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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">98</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:3686FA2D-1003-5130-A1A9-6F0A4E63DF4D</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:A3B9B45C-7F69-42DF-85B6-B3189A5B43F2</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Caucasiana</journal-title>
        <abbrev-journal-title xml:lang="en">Caucasiana</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="epub">2667-9809</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/caucasiana.5.e204607</article-id>
      <article-id pub-id-type="publisher-id">204607</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Short Communication</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Colubridae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biodiversity &amp; Conservation</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>A melanistic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">Eirenis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic> (Martin, 1838) from the Anatolian mainland: first record outside island populations</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Korkmaz</surname>
            <given-names>Ahmet Gökay</given-names>
          </name>
          <email xlink:type="simple">ahmetgokay.korkmaz@deu.edu.tr</email>
          <uri content-type="orcid">https://orcid.org/0009-0009-8774-9156</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Candan</surname>
            <given-names>Kamil</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-6934-3971</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Hastürk</surname>
            <given-names>Ecem Büşra</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0000-2564-8733</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Gül</surname>
            <given-names>Serkan</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-0372-7462</uri>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Şahin</surname>
            <given-names>Mehmet Kürşat</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-0834-5081</uri>
          <xref ref-type="aff" rid="A3">3</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Caynak</surname>
            <given-names>Elif Yıldırım</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-6794-2527</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Kumlutaş</surname>
            <given-names>Yusuf</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-1154-6757</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Dokuz Eylül University, İzmir, Türkiye</addr-line>
        <institution>Dokuz Eylül University</institution>
        <addr-line content-type="city">İzmir</addr-line>
        <country>Türkiye</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Recep Tayyip Erdoğan University, Rize, Türkiye</addr-line>
        <institution>Recep Tayyip Erdoğan University</institution>
        <addr-line content-type="city">Rize</addr-line>
        <country>Türkiye</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Hacettepe University, Ankara, Türkiye</addr-line>
        <institution>Hacettepe University</institution>
        <addr-line content-type="city">Ankara</addr-line>
        <country>Türkiye</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Ahmet Gökay Korkmaz (<email xlink:type="simple">ahmetgokay.korkmaz@deu.edu.tr)</email></p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: David Tarkhnishvili</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>16</day>
        <month>09</month>
        <year>2026</year>
      </pub-date>
      <volume>5</volume>
      <fpage>139</fpage>
      <lpage>146</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/5937F638-A9C8-5362-B7C5-6DF4F4265DA8">5937F638-A9C8-5362-B7C5-6DF4F4265DA8</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/64800B22-699C-4195-A8AC-E5BA0C64462A">64800B22-699C-4195-A8AC-E5BA0C64462A</uri>
      <history>
        <date date-type="received">
          <day>18</day>
          <month>06</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>04</day>
          <month>07</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Ahmet Gökay Korkmaz, Kamil Candan, Ecem Büşra Hastürk, Serkan Gül, Mehmet Kürşat Şahin, Elif Yıldırım Caynak, Yusuf Kumlutaş</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/64800B22-699C-4195-A8AC-E5BA0C64462A</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>Melanism is among the most ecologically significant chromatic anomalies reported in squamates, and yet its evolutionary drivers remain a topic of debate. Two principal hypotheses have been proposed: thermal melanism, which holds that darker individuals heat more rapidly in cool climates, and Gloger's rule, which predicts greater melanin deposition in warm, humid environments. Melanistic individuals of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">Eirenis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic> are a rarely documented phenomenon given the species' characteristically pale dorsal coloration; to date, every previously published record of this dark morph had come from island populations. Here we report the first mainland record of melanism in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic>, based on a single specimen collected on 9 May 2024 near Çağman–Demre, Antalya Province, southwestern Türkiye. The individual exhibited entirely black dorsal coloration from head to tail, with greyish-white ventral scales darkening progressively towards the tail. Pholidotic characters were consistent with those previously documented for the species. The Çağman–Demre locality, like all previously reported sites, is characterised by relatively high humidity and a Mediterranean coastal climate, a pattern more consistent with Gloger's rule than with thermal melanism. Further sampling across the Anatolian mainland and adjacent islands is needed to assess the frequency and geographic distribution of this morph.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="family" reg="Colubridae">Colubridae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>color polymorphism</kwd>
        <kwd>Gloger's rule</kwd>
        <kwd>thermal melanism</kwd>
      </kwd-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="sec1">
        <title>Citation</title>
        <p>Korkmaz AG, Candan K, Hastürk EB, Gül S, Şahin MK, Caynak EY, Kumlutaş Y (2026) A melanistic Eirenis modestus (Martin, 1838) from the Anatolian mainland: first record outside island populations. Caucasiana 5: 139–146. <ext-link xlink:type="simple" ext-link-type="doi" xlink:href="10.3897/caucasiana.5.e204607">https://doi.org/10.3897/caucasiana.5.e204607</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec2">
      <title>Introduction</title>
      <p>Body coloration in reptiles results from a complex interaction between pigmentary and structural components. It serves diverse functions, including crypsis, aposematism, social signalling, and thermoregulation (<xref ref-type="bibr" rid="B6">Bogert 1949</xref>; <xref ref-type="bibr" rid="B11">Cuthill et al. 2017</xref>). Among chromatic anomalies in squamates, melanism is the most frequently reported and is considered particularly ecologically significant. Melanism is defined as the excessive deposition of dark eumelanin-based pigments. In reptiles, melanism occurs across many lineages and has been linked to thermoregulation, camouflage against dark substrates, protection from ultraviolet radiation, and predator-prey interactions, so that dark morphs provide useful natural models for testing ecological and evolutionary hypotheses about color variation (<xref ref-type="bibr" rid="B9">Clusella-Trullas et al. 2007</xref>; <xref ref-type="bibr" rid="B11">Cuthill et al. 2017</xref>). Two main adaptive hypotheses have been proposed to account for its geographic distribution. The thermal melanism hypothesis was formalised by Clusella-Trullas, et al. (<xref ref-type="bibr" rid="B9">2007</xref>) following <xref ref-type="bibr" rid="B6">Bogert (1949)</xref>. It proposes that darker individuals are favoured in cooler climates because they heat more rapidly and attain higher equilibrium body temperatures than lighter individuals under equivalent solar radiation. By contrast, Gloger’s rule predicts that darker pigmentation should predominate in warm, humid environments (<xref ref-type="bibr" rid="B22">Rensch 1929</xref>; <xref ref-type="bibr" rid="B12">Delhey 2019</xref>). However, whether this thermal advantage is sufficient to drive melanism evolution in specific lineages remains unresolved.</p>
      <p>Melanism in snakes has frequently been used as a model system for studying color polymorphism, owing to its broad taxonomic occurrence and pronounced geographic variation (<xref ref-type="bibr" rid="B23">Sahlean et al. 2025</xref>). Field and intraspecific studies indicate that melanistic individuals may exhibit higher growth rates and improved body condition. They may also have longer activity periods and enhanced locomotor performance compared to normally colored individuals (<xref ref-type="bibr" rid="B1">Andrén and Nilson 1981</xref>; <xref ref-type="bibr" rid="B9">Clusella-Trullas et al. 2007</xref>; <xref ref-type="bibr" rid="B7">Castella et al. 2013</xref>). At the macroevolutionary level, <xref ref-type="bibr" rid="B18">Martínez-Freiría et al. (2020)</xref> provided the first phylogenetic comparative evidence supporting the thermal melanism hypothesis in Eurasian vipers. <xref ref-type="bibr" rid="B26">Storniolo et al. (2025)</xref> reported similar findings for elapids. However, the evidence remains inconsistent. A recent meta-analysis of polymorphic terrestrial snakes found that the prevalence of melanism correlated more strongly with annual precipitation than with temperature. This suggests that Gloger’s rule may better explain the distribution of dark morphs than thermal advantage alone (<xref ref-type="bibr" rid="B23">Sahlean et al. 2025</xref>). Predation pressure adds further complexity, as dark snakes lacking species-typical patterns may be more conspicuous to visually oriented predators (<xref ref-type="bibr" rid="B18">Martínez-Freiría et al. 2020</xref>). The adaptive significance of melanism in snakes remains debated. Melanism has been recorded in several snake genera such as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Vipera">Vipera</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B18">Martínez-Freiría et al. 2020</xref>; <xref ref-type="bibr" rid="B24">Schmid et al. 2023</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dolichophis">Dolichophis</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B14">Göçmen et al. 2015</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hierophis">Hierophis</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B28">Zuffi 2008</xref>; <xref ref-type="bibr" rid="B20">Rato et al. 2009</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Natrix">Natrix</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B19">Mebert 2011</xref>; <xref ref-type="bibr" rid="B15">Habiboğlu et al. 2016</xref>; <xref ref-type="bibr" rid="B27">Yenmiş et al. 2022</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Coronella">Coronella</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B4">Barbadillo et al. 1997</xref>; <xref ref-type="bibr" rid="B21">Reading and Pernetta 2009</xref>), and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Zamenis">Zamenis</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B10">Colla et al. 2025</xref>).</p>
      <p>Asia Minor Dwarf Racer, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">Eirenis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic> (Martin, 1838), is a small to medium-sized colubrid. It rarely exceeds 600 mm in total length. Its range includes Armenia, Azerbaijan, eastern Georgia, and southern Russia, as well as north-western Iran and several Greek islands in the nominal subspecies <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">m.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">m.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="semimaculatus">semimaculatus</tp:taxon-name-part></tp:taxon-name></italic> (Boettger, 1876) in central and western Anatolia, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">m.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="cilicius">cilicius</tp:taxon-name-part></tp:taxon-name></italic> (Schmidtler, 1993) in southern Türkiye (<xref ref-type="bibr" rid="B17">Mahlow et al. 2013</xref>). This species is typically found in dry, rocky habitats with sparse vegetation, such as maquis, steppe, fallow fields, and ruins. It shelters under flat stones during the day and emerges at twilight to forage. Its diet consists primarily of terrestrial arthropods, particularly beetles, orthopterans, and centipedes. Occasionally, it preys on small lizards (<xref ref-type="bibr" rid="B8">Çiçek and Mermer 2007</xref>; <xref ref-type="bibr" rid="B3">Baran et al. 2021</xref>). The dorsum is generally uniform, light brown to greyish, often with a darker collar and a characteristic dark pileus pattern that fades with age. Populations from southern Anatolia and parts of the Aegean exhibit variable rows of dark blotches on the anterior body, especially within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">m.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="semimaculatus">semimaculatus</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B17">Mahlow et al. 2013</xref>).</p>
      <p>Set against a backdrop of generally light coloration, melanistic individuals in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic> are a rarely documented phenomenon. The earliest published account is by <xref ref-type="bibr" rid="B2">Baran (1986)</xref>, who described island populations from Yassıca and Sarıot in the Muğla Province of south-western Türkiye that included dark individuals. Three decades later, <xref ref-type="bibr" rid="B16">Kalaentzis et al. (2018)</xref> reported two melanistic specimens – a juvenile and an adult – from the Greek island of Kastellorizo (Megisti) in the south-eastern Aegean Sea. More recently, <xref ref-type="bibr" rid="B25">Searle-Webb and Kuttalam (2025)</xref> documented a single melanistic adult from Symi, also in the Dodecanese. This individual had a dark grey dorsum that contrasted only faintly with the head pattern. Despite these scattered records, melanism in E. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis"/><tp:taxon-name-part taxon-name-part-type="species">modestus</tp:taxon-name-part></tp:taxon-name> remains largely unstudied. There is no published assessment of its frequency within populations, nor information on its genetic basis, and no formal evaluation of whether it follows the thermoregulatory or moisture-related patterns observed in other snakes (<xref ref-type="bibr" rid="B16">Kalaentzis et al. 2018</xref>). Continued documentation of additional cases, especially from the Anatolian mainland where the species is most abundant, is essential to clarify the geographic distribution of dark morphs and to support future ecological and evolutionary analyses.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec3">
      <title>Materials and methods</title>
      <p>The melanistic specimen was collected on 9 May 2024 during fieldwork, between Çağman and Demre, Antalya Province, Türkiye (<xref ref-type="fig" rid="F1">Fig. 1</xref>). The specimen was discovered beneath a rock at approximately 17:30 h. The exact locality of the specimen was determined using a GPS. The specimen was anesthetised with ether, fixed with a 96% ethanol injection, and deposited in 96% ethanol in the Zoology Lab. of the Department of Biology at the Faculty of Science, Dokuz Eylül University. The specimen was photographed using a Nikon D80 DSLR camera in order to determine color and pattern characteristics. Mensural, meristic, and qualitative data were recorded following the system of <xref ref-type="bibr" rid="B5">Başoğlu and Baran (1980)</xref> and <xref ref-type="bibr" rid="B17">Mahlow et al. (2013)</xref>. Meristic (scale) counts and pholidosis were examined under a Leica stereomicroscope, whereas mensural (metric) measurements were taken with digital callipers. The ventral plates were counted according to <xref ref-type="bibr" rid="B13">Dowling (1951)</xref>.</p>
      <fig id="F1">
        <object-id content-type="doi">10.3897/caucasiana.5.e204607.figure1</object-id>
        <object-id content-type="arpha">DBCA0BE4-41D6-52A8-9BCA-F4886ABFD212</object-id>
        <label>Figure 1.</label>
        <caption>
          <p>Map showing previous records and the new record of the melanistic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">Eirenis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic>.</p>
        </caption>
        <graphic xlink:href="caucasiana-05-139-g001.jpg" id="oo_1772740.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1772740</uri>
        </graphic>
      </fig>
    </sec>
    <sec sec-type="Results" id="sec4">
      <title>Results</title>
      <p>Fieldwork in the area yielded no additional individuals of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic>, and the findings reported here are therefore based on this single specimen. The individual had a snout-vent length of 155 mm and a tail length of 47.3 mm, with 17 dorsal scale rows and 161 ventral scales. Pileus length and width measured 7.55 mm and 4.27 mm, respectively; frontal length and width were 2.66 mm and 1.63 mm. The preocular and postocular counts of 1 and 2, respectively, fall within the range documented for the species (<xref ref-type="bibr" rid="B17">Mahlow et al. 2013</xref>). <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Supralabialia">Supralabialia</tp:taxon-name-part></tp:taxon-name> were 7 on both sides (<xref ref-type="fig" rid="F2">Fig.  2</xref>). This configuration is consistent with the typical count of 7 reported for the species and the melanistic individual from Yassıcaada (<xref ref-type="bibr" rid="B2">Baran 1986</xref>). <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Sublabialia">Sublabialia</tp:taxon-name-part></tp:taxon-name> numbered 8 on both sides, while the usual count was 8, rarely 9, reported for <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name>. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Subcaudalia">Subcaudalia</tp:taxon-name-part></tp:taxon-name> totalled 72, consistent with the species range of 51–86. Taken together, these pholidotic characters (17 dorsal scale rows at midbody, a single preocular and two postoculars, seven supralabials, eight sublabials, 161 ventrals and 72 subcaudals), together with the characteristic dark pileus and nuchal-collar pattern, identify the specimen as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic> and distinguish it from other small <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">Eirenis</tp:taxon-name-part></tp:taxon-name></italic> species occurring in southern Anatolia (<xref ref-type="bibr" rid="B17">Mahlow et al. 2013</xref>).</p>
      <fig id="F2">
        <object-id content-type="doi">10.3897/caucasiana.5.e204607.figure2</object-id>
        <object-id content-type="arpha">C2C3A232-12FC-5C7F-A8A8-7819A6B55239</object-id>
        <label>Figure 2.</label>
        <caption>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">Eirenis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic>; <bold>A</bold>: Dorsal view of the head showing uniform black pigmentation and characteristic pileus pattern; <bold>B</bold>: Lateral view of the head; <bold>C</bold>: Lateral view of the anterior body showing the overall coloration; note the greyish-white ventral surface and the gradual darkening of lateral scales toward the dorsum.</p>
        </caption>
        <graphic xlink:href="caucasiana-05-139-g002.jpg" id="oo_1772741.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1772741</uri>
        </graphic>
      </fig>
      <p>The dorsum was entirely black from the head to the tip of the tail (<xref ref-type="fig" rid="F3">Fig.  3</xref>). In this comparatively young individual, greyish-white scales were scattered across the neck region, fading rapidly toward midbody, with white scales gradually increasing from the lateral surfaces toward the ventrals. The ventral surface was greyish-white, becoming progressively darker toward the tail.</p>
      <fig id="F3">
        <object-id content-type="doi">10.3897/caucasiana.5.e204607.figure3</object-id>
        <object-id content-type="arpha">94AA224C-3274-5280-8D87-27BB6749A62B</object-id>
        <label>Figure 3.</label>
        <caption>
          <p>Overal view of the melanistic specimen of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">Eirenis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic> that was collected on 9 May 2024 during fieldwork.</p>
        </caption>
        <graphic xlink:href="caucasiana-05-139-g003.jpg" id="oo_1772742.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/1772742</uri>
        </graphic>
      </fig>
      <p>The meta-analysis by <xref ref-type="bibr" rid="B23">Sahlean et al. (2025)</xref> suggests that melanistic snakes are more frequently encountered in coastal regions and on islands than in mainland populations. Therefore, this mainland record represents a noteworthy finding for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic>. As in previous records (<xref ref-type="bibr" rid="B2">Baran 1986</xref>; <xref ref-type="bibr" rid="B16">Kalaentzis et al. 2018</xref>; <xref ref-type="bibr" rid="B25">Searle-Webb and Kuttalam 2025</xref>), melanistic individuals of this species have been documented exclusively from islands. This may be attributable to adaptation to coastal environments or to the constraints associated with island living. Melanistic individuals generally inhabit cooler regions than their lighter-colored counterparts (<xref ref-type="bibr" rid="B9">Clusella-Trullas et al. 2007</xref>; <xref ref-type="bibr" rid="B18">Martínez-Freiría et al. 2020</xref>; <xref ref-type="bibr" rid="B24">Schmid et al. 2023</xref>). On the other hand, according to Gloger's rule, animals in warmer and more humid environments tend to deposit greater amounts of melanin (<xref ref-type="bibr" rid="B12">Delhey 2019</xref>). Çağman-Demre and the previously reported island localities are all characterised by relatively high humidity. Such environments harbour denser vegetation and soils with higher organic matter content. In these settings, background coloration is generally darker and light conditions are reduced, which favours darker pigmentation for camouflage. Our single observation is therefore more consistent with the pattern predicted by Gloger's rule than with thermal melanism, although a single record cannot by itself test either hypothesis. Because this finding is based on one individual, its dark phenotype could equally reflect a rare, non-adaptive mutation rather than a locally adaptive trait: if melanism were adaptive in this region, a substantial proportion of the local <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic> would be expected to be dark, whereas the typical pale morph in fact predominates throughout southern Anatolia (<xref ref-type="bibr" rid="B17">Mahlow et al. 2013</xref>; <xref ref-type="bibr" rid="B3">Baran et al. 2021</xref>).</p>
      <p>Further research in Anatolia and nearby islands is needed. This will help determine whether melanism in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eirenis">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="modestus">modestus</tp:taxon-name-part></tp:taxon-name></italic> and other snake species is truly limited to humid coastal and island habitats. Also, studying the molecular mechanisms of melanism, along with broad geographic and climate data, may clarify the evolutionary advantages and disadvantages of this trait, and the relative importance of thermal melanism and Gloger's rule.</p>
    </sec>
  </body>
  <back>
    <sec sec-type="Additional information" id="sec5">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec6">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec7">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec8">
        <title>Artificial Intelligence (AI) use</title>
        <p>The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI. No AI tools were used in the preparation of this manuscript.</p>
      </sec>
      <sec sec-type="Funding" id="sec9">
        <title>Funding</title>
        <p>The study was partially funded by the Shota Rustaveli National Science Foundation of Georgia (award no. FR-23-17324) and by the Federal Ministry of Education and Research of Georgia under grant number 01DK20014A.</p>
      </sec>
      <sec sec-type="Author contributions" id="sec10">
        <title>Author contributions</title>
        <p>AGK: Conceptualization, fieldwork, specimen preparation, morphological data collection, map preparation, writing – original draft, writing – review and editing. KC: Fieldwork, morphological data collection, writing – review and editing. EBH: Morphological data collection, writing – review and editing. SG: Fieldwork, writing – review and editing. MKS: Fieldwork, Writing – review and editing. EYC: Writing – review and editing. YK: Conceptualization, photography, supervision, writing – review and editing.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec11">
        <title>Author ORCIDs</title>
        <p>Ahmet Gökay Korkmaz <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0009-0009-8774-9156">https://orcid.org/0009-0009-8774-9156</ext-link></p>
        <p>Kamil Candan <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0002-6934-3971">https://orcid.org/0000-0002-6934-3971</ext-link></p>
        <p>Ecem Büşra Hastürk <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0009-0000-2564-8733">https://orcid.org/0009-0000-2564-8733</ext-link></p>
        <p>Serkan Gül <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0002-0372-7462">https://orcid.org/0000-0002-0372-7462</ext-link></p>
        <p>Mehmet Kürşat Şahin <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0003-0834-5081">https://orcid.org/0000-0003-0834-5081</ext-link></p>
        <p>Elif Yıldırım Caynak <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0001-6794-2527">https://orcid.org/0000-0001-6794-2527</ext-link></p>
        <p>Yusuf Kumlutaş <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0003-1154-6757">https://orcid.org/0000-0003-1154-6757</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec12">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text or Supplementary Information.</p>
      </sec>
    </sec>
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