec4">
        <title>Morphological characters</title>
        <p>The description of the two new species herein proposed involved recording meristic, mensural and categorical morphological characters routinely used in the taxonomy of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scincidae">Scincidae</tp:taxon-name-part></tp:taxon-name>, such as scale counts, presence or absence of homologous scale fusions, or color patterns (cf. <xref ref-type="bibr" rid="B2">Andreone and Greer 2002</xref>; <xref ref-type="bibr" rid="B24">Miralles et al. 2011a</xref>, <xref ref-type="bibr" rid="B26">2011b</xref>, <xref ref-type="bibr" rid="B27">2011c</xref>, <xref ref-type="bibr" rid="B25">2016</xref>). Ventral scales are counted in a single row from the postmental scales to the preanal scales, which both are included in the count, while the mental scale is excluded. The paravertebral scales are counted in a single row, starting from the first scale located immediately behind a line drawn perpendicular to the longitudinal axis of the body and connecting the posterior edges of the thighs, and continuing forward up to and including the nuchal scales. Nuchal scales (sensu <xref ref-type="bibr" rid="B21">Miralles 2006</xref>) are defined as enlarged scales of the nape, occupying transversally the place of two or more rows of dorsal cycloid scale. The frontal scale is considered hourglass-shaped when constricted by the first supraocular, bell-shaped otherwise. Measurements of specimens were recorded to the nearest 0.1 mm using a dial caliper. Ranges are given for each meristic and mensural character, followed by the mean ± the standard deviation, with sample size in parentheses. For some bilateral characters, the sample size has been noted as the number of sides rather than specimens (indicated after sample size).</p>
      </sec>
      <sec sec-type="Molecular analysis" id="sec5">
        <title>Molecular analysis</title>
        <p>The de novo molecular dataset consists of eight sequences produced by Sanger sequencing for two individuals of the two new species under description. These sequences were combined with the extensive multilocus dataset previously published by <xref ref-type="bibr" rid="B29">Miralles and Vences (2013)</xref>, which comprises six loci: two mitochondrial genes (<abbrev xlink:title="16S rRNA,">16S</abbrev> rRNA, <bold><abbrev xlink:title="16S rRNA,">16S</abbrev></bold>; and NADH Dehydrogenase Subunit 1, <bold><abbrev xlink:title="NADH Dehydrogenase Subunit 1">ND1</abbrev></bold>) and four nuclear-encoded protein-coding genes (Brain-derived Neurotrophic Factor, <bold><abbrev xlink:title="Brain-derived Neurotrophic Factor">BDNF</abbrev></bold>; Recombination-Activating Gene 2, <bold><abbrev xlink:title="Recombination-Activating Gene 2">RAG2</abbrev></bold>; Oocyte Maturation factor mos, <bold><abbrev xlink:title="Oocyte Maturation factor mos">CMOS</abbrev></bold>; Phosducin, <bold><abbrev xlink:title="Phosducin">PDC</abbrev></bold>). See <xref ref-type="bibr" rid="B29">Miralles and Vences (2013)</xref> for detailed descriptions of all the molecular procedures including primers and PCR protocols. We used CodonCode Aligner 6.0.2 (CodonCode Corporation) to verify sequence quality of chromatograms and eliminate regions of poor read quality. New sequences were submitted to GenBank (accession numbers <ext-link ext-link-type="gen" xlink:href="PX853109" xlink:type="simple">PX853109</ext-link> to <ext-link ext-link-type="gen" xlink:href="PX853114" xlink:type="simple">PX853114</ext-link>, and <ext-link ext-link-type="gen" xlink:href="PX854599" xlink:type="simple">PX854599</ext-link> to <ext-link ext-link-type="gen" xlink:href="PX854600" xlink:type="simple">PX854600</ext-link>). The resulting data matrix is 98.5% complete, missing only 15 sequences from a possible total of 966; see File S2 for a detailed list of samples and GenBank accession numbers.</p>
        <p>In total, three Bayesian inference (<bold><abbrev xlink:title="Bayesian inference">BI</abbrev></bold>) phylogenetic analyses were carried out: (1) the main phylogenetic analysis involved the whole concatenated molecular dataset and was carried out using MrBayes 3.1.2 (<xref ref-type="bibr" rid="B36">Ronquist and Huelsenbeck 2003</xref>). Models of evolution were determined for each gene by AIC in MrModeltest 2.3 (<xref ref-type="bibr" rid="B30">Nylander 2004</xref>): GTR+G+I for <abbrev xlink:title="NADH Dehydrogenase Subunit 1">ND1</abbrev> and <abbrev xlink:title="16S rRNA,">16S</abbrev> rRNA, GTR+G for <abbrev xlink:title="Oocyte Maturation factor mos">CMOS</abbrev>, K80+I for <abbrev xlink:title="Brain-derived Neurotrophic Factor">BDNF</abbrev>, K80+G for <abbrev xlink:title="Phosducin">PDC</abbrev>, and HKY for <abbrev xlink:title="Recombination-Activating Gene 2">RAG2</abbrev>. We performed one run of 10 million generations (started on random trees) and four incrementally heated Markov chains (using default heating values) each, sampling the Markov chains at intervals of 1000 generations. The first 2.5 million generations were conservatively discarded and 2500 trees were retained post burn-in and summed to generate a 50%-majority rule consensus tree. As hierarchical outgroups, we used <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Flexiseps">Flexiseps</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="meva">meva</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paracontias">Paracontias</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="fasika">fasika</tp:taxon-name-part></tp:taxon-name></italic>, the latter belonging to the sister clade of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B11">Erens et al. 2017</xref>; <xref ref-type="bibr" rid="B3">Belluardo et al. 2023</xref>). Additionally, two complementary Bayesian analyses were carried out to detect possible mitonuclear phylogenetic discordance: (2) a <abbrev xlink:title="Bayesian inference">BI</abbrev> combining only the four concatenated genes of the nDNA data set, and (3) another one combining only the two concatenated genes of the mtDNA data set. We relied on divergence estimates from already published time-calibrated phylogenetic trees (<xref ref-type="bibr" rid="B23">Miralles et al. 2015</xref>; <xref ref-type="bibr" rid="B3">Belluardo et al. 2023</xref>; <xref ref-type="bibr" rid="B28">Miralles et al. 2025</xref>).</p>
      </sec>
    </sec>
    <sec sec-type="Results" id="sec6">
      <title>Results</title>
      <sec sec-type="Molecular analyses" id="sec7">
        <title>Molecular analyses</title>
        <p>The three Bayesian phylogenetic inferences (mtDNA, nDNA and mtDNA+nDNA) were congruent with each other and with previous studies focusing on <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic>, in recovering five main clades which correspond to: (1) the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group (i.e., composed of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="polleni">polleni</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="miafina">miafina</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="arenicola">arenicola</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stumpffi">stumpffi</tp:taxon-name-part></tp:taxon-name></italic>), (2) the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">igneocaudatus</tp:taxon-name-part></tp:taxon-name> group (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="igneocaudatus">igneocaudatus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pyrurus">pyrurus</tp:taxon-name-part></tp:taxon-name></italic>), (3) <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mouroundavae">mouroundavae</tp:taxon-name-part></tp:taxon-name></italic>, (4) the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> group (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="melanopleura">melanopleura</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minutus">minutus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ankodabensis">ankodabensis</tp:taxon-name-part></tp:taxon-name></italic>) and (5) the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">nanus</tp:taxon-name-part></tp:taxon-name> group. In the same line, they also recovered the monophyly of a larger clade gathering the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">igneocaudatus</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">mouroundavae</tp:taxon-name-part></tp:taxon-name> groups together, and the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">nanus</tp:taxon-name-part></tp:taxon-name> group as the most basally connected branch of the genus tree (i.e., sister lineage to the clade comprising all the other species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> (Figs <xref ref-type="fig" rid="F1">1</xref>, <xref ref-type="fig" rid="F2">2A</xref>)).</p>
        <fig id="F1">
          <object-id content-type="doi">10.3897/vz.76.e176241.figure1</object-id>
          <object-id content-type="arpha">03714DFB-1718-57BF-8062-2385DF17E311</object-id>
          <label>Figure 1.</label>
          <caption>
            <p>Phylogenetic tree obtained by Bayesian Inference, based on the concatenated dataset (<abbrev xlink:title="Brain-derived Neurotrophic Factor">BDNF</abbrev>, <abbrev xlink:title="Oocyte Maturation factor mos">CMOS</abbrev>, <abbrev xlink:title="Phosducin">PDC</abbrev>, <abbrev xlink:title="Recombination-Activating Gene 2">RAG2</abbrev>, <abbrev xlink:title="NADH Dehydrogenase Subunit 1">ND1</abbrev> and <abbrev xlink:title="16S rRNA,">16S</abbrev>). The molecular dating of the divergence (1) between the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group and the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">igneocaudatus</tp:taxon-name-part></tp:taxon-name> group, (2) at the base of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group, (3) between the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> group and its sister clade, and (4) at the base of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> clade was added based on <xref ref-type="bibr" rid="B23">Miralles et al. (2015)</xref> and <xref ref-type="bibr" rid="B3">Belluardo et al. (2023)</xref>.</p>
          </caption>
          <graphic xlink:href="vertebrate-zoology-76-135-g001.jpg" id="oo_1556038.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1556038</uri>
          </graphic>
        </fig>
        <fig id="F2">
          <object-id content-type="doi">10.3897/vz.76.e176241.figure2</object-id>
          <object-id content-type="arpha">5811F6B6-2309-50D9-B0DF-4E0FBB6E935D</object-id>
          <label>Figure 2.</label>
          <caption>
            <p>Molecular and morphological diversification within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic>. <bold>A</bold> Comparison of the phylogenetic trees based on nDNA (<abbrev xlink:title="Brain-derived Neurotrophic Factor">BDNF</abbrev>, <abbrev xlink:title="Oocyte Maturation factor mos">CMOS</abbrev>, <abbrev xlink:title="Phosducin">PDC</abbrev> and <abbrev xlink:title="Recombination-Activating Gene 2">RAG2</abbrev>) and mtDNA (<abbrev xlink:title="NADH Dehydrogenase Subunit 1">ND1</abbrev> and <abbrev xlink:title="16S rRNA,">16S</abbrev>). <bold>B</bold>–<bold>D</bold> Graphical diagnosis of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. displaying various morphospaces based on the ranges of intraspecific variation (min–max) of four traits: <bold>B</bold> Number of ventral scales and scale counts around midbody. <bold>C</bold> Number of subdigital lamellae on fourth fingers and fourth toes. <bold>D</bold> Comparison of the snout–vent length (<abbrev xlink:title="snout–vent length">SVL</abbrev>) variation for each species. Juvenile specimens have been removed, by using an arbitrary threshold (only specimens with <abbrev xlink:title="snout–vent length">SVL</abbrev> ≥ 50% of the maximum <abbrev xlink:title="snout–vent length">SVL</abbrev> value measured for each species were included).</p>
          </caption>
          <graphic xlink:href="vertebrate-zoology-76-135-g002.jpg" id="oo_1556039.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1556039</uri>
          </graphic>
        </fig>
        <p>In every analysis, the trees congruently retrieved the new lineages from Nosy Be and Namoroka as long branches deeply nested within the genus. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. was retrieved, with relatively good support in both the combined and the mtDNA tree, as sister species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group (<abbrev xlink:title="Bayesian inference">BI</abbrev>: 0.91 and 0.95, respectively). In contrast, its position is less resolved in the nDNA tree, as it is connected (with a weak support of 0.53) at a polytomous node, together with the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group, the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">igneocaudatus</tp:taxon-name-part></tp:taxon-name> group, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mouroundavae">mouroundavae</tp:taxon-name-part></tp:taxon-name></italic>. The uncorrected p-distances calculated between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. and the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group ranged from 7.4% to 9.3% for the <abbrev xlink:title="16S rRNA,">16S</abbrev> gene and from 16.4% to 18.5% for the <abbrev xlink:title="NADH Dehydrogenase Subunit 1">ND1</abbrev> gene. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. was congruently retrieved in all analyses and with strong support (<abbrev xlink:title="Bayesian inference">BI</abbrev>: 1.0 in both the all-marker tree and in the mtDNA tree, 0.93 in the nDNA tree) as sister species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> group. The uncorrected p-distances calculated between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. and the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> group ranged from 9.0% to 10.9% for the <abbrev xlink:title="16S rRNA,">16S</abbrev> gene and from 11.9% to 15.9% for the <abbrev xlink:title="NADH Dehydrogenase Subunit 1">ND1</abbrev> gene.</p>
        <p>According to the topology obtained from the analysis of the all-marker dataset, it can be deduced that the ancestral lineage of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. began to diverge after the differentiation between the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group and the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">igneocaudatus</tp:taxon-name-part></tp:taxon-name> group, and before the diversification within the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group (Fig. <xref ref-type="fig" rid="F1">1</xref>). This represents an interval ranging between 26.9/24.6 million years (My) and 11.2/8.8 My according to <xref ref-type="bibr" rid="B3">Belluardo et al. 2023</xref> and <xref ref-type="bibr" rid="B23">Miralles et al. 2015</xref>, respectively, which corresponds to the first half of the Miocene. Following the same rationale, the ancestral lineage of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. began to diverge after the divergence of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> group lineage, but before the diversification within that same group. This represents an interval ranging between 33.9/27.7 My and 24.9/16.0 My (also according to <xref ref-type="bibr" rid="B3">Belluardo et al. 2023</xref> and <xref ref-type="bibr" rid="B23">Miralles et al. 2015</xref>, respectively) which corresponds to the Oligocene.</p>
      </sec>
      <sec sec-type="Morphological results" id="sec8">
        <title>Morphological results</title>
        <p>Morphological results are summarized in Table <xref ref-type="table" rid="T1">1</xref>, and the most diagnostic traits are graphically represented in Figure <xref ref-type="fig" rid="F2">2B</xref><bold>–</bold>D. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. are both unambiguously diagnosable from all species of their respective sister clades (i.e., species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group and species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> group, respectively). More precisely, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. is distinguishable from all the species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group (but also from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mouroundavae">mouroundavae</tp:taxon-name-part></tp:taxon-name></italic> and species of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">igneocaudatus</tp:taxon-name-part></tp:taxon-name></italic> group) by a lower number of ventral scales, a dark and uniform colouration, a significantly smaller size and a very distinct head shape, characterized by a rather short and pointed snout and large eyes. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. differs from the species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> group by a higher number of ventral scales and a particular colouration pattern (see species diagnoses for more details).</p>
        <table-wrap id="T1" position="float" orientation="portrait">
          <label>Table 1.</label>
          <caption>
            <p>Comparison of the most relevant morphological characters in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> species. Ranges are given for meristic and mensural characters, followed by the mean ± the standard deviation, with sample size in parentheses. For bilateral characters, the sample size was noted as the number of sides rather than specimens. Data from <xref ref-type="bibr" rid="B2">Andreone and Greer (2002)</xref>, <xref ref-type="bibr" rid="B14">Glaw and Vences (2007)</xref>, Miralles et al. (<xref ref-type="bibr" rid="B24">2011a</xref>, <xref ref-type="bibr" rid="B25">2016</xref>), <xref ref-type="bibr" rid="B29">Miralles and Vences (2013)</xref>, <xref ref-type="bibr" rid="B33">Porcel et al. (2021)</xref>. Spec. <bold>–</bold> spectacled; vivi. <bold>–</bold> viviparous; ovi. – oviparous.</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <th rowspan="1" colspan="1"/>
                <th rowspan="1" colspan="1"/>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">igneocaudatus</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">pyrurus</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">minutus</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">ankodabensis</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">mouroundavae</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">nanus</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                  <italic/>
                  <bold>group</bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">arenicola</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">miafina</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1">
                  <bold>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/>
                      <tp:taxon-name-part taxon-name-part-type="species">stumpffi</tp:taxon-name-part>
                    </tp:taxon-name>
                  </bold>
                </th>
                <th rowspan="1" colspan="1"><bold><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">minotaurus</tp:taxon-name-part></tp:taxon-name> sp. nov</bold>.</th>
                <th rowspan="1" colspan="1"><bold><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">irery</tp:taxon-name-part></tp:taxon-name> sp. nov</bold>.</th>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">N lamellae under 4<sup>th</sup> finger</td>
                <td rowspan="1" colspan="1">min–max: mean ± SD: n sides:</td>
                <td rowspan="1" colspan="1">8–11 9.0 ± 0.9 (50)</td>
                <td rowspan="1" colspan="1">8–11 9.1 ± 0.9 (14)</td>
                <td rowspan="1" colspan="1">5–7 6.0 ± 0.8 (13)</td>
                <td rowspan="1" colspan="1">5–8 7.0 ± 0.6 (40)</td>
                <td rowspan="1" colspan="1">5–8 6.3 ± 1.0 (9)</td>
                <td rowspan="1" colspan="1">8–11 9.8 ± 0.7 (16)</td>
                <td rowspan="1" colspan="1">3–5 3.9 ± 0.6 (8)</td>
                <td rowspan="1" colspan="1">6–7 6.4 ± 0.5 (11)</td>
                <td rowspan="1" colspan="1">7–8 7.6 ± 0.5 (22)</td>
                <td rowspan="1" colspan="1">6–9 7.5 ± 0.7 (26)</td>
                <td rowspan="1" colspan="1">6–9 7.3 ± 0.8 (27)</td>
                <td rowspan="1" colspan="1">7 — (1)</td>
                <td rowspan="1" colspan="1">8 8.0± 0.0 (3)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">N lamellae under 4<sup>th</sup> toe</td>
                <td rowspan="1" colspan="1">min–max: mean ± SD: n sides:</td>
                <td rowspan="1" colspan="1">15–22 18.1 ± 1.4 (54)</td>
                <td rowspan="1" colspan="1">15–18 16.4 ± 1.3 (13)</td>
                <td rowspan="1" colspan="1">9–13 11.3 ± 1.5 (10)</td>
                <td rowspan="1" colspan="1">12–16 14.1 ± 1.2 (52)</td>
                <td rowspan="1" colspan="1">12–15 13.8 ± 1.1 (12)</td>
                <td rowspan="1" colspan="1">16–20 17.5 ± 1.2 (15)</td>
                <td rowspan="1" colspan="1">5–8 6.8 ± 1.3 (9)</td>
                <td rowspan="1" colspan="1">16–19 17.5 ± 0.8 (13)</td>
                <td rowspan="1" colspan="1">18–23 20.6 ± 1.3 (20)</td>
                <td rowspan="1" colspan="1">16–22 18.5 ± 1.5 (22)</td>
                <td rowspan="1" colspan="1">15–20 17.9 ± 1.2 (28)</td>
                <td rowspan="1" colspan="1">15 — (1)</td>
                <td rowspan="1" colspan="1">12–15 13.3 ± 1.5 (3)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">N ventral scale rows</td>
                <td rowspan="1" colspan="1">min–max: mean ± SD: n:</td>
                <td rowspan="1" colspan="1">68–83 76.7 ± 4.4 (21)</td>
                <td rowspan="1" colspan="1">73–78 75.7 ± 1.8 (7)</td>
                <td rowspan="1" colspan="1">55–63 58.3 ± 3.0 (7)</td>
                <td rowspan="1" colspan="1">56–61 58.8 ± 1.2 (27)</td>
                <td rowspan="1" colspan="1">59–63 60.2 ± 1.5 (6)</td>
                <td rowspan="1" colspan="1">63–66 64.3 ± 1.0 (8)</td>
                <td rowspan="1" colspan="1">52–60 57.6 ± 3.3 (5)</td>
                <td rowspan="1" colspan="1">75–80 77.9 ± 1.6 (7)</td>
                <td rowspan="1" colspan="1">65–73 68.7 ± 2.1 (14)</td>
                <td rowspan="1" colspan="1">74–78 75.8 ± 1.2 (12)</td>
                <td rowspan="1" colspan="1">70–88 81. 3 ± 4.0 (16)</td>
                <td rowspan="1" colspan="1">68 — (1)</td>
                <td rowspan="1" colspan="1">55 55.0 ± 0.0 (2)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">N paravertebral scale rows</td>
                <td rowspan="1" colspan="1">min–max: mean ± SD: n:</td>
                <td rowspan="1" colspan="1">69–80 74.7 ± 3.0 (26)</td>
                <td rowspan="1" colspan="1">71–79 74.6 ± 3.7 (7)</td>
                <td rowspan="1" colspan="1">57–65 59.7 ± 3.4 (7)</td>
                <td rowspan="1" colspan="1">51–62 55.9 ± 2.9 (28)</td>
                <td rowspan="1" colspan="1">52–62 57.7 ± 3.1 (7)</td>
                <td rowspan="1" colspan="1">60–65 62.6 ± 2.1 (8) –</td>
                <td rowspan="1" colspan="1">50–57 53.6 ± 2.5 (5)</td>
                <td rowspan="1" colspan="1">74–81 79.0 ± 2.3 (7)</td>
                <td rowspan="1" colspan="1">65–79 68.7 ± 3.3 (14)</td>
                <td rowspan="1" colspan="1">71–81 77.9 ± 2.6 (13)</td>
                <td rowspan="1" colspan="1">76–88 82.7 ± 3.2 (15)</td>
                <td rowspan="1" colspan="1">69 — (1)</td>
                <td rowspan="1" colspan="1">53–56 54.5 ± 2.12 (2)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">N longitudinal scale rows at midbody</td>
                <td rowspan="1" colspan="1">min–max: mean ± SD: n:</td>
                <td rowspan="1" colspan="1">24–26 24.2 ± 0.6 (28)</td>
                <td rowspan="1" colspan="1">22–24 23.3 ± 1.0 (7)</td>
                <td rowspan="1" colspan="1">22–26 24.0 ± 1.2 (7)</td>
                <td rowspan="1" colspan="1">24–26 24.1 ± 0.4 (27)</td>
                <td rowspan="1" colspan="1">22–26 23.7 ± 1.5 (6)</td>
                <td rowspan="1" colspan="1">28–30 29.0 ±4 2.1 (8)</td>
                <td rowspan="1" colspan="1">18–20 19.6 ± 0.9 (5)</td>
                <td rowspan="1" colspan="1">26 26.0 ± 0 (7)</td>
                <td rowspan="1" colspan="1">24–26 24.1 ± 0.5 (14)</td>
                <td rowspan="1" colspan="1">24–26 25.4 ± 0.9 (13)</td>
                <td rowspan="1" colspan="1">30–32 31.6 ± 0.8 (16)</td>
                <td rowspan="1" colspan="1">20 — (1)</td>
                <td rowspan="1" colspan="1">24 24.0 ± 0.0 (2)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Enlarged nuchal scales</td>
                <td rowspan="1" colspan="1">absent: one row: two rows: three rows: four rows: n sides:</td>
                <td rowspan="1" colspan="1">— — 23.2% 71.4% 5.4% (56)</td>
                <td rowspan="1" colspan="1">— — 21.4% 78.6% — (14)</td>
                <td rowspan="1" colspan="1">— — 28.6% 57.1% 14.3% (14)</td>
                <td rowspan="1" colspan="1">— 2% 50% 48% — (58)</td>
                <td rowspan="1" colspan="1">— 7.1% 35.8% 57.1% — (14)</td>
                <td rowspan="1" colspan="1">— 100% — — — (16)</td>
                <td rowspan="1" colspan="1">— 40.0% 20.0% 40.0% — (10)</td>
                <td rowspan="1" colspan="1">42.9% 57.1% — — — (14)</td>
                <td rowspan="1" colspan="1">92.3% 7.7% — — — (26)</td>
                <td rowspan="1" colspan="1">56.3% 37.5% 6.2% — — (22)</td>
                <td rowspan="1" colspan="1">81.3% 18.7% — — — (32)</td>
                <td rowspan="1" colspan="1">— — 100% — — (2)</td>
                <td rowspan="1" colspan="1">100% — — — — (4)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Postnasal</td>
                <td rowspan="1" colspan="1">present: absent: n sides:</td>
                <td rowspan="1" colspan="1">100% — (56)</td>
                <td rowspan="1" colspan="1">100% — (14)</td>
                <td rowspan="1" colspan="1">100% — (14)</td>
                <td rowspan="1" colspan="1">100% — (58)</td>
                <td rowspan="1" colspan="1">100% — (14)</td>
                <td rowspan="1" colspan="1">100% — (16)</td>
                <td rowspan="1" colspan="1">100% — (10)</td>
                <td rowspan="1" colspan="1">— 100% (14)</td>
                <td rowspan="1" colspan="1">89.3% 10.7% (28)</td>
                <td rowspan="1" colspan="1">100% — (26)</td>
                <td rowspan="1" colspan="1">94.4% 5.6% (36)</td>
                <td rowspan="1" colspan="1">100% — (2)</td>
                <td rowspan="1" colspan="1">100% — (4)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Frontal and interparietal</td>
                <td rowspan="1" colspan="1">fused: separated: n:</td>
                <td rowspan="1" colspan="1">— 100% (28)</td>
                <td rowspan="1" colspan="1">— 100% (14)</td>
                <td rowspan="1" colspan="1">— 100% (7)</td>
                <td rowspan="1" colspan="1">— 100% (28)</td>
                <td rowspan="1" colspan="1">— 100% (7)</td>
                <td rowspan="1" colspan="1">87.5% 12.5% (8)</td>
                <td rowspan="1" colspan="1">— 100% (10)</td>
                <td rowspan="1" colspan="1">— 100% (7)</td>
                <td rowspan="1" colspan="1">— 100% (14)</td>
                <td rowspan="1" colspan="1">— 100% (13)</td>
                <td rowspan="1" colspan="1">— 100% (16)</td>
                <td rowspan="1" colspan="1">— 100% (1)</td>
                <td rowspan="1" colspan="1">— 100% (1)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Frontal</td>
                <td rowspan="1" colspan="1">bell-shaped: hour–glass shaped: n:</td>
                <td rowspan="1" colspan="1">100% — — (23)</td>
                <td rowspan="1" colspan="1">100% — — (12)</td>
                <td rowspan="1" colspan="1">— 100% — (14)</td>
                <td rowspan="1" colspan="1">— 100% — (29)</td>
                <td rowspan="1" colspan="1">— 100% — (14)</td>
                <td rowspan="1" colspan="1">— 100% — (8)</td>
                <td rowspan="1" colspan="1">— 100% — (10)</td>
                <td rowspan="1" colspan="1">100% — — (7)</td>
                <td rowspan="1" colspan="1">100% — — (14)</td>
                <td rowspan="1" colspan="1">100% — — (13)</td>
                <td rowspan="1" colspan="1">47.2% 52.8% — (18)</td>
                <td rowspan="1" colspan="1">100% — — (1)</td>
                <td rowspan="1" colspan="1">— 100% — (2)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Snout–vent length (mm)</td>
                <td rowspan="1" colspan="1">max: mean ± SD: n:</td>
                <td rowspan="1" colspan="1">73.0 56.3 ± 11.6 (9)</td>
                <td rowspan="1" colspan="1">54.2 52.3 ± 2.1 (4)</td>
                <td rowspan="1" colspan="1">47.4 42.0 ± 5.1 (7)</td>
                <td rowspan="1" colspan="1">53.5 49.5±2.5 (21)</td>
                <td rowspan="1" colspan="1">50.5 48.0 ± 2.4 (5)</td>
                <td rowspan="1" colspan="1">68.5 60.1 ± 9.6 (7)</td>
                <td rowspan="1" colspan="1">33.6 27.8 ± 8.2 (6)</td>
                <td rowspan="1" colspan="1">81.7 72.3 ± 6.1 (7)</td>
                <td rowspan="1" colspan="1">61 54.9 ± 3.1 (14)</td>
                <td rowspan="1" colspan="1">75 66.0 ± 7.1 (13)</td>
                <td rowspan="1" colspan="1">114.0 89.6 ± 10.8 (14)</td>
                <td rowspan="1" colspan="1">39.9 — (1)</td>
                <td rowspan="1" colspan="1">39.9 36.9 ± 4.1 (2)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Supraciliaries</td>
                <td rowspan="1" colspan="1">five: six: seven: eight: n sides:</td>
                <td rowspan="1" colspan="1">— 98.2% 1.8% — (56)</td>
                <td rowspan="1" colspan="1">7.1% 92.9% — — (14)</td>
                <td rowspan="1" colspan="1">7.1% 57.1% 35.8% (14)</td>
                <td rowspan="1" colspan="1">— 8.6% 79.3% 12.7% (58)</td>
                <td rowspan="1" colspan="1">— 35.7% 50.0% 14.3% (14)</td>
                <td rowspan="1" colspan="1">— 93.8% 6.2% — (16)</td>
                <td rowspan="1" colspan="1">— 66.6% 33.3% — (6)</td>
                <td rowspan="1" colspan="1">— 92.9% 7.1% — (14)</td>
                <td rowspan="1" colspan="1">— 100% — — (2)</td>
                <td rowspan="1" colspan="1">— 87.5% 12.5% — (8)</td>
                <td rowspan="1" colspan="1"> n/a</td>
                <td rowspan="1" colspan="1">— 100% — — (2)</td>
                <td rowspan="1" colspan="1">— 100% — — (2)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Subocular</td>
                <td rowspan="1" colspan="1">third <abbrev xlink:title="supralabial">SL</abbrev>: fourth <abbrev xlink:title="supralabial">SL</abbrev>: n sides:</td>
                <td rowspan="1" colspan="1">1.8% 98.2% (56)</td>
                <td rowspan="1" colspan="1">— 100% (14)</td>
                <td rowspan="1" colspan="1">— 100% (14)</td>
                <td rowspan="1" colspan="1">— 100% (58)</td>
                <td rowspan="1" colspan="1">7.1% 92.9% (14)</td>
                <td rowspan="1" colspan="1">— 100% (24)</td>
                <td rowspan="1" colspan="1">100% — (10)</td>
                <td rowspan="1" colspan="1">— 100% (14)</td>
                <td rowspan="1" colspan="1">3.6% 96.4% (28)</td>
                <td rowspan="1" colspan="1">— 100% (24)</td>
                <td rowspan="1" colspan="1">— 100% (32)</td>
                <td rowspan="1" colspan="1">— 100% (2)</td>
                <td rowspan="1" colspan="1">— 100% (4)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Lower eyelid window</td>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1">spec.</td>
                <td rowspan="1" colspan="1">spec.</td>
                <td rowspan="1" colspan="1">spec.</td>
                <td rowspan="1" colspan="1">spec.</td>
                <td rowspan="1" colspan="1">spec.</td>
                <td rowspan="1" colspan="1">scaly</td>
                <td rowspan="1" colspan="1">scaly</td>
                <td rowspan="1" colspan="1">scaly</td>
                <td rowspan="1" colspan="1">scaly</td>
                <td rowspan="1" colspan="1">scaly</td>
                <td rowspan="1" colspan="1">scaly</td>
                <td rowspan="1" colspan="1">spec.</td>
                <td rowspan="1" colspan="1">scaly</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Reproduction*</td>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1">vivi.</td>
                <td rowspan="1" colspan="1">ovi.</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">ovi.</td>
                <td rowspan="1" colspan="1">ovi.</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">?</td>
                <td rowspan="1" colspan="1">ovi?</td>
                <td rowspan="1" colspan="1">?</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">Elevational range</td>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1">≤ 500 m</td>
                <td rowspan="1" colspan="1">≥ 1500 m</td>
                <td rowspan="1" colspan="1">≤ 1000 m</td>
                <td rowspan="1" colspan="1">≤ 1000 m</td>
                <td rowspan="1" colspan="1">≤ 1000 m</td>
                <td rowspan="1" colspan="1">≤ 1000</td>
                <td rowspan="1" colspan="1">500–1500 m</td>
                <td rowspan="1" colspan="1"><bold>≤</bold> 500 m</td>
                <td rowspan="1" colspan="1">≤ 500 m</td>
                <td rowspan="1" colspan="1">≤ 500 m</td>
                <td rowspan="1" colspan="1">≤ 500 m</td>
                <td rowspan="1" colspan="1">≤ 500 m</td>
                <td rowspan="1" colspan="1">≤ 500 m</td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
        <p>This consistent morphological differentiation, in concert with isolated phylogenetic positions and high genetic divergences of the two new lineages unambiguously support their species status, and we therefore formally name and describe them in the following.</p>
      </sec>
    </sec>
    <sec sec-type="Taxonomy" id="sec9">
      <title>Taxonomy</title>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order">Squamata</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family">Scincidae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <tp:taxon-name><object-id content-type="arpha">46AE6986-14A2-5638-AA4F-42F54C767A45</object-id>
                		<tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part>
                	
                		<object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/02B6EDA4-CC86-4BA9-9720-AADCC7622D1E</object-id>
                	</tp:taxon-name>
          <tp:taxon-status>sp. nov.</tp:taxon-status>
          <xref ref-type="fig" rid="F3">Figures 3</xref>
          <xref ref-type="fig" rid="F4">, 4</xref>
          <xref ref-type="fig" rid="F5">, 5</xref>
          <xref ref-type="fig" rid="F6">, 6</xref>
          <xref ref-type="fig" rid="F7">, 7</xref>
          <xref ref-type="fig" rid="F8">, 8</xref>
          <xref ref-type="fig" rid="F9">, 9</xref>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="Holotype">
          <title>Holotype.</title>
          <p>MADAGASCAR – <bold>Boeny Region</bold> • 1 adult ♀; Tsingy de Namoroka National Park, Petit Tsingy; <named-content content-type="dwc:verbatimCoordinates">16°26’7”S, 45°22’6”E</named-content>; 125 m a.s.l.; 08 Oct. 2023, 07:00–09:00 a.m.; A. Miralles, N.A. Rahagalala, A. Rakotoarison, D. Razafimanafo and A. Razafimanantsoa leg.; <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 116/2023 (ZCMV 15819).</p>
          <fig id="F3">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure3</object-id>
            <object-id content-type="arpha">A7813430-FA0C-55BC-9EC2-A007AB422972</object-id>
            <label>Figure 3.</label>
            <caption>
              <p>Photos of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. in life. <bold>A</bold>, <bold>B</bold> Holotype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 116/2023, from Tsingy de Namoroka National Park. <bold>C</bold> Uncollected specimen from Grand Tsingy. Photographs A and B by AM, C by Ivan Ineich.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g003.jpg" id="oo_1556040.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556040</uri>
            </graphic>
          </fig>
          <fig id="F4">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure4</object-id>
            <object-id content-type="arpha">A957992C-FB91-56A3-8EF4-FB2F98F1883D</object-id>
            <label>Figure 4.</label>
            <caption>
              <p>Preserved holotype of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 116/2023, from Tsingy de Namoroka National Park. <bold>A</bold>, <bold>B</bold> photographs and <bold>C</bold> head drawings in lateral (above) and dorsal view (below). Abbreviations used in the drawings are explained in the Materials and Methods section.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g004.jpg" id="oo_1556041.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556041</uri>
            </graphic>
          </fig>
          <fig id="F5">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure5</object-id>
            <object-id content-type="arpha">788C15E6-FA28-5F8F-95EA-17543FCF278B</object-id>
            <label>Figure 5.</label>
            <caption>
              <p>Habitat of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. at Tsingy de National Park. <bold>A</bold> View of the Grand Tsingy from the surrounding savanna. <bold>B</bold> View from inside a canyon at the Grand Tsingy. <bold>C</bold> Campsite at the border of Grand Tsingy, showing the leaf litter micro-habitat where the uncollected specimen (sample MIRZC 1217) was found. <bold>D</bold> Petit Tsingy, exact position of the type locality. <bold>E</bold> Diagram representing a transverse view of Grand Tsingy during the dry season (during the wet season, the depressed ground is heavily flooded). Specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. have been encountered in the leaf-litter located in the border of the karts system (corresponding to the orange surface on the left side of the diagram), although the species is likely present in the corridors too.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g005.jpg" id="oo_1556042.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556042</uri>
            </graphic>
          </fig>
          <fig id="F6">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure6</object-id>
            <object-id content-type="arpha">6D595353-DE63-562D-9A8B-499E2947B549</object-id>
            <label>Figure 6.</label>
            <caption>
              <p>Satellite views of the Tsingy de Namoroka. <bold>A</bold> Namoroka National Park and surrounding areas. <bold>B</bold> Details of the main karstic area. <bold>C</bold>, <bold>D</bold> Details of areas are here interpreted as vestigial karstic fragments located outside of the national park. Karstic fragments (i.e., showing the typical network of canyons/fissures arranged perpendicular to each other and filled with vegetation) are delimited in white, while the perimeter of the national park is delimited in green. Turquoise dots correspond to the known localities of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. Satellite imagery from Airbus, via Google Earth.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g006.jpg" id="oo_1556043.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556043</uri>
            </graphic>
          </fig>
          <fig id="F7">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure7</object-id>
            <object-id content-type="arpha">7B176930-49AC-5385-9C11-DFCFA5C7AEAB</object-id>
            <label>Figure 7.</label>
            <caption>
              <p>Drawings of the heads in dorsal and lateral views of most of the species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic>. <bold>A</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="arenicola">arenicola</tp:taxon-name-part></tp:taxon-name></italic>, holotype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 1565/2008, Baie des Sakalava. <bold>B</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="miafina">miafina</tp:taxon-name-part></tp:taxon-name></italic>, holotype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 1562/2008, Ankarana Special Reserve. <bold>C</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stumpffi">stumpffi</tp:taxon-name-part></tp:taxon-name></italic>, holotype <named-content content-type="dwc:institutional_code" xlink:title="Forschungsinstitut und Naturmuseum Senckenberg" xlink:href="https://scientific-collections.gbif.org/institution/16946ec1-8db3-45b8-b084-7644384cc5f5">SMF</named-content> 16019, “<italic>Nossibé</italic>” (= Nosy Be). <bold>D</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="polleni">polleni</tp:taxon-name-part></tp:taxon-name></italic>, holotype <named-content content-type="dwc:institutional_code" xlink:title="MNHN - Museum national d'Histoire naturelle" xlink:href="https://scientific-collections.gbif.org/institution/6a6ac6c5-1b8a-48db-91a2-f8661274ff80">MNHN</named-content> 1895.210, “Mouroundava” (= Morondava). <bold>E</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., holotype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 135/2019, Nosy Be. <bold>F</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pyrurus">pyrurus</tp:taxon-name-part></tp:taxon-name></italic>, holotype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 520/2001, Mont Ibity. <bold>G</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="igneocaudatus">igneocaudatus</tp:taxon-name-part></tp:taxon-name></italic>, <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 1600/2010, Anakao. <bold>H</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mouroundavae">mouroundavae</tp:taxon-name-part></tp:taxon-name></italic>, <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 13/2005, Andasibe. <bold>I</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ankodabensis">ankodabensis</tp:taxon-name-part></tp:taxon-name></italic>, <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 355/2006, Ranomafana. <bold>J</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="melanopleura">melanopleura</tp:taxon-name-part></tp:taxon-name></italic>, <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 20/2005, Andasibe. <bold>K</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minutus">minutus</tp:taxon-name-part></tp:taxon-name></italic>, <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 400/2005, Nosy Mangabe. <bold>L</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., holotype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 116/2023, Namoroka. Dwarf species, namely <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nanus">nanus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="macrolepis">macrolepis</tp:taxon-name-part></tp:taxon-name></italic>, not represented. Scale bars = 2 mm.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g007.jpg" id="oo_1556044.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556044</uri>
            </graphic>
          </fig>
          <fig id="F8">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure8</object-id>
            <object-id content-type="arpha">757D556A-5670-55A8-8A9A-D61121BEBFA3</object-id>
            <label>Figure 8.</label>
            <caption>
              <p>Plate showing photographs in life of some of the recognized species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> (not to scale).</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g008.jpg" id="oo_1556045.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556045</uri>
            </graphic>
          </fig>
          <fig id="F9">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure9</object-id>
            <object-id content-type="arpha">E1465D55-399F-5B6A-9698-2A4725BCD88A</object-id>
            <label>Figure 9.</label>
            <caption>
              <p>Schematic maps for Madagascar showing known distribution for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> species. For reasons of readability, the different species are shown on six separate maps. Colored dots represent localities confirmed by DNA sequences, whereas white dots represent localities of specimens only identified by morphology (more details in <xref ref-type="bibr" rid="B25">Miralles et al. 2016</xref>). The supplementary map located at the bottom of the figure, employs minimum convex polygons to compare the distribution ranges of all species.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g009.jpg" id="oo_1556046.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556046</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Other material examined">
          <title>Other material examined.</title>
          <p>MADAGASCAR – <bold>Boeny Region</bold> • 1 tissue sample (MIRZC 1217, voucher not collected); Tsingy de Namoroka National Park, Grand Tsingy, south of the Tsingy massif, Camp 2; <named-content content-type="dwc:verbatimCoordinates">16°28’9”S, 45°20’54”E</named-content>, 128 m a.s.l.; 10 Oct. 2023, 03:00–07:00 p.m.; A. Miralles, N. A. Rahagalala, A. Rakotoarison, D. Raza­fi­manafo and A. Razafimanantsoa leg. • 1 specimen pho­to­graphed (unsampled and uncollected); Tsingy de Na­mo­roka National Park, Grand Tsingy; <named-content content-type="dwc:verbatimCoordinates">16°28’1”S, 45°21’0”E</named-content>; photographed by Ivan Ineich on 3 Sept. 2012.</p>
          <p>Furthermore, four specimens deposited in the UMMZ collection and registered as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="intermedius">intermedius</tp:taxon-name-part></tp:taxon-name></italic> (junior synonym of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="polleni">polleni</tp:taxon-name-part></tp:taxon-name></italic>) have been collected only 50 m from the sample MIRZC 1217. Given their collection locality and the superficial resemblance between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="polleni">polleni</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., we hypothesize that these specimens likely correspond to the new species: UMMZ 222106–222109, from Namoroka, camp 1, at the edge of Tsingy (<named-content content-type="dwc:verbatimCoordinates">16°28’11”S, 45°20’54”E</named-content>); Dec. 1996; C.J. Raxworthy, J.B. Ramanamanjato, A. ­Razafimanantsoa and J. Rafanomezantsoa leg. These specimens/samples are not designated as paratypes because the specimens were not studied.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Diagnosis">
          <title>Diagnosis.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. differs from all other species in the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> by the combination of a lower number of scales around midbody and a higher number of ventrals. It differs from the species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">melanopleura</tp:taxon-name-part></tp:taxon-name> group (its sister clade formed by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="melanopleura">melanopleura</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ankodabensis">ankodabensis</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minutus">minutus</tp:taxon-name-part></tp:taxon-name></italic>) by a distinctly higher number of ventrals (68, versus 55 to 63) and paravertebral scales (69, versus 51 to 65), by a bell-shaped frontal scale (versus hourglass-shaped, i.e., frontal constricted by the first pair of supraoculars) and a distinct, lighter and warmer coloration (versus darker and brownish).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Description of the holotype">
          <title>Description of the holotype.</title>
          <p>(Figs <xref ref-type="fig" rid="F3">3</xref>, <xref ref-type="fig" rid="F4">4</xref>) An adult female, with a single large egg in the abdominal cavity. Good state of preservation, with exception of the left arm and the left leg that have been removed for molecular analyses. <abbrev xlink:title="snout–vent length">SVL</abbrev> (39.9 mm) 7 times head length (5.9 mm), almost as long as tail (41.0 mm, either entire or completely regenerated). Limbs very short, <abbrev xlink:title="snout–vent length">SVL</abbrev> 7.1 times forelimb length (5.5 mm) and 4.3 times hind limb length (8.9 mm). Snout short and rounded on lateral aspect, with a rostral tip bluntly rounded in dorsal aspect. Rostral wider than long, contacting first supralabial scales, nasals, and supranasal scales. Paired supranasal scales in median contact, contacting loreals and postnasal scales. Frontonasal scales roughly triangular, wider than long, contacting loreals, first supraciliaries and first supraoculars. Prefrontals absent. Frontal longer than wide, wider posteriorly, in contact with frontonasal, supraoculars, parietals and interparietal. Four supraoculars, all of them in contact with frontal; second pair larger than posteriormost pair; first pair not constricting frontal (frontal bell-shaped sensu <xref ref-type="bibr" rid="B2">Andreone and Greer 2002</xref>). Frontoparietals absent. Interparietal as long as wide, well separated from supraoculars; parietal eyespot present with parietal eye evident. Parietals contact posterior to interparietal. Two pairs of enlarged nuchals, each nuchal scale width corresponding to two transverse rows of adjacent cycloid scales. Nasals slightly larger than nostrils; contacting rostral, first supralabials, postnasals and supranasals. Postnasals present, separating supranasals from first supralabials, and nasals from loreals. Loreal single, slightly higher than long. Preocular trapezoidal, as long as high, single. Presubocular roughly square, single. Six supraciliaries on both sides, in continuous row; first and last pairs significantly larger than the intermediate ones; last pair projecting onto supraocular shelf. Pretemporals single, contacted by parietal. Two postsuboculars; upper contacting pretemporal; both contacting penultimate supralabial. Lower eyelid moveable, with a spectacled translucent window; lower palpebrals small, longer than high. Contact between upper palpebrals and supraciliaries seemingly direct but flexible, i.e., palpebral cleft narrow. Primary temporal single. Two secondary temporals; upper long, contacting pretemporal anteriorly and overlapping lower secondary temporal ventrally. Six supralabials, with the fourth being the enlarged subocular, contacting scales of the lower eyelid. External ear opening rounded, without lobules. Mental wider than long, posterior margin convex. Postmental wider than long, contacting first two pairs of infralabials. Six infralabials. Three pairs of large chin scales, both members of first pair in contact, both members of second pair separated by a single median scale, and members of third pair separated by three scale rows. No scales extending between infralabials and large chin scales. Gulars similar in size and outline to ventrals. All scales, except head shields and scales on palms, soles, and digits, cycloid, smooth, and imbricate; 20 longitudinal scale rows at midbody; 69 paravertebrals, similar in size to adjacent scales; 68 ventrals. Inner preanals overlapped by outer. Both anterior and posterior limbs pentadactyl; fingers and toes relatively short, clawed, except the fifth finger which does not bear a claw; relative length of toes in the following order: I&lt;II&lt;III&lt;V&lt;IV. Subdigital lamellae smooth, single, with 7 under right fourth finger, 15 under right fourth toe.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Coloration of the holotype">
          <title>Coloration of the holotype.</title>
          <p>In preservative, specimen with a pair of lateral dark brown stripes (about one scale wide on the neck), well defined anteriorly (overlapping rostral, mental, first four supralabials, loreals, and presuboculars), then progressively breaking up into a thin dashed lines posteriorly to forelimbs, hardly distinguishable from the rest of the dark dots covering the dorsum and the flanks. Presence on the anterior part of the body of a pair of short light cream dorsolateral stripes, separating the dark lateral stripe from the bronze dorsal field, well-visible from supranasals, supraciliaries, temporal area and the neck to the level of the insertion of the forelimbs, but progressively fading and disappearing posteriorly. Dorsum and dorsal sides of forelimbs, hindlimbs and tail light bronze. The bronze dorsal field and the slightly lighter flanks are covered by numerous little dark dots, with each cycloid presenting a distinctive lunular at their insertion point and a cloud of tiny dark pigmentations at their middle, resulting in many thin dash lines (14 at midbody). No distinct border between the background coloration of the darker dorsal and the lighter ventral sides. Immaculate light cream ventral field extending from lower side of head (mental excluded), throat, lower side of limbs and venter, to ventral side of tail. Palms and soles greyish, darker than venter. In life, the coloration pattern was relatively similar to that in preservative, but with a slightly brighter and warmer overall aspect, evoking more a copper than bronze background coloration. Light dorsolateral stripes on the head and the neck were more discernable and extending along the entire length of the body until hindlimbs insertion and presenting a brighter orange tint. Presence of slightly iridescent glints of scales (cf. Fig. <xref ref-type="fig" rid="F3">3A, B</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Variation">
          <title>Variation.</title>
          <p>The only available additional information concerns a specimen photographed by Ivan Ineich at the Grand Tsingy; it presents a color pattern identical to that of the holotype (Fig. <xref ref-type="fig" rid="F3">3C</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Etymology">
          <title>Etymology.</title>
          <p>The specific epithet minotaurus, invariable noun in apposition, is derived from Minṓtauros (ancient Greek: Μινώταυρος), a taurine creature of the Greek mythology. It alludes to the labyrinthine structure of the Tsingy de Namoroka, within which the new species, like the Minotaur in Daedalus’s construction, appears unable to escape.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Distribution">
          <title>Distribution.</title>
          <p>(Figs <xref ref-type="fig" rid="F5">5</xref>, <xref ref-type="fig" rid="F6">6</xref>, <xref ref-type="fig" rid="F9">9</xref>) Only known from the ­Tsingy area of the Namoroka National Park, North West of Madagascar, and probably distributed all over this karstic massif and surrounding patches of deciduous forest. The holotype was collected at the border of the Petit Tsingy (eastern part of the Namoroka massif), at the limit between the rock corridor and the adjacent dry deciduous forested area. The sample MIRZC 1217 was found on the border of the Grand Tsingy (southern part of the massif).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Ecology">
          <title>Ecology.</title>
          <p>The two specimens observed in 2023 were foraging in dense leaf-litter layers in shaded areas (by day) or active at dusk.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Conservation status">
          <title>Conservation status.</title>
          <p>Considering that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. is likely widely distributed across the Namoroka karst massif, and based on satellite imagery, it is possible to conclude that its EOO might at least range from 50 to 200 km², depending on whether the species is present in peripheral karstic isolates too (cf. Fig. <xref ref-type="fig" rid="F6">6</xref>). The possibility of a broader distribution within the region for this species remains however plausible and cannot be dismissed in the absence of additional data. Although the Namoroka massif is well preserved and currently shows no significant signs of anthropogenic disturbance, the new species’ habitat is likely highly fragmented. Pending further information to confirm this species’ range, population status, and potential threats, we recommended assigning the species the status VU (Vulnerable) based on the criteria D2 of the <xref ref-type="bibr" rid="B18">IUCN Standards and Petitions Committee (2024)</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">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order">Squamata</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family">Scincidae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <tp:taxon-name><object-id content-type="arpha">7E62E052-57DD-53F0-96F0-AC92D64CC1B8</object-id>
                		<tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part>
                	
                		<object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/C416E285-B102-46BD-B085-7B89ED840C0E</object-id>
                	</tp:taxon-name>
          <tp:taxon-status>sp. nov.</tp:taxon-status>
          <xref ref-type="fig" rid="F7">Figures 7</xref>
          <xref ref-type="fig" rid="F8">, 8</xref>
          <xref ref-type="fig" rid="F9">, 9</xref>
          <xref ref-type="fig" rid="F10">, 10</xref>
          <xref ref-type="fig" rid="F11">, 11</xref>
          <xref ref-type="fig" rid="F12">, 12</xref>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="Holotype">
          <title>Holotype.</title>
          <p>MADAGASCAR – <bold>Diana Region</bold> • unsexed specimen; Nosy Be, Lokobe National Park; <named-content content-type="dwc:verbatimCoordinates">13°24’29”S, 48°18’31”E</named-content>; ca. 210 m a.s.l.; 28 Apr. 2019, ca. 12:00 a.m.; M.D. Scherz, F. Glaw, A. Razafimanantsoa and J.H. Razafindraibe leg.; <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 135/2019 (FGZC 5612).</p>
          <fig id="F10">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure10</object-id>
            <object-id content-type="arpha">F79E0D8F-3F31-516B-B5BD-4626C423487E</object-id>
            <label>Figure 10.</label>
            <caption>
              <p>Alive picture of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov. A</bold>, <bold>B</bold>, <bold>C</bold> Holotype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 135/2019, from Lokobe National Park, Nosy Be. Photographs by MDS.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g010.jpg" id="oo_1556047.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556047</uri>
            </graphic>
          </fig>
          <fig id="F11">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure11</object-id>
            <object-id content-type="arpha">3760F173-D028-54D3-A1FE-9BA0892640B6</object-id>
            <label>Figure 11.</label>
            <caption>
              <p>Preserved specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov. A</bold> Anterior body of the paratype, <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 86/2015 in dorsal view. <bold>B</bold> dorsal view of the entire body, <bold>C</bold> lateral and dorsal view of the head, <bold>D</bold> drawing of head in lateral and dorsal views of holotype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 135/2019 from Lokobe National Park. Abbreviations used in the drawings are explained in the Materials and Methods section.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g011.jpg" id="oo_1556048.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556048</uri>
            </graphic>
          </fig>
          <fig id="F12">
            <object-id content-type="doi">10.3897/vz.76.e176241.figure12</object-id>
            <object-id content-type="arpha">4F848C58-FBF7-5341-A6DE-795B64691935</object-id>
            <label>Figure 12.</label>
            <caption>
              <p><bold>A</bold> Map indicating known localities of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. on Nosy Be (red dots). <bold>B</bold> Forest at the Lokobe National Park. <bold>C</bold> Detail of the habitat where the holotype of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. was collected.</p>
            </caption>
            <graphic xlink:href="vertebrate-zoology-76-135-g012.jpg" id="oo_1556049.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1556049</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Paratype">
          <title>Paratype.</title>
          <p>MADAGASCAR – <bold>Diana Region</bold> • 1 unsexed specimen; Nosy Be, Lokobe National Park; <named-content content-type="dwc:verbatimCoordinates">13°24’31”S, 48°20’8”E</named-content>; ca. 90 m a.s.l.; 18 Feb. 2015; “Frontier Mada­gascar” team leg.; <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 86/2015.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Diagnosis">
          <title>Diagnosis.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. differs from all other species in the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> (except those of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">nanus</tp:taxon-name-part></tp:taxon-name> group) by a lower number of ventral scales (55, versus 56 to 88), a short and pointed snout with visibly enlarged eyes and a dark and uniform coloration (versus lighter coloration, most often with dark lateral stripes variable in contrast and length). Additionaly, it differs from the species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">nanus</tp:taxon-name-part></tp:taxon-name> group by a higher number of scales around midbody (24, versus 18 to 20), and more elongated digits, with both a higher number of subdigital lamellae under the fourth finger (8, versus 3 to 5) and the fourth toe (12 to 15, versus 5 to 8). It differs from the species of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> group (its sister clade, formed by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="polleni">polleni</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="arenicola">arenicola</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stumpffi">stumpffi</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="miafina">miafina</tp:taxon-name-part></tp:taxon-name></italic>) by a distinctly lower number of ventral scales (55, versus 65 to 88) and of paravertebral scales (53 to 56, versus 65 to 88), a smaller size (33.8 to 39.9 mm, versus 54.9 to 89.6 mm), an hourglass-shaped frontal, i.e., constricted by the first pair of supraocular (versus always bell-shaped in all species except in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stumpffi">stumpffi</tp:taxon-name-part></tp:taxon-name></italic>, where this state of character is variable), and a lower number of subdigital lamellae under the fourth toe (12 to 15, versus 15 to 23).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Description of the holotype">
          <title>Description of the holotype.</title>
          <p>(Figs <xref ref-type="fig" rid="F10">10</xref>, <xref ref-type="fig" rid="F11">11</xref>) Presumably adult specimen, in an overall good state of preservation, with exception of the right hindlimb, which has been cut off as tissue sample, and a longitudinal row of approximately 20 scales ripped out on the left side of the body. A rather small <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> species with relatively large eyes and short and pointed snout. <abbrev xlink:title="snout–vent length">SVL</abbrev> (33.8 mm) 7 times head length (5.9 mm), almost as long as tail (41.0 mm, apparently entire and not regenerated). Limbs very short: <abbrev xlink:title="snout–vent length">SVL</abbrev> 7.1 times forelimb length (5.5 mm) and 4.3 times hind limb length (8.9 mm). Snout short and rounded in lateral aspect and dorsal aspect. Rostral wider than high/long, contacting first supralabial scales, nasals, and supranasal scales. Paired supranasal scales in median contact, contacting loreals and postnasals. Frontonasal roughly triangular, wider than long, contacting loreals, first supraciliaries and first suprocular scales. Prefrontal scales absent. Frontal longer than wide, wider posteriorly, in contact with frontonasal, supraoculars, parietals and interparietal. Four supraoculars, all of them in contact with frontal; the second and third pairs larger than the anteriormost and posteriormost pairs; the first pair weakly constricting frontal (frontal hourglass-shaped sensu <xref ref-type="bibr" rid="B2">Andreone and Greer 2002</xref>). Frontoparietals absent. Interparietal longer than wide, well separated from supraoculars; parietal eyespot present with parietal eye evident. Parietals contact posterior to interparietal. Enlarged nuchals absent. Nasals slightly larger than nostrils; contacting rostral, first supralabials, postnasals and supranasals. Postnasals present, separating supranasals from first supralabials, and nasals from loreals. Loreal single, slightly higher than long. Preocular trapezoidal, as longer as high, single. Presubocular roughly triangular, elongated, single. Seven supraciliaries on both sides, in continuous row; first and last pairs significantly larger than the intermediate ones; last pair projecting onto supraocular shelf. Two pretemporals. Two postsuboculars; upper contacting lower pretemporal; both contacting penultimate supralabial. Lower eyelid moveable, scaly; lower palpebrals small, longer than high, interdigitating with large polygonal scales of central eyelid. Contact between upper palpebrals and supraciliaries seemingly direct but flexible, i.e., palpebral cleft narrow. Primary temporal single. Two secondary temporals; upper contacting lower pretemporal anteriorly and overlapping lower secondary temporal ventrally. Six supralabials, with the fourth being the enlarged subocular, contacting scales of the lower eyelid. External ear opening small, higher than wide, without lobules. Mental wider than long, posterior margin convex. Postmental wider than long, contacting first two pairs of infralabials. Six infralabials. Three pairs of large chin scales, both members of first pair in contact, both members of second pair separated by a single median scale, and members of third pair separated by three scale rows. No scales extending between infralabials and large chin scales. Gulars similar in size and outline to ventrals. All scales, except head shields and scales on palms, soles, and digits, cycloid, smooth, and imbricate; 24 longitudinal scale rows at midbody; 53 paravertebrals, similar in size to adjacent scales; 55 ventrals. Inner preanals overlapped by outer. Both anterior and posterior limbs pentadactyl; fingers and toes clawed; relative length of toes in the following order: I&lt;II&lt;III&lt;V&lt;IV. Subdigital lamellae smooth, single, with eight under both fourth finger, 12 under left fourth toe.</p>
          <p>Coloration of the holotype. In preservative, specimen with a relatively uniform bronze coloration on the dorsal side from head to tail, slightly darker along two barely visible paravertebral lines, progressively fading on the flanks to become an immaculate light cream coloration on the ventral side. Palms and soles barely darker than the rest of the ventral side. In life, the coloration pattern was relatively similar to that in preservative, with however a slightly brighter and warmer overall aspect. The scales in life were iridescent (Fig. <xref ref-type="fig" rid="F10">10</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Variation">
          <title>Variation.</title>
          <p>The paratype <abbrev content-type="institution" xlink:title="Zoologische Staatssammlung München">ZSM</abbrev> 0086/2015 is generally very similar to the holotype (Fig. <xref ref-type="fig" rid="F11">11</xref>). It has the following traits: <abbrev xlink:title="snout–vent length">SVL</abbrev> 39.9 mm, width at midbody 5.3 mm, tail length 52 mm, hindlimb 10.5 and 9.5 mm (right and left, respectively), 13 and 15 subdigital lamellae under the fourth toes, 55 ventrals, 56 dorsals, 24 scales around midbody.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Etymology">
          <title>Etymology.</title>
          <p>The specific epithet irery, invariable noun in apposition, is derived from the Malagasy language, and means “alone” or “solitary”. It refers both to the assumption that this species seems to be isolated on the small island of Nosy Be and the fact that it represents a distinct long phylogenetic branch.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Distribution">
          <title>Distribution.</title>
          <p>Only known from Lokobe National Park (east and west, and therefore likely widespread across the park) on Nosy Be (Fig. <xref ref-type="fig" rid="F12">12</xref>). Surveys on the adjacent island of Nosy Komba failed to find this species (<xref ref-type="bibr" rid="B16">Hyde Roberts and Daly 2014</xref>; <xref ref-type="bibr" rid="B4">Blumgart et al. 2017</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Ecology">
          <title>Ecology.</title>
          <p>In 2019, the holotype of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. was collected active around noon on 28 April, on a boulder field next to a small stream on the Circuit Mitsinjo within Lokobe National Park. The microhabitat was characterized by large boulders covered in moss and liverworts (Fig. <xref ref-type="fig" rid="F12">12C</xref>). This kind of habitat occurs in several locations within Lokobe, which is comprised mostly of lowland moist evergreen forest, but our brief survey did not find the species elsewhere. Far more comprehensive fieldwork is needed to establish the spread of the species within the park, and surveys in other forest fragments on the island should be carried out to establish whether it occurs outside the park.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Conservation status">
          <title>Conservation status.</title>
          <p>Considering that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name> ­irery</italic><bold>sp. nov</bold>. is likely widely distributed across the Lokobe reserve, its EOO might be at least of 7.4 km² (<xref ref-type="bibr" rid="B1">Andreone et al. 2003</xref>). The possibility of a broader distribution within Nosy Be island for this species remains however plausible and cannot be dismissed in the absence of additional data. The Lokobe area exhibits a largely intact forest cover, and direct anthropogenic pressures in its central protected areas remain limited. However, the distribution range of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. is extremely restricted (possibly less than 10 km²), warranting particular attention and regular monitoring to assess its global risk of extinction. Pending further information to confirm this species’ range, population status, and potential threats, we recommend to assign the species the status <abbrev xlink:title="Vulnerable">VU</abbrev> (Vulnerable) according the criteria D2 of the <xref ref-type="bibr" rid="B18">IUCN Standards and Petitions Committee (2024)</xref>.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
    <sec sec-type="Discussion" id="sec10">
      <title>Discussion</title>
      <sec sec-type="Integrative species delimitation" id="sec11">
        <title>Integrative species delimitation</title>
        <p>The two new species described here are known from a very limited sampling (one sequenced voucher specimen for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic> and two for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic>). Nevertheless, their recognition as distinct species is robustly supported by independent lines of evidence. Each species represents a highly divergent and likely ancient lineage within the genus, as supported by the substantial branch lengths obtained independently in the mitochondrial and in the nuclear phylogenetic trees. Accordingly, the <abbrev xlink:title="16S rRNA,">16S</abbrev> uncorrected p-distances calculated between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. and the species in its sister clade (7.4–9.3%), and those between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. and its sister clade (9.0–10.9%) correspond to those already obtained between recognized species in the genus (mean = 8.5%, min–max = 2.8–12.5%, N = 55 species pair comparisons), and are two to three times higher than intraspecific distances which, for the same genus and the same genetic marker never exceed 3.1% (N = 11 species, data from <xref ref-type="bibr" rid="B25">Miralles et al. 2016</xref>). Both species also exhibit several morphological characteristics that unambiguously differentiate them from all other species known in the genus in general, and from their closest relatives (sister clades) in particular.</p>
      </sec>
      <sec sec-type="Biogeography" id="sec12">
        <title>Biogeography</title>
        <p>The completion of a full inventory of life on Earth is inherently challenging, as it is impossible to anticipate with certainty where undescribed species remain to be found. Taxonomic discoveries, however, do not rely solely on chance. Exploration targeting remote, isolated and understudied regions that possess distinct ecological characteristics in contrast to those prevailing in the surrounding areas holds the potential for the discovery of ancient or relict lineages. The more ancient and ecologically isolated these regions are, the more likely they may have served as refugia where evolutionary processes followed unique and divergent trajectories. This assumption is exemplified by the present study, which highlights the existence of two new species found opportunistically, each representing deeply divergent phylogenetic lineages within the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> and appearing to have highly restricted geographic distributions. As such, they provide novel insights into the biogeographic history of the genus:</p>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="irery">irery</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. might be endemic to Nosy Be (but should also be sought on adjacent islands and in adjacent areas of the Sambirano region on mainland Madagascar), while its sister group, the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> clade, is primarily confined to the northern part of mainland Madagascar (except for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="polleni">polleni</tp:taxon-name-part></tp:taxon-name></italic>, which is found slightly further south, in the North West, Fig. <xref ref-type="fig" rid="F9">9</xref>). Although some important early taxonomic works on the Malagasy herpetofauna were based on material sent to Europe from Nosy Be (<xref ref-type="bibr" rid="B5">Boettger 1880</xref>, <xref ref-type="bibr" rid="B6">1881</xref>; <xref ref-type="bibr" rid="B32">Peters 1880</xref>; <xref ref-type="bibr" rid="B39">Vaillant 1885</xref>), scientific exploration of the island—which today is a hotspot for tourism with an international airport—has been limited (<xref ref-type="bibr" rid="B1">Andreone et al. 2003</xref>). This new <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> species represents the second to be reported from Nosy Be, alongside <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="stumpffi">stumpffi</tp:taxon-name-part></tp:taxon-name></italic>, which appears to be more opportunistic, with a broad distribution across multiple localities throughout this island (<xref ref-type="bibr" rid="B1">Andreone et al. 2003</xref>; iNaturalist). These results support the hypothesis according to which the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus"/><tp:taxon-name-part taxon-name-part-type="species">polleni</tp:taxon-name-part></tp:taxon-name> clade most likely originated and diversified in northern Madagascar.</p>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. is only known from the Tsingy de Namoroka, a karstic system located in the dry West of Madagascar. Its sister clade (the <italic>melano­pleura</italic> group) is contrastingly distributed across the entire humid eastern part of the island (and likely has an origin in the North, Fig. <xref ref-type="fig" rid="F9">9</xref>). Interestingly, a similar biogeographic pattern can be found in the Velvet geckos (genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Blaesodactylus">Blaesodactylus</tp:taxon-name-part></tp:taxon-name></italic> Boettger, 1893), with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Blaesodactylus">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="victori">victori</tp:taxon-name-part></tp:taxon-name></italic> Ineich, Glaw &amp; Vences, 2016, also endemic to Namoroka, which has been recovered as sister group of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Blaesodactylus">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="microtuberculatus">microtuberculatus</tp:taxon-name-part></tp:taxon-name></italic> Jono, Bauer, Brennan &amp; Mori, 2015, a species only found in the North of the island (<xref ref-type="bibr" rid="B40">Vences et al. 2025</xref>). Such biogeographical affinities between Namoroka and the North suggest these two regions might have been connected in the past, during a period when the climate was more humid. The Tsingy de Namoroka massif appears to offer a distinct microclimate—cooler and more humid than the surrounding Mahajanga region, which is otherwise characterized by a prolonged dry season. This unique set of conditions may have served as a climatic refuge for the local fauna. Likely contributing to this microclimate, the area indeed experiences substantial flooding during the rainy season and benefits year-round from a natural cooling effect generated by an extensive network of caves, where water reservoirs persist even through the dry season. (A.M. pers. obs., Fig. <xref ref-type="fig" rid="F5">5E</xref>; see also <xref ref-type="bibr" rid="B40">Vences et al. 2025</xref> for more details on the Tsingy de Namoroka habitat).</p>
        <p>The Tsingy of Namoroka represents a region of high interest to the scientific community due to the important number of apparently micro-endemic species hosted by these karst formations. This remarkable level of endemism appears to affect a wide range of taxonomic groups, including plants (<xref ref-type="bibr" rid="B41">Wilkin et al. 2002</xref>; <xref ref-type="bibr" rid="B13">Gautier et al. 2022</xref>), insects (<xref ref-type="bibr" rid="B31">Paulian and Grjebine 1953</xref>; <xref ref-type="bibr" rid="B38">Synave 1953</xref>; <xref ref-type="bibr" rid="B35">Risbec 1956</xref>; <xref ref-type="bibr" rid="B9">Chłond et al. 2018</xref>; <xref ref-type="bibr" rid="B12">Faille and Lecoq 2018</xref>; <xref ref-type="bibr" rid="B15">Guilbert 2020</xref>), arachnids (<xref ref-type="bibr" rid="B19">Lourenço and Goodman 2004</xref>; <xref ref-type="bibr" rid="B42">Wood 2008</xref>; <xref ref-type="bibr" rid="B20">Lourenço and Wilmé 2019</xref>) and squamates (<xref ref-type="bibr" rid="B17">Ineich et al. 2016</xref>; <xref ref-type="bibr" rid="B34">Rakotoarison et al. 2025</xref>; <xref ref-type="bibr" rid="B40">Vences et al. 2025</xref>). Satellite imagery analysis indicates that the Namoroka karst system extends well beyond the current boundaries of the national park (Fig. <xref ref-type="fig" rid="F6">6</xref>). In the northwestern direction, a wide continuous karstified band stretches over approximately 17 km, suggesting significant continuity (Fig. <xref ref-type="fig" rid="F6">6A</xref>). To the southeast, several isolated karst features, each spanning from two to ten kilometers long, are scattered over a distance of roughly 100 km, following a roughly linear alignment reaching the Mahamay river. Although the southernmost formations appear to be more heavily eroded, the presence of densely vegetated canyons intersecting at near-perpendicular angles is a distinctive morphological feature, closely resembling the structure observed in the main Namoroka karst area (Fig. <xref ref-type="fig" rid="F6">6C, D</xref>). Given that these distinct karst outcrops were likely connected in the past, it is plausible that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minotaurus">minotaurus</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., along with other Namoroka endemic species, might also persist in these areas. Due to their relative inaccessibility, these isolated areas remain poorly explored, and comprehensive field surveys are warranted to evaluate their local biodiversity and to identify potential conservation priorities associated with these remote environments.</p>
      </sec>
    </sec>
    <sec sec-type="Conclusion" id="sec13">
      <title>Conclusion</title>
      <p>This paper increases the number of described species in the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> from 12 to 14. However, the species inventory for this genus is not yet complete, as suggested by the identification of at least three credible, as-yet-undescribed candidate species within the genus in previous studies: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> sp. “baeus” (cf. <xref ref-type="bibr" rid="B14">Glaw and Vences 2007</xref>) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> sp. “Betampona” (<xref ref-type="bibr" rid="B3">Belluardo et al. 2023</xref>; work in preparation), two candidate species belonging to the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nanus">nanus</tp:taxon-name-part></tp:taxon-name></italic> group, as well as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">Madascincus</tp:taxon-name-part></tp:taxon-name></italic> sp. “vitreus”, a candidate species identified in the Morondava region, and likely belonging to the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="igneocaudatus">igneocaudatus</tp:taxon-name-part></tp:taxon-name></italic> group (<xref ref-type="bibr" rid="B14">Glaw and Vences 2007</xref>). Furthermore, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Madascincus">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="minutus">minutus</tp:taxon-name-part></tp:taxon-name></italic>, located at the northern tip of the island, might represent a complex of several morphologically cryptic species, given the considerable genetic diversity observed within this taxon (<xref ref-type="bibr" rid="B29">Miralles and Vences 2013</xref>; <xref ref-type="bibr" rid="B25">Miralles et al. 2016</xref>). Finally, as illustrated by the present study, it is also possible that species yet to be collected remain undiscovered in various parts of the island, even those heavily visited by tourists and local researchers.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgments</title>
      <p>We are grateful to Ivan Ineich, Gabriele Keunecke, Ny Ando Tsiky Rahagalala, Angeluc Razafimanantsoa, Mathilde Aladini, Jérôme Courtois, Stéphane Grosjean, Didier Geffard-Kuriyama, Jary H. Razafindraibe, and Fanomezana M. Ratsoavina for advice and/or help with logistics, field work or laboratory work. Fieldwork was carried out in the framework of a collaboration accord between the Technische Universität Braunschweig, the Université d’Antananarivo, and the Ministry of the Environment and of the Sustainable Development. Many thanks to Ishan Agarwal, Uwe Fritz, Daniel Jablonski and Jörn Köhler, who provided valuable comments that helped to improve the manuscript. AM and MV acknowledges support from the Deutsche Forschungsgemeinschaft (grant MI 2748/1-1) and the Agence Nationale de la Recherche (ANR-24-CPJ1-0129-01). Thanks are due to the Malagasy authorities, in particular to the Ministry of the Environment and of the Sustainable Development and Madagascar National Parks, for research, collection and export permits (research permit 315/23/MEDD/SG/DGGE/DAPRNE/SCBE.Re; MNP access permit to Tsingy de Namoroka National Park 158/2023). Authors are also gratefull to the Atelier Iconographie Scientifique, UAR 2700 2AD, BAOBAB facilities supported by DIM-MAP Île-de-France, CNRS and <abbrev xlink:title="Muséum National d’Histoire Naturelle, Paris">MNHN</abbrev>.</p>
    </ack>
    <ref-list>
      <title>References</title>
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      </ref>
      <ref id="B2">
        <mixed-citation>Andreone F, Greer AE (2002) Malagasy scincid lizards: Descriptions of nine new species, with notes on the morphology, reproduction and taxonomy of some previously described species (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superclass">Reptilia</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Squamata</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scincidae</tp:taxon-name-part></tp:taxon-name>). Journal of Zoology 258: 139–181. <ext-link xlink:href="10.1017/S0952836902001280" ext-link-type="doi">https://doi.org/10.1017/S0952836902001280</ext-link></mixed-citation>
      </ref>
      <ref id="B3">
        <mixed-citation>Belluardo F, Muñoz-Pajares AJ, Miralles A, Silvestro D, Cocca W, Ratsoavina FM, Villa A, Hyde Roberts S, Mezzasalma M, Zizka A, Antonelli A, Crottini A (2023) Slow and steady wins the race: Diversification rate is independent from body size and lifestyle in Malagasy skinks (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Squamata</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scincidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Scincinae</tp:taxon-name-part></tp:taxon-name>). Molecular Phylogenetics and Evolution 178: 107635. <ext-link xlink:href="10.1016/j.ympev.2022.107635" ext-link-type="doi">https://doi.org/10.1016/j.ympev.2022.107635</ext-link></mixed-citation>
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    <sec sec-type="supplementary-material">
      <title>Supplementary materials</title>
      <supplementary-material id="S1" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/vz.76.e176241.suppl1</object-id>
        <object-id content-type="arpha">9B60DCE7-75C3-5D04-92F1-77B419249659</object-id>
        <label>Supplementary Material 1</label>
        <caption>
          <p>Files S1, S2</p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p><bold/>: .zip</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation notes</label>
          <p><bold>File S1</bold>. List of specimens examined morphologically [docx file]. — <bold>File S2</bold>. GenBank accession numbers, and localities [docx file].</p>
        </statement>
        <media xlink:href="vertebrate-zoology-76-135-s001.zip" mimetype="application" mime-subtype="zip" position="float" orientation="portrait" id="oo_1556050.zip">
          <uri content-type="original_file">https://binary.pensoft.net/file/1556050</uri>
        </media>
        <permissions>
          <license>
            <license-p>This dataset is made available under the Open Database License (<ext-link ext-link-type="uri" xlink:href="http://opendatacommons.org/licenses/odbl/1.0">http://opendatacommons.org/licenses/odbl/1.0</ext-link>). 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"> Miralles A, Scherz MD, Hyde Roberts S, Rakotoarison A, Glaw F, Vences M (2026)</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
