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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">66</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:5F805F0D-E48A-55F8-A459-212B9124D765</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:0535FF1B-CC50-4DC5-A25D-FE8188C9D477</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Alpine Entomology</journal-title>
        <abbrev-journal-title xml:lang="en">AlpEnt</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="epub">2535-0889</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/alpento.10.207817</article-id>
      <article-id pub-id-type="publisher-id">207817</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Crustacea</subject>
          <subject>Malacostraca</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Aquatic biology</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Alps</subject>
          <subject>Europe</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>First record of the Ponto-Caspian amphipod <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> (Martynov, 1932) (= <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Echinogammarus">Echinogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic>) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subphylum" reg="Crustacea">Crustacea</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Gammaridae">Gammaridae</tp:taxon-name-part></tp:taxon-name>) in Lake Geneva</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Marle</surname>
            <given-names>Pierre</given-names>
          </name>
          <email xlink:type="simple">pierre.marle@unil.ch</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-4464-9795</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <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/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
          <role content-type="http://credit.niso.org/contributor-roles/validation/">Validation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Stoffel</surname>
            <given-names>Céline</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0002-4538-9518</uri>
          <xref ref-type="aff" rid="A2">2</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/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/validation/">Validation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Renahy</surname>
            <given-names>Simon</given-names>
          </name>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</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/methodology/">Methodology</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Department of Ecology and Evolution, University of Lausanne, Lausanne, Switzerland</addr-line>
        <institution>Department of Ecology and Evolution, University of Lausanne</institution>
        <addr-line content-type="city">Lausanne</addr-line>
        <country>Switzerland</country>
        <uri content-type="ror">https://ror.org/019whta54</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Department of Zoology, State Museum of Natural Sciences, Lausanne, Switzerland</addr-line>
        <institution>Department of Zoology, State Museum of Natural Sciences</institution>
        <addr-line content-type="city">Lausanne</addr-line>
        <country>Switzerland</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Sage Environnement, Annecy le Vieux, France</addr-line>
        <institution>Sage Environnement</institution>
        <addr-line content-type="city">Annecy le Vieux</addr-line>
        <country>France</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Pierre Marle (<email xlink:type="simple">pierre.marle@unil.ch</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Oliver Martin</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>25</day>
        <month>09</month>
        <year>2026</year>
      </pub-date>
      <volume>10</volume>
      <fpage>163</fpage>
      <lpage>168</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/6FF53407-9977-5D95-8A1D-FBA81D81C761">6FF53407-9977-5D95-8A1D-FBA81D81C761</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/829EA181-7232-4B10-9C6C-1CC209E30EA7">829EA181-7232-4B10-9C6C-1CC209E30EA7</uri>
      <history>
        <date date-type="received">
          <day>14</day>
          <month>07</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>17</day>
          <month>09</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Pierre Marle, Céline Stoffel, Simon Renahy</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/829EA181-7232-4B10-9C6C-1CC209E30EA7</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>The Ponto-Caspian amphipod <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> was recorded for the first time in Lake Geneva. The identification of the collected specimens was established through morphological examination and independently confirmed by DNA barcoding based on the mitochondrial <abbrev xlink:title="mitochondrial cytochrome c oxidase subunit I">COI</abbrev> gene. The present observation in Lake Geneva is noteworthy, as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> has only rarely been reported from lake ecosystems, most records originating from large rivers. In addition to addressing sampling aspects, we describe the habitat of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> in Lake Geneva, detail the morphological identification criteria used to distinguish the species, and discuss the probable implications of this newly recorded species for the interpretation of previous findings in the lake.</p>
      </abstract>
      <kwd-group>
        <label>Key Words</label>
        <kwd>Alien species</kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="order" reg="Amphipoda">Amphipoda</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>bryophytes</kwd>
        <kwd>DNA</kwd>
        <kwd>exotic species</kwd>
        <kwd>invasive species</kwd>
        <kwd>non-native species</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec1">
      <title>Introduction</title>
      <p>The native distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> extends across the Ponto-Caspian region, including the lagoons and estuarine habitats of the northwestern Black Sea (<xref ref-type="bibr" rid="B19">Rewicz et al. 2016</xref>; <xref ref-type="bibr" rid="B8">Copilaș-Ciocianu et al. 2023</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> was first reported as an alien species in Germany in 1996 (<xref ref-type="bibr" rid="B22">Weinzierl et al. 1997</xref>) and has since spread throughout the Danube basin (<xref ref-type="bibr" rid="B4">Boets et al. 2012</xref>) and several catchments in northeastern France (<xref ref-type="bibr" rid="B15">Labat et al. 2011</xref>). In Switzerland, the species has been recorded in the Rhine basin since 2003 and, more recently, in the Aare basin in 2025 (Info fauna 2026). It has also been reported from the upper Dnieper River in Belarus (<xref ref-type="bibr" rid="B16">Lipinskaya et al. 2018</xref>) and the Oder catchment in Central Europe (<xref ref-type="bibr" rid="B8">Copilaș-Ciocianu et al. 2023</xref>).</p>
      <p>In Lake Geneva, non-native crustaceans now constitute almost the entire amphipod fauna. Native amphipods, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pulex">pulex</tp:taxon-name-part></tp:taxon-name></italic> (Linnaeus, 1758) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="fossarum">fossarum</tp:taxon-name-part></tp:taxon-name></italic> Koch, 1836, are progressively displaced by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="villosus">villosus</tp:taxon-name-part></tp:taxon-name></italic> Sowinsky, 1894 (<xref ref-type="bibr" rid="B5">Bollache 2004</xref>; <xref ref-type="bibr" rid="B17">Lods-Crozet and Reymond 2006</xref>) and are only rarely found in surveys. The limited sampling effort conducted along the French shoreline during the last two decades has reduced the likelihood of detecting newly established alien amphipods, whereas regular monitoring has continued on the Swiss shore (i.e. CIPEL monitoring).</p>
      <p>In 2024, samplings conducted along the French littoral of Lake Geneva led to the first detection of the Ponto-Caspian amphipod <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> in the lake. Here, we report the first record of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> in Lake Geneva, describe the morphological characters that distinguish it from other amphipods occurring in the lake, and confirm its identification using mitochondrial <abbrev xlink:title="mitochondrial cytochrome c oxidase subunit I">COI</abbrev> DNA barcoding. We also characterize the habitat in which the species was collected and discuss its possible introduction pathway, as well as the implications of this finding for interpreting previous surveys and guiding future monitoring in Lake Geneva.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec2">
      <title>Material and methods</title>
      <sec sec-type="Sampling" id="sec3">
        <title>Sampling</title>
        <p>The Evian-les-Bains site was sampled on 12 August 2024 (GPS coordinates: <named-content content-type="dwc:verbatimCoordinates">46.40098197, 6.583298966</named-content>). Multiple samples representing different lake substrates were collected using a Surber net (mesh size 500 μm, aperture 20 cm × 25 cm). The riparian habitat at the site is characterized by a vertical quay wall along the shoreline and a substrate composed primarily of boulders and cobbles (Fig. <xref ref-type="fig" rid="F1">1A</xref>), partially covered by quagga, zebra mussels and bryophytes (Fig. <xref ref-type="fig" rid="F1">1B-D</xref>). The shoreline is intermittently reinforced with short sections of riprap. Water depth at the base of the shoreline ranges from approximately 1 to 1.5 m. The lakebed has a moderate slope extending to about 80 m from the shore, where the depth reaches 6–7 m, beyond which it becomes markedly steeper.</p>
        <fig id="F1">
          <object-id content-type="doi">10.3897/alpento.10.207817.figure1</object-id>
          <object-id content-type="arpha">1C8A4593-0589-5686-9D57-7CE8624D94BB</object-id>
          <label>Figure 1.</label>
          <caption>
            <p>Photographs of the sampling site (<bold>A</bold>) and the sampled habitat (<bold>B–D</bold>), taken on the day of sampling where <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> was detected. Photos: S. Renahy.</p>
          </caption>
          <graphic xlink:href="alpineentomology-10-163-g001.jpg" id="oo_1790820.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1790820</uri>
          </graphic>
        </fig>
        <p>Amphipods were collected from a bryophyte sample growing on rocks at a water depth of 0.2 m, within the “wave-splash” zone where riparian bryophytes typically occur in Lake Geneva. Bryophyte development is generally absent or very limited below the “wave-splash” zone, as these species (e.g. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Orthotrichum">Orthotrichum</tp:taxon-name-part></tp:taxon-name></italic> sp.) are essentially riparian bryophytes that can tolerate periodic submersion rather than truly aquatic species (e.g. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Fontinalis">Fontinalis</tp:taxon-name-part></tp:taxon-name></italic> sp.).</p>
        <p>The sample was sorted and preserved in approximately 70% ethanol. Two amphipod specimens were identified as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> based on morphological characters following the identification key provided by <xref ref-type="bibr" rid="B1">Altermatt et al. (2019)</xref>. The identification was also verified using the characters described by <xref ref-type="bibr" rid="B9">Eiseler (2010)</xref> which include the morphology of the uropods, urosome segments, antennae, and gnathopods.</p>
      </sec>
      <sec sec-type="Laboratory protocols, sequencing and alignment" id="sec4">
        <title>Laboratory protocols, sequencing and alignment</title>
        <p>Due to the uncertainty regarding morphological identification of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic>, the two specimens collected in 2024 were checked using molecular techniques in 2025. GBIF code was added to both samples, GBIFCH00967050 for the first one and GBIFCH00967051 for the second one.</p>
        <p>Total genomic DNA was extracted using the BioSprint 96 extraction robot (Qiagen Inc., Hilden, Germany), following the supplier’s instructions. The non-destructive protocol described in <xref ref-type="bibr" rid="B21">Vuataz et al. (2011)</xref>, which enables post-extraction morphological study of specimens, was implemented. A portion (181 bp) of the mitochondrial cytochrome c oxidase subunit I (<abbrev xlink:title="mitochondrial cytochrome c oxidase subunit I">COI</abbrev>), was amplified by using the LCO1490 (5’-GGTCAACAAATCATAAAGATATTGG-3’, <xref ref-type="bibr" rid="B10">Folmer et al. 1994</xref>) and CO1‐CFMRb (5'‐GGNACTAATCAATTHCCAAATCC‐3', <xref ref-type="bibr" rid="B14">Jusino et al. 2019</xref>) primers. Polymerase Chain Reaction (<abbrev xlink:title="Polymerase Chain Reaction">PCR</abbrev>) was conducted in a volume of 25 μl, consisting of 2 μl (unknown concentration) of template DNA, 1.3 μl (10 μM) of each primer, 2 μl (2.5 mM) of dNTP solution (Carl-Roth GmbH), 5 μl (5X) of buffer (Promega) containing 7.5 mM of MgCl<sub>2</sub>, 0.5 μl (10 mg/ml) of BSA (Roche), 1 U of GoTaq G2 DNA Polymerase (Promega), and 12.8 μl of sterile ddH<sub>2</sub>O. Optimized <abbrev xlink:title="Polymerase Chain Reaction">PCR</abbrev> conditions included initial denaturation at 95 °C for 5 min, 5 cycles of denaturation at 95 °C for 30 s, annealing at 45 °C for 30 s, and extension at 72 °C for 45 s, and 35 cycles of denaturation at 95 °C for 30 s, annealing at 50 °C for 30 s, and extension at 72 °C for 45 s, and with final extension at 72 °C for 7 min. Purification and Sanger sequencing was carried out by Microsynth (Balgach, Switzerland).</p>
        <p>Both sequences were identified as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic>. They clustered in a single clade with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> reference sequences from BOLD, with 100% support. The FASTA sequences are provided as Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>. The two specimens are housed in the Department of Zoology, Naturéum Lausanne, Switzerland.</p>
      </sec>
    </sec>
    <sec sec-type="Results" id="sec5">
      <title>Results</title>
      <p>On 12 August 2024, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> was detected for the first time in Lake Geneva at the Evian-les-Bains site. Species identification of the two specimens was carried out by the authors using morphological examination and was further validated by DNA barcoding based on the mitochondrial cytochrome c oxidase subunit I (<abbrev xlink:title="mitochondrial cytochrome c oxidase subunit I">COI</abbrev>) gene.</p>
      <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">Amphipoda</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family">Gammaridae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <tp:taxon-name><object-id content-type="arpha">5F3698E9-047E-5802-A148-7623678DD54C</object-id>
                		<tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part>
                	</tp:taxon-name>
          <tp:taxon-authority>Martynov, 1932</tp:taxon-authority>
          <xref ref-type="fig" rid="F2">Fig. 2</xref>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="Synonymy">
          <title>Synonymy.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Echinogammarus">Echinogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> (Martynov, 1932),</p>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trichogammarus">Trichogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> (Martynov, 1932).</p>
          <p><bold>Material examined</bold>. GBIFCH00967050 &amp; GBIFCH00967051, 2 specimens, France, Lake Geneva, Evian-les-Bains, <named-content content-type="dwc:verbatimCoordinates">46.40098197°N, 6.583298966°E</named-content>, 372 m, 12.08.2024, exoskeleton well preserved, soft tissues used for <abbrev xlink:title="mitochondrial cytochrome c oxidase subunit I">COI</abbrev> gene sequencing, leg. S. Renahy.</p>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part></tp:taxon-name></italic> can easily be distinguished from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part></tp:taxon-name></italic> by the morphology of the third uropod and the first and second urosome segments. In the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part></tp:taxon-name></italic>, the inner ramus of the third uropod is very short (Fig. <xref ref-type="fig" rid="F2">2B</xref>), whereas it is well developed in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part></tp:taxon-name></italic>. Furthermore, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part></tp:taxon-name></italic> lack the paired dorsal protuberances bearing spines on the first and second urosome segments (Fig. <xref ref-type="fig" rid="F2">2A</xref>), a characteristic feature of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part></tp:taxon-name></italic> species.</p>
          <fig id="F2">
            <object-id content-type="doi">10.3897/alpento.10.207817.figure2</object-id>
            <object-id content-type="arpha">CE7B7826-1FA7-572C-BDB9-200762982565</object-id>
            <label>Figure 2.</label>
            <caption>
              <p>Adult specimen of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> collected at the Evian-les-Bains site (Lake Geneva) on 12 August 2024. The figure shows (<bold>A</bold>) a lateral view of the entire specimen and (<bold>B</bold>) a dorsal view of the third uropod. Photos: S. Renahy.</p>
            </caption>
            <graphic xlink:href="alpineentomology-10-163-g002.jpg" id="oo_1790821.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1790821</uri>
            </graphic>
          </fig>
          <p>Compared with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ischnus">ischnus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> is characterized by a third uropod densely covered with recurved setae (Fig. <xref ref-type="fig" rid="F2">2B</xref>). In females of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic>, these recurved setae are at least densely developed at the apex of the third uropod.</p>
          <p>The identification key provided below covers amphipod taxa commonly encountered in Lake Geneva but does not include <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chelicorophium">Chelicorophium</tp:taxon-name-part></tp:taxon-name></italic> species.</p>
          <sec sec-type="Identification key" id="sec6">
            <title>Identification key</title>
            <table-wrap content-type="key" position="anchor" orientation="portrait">
              <table>
                <tbody>
                  <tr>
                    <td>1</td>
                    <td>Inner ramus of third uropod well developed</td>
                    <td>
                      <italic>
                        <tp:taxon-name>
                          <tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part>
                        </tp:taxon-name>
                      </italic>
                    </td>
                  </tr>
                  <tr>
                    <td>–</td>
                    <td>Inner ramus of third uropod reduced</td>
                    <td>2</td>
                  </tr>
                  <tr>
                    <td>2</td>
                    <td>Urosome segments 1 and 2 with dorsal protuberances bearing spines</td>
                    <td>
                      <italic>
                        <tp:taxon-name>
                          <tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part>
                        </tp:taxon-name>
                      </italic>
                    </td>
                  </tr>
                  <tr>
                    <td>–</td>
                    <td>Urosome segments 1 and 2 without dorsal protuberances bearing spines</td>
                    <td>3</td>
                  </tr>
                  <tr>
                    <td>3</td>
                    <td>Third uropod densely covered with recurved setae</td>
                    <td>
                      <italic>
                        <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name>
                      </italic>
                    </td>
                  </tr>
                  <tr>
                    <td>–</td>
                    <td>Third uropod lacking dense recurved setae</td>
                    <td>
                      <italic>
                        <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ischnus">ischnus</tp:taxon-name-part></tp:taxon-name>
                      </italic>
                    </td>
                  </tr>
                </tbody>
              </table>
            </table-wrap>
          </sec>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
    <sec sec-type="Discussion" id="sec7">
      <title>Discussion</title>
      <p>The native distribution region of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> includes the Black Sea, the Azov Sea, their deltas as well as the coastal lakes of Bulgaria and Romania (<xref ref-type="bibr" rid="B7">Cărăuşu et al. 1955</xref>; <xref ref-type="bibr" rid="B18">Mordukhaj-Boltovskoj 1969</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> is characterized by a relatively short life cycle and generation time, brood protection, and an omnivorous feeding strategy (<xref ref-type="bibr" rid="B20">Schmidt-Kloiber and Hering 2026</xref>). The species exhibits high ecological plasticity, being able to tolerate a broad range of environmental conditions, including wide salinity gradients (<xref ref-type="bibr" rid="B3">bij de Vaate et al. 2002</xref>). Outside its native range, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> has successfully colonized several European river systems, including the Upper Danube and the Lower Rhine, probably via the Main–Danube Canal (<xref ref-type="bibr" rid="B12">Holdich and Pöckl 2007</xref>). Despite its ecological plasticity, the species is only infrequently recorded in lake environments (but see <xref ref-type="bibr" rid="B19">Rewicz et al. 2016</xref>) and is generally considered to be more closely associated with large rivers (Labat et al. 2011; Info fauna 2026) where it occupies a variety of habitats, ranging from artificial riprap structures to more natural environments such as gravel banks, mud, roots, and stony substrates (<xref ref-type="bibr" rid="B6">Borza 2009</xref>).</p>
      <p>In the Rhone River Basin, records from the lower and middle French Rhone River suggest an ongoing expansion, with occurrences reported downstream of Lyon (France) at Péage-de-Roussillon, Baix-Logis-Neuf, Donzère-Mondragon and Montélimar between 2017 and 2022 in the main river channel and two active secondary channels (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Forcellini">Forcellini</tp:taxon-name-part></tp:taxon-name> pers. comm.). This indicates a slow and progressive upstream colonization of the Rhone River basin prior to its detection in Lake Geneva. Nevertheless, because the species has not been recorded in the French Upper Rhone, it is unlikely that its arrival in Lake Geneva originated from upstream dispersal through the French Rhone corridor. This hypothesis is further supported by the absence of records from the littoral zone of the western basin (i.e. small lake) of Lake Geneva, despite regular monitoring efforts since more than ten years. According to <xref ref-type="bibr" rid="B11">Gallardo and Aldridge (2013)</xref>, proximity to ports is a key predictor of the distribution of several Ponto-Caspian invasive species, including <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic>. Consequently, human-mediated transport appears to be the most plausible explanation for this finding.</p>
      <p>In newly colonized ecosystems, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> frequently co-occurs with other Ponto-Caspian amphipods, including <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ischnus">ischnus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="villosus">villosus</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="haemobaphes">haemobaphes</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chelicorophium">Chelicorophium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="robustum">robustum</tp:taxon-name-part></tp:taxon-name></italic>, as documented in the Rhine River (<xref ref-type="bibr" rid="B15">Labat et al. 2011</xref>). The present finding suggests that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> can coexist with related crustacean species already present in Lake Geneva as e.g. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="villosus">villosus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chelicorophium">Chelicorophium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="curvispinum">curvispinum</tp:taxon-name-part></tp:taxon-name></italic>. However, its establishment may also lead to future changes in community structure (<xref ref-type="bibr" rid="B6">Borza 2009</xref>). Competitive interactions are likely to occur within the bryophyte mats covering riprap and cobble habitats colonized by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic>, potentially affecting populations of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chelicorophium">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="curvispinum">curvispinum</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part></tp:taxon-name></italic> spp., as well as other bryophyte-associated aquatic macroinvertebrates such as insect larvae and small gastropods. Nevertheless, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chelicorophium">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="curvispinum">curvispinum</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dikerogammarus">Dikerogammarus</tp:taxon-name-part></tp:taxon-name></italic> spp. also inhabit deeper soft substrates and macrophytes. They generally occur there in lower numbers, but these habitats may provide alternative refuges and thereby limit the impact of competition with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic>.</p>
      <p>Beyond Lake Geneva and the Rhone River, the present record raises the possibility of further colonization of other peri-alpine and mountain lakes. Lake Geneva is surrounded by numerous lakes supporting recreational boating, with watercraft occasionally transferred between water bodies. If introduced, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> may find suitable habitats along bryophyte-covered rocky shores that are common in alpine lakes. This may be of particular concern for alpine lakes supporting glacial relict populations of native amphipods, such as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lacustris">lacustris</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B2">Alther et al. 2017</xref>), where the introduction of non-native species could threaten these conservation-important populations.</p>
      <p>The present finding has practical implications for the interpretation of previous surveys and future monitoring in Lake Geneva. Small individuals of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> may easily be mistaken for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part></tp:taxon-name></italic> spp. when examined only in lateral view, particularly during routine sorting of lake samples. Careful examination of the third uropod is therefore essential to distinguish <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">Chaetogammarus</tp:taxon-name-part></tp:taxon-name></italic> from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Gammarus">Gammarus</tp:taxon-name-part></tp:taxon-name></italic>, while the dense recurved setae on this appendage allows <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> to be separated from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ischnus">ischnus</tp:taxon-name-part></tp:taxon-name></italic>. Re-examination of preserved material could help determine whether <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chaetogammarus">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="trichiatus">trichiatus</tp:taxon-name-part></tp:taxon-name></italic> had been present in Lake Geneva before its first detection in 2024. In addition, targeted monitoring at sites located near the major ports of Lake Geneva may be particularly valuable for the early detection of newly introduced exotic species. The identification key provided here should facilitate the routine recognition of the species during macroinvertebrate surveys and reduce the risk of future misidentifications.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>The authors gratefully acknowledge all members of the Sage Environnement team for their assistance with field sampling, sample sorting, and specimen identification.</p>
    </ack>
    <ref-list>
      <title>References</title>
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        <mixed-citation>Cărăuşu S, Dobreanu E, Manolache C (1955) <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Amphipoda</tp:taxon-name-part></tp:taxon-name> forme salmastre şi de apă dulce. Fauna Republicii Populare Romîne, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subphylum">Crustacea</tp:taxon-name-part></tp:taxon-name> 4(4): 1–407.</mixed-citation>
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    <sec sec-type="Additional information" id="sec8">
      <title>Additional information</title>
      <sec sec-type="Conflict of interest" id="sec9">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="sec10">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Artificial Intelligence (AI) use" id="sec11">
        <title>Artificial Intelligence (AI) use</title>
        <p>The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.</p>
        <p>No AI tools were used in the preparation of this manuscript.</p>
      </sec>
      <sec sec-type="Funding" id="sec12">
        <title>Funding</title>
        <p>No funding was reported.</p>
      </sec>
      <sec sec-type="Author contributions" id="sec13">
        <title>Author contributions</title>
        <p>Conceptualization: SR. Data curation: CS. Investigation: PM. Methodology: SR, CS. Project administration: PM. Validation: PM, CS. Writing - original draft: PM. Writing - review and editing: SR, CS.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="sec14">
        <title>Author ORCIDs</title>
        <p>P. Marle <ext-link xlink:href="https://orcid.org/0000-0003-4464-9795" ext-link-type="uri">https://orcid.org/0000-0003-4464-9795</ext-link></p>
        <p>C. Stoffel <ext-link xlink:href="https://orcid.org/0009-0002-4538-9518" ext-link-type="uri">https://orcid.org/0009-0002-4538-9518</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="sec15">
        <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>
    <sec sec-type="supplementary-material">
      <title>Supplementary materials</title>
      <supplementary-material id="S1" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="arpha">81F9E117-A21F-56FB-8066-8CE3E36C29E9</object-id>
        <label>Supplementary material 1</label>
        <caption>
          <p>Fasta sequences</p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>docx</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p>Text.</p>
        </statement>
        <media xlink:href="alpineentomology-10-163-s001.docx" mimetype="application" mime-subtype="vnd.openxmlformats-officedocument.wordprocessingml.document" position="float" orientation="portrait" id="oo_1790822.docx">
          <uri content-type="original_file">https://binary.pensoft.net/file/1790822</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"> Pierre Marle, Céline Stoffel, Simon Renahy</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
