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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">2</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">ZooKeys</journal-title>
        <abbrev-journal-title xml:lang="en">ZK</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1313-2989</issn>
      <issn pub-type="epub">1313-2970</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/zookeys.1208.119147</article-id>
      <article-id pub-id-type="publisher-id">119147</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Achatinidae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Faunistics &amp; Distribution</subject>
          <subject>Nomenclature</subject>
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Neogene</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Italy</subject>
          <subject>World</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>﻿Five alien achatinid land snails (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Gastropoda</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="suborder">Eupulmonata</tp:taxon-name-part></tp:taxon-name>) first reported in greenhouses of Italian botanical gardens</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Manganelli</surname>
            <given-names>Giuseppe</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-8453-280X</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Benocci</surname>
            <given-names>Andrea</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0003-3145-215X</uri>
          <xref ref-type="aff" rid="A3">3</xref>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Barbato</surname>
            <given-names>Debora</given-names>
          </name>
          <email xlink:type="simple">debora.barbato@unisi.it</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-1105-1711</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Giusti</surname>
            <given-names>Folco</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-8722-4653</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Dipartimento di Scienze Fisiche, della Terra e dell’Ambiente, Università di Siena, Via Mattioli 4, 53100 Siena, Italy</addr-line>
        <institution>Università di Siena</institution>
        <addr-line content-type="city">Siena</addr-line>
        <country>Italy</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">NBFC (National Biodiversity Future Center), Palermo, Italy</addr-line>
        <institution>NBFC (National Biodiversity Future Center)</institution>
        <addr-line content-type="city">Palermo</addr-line>
        <country>Italy</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Museo di Storia Naturale dell’Accademia dei Fisiocritici, Piazzetta S. Gigli 2, 53100 Siena, Italy</addr-line>
        <institution>Museo di Storia Naturale dell’Accademia dei Fisiocritici</institution>
        <addr-line content-type="city">Siena</addr-line>
        <country>Italy</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Debora Barbato (<email xlink:type="simple">debora.barbato@unisi.it)</email></p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Eike Neubert</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2024</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>29</day>
        <month>07</month>
        <year>2024</year>
      </pub-date>
      <volume>1208</volume>
      <fpage>99</fpage>
      <lpage>132</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/DB1B34B8-CC15-5B53-8857-2128652E2F3E">DB1B34B8-CC15-5B53-8857-2128652E2F3E</uri>
      <uri content-type="zoobank" xlink:href="http://zoobank.org/BEF04EEA-B9D0-4220-9BC4-84208488CCF2">BEF04EEA-B9D0-4220-9BC4-84208488CCF2</uri>
      <history>
        <date date-type="received">
          <day>19</day>
          <month>01</month>
          <year>2024</year>
        </date>
        <date date-type="accepted">
          <day>04</day>
          <month>06</month>
          <year>2024</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Giuseppe Manganelli, Andrea Benocci, Debora Barbato, Folco Giusti</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">http://zoobank.org/BEF04EEA-B9D0-4220-9BC4-84208488CCF2</self-uri>
      <abstract>
        <label>﻿Abstract</label>
        <p>Plant trade and exchange for horticulture, recreation or research play a significant role in the dispersal of molluscs. Alien slugs and snails accidentally introduced into Europe have established rich communities in several countries, but although these introductions could have ecological and economic implications, mollusc xenodiversity in Italian botanical gardens, plant nurseries, and greenhouses has never been investigated. Facilities throughout the country were therefore visited between 2017 and 2023. Here the list of the achatinoidean species so far recorded from Italian greenhouses is provided, giving a short description of their diagnostic characters. The greenhouses of Trento and Padua host interesting assemblages of achatinoideans: the results of this study include the first reports of four achatinids, namely <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceum">achatinaceum</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> from Italy and the first record of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> from Europe. Reproductive anatomy of all species except <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> is illustrated in detail, integrating anatomical knowledge of this group of tiny molluscs, some of which are known mainly from their shell characters. The systematics of all the species is discussed, highlighting topics for future research (e.g., status of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mauritianum">mauritianum</tp:taxon-name-part></tp:taxon-name></italic>, real identity of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Helix">Helix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic>, anatomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic>, species-level taxonomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part></tp:taxon-name></italic>, systematic relationships and species-level taxonomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic>).</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>Geographical distribution</kwd>
        <kwd>non-indigenous molluscs</kwd>
        <kwd>shell and genital morphology</kwd>
        <kwd>taxonomy</kwd>
        <kwd>translocated species</kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>European Union - NextGenerationEU through the Italian Ministry of University and Research under PNRR - M4C2-I1.4 Project code: CN00000033, Title: NBFC - National Biodiversity Future Center</funding-statement>
      </funding-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="SECID0EJAAC">
        <title>Citation</title>
        <p>Manganelli G, Benocci A, Barbato D, Giusti F (2024) Five alien achatinid land snails (Gastropoda, Eupulmonata) first reported in greenhouses of Italian botanical gardens. ZooKeys 1208: 99–132. <ext-link xlink:type="simple" ext-link-type="doi" xlink:href="10.3897/zookeys.1208.119147">https://doi.org/10.3897/zookeys.1208.119147</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="﻿Introduction" id="SECID0E5AAC">
      <title>﻿Introduction</title>
      <p>An increasing number of organisms of limited mobility are rapidly spreading outside their native range and habitat (<xref ref-type="bibr" rid="B7">Bergey et al. 2014</xref>; <xref ref-type="bibr" rid="B108">Pergl et al. 2017</xref>; <xref ref-type="bibr" rid="B71">Hulme 2021</xref>) due to the global expansion of trade and transport of goods (<xref ref-type="bibr" rid="B69">Hulme 2009</xref>; <xref ref-type="bibr" rid="B143">Seebens et al. 2017</xref>). This may have dramatic implications for local biodiversity, food crops, human health, ecosystem processes and habitat degradation (<xref ref-type="bibr" rid="B77">Kenis et al. 2009</xref>; <xref ref-type="bibr" rid="B76">Keller et al. 2011</xref>). In temperate countries, a major component of xenodiversity consists of tropical species unwittingly introduced with plants and substrates for horticulture or recreation. Botanical gardens and hothouses play a major role in the accidental introduction of foreign species (e.g., <xref ref-type="bibr" rid="B157">Teodorescu and Matei 2010</xref>; <xref ref-type="bibr" rid="B62">Heywood 2011</xref>; <xref ref-type="bibr" rid="B155">Szczepkowski et al. 2014</xref>; <xref ref-type="bibr" rid="B70">Hulme 2015</xref>; <xref ref-type="bibr" rid="B81">Kolicka et al. 2015</xref>; <xref ref-type="bibr" rid="B174">Wang et al. 2015</xref>; <xref ref-type="bibr" rid="B74">Jaskuła et al. 2019</xref>).</p>
      <p>Alien slugs and snails have already established rich communities in European plant nurseries, greenhouses, and botanical gardens (e.g., <xref ref-type="bibr" rid="B37">Da Sois 2015</xref>; <xref ref-type="bibr" rid="B128">Reischütz et al. 2018</xref>; <xref ref-type="bibr" rid="B82">Kwitt et al. 2019</xref>) and new species arrive every year (e.g., <xref ref-type="bibr" rid="B172">von Proschwitz 2004</xref>; <xref ref-type="bibr" rid="B122">Preece and Naggs 2014</xref>; <xref ref-type="bibr" rid="B74">Jaskuła et al. 2019</xref>; <xref ref-type="bibr" rid="B129">Richling and von Proschwitz 2021</xref>). Land molluscs are easily transported with living plants via eggs in root balls or hatched individuals on leaves, stems, moss on pots, decaying plant matter on the soil surface or in the substrate (<xref ref-type="bibr" rid="B131">Robinson 1999</xref>; <xref ref-type="bibr" rid="B7">Bergey et al. 2014</xref>). However, despite the important ecological and economic implications of the introduction of alien gastropods, the non-native mollusc communities of Italian botanical gardens have not been thoroughly investigated. Therefore, our research group tried to fill this gap, visiting facilities throughout the country: here we report the rich assemblage of achatinoideans found in the first years of this study, including the first European record of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> (Swainson, 1840).</p>
      <p>Achatinoideans include several travelling snails (e.g., species that have the ecological flexibility to survive long journeys and spread and prosper in their new habitats; <xref ref-type="bibr" rid="B148">Smith 1989</xref>) that have been unintentionally introduced outside their native range through plant trade and exchanges: some have now achieved a pantropical distribution, while others, widespread in greenhouses and plant nurseries, have become cosmopolitan and occur in very high densities in such anthropogenic habitats. So far seven species have been recorded from European greenhouses and hothouses: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> (Potiez &amp; Michaud, 1838), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gracile">gracile</tp:taxon-name-part></tp:taxon-name></italic> (Hutton, 1834), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leptinaria">Leptinaria</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="unilamellata">unilamellata</tp:taxon-name-part></tp:taxon-name></italic> (d’Orbigny, 1838), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic> (Rang, 1831), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceum">achatinaceum</tp:taxon-name-part></tp:taxon-name></italic> (Pfeiffer, 1846), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> (Bruguière, 1789), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="striatella">striatella</tp:taxon-name-part></tp:taxon-name></italic> (Rang, 1831) (<xref ref-type="bibr" rid="B66">Horsák et al. 2020</xref>).</p>
      <p>As pointed out by <xref ref-type="bibr" rid="B66">Horsák et al. (2020)</xref> and <xref ref-type="bibr" rid="B129">Richling and von Proschwitz (2021)</xref>, it is not easy to distinguish these small similar-looking snails, especially for scientists in countries where such genera and species are not native. Tiny achatinoideans include many described taxa, the great majority of which live in tropical areas relatively neglected by research biologists. The taxonomy of this speciose group is still being defined and shell-based identifications are not considered very reliable (<xref ref-type="bibr" rid="B151">Solem 1989</xref>; <xref ref-type="bibr" rid="B100">Naggs 1994</xref>; <xref ref-type="bibr" rid="B171">von Proschwitz 1994</xref>; <xref ref-type="bibr" rid="B27">Cowie 1997</xref>; <xref ref-type="bibr" rid="B158">Thompson 2011</xref>; <xref ref-type="bibr" rid="B17">Brodie and Barker 2012</xref>; <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>; <xref ref-type="bibr" rid="B21">Budha et al. 2015</xref>; <xref ref-type="bibr" rid="B66">Horsák et al. 2020</xref>; <xref ref-type="bibr" rid="B129">Richling and von Proschwitz 2021</xref>). There is evidence that a conchological approach has often caused generic misallocations and species misidentifications (<xref ref-type="bibr" rid="B27">Cowie 1997</xref>, <xref ref-type="bibr" rid="B28">2000</xref>; <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>; <xref ref-type="bibr" rid="B21">Budha et al. 2015</xref>).</p>
      <p>Anatomical dissection of small achatinoideans is not easy: the proximal part of the penial complex is difficult to interpret since its components are so reduced and closely packed as to appear a single structure. Nor are anatomical differences between extant genera very evident. Nevertheless, although anatomical study may only provide partial solutions, it is absolutely necessary because generic taxa were defined on anatomical characters so that identification is only possible by this examination. The aim of the paper is to list the species of achatinoideans so far recorded from Italy, giving a short description of their diagnostic shell and anatomical characters in order to facilitate identification, and understanding of their taxonomic and nomenclatural framework.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0EHJAC">
      <title>﻿Materials and methods</title>
      <p>Between 2017 and 2023, the tropical greenhouses of seven facilities in central and northern Italy (<named-content xlink:type="simple" content-type="institution" xlink:href="http://grbio.org/institution/museo-delle-scienze-0" id="NCID0EDJAC">Science Museum of Trento</named-content> (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>); botanical gardens of Siena, Florence, Pisa, Padua, Bologna, and Milan) were visited and inspected for alien molluscs as part of the XenoDOMuS project on xenodiversity in tropical greenhouses of Italian botanical gardens and scientific museums.</p>
      <p>Snails and slugs were detected by visual search and by collecting leaf litter and surface soil. The visual search enabled detection of larger specimens, while collection of litter and soil revealed smaller species. The litter and soil were dried and sieved through decreasing mesh sizes. The coarser fraction was sorted visually, the others under a stereomicroscope. Although this method involves a huge investment in terms of sampling effort and time, it ensures more efficient collection of small mollusc species than a visual search.</p>
      <p>Identifications were based on morphological features (i.e., shell and anatomical characters). Live specimens were drowned in water, then fixed and preserved in 75% ethanol buffered with sodium carbonate. The bodies were dissected under a light microscope using fine-pointed watchmaker’s forceps. Anatomical organs were drawn using a camera lucida. Anatomical nomenclature usually followed the standard references for <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="suborder">Eupulmonata</tp:taxon-name-part></tp:taxon-name> (e.g., <xref ref-type="bibr" rid="B86">Manganelli et al. 2022</xref> and references therein); as regards the proximal penial complex, the terms epiphallus and penial caecum are in customary use in the “subulinids” (e.g., <xref ref-type="bibr" rid="B100">Naggs 1994</xref>; <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>).</p>
      <p>The descriptive terms (e.g., short vs long, slender vs wide) refer to comparisons of the same section of the genitalia in different taxa of achatinids. The directional terms proximal, basal, and initial denote the part closer to the gonad, whereas distal, apical, final, and terminal denote the part closer to the gonopore in the case of ducts of the main axis of the genitalia (e.g., free oviduct, vagina, vas deferens, penis). The same terms denote the part closer to (proximal, basal, or initial) or further from (distal, apical, final, or terminal) the main axis of the genitalia in the case of blind structures radiating from the main axis of the genitalia (bursa copulatrix, penial/vaginal/atrial appendix, etc.).</p>
      <p>The material is stored in the G. Manganelli collection (GMC; Dipartimento di Scienze Fisiche, della Terra e dell’Ambiente, Università di Siena, Italy).</p>
      <p>Acronyms of the shell variables:</p>
      <p><bold><abbrev xlink:title="aperture height" id="ABBRID0ELKAC">AH</abbrev></bold> aperture height,</p>
      <p><bold><abbrev xlink:title="aperture width" id="ABBRID0ESKAC">AW</abbrev></bold> aperture width,</p>
      <p><bold><abbrev xlink:title="shell diameter" id="ABBRID0EZKAC">SD</abbrev></bold> shell diameter,</p>
      <p><bold><abbrev xlink:title="shell height" id="ABBRID0EALAC">SH</abbrev></bold> shell height.</p>
      <p>Acronyms of the anatomical organs:</p>
      <p><bold><abbrev xlink:title="albumen gland" id="ABBRID0EILAC">AG</abbrev></bold> albumen gland,</p>
      <p><bold><abbrev xlink:title="bursa copulatrix (gametolytic gland)" id="ABBRID0EPLAC">BC</abbrev></bold> bursa copulatrix (gametolytic gland),</p>
      <p><bold><abbrev xlink:title="body wall" id="ABBRID0EWLAC">BW</abbrev></bold> body wall,</p>
      <p><bold><abbrev xlink:title="duct of bursa copulatrix" id="ABBRID0E4LAC">DBC</abbrev></bold> duct of bursa copulatrix,</p>
      <p><bold><abbrev xlink:title="distal penis" id="ABBRID0EEMAC">DP</abbrev></bold> distal penis,</p>
      <p><bold><abbrev xlink:title="distal vagina" id="ABBRID0ELMAC">DV</abbrev></bold> distal vagina,</p>
      <p><bold><abbrev xlink:title="egg/eggs in the uterine spermoviduct" id="ABBRID0ESMAC">Eg</abbrev></bold> egg/eggs in the uterine spermoviduct,</p>
      <p><bold><abbrev xlink:title="epiphallus" id="ABBRID0EZMAC">Ep</abbrev></bold> epiphallus,</p>
      <p><bold><abbrev xlink:title="first hermaphrodite duct" id="ABBRID0EANAC">FHD</abbrev></bold> first hermaphrodite duct,</p>
      <p><bold><abbrev xlink:title="free oviduct" id="ABBRID0EHNAC">FO</abbrev></bold> free oviduct,</p>
      <p><bold><abbrev xlink:title="genital atrium" id="ABBRID0EONAC">GA</abbrev></bold> genital atrium,</p>
      <p><bold><abbrev xlink:title="vaginal lateral bulge" id="ABBRID0EVNAC">LB</abbrev></bold> vaginal lateral bulge,</p>
      <p><bold>P</bold> penis,</p>
      <p><bold><abbrev xlink:title="penial caecum" id="ABBRID0EAOAC">PC</abbrev></bold> penial caecum,</p>
      <p><bold><abbrev xlink:title="proximal penis" id="ABBRID0EHOAC">PP</abbrev></bold> proximal penis,</p>
      <p><bold><abbrev xlink:title="penial retractor" id="ABBRID0EOOAC">PR</abbrev></bold> penial retractor,</p>
      <p><bold><abbrev xlink:title="penial sheath" id="ABBRID0EVOAC">PS</abbrev></bold> penial sheath,</p>
      <p><bold><abbrev xlink:title="prostatic spermoviduct" id="ABBRID0E3OAC">PSO</abbrev></bold> prostatic spermoviduct,</p>
      <p><bold><abbrev xlink:title="spermoviduct (ovispermiduct; second hermaphrodite duct)" id="ABBRID0EDPAC">SOD</abbrev></bold> spermoviduct (ovispermiduct; second hermaphrodite duct),</p>
      <p><bold><abbrev xlink:title="uterine spermoviduct" id="ABBRID0EKPAC">USO</abbrev></bold> uterine spermoviduct,</p>
      <p><bold>V</bold> vagina,</p>
      <p><bold><abbrev xlink:title="vas deferens" id="ABBRID0EVPAC">VD</abbrev></bold> vas deferens.</p>
    </sec>
    <sec sec-type="﻿Results" id="SECID0EZPAC">
      <title>﻿Results</title>
      <p>Non-native molluscs were observed in all the facilities investigated, but achatinids were only found in Padua (3 species: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceum">achatinaceum</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic>) and Trento (4 species: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic>). In these two greenhouses, high densities of small achatinids and other exotic slugs, snails, and soil invertebrates were found. The systematics, morphology, ecology, and distribution of the five species of achatinids are described below. A small undescribed slug of the little-known systellommatophoran family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rathouisiidae</tp:taxon-name-part></tp:taxon-name> found in the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>) has already been described (<xref ref-type="bibr" rid="B86">Manganelli et al. 2022</xref>).</p>
      <sec sec-type="﻿Family Achatinidae Swainson, 1840" id="SECID0E1CAE">
        <title>﻿Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name> Swainson, 1840</title>
        <p>The superfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Achatinoidea</tp:taxon-name-part></tp:taxon-name> Swainson, 1840 has a complex and still unresolved systematic framework. <xref ref-type="bibr" rid="B102">Nordsieck (1986)</xref> included five families in this group (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name> Swainson, 1840, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name> Bourguignat, 1883, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Subulinidae</tp:taxon-name-part></tp:taxon-name> Fischer &amp; Crosse, 1877, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Coeliaxidae</tp:taxon-name-part></tp:taxon-name> Pilsbry, 1907, and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Thyrophorellidae</tp:taxon-name-part></tp:taxon-name> Girard, 1895), while <xref ref-type="bibr" rid="B11">Bouchet and Rocroi (2005)</xref> listed only four (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Subulinidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Micractaeonidae</tp:taxon-name-part></tp:taxon-name> Schileyko, 1999). On the other hand, <xref ref-type="bibr" rid="B139">Schileyko (1999)</xref> only included the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name> in the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Achatinoidea</tp:taxon-name-part></tp:taxon-name>, and grouped <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Subulinidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Glessulidae</tp:taxon-name-part></tp:taxon-name> Godwin-Austen, 1920, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Micractaeonidae</tp:taxon-name-part></tp:taxon-name>, and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name> in their own superfamily, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Subulinoidea</tp:taxon-name-part></tp:taxon-name> Fischer &amp; Crosse, 1877. <xref ref-type="bibr" rid="B48">Fontanilla et al. (2017)</xref> published a multi-gene phylogenetic analysis of the achatinoideans examining 24 taxa from five achatinoid families (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Coeliaxidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Subulinidae</tp:taxon-name-part></tp:taxon-name>, and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Thyrophorellidae</tp:taxon-name-part></tp:taxon-name>). The results suggest that the family level systematics requires a radical re-evaluation because among the traditionally recognised families, only <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name> are monophyletic whereas <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Coeliaxidae</tp:taxon-name-part></tp:taxon-name>, and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Subulinidae</tp:taxon-name-part></tp:taxon-name> are polyphyletic or unresolved and members of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Coeliaxidae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Thyrophorellidae</tp:taxon-name-part></tp:taxon-name> cluster among the subulinids.</p>
        <p>In their “Revised classification, nomenclator and typification of gastropod and monoplacophoran families”, <xref ref-type="bibr" rid="B12">Bouchet et al. (2017)</xref>, following <xref ref-type="bibr" rid="B48">Fontanilla et al. (2017)</xref>, moved the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Subulinidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Coeliaxidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Thyrophorellidae</tp:taxon-name-part></tp:taxon-name>, and the ferussaciid genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> Férussac, 1814 to the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name>. However, in the absence of a more comprehensive molecular study of members of all the families and subfamilies previously recognised in the achatinoideans, they proposed a preliminary classification in which most of the subfamilies listed by <xref ref-type="bibr" rid="B139">Schileyko (1999)</xref> were treated as valid and allocated <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> to a distinct subfamily (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Cecilioidinae</tp:taxon-name-part></tp:taxon-name> Mörch, 1864). It is nonetheless possible that the removal of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> from the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name> was based on a misidentification of the species examined by <xref ref-type="bibr" rid="B48">Fontanilla et al. (2017)</xref> (E. Neubert, pers. comm. 7 May 2024). Indeed MolluscaBase (<xref ref-type="bibr" rid="B95">MolluscaBase eds 2024a</xref>) still lists <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> in the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name>, albeit citing <xref ref-type="bibr" rid="B12">Bouchet et al. (2017)</xref> as the source for the page dealing with <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name>. Thus, in the absence of a more consistent classification at subfamily level, the species considered in the present study are simply taken in alphabetical rather than systematic order, with the exception of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic> Crosse, 1867, traditionally considered a ferussaciid based solely on shell characters, which we consider last. The inclusion of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic> in the ferussaciids is based on its close shell similarity with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic>, but without anatomical and molecular evidence, its attribution to a different group/superfamily cannot be excluded.</p>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Stylommatophora</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Subulinidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">AC961840-E0DA-51D1-9C3B-7DCD7D15F454</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>(Potiez &amp; Michaud, 1838)</tp:taxon-authority>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Bulimus">Bulimus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinus">clavulinus</tp:taxon-name-part></tp:taxon-name>
                <comment>Potiez &amp; Michaud, 1838, 1: 136, pl. 14, figs 9, 10. Type locality: “L’île Bourbon”, namely Réunion Island, Mascarene Archipelago. Type material: lost (<xref ref-type="bibr" rid="B149">Smith 1992</xref>: 309).</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="material" id="SECID0ELNAE">
            <title>Material examined.</title>
            <p>Italy • 6 shells and 6 spirit specimens; Trento, Tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>); <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[11.113911,46.062544]}" id="NCID0EYNAE">46°03'45.16"N, 11°06'50.08"E</named-content></named-content>; 14 Dec. 2017; A. Benocci, G. Manganelli, D. Miserocchi leg.; GMC 47556 • 623 shells, 10 spirit specimens; same locality; 04 Jan. 2019, 10 Feb. 2019, 04 May 2019; D. Barbato, G. Bolzonella leg.; GMC 51194 • 418 shells, 2 spirit specimens; same locality; 01 Feb. 2022; D. Barbato, A. Benocci leg.; GMC 51184 • 133 shells; same locality; 02 Feb. 2023; F. Rossi leg.; GMC 57343 • 155 shells; same locality; 9 Feb. 2023; D. Barbato, A. Benocci leg.; GMC 57350 • 145 shells and 1 spirit specimen; Padua, Biodiversity Garden (Botanical Garden of Padua); <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[11.880658,45.397942]}" id="NCID0EAOAE">45°23'52.59"N, 11°52'50.37"E</named-content></named-content>; 06 Mar. 2019; D. Barbato leg.; GMC 57373.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EFOAE">
            <title>Description.</title>
            <p><bold><italic>Shell</italic></bold> (Figs <xref ref-type="fig" rid="F1">1–6</xref>). Dextral, small, minutely perforate to imperforate, elongate, slender, conical, rather robust, pearly off-white, glossy and sub-transparent when fresh, with 5–7 slightly convex whorls, separated by moderately deep sutures. Apex obtuse, rounded, and smooth. Last whorl ~ 1/2 of shell height. Aperture small, ~ 1/3 of shell height, obliquely pyriform, slightly prosocline. Peristome interrupted, not thickened, only slightly reflected on columella, sometimes with slightly evident callous rim on parietum; columella straight; outer margin sinuous in lateral view (approximately inverse S-shaped). Protoconch smooth; teleoconch with thin and irregular collabral growth lines. Shell dimensions: <abbrev xlink:title="shell height" id="ABBRID0EUOAE">SH</abbrev> 5.5–6.8 mm; <abbrev xlink:title="shell diameter" id="ABBRID0EYOAE">SD</abbrev> 2.2–2.6 mm; <abbrev xlink:title="aperture height" id="ABBRID0E3OAE">AH</abbrev> 1.9–2.3 mm; <abbrev xlink:title="aperture width" id="ABBRID0EAPAE">AW</abbrev> 1.2–1.4 mm.</p>
            <fig id="F1" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures1-8</object-id>
              <object-id content-type="arpha">5C758374-160B-5D67-A4C2-041DC4733B31</object-id>
              <label>Figures 1–8.</label>
              <caption>
                <p>Shells of alien achatinids in Italian greenhouses: <bold>1–6</bold> shells of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 May 2019 <bold>7, 8</bold> shells of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 May 2019.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103567.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103567</uri>
              </graphic>
            </fig>
            <p><bold><italic>Female distal genitalia</italic></bold> (Figs <xref ref-type="fig" rid="F2">9</xref>–<xref ref-type="fig" rid="F3">12</xref>). Free oviduct long and wide. Bursa copulatrix sac-like, oval with long slender duct (slightly longer than bursa copulatrix), sometimes initially flared. Vagina short and wide (approximately as long as free oviduct) with small lateral bulge.</p>
            <fig id="F2" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures9-11</object-id>
              <object-id content-type="arpha">7649642A-49E5-5145-88EF-22120B7BF3A2</object-id>
              <label>Figures 9–11.</label>
              <caption>
                <p>Genital anatomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 May 2019: <bold>9, 10</bold> genitalia (hermaphrodite gonad excluded) <bold>11</bold> distal genitalia.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103568.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103568</uri>
              </graphic>
            </fig>
            <p><bold><italic>Male distal genitalia</italic></bold> (Figs <xref ref-type="fig" rid="F2">9</xref>–<xref ref-type="fig" rid="F3">13</xref>, <xref ref-type="fig" rid="F4">15</xref>, <xref ref-type="fig" rid="F4">16</xref>). Vas deferens almost uniform in diameter (very thin to thin along its entire length), entering penial complex at its proximal end. Penial complex consisting of epiphallus, penial caecum and penis. Epiphallus very short. Penial caecum very short (as long as epiphallus). Penis short to rather long, almost uniformly cylindrical, undivided, with penial sheath enveloping its distal tract. Penial retractor muscle bifid, one branch inserted on proximal end of epiphallus, one branch on tip of penial caecum.</p>
            <fig id="F3" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures12-14</object-id>
              <object-id content-type="arpha">D1195957-08D5-552D-9A51-4E822225B94E</object-id>
              <label>Figures 12–14.</label>
              <caption>
                <p>Genital anatomy of alien achatinids in Italian greenhouses: <bold>12, 13</bold> genitalia (hermaphrodite gonad excluded) and diagram of the proximal penial complex of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 May 2019 <bold>14</bold> genitalia (hermaphrodite gonad excluded) of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 May 2019.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103569.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103569</uri>
              </graphic>
            </fig>
            <fig id="F4" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures15-16</object-id>
              <object-id content-type="arpha">60294E14-7E76-5659-A11F-D50BF0C7439F</object-id>
              <label>Figures 15, 16.</label>
              <caption>
                <p>Proximal penial complex of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 May 2019.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g004.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103570.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103570</uri>
              </graphic>
            </fig>
            <p><bold><italic>Genital atrium</italic></bold> (Figs <xref ref-type="fig" rid="F2">9</xref>–<xref ref-type="fig" rid="F3">12</xref>). Rather long.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="remarks" id="SECID0ETVAE">
            <title>Remarks.</title>
            <p>Although <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> is a well-known greenhouse snail (<xref ref-type="bibr" rid="B78">Kerney and Cameron 1979</xref>), its taxonomic and systematic placement is still not definitive, nor is its native range clear. The species was first described from Réunion Island (as Île Bourbon), Mascarene archipelago, in the western Indian Ocean (<xref ref-type="bibr" rid="B121">Potiez and Michaud 1838</xref>), but <xref ref-type="bibr" rid="B57">Griffiths and Florens (2006)</xref> thought it an East African species introduced into the Mascarene islands. The hypothesis that the species is native to Africa or East Africa was also maintained by <xref ref-type="bibr" rid="B78">Kerney and Cameron (1979)</xref>, <xref ref-type="bibr" rid="B131">Robinson (1999)</xref>, <xref ref-type="bibr" rid="B124">Probst (2001)</xref>, <xref ref-type="bibr" rid="B144">Shea (2007)</xref>, <xref ref-type="bibr" rid="B30">Cowie et al. (2008)</xref>, <xref ref-type="bibr" rid="B153">Stanisic et al. (2010)</xref>, and <xref ref-type="bibr" rid="B49">Foon et al. (2017)</xref>. However <xref ref-type="bibr" rid="B135">Rowson et al. (2010)</xref> observed that Verdcourt, familiar with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> in botanic gardens of the UK, never reported it from East Africa (e.g., <xref ref-type="bibr" rid="B164">Verdcourt 1983</xref>, <xref ref-type="bibr" rid="B166">2000</xref>, <xref ref-type="bibr" rid="B167">2006</xref>). Alternatively an Asian / south-east Asian origin was proposed by <xref ref-type="bibr" rid="B19">Brook et al. (2010)</xref> and <xref ref-type="bibr" rid="B136">Rumi et al. (2010)</xref>. Support for a south-east Asian origin could come from the putative finding of a shell in the Holocene of Thailand (<xref ref-type="bibr" rid="B130">Robba et al. 2007</xref>) and its membership of a molecularly based monophyletic group, including other species from Sri Lanka (<xref ref-type="bibr" rid="B48">Fontanilla et al. 2017</xref>).</p>
            <p>Today, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> occurs in humid tropical and subtropical lowlands across the world. It is reported from the West Indies, South America, West Indian Ocean islands, South and South-east Asia, New Guinea, Australia, and Pacific islands. Outside the tropics and subtropics it only occurs in heated greenhouses or very disturbed habitats, mainly in the northern hemisphere. Indeed it has been reported from North America, Europe, the Middle East, and New Zealand (see Table <xref ref-type="table" rid="T1">1</xref> for details and references). An alleged distinct subspecies <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="kyotoense">kyotoense</tp:taxon-name-part></tp:taxon-name></italic> (Pilsbry &amp; Hirase, 1904) is reported from Korea and Japan (<xref ref-type="bibr" rid="B91">Minato 1988</xref>; <xref ref-type="bibr" rid="B103">Noseworthy et al. 2007</xref>).</p>
            <table-wrap id="T1" position="float" orientation="portrait">
              <label>Table 1.</label>
              <caption>
                <p>Geographical distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic>. Asterisks indicate countries / islands where the species has been recorded only in greenhouses or very disturbed anthropogenic habitats.</p>
              </caption>
              <table id="TID0EAQDI" rules="all">
                <tbody>
                  <tr>
                    <th rowspan="1" colspan="1">Regions</th>
                    <th rowspan="1" colspan="1">Countries / Islands</th>
                    <th rowspan="1" colspan="1">References</th>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>North America</bold>
                    </td>
                    <td rowspan="1" colspan="1">United States*</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B40">Dundee (1974)</xref>, <xref ref-type="bibr" rid="B101">Nekola (2014)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>West Indies</bold>
                    </td>
                    <td rowspan="1" colspan="1">Hispaniola</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B44">Espinosa and Robinson (2021)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South America</bold>
                    </td>
                    <td rowspan="1" colspan="1">Brazil and Suriname</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B87">Marcus and Marcus (1968)</xref>, <xref ref-type="bibr" rid="B163">van Regteren Altena (1975)</xref>, <xref ref-type="bibr" rid="B145">Simone (2006)</xref>, <xref ref-type="bibr" rid="B136">Rumi et al. (2010)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Europe</bold>
                    </td>
                    <td rowspan="1" colspan="1">Austria*, Czech Republic*, Finland*, Germany*, Great Britain*, Ireland*, Italy*, Netherlands* and Sweden*</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B78">Kerney and Cameron (1979)</xref>, <xref ref-type="bibr" rid="B171">von Proschwitz (1994)</xref>, <xref ref-type="bibr" rid="B83">Leiss and Reischütz (1996)</xref>, <xref ref-type="bibr" rid="B65">Horsák et al. (2004</xref>, <xref ref-type="bibr" rid="B66">2020</xref>), <xref ref-type="bibr" rid="B37">Da Sois (2015)</xref>, <xref ref-type="bibr" rid="B128">Reischütz et al. (2018)</xref>, <xref ref-type="bibr" rid="B1">Anderson and Rowson (2020)</xref>, this paper</td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Indian Ocean</bold>
                    </td>
                    <td rowspan="1" colspan="1">Madagascar, Mascarene, and Seychelles</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B124">Probst (2001)</xref>, <xref ref-type="bibr" rid="B51">Gerlach (2006)</xref>, <xref ref-type="bibr" rid="B57">Griffiths and Florens (2006)</xref>, <xref ref-type="bibr" rid="B6">Bank and Menkhorst (2008)</xref>, <xref ref-type="bibr" rid="B42">Emberton et al. (2010)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>West Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Israel*</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B89">Mienis et al. (2012)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Nepal</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B21">Budha et al. (2015)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South-East Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Indonesia, Peninsular Malaysia, Philippines, Sabah in Malaysian Borneo and Singapore</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B63">Ho (1995)</xref>, <xref ref-type="bibr" rid="B49">Foon et al. (2017)</xref>, <xref ref-type="bibr" rid="B113">Phung et al. (2017)</xref>, <xref ref-type="bibr" rid="B104">Nurinsiyaha and Hausdorf (2019)</xref>, <xref ref-type="bibr" rid="B107">Parcon et al. (2020)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Oceania</bold>
                    </td>
                    <td rowspan="1" colspan="1">Australia, New Guinea, New Zealand* and Pacific islands (American Samoa, Chilean Islands: Rapa Nui, Cook Islands, Fiji, French Polynesia, Hawaii, Norfolk Island, Pitcairn Islands and Tonga)</td>
                    <td rowspan="1" colspan="1">Anonymous undated, <xref ref-type="bibr" rid="B159">van Benthem Jutting (1964)</xref>, <xref ref-type="bibr" rid="B151">Solem (1989)</xref>, <xref ref-type="bibr" rid="B28">Cowie (2000</xref>, <xref ref-type="bibr" rid="B29">2001</xref>), <xref ref-type="bibr" rid="B144">Shea (2007)</xref>, <xref ref-type="bibr" rid="B152">Stanisic et al. (2007</xref>, <xref ref-type="bibr" rid="B153">2010</xref>), <xref ref-type="bibr" rid="B19">Brook et al. (2010)</xref>, <xref ref-type="bibr" rid="B18">Brook (2014)</xref>, <xref ref-type="bibr" rid="B31">Cowie et al. (2017)</xref>, <xref ref-type="bibr" rid="B88">Maynard et al. (2018)</xref>, <xref ref-type="bibr" rid="B106">Osorio (2018)</xref></td>
                  </tr>
                </tbody>
              </table>
            </table-wrap>
            <p>The genital anatomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part></tp:taxon-name></italic> species is poorly understood. Earlier studies have been performed on specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gracile">gracile</tp:taxon-name-part></tp:taxon-name></italic> (Hutton, 1834) from Puerto Rico (<xref ref-type="bibr" rid="B5">Baker 1945</xref>; Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>) and Suriname (<xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> from Scotland (<xref ref-type="bibr" rid="B5">Baker 1945</xref>; Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>) and Brazil (<xref ref-type="bibr" rid="B87">Marcus and Marcus 1968</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mauritianum">mauritianum</tp:taxon-name-part></tp:taxon-name></italic> (Pfeiffer, 1853) from Mauritius (<xref ref-type="bibr" rid="B5">Baker 1945</xref>; Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part></tp:taxon-name></italic> “spec. 2 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part></tp:taxon-name></italic> spec. 3” from Pemba (<xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>). The overall distal genitalia organisation of our specimens (Figs <xref ref-type="fig" rid="F2">9</xref>–<xref ref-type="fig" rid="F3">12</xref>) is consistent with that described in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gracile">gracile</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B5">Baker 1945</xref>: 88–89; Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: 178, fig. 84.9; <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>: 255, fig. 10), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> (Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: fig. 84.6; <xref ref-type="bibr" rid="B87">Marcus and Marcus 1968</xref>: fig. 9), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part></tp:taxon-name></italic> spec. 2 (<xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>: 254–255, fig. 11) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part></tp:taxon-name></italic> spec. 3 (<xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>: 255, fig. 12). However the relationships between the penial sheath and the vas deferens seem different from what was illustrated by <xref ref-type="bibr" rid="B87">Marcus and Marcus (1968)</xref>, the only authors to describe them: according to <xref ref-type="bibr" rid="B87">Marcus and Marcus (1968</xref>: fig. 9) the vas deferens runs externally to the penial sheath whereas we found that it runs internally. The proximal penial complex consists of a short bulbous epiphallus and thin walled penial caecum with a branch of the penial retractor on the proximal tip of each (Figs <xref ref-type="fig" rid="F3">13</xref>, <xref ref-type="fig" rid="F4">15</xref>, <xref ref-type="fig" rid="F4">16</xref>). The epiphallus and the penial caecum are so closely juxtaposed as to resemble a usual proximal penis tip with an undivided penial retractor joined to it (Figs <xref ref-type="fig" rid="F2">9</xref>–<xref ref-type="fig" rid="F3">12</xref>). This arrangement matches that already described in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gracile">gracile</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B5">Baker 1945</xref>: 88; Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: 178, fig. 84.10), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B5">Baker 1945</xref>: 90; Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: 180, fig. 84.6; <xref ref-type="bibr" rid="B87">Marcus and Marcus 1968</xref>: fig. 9), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mauritianum">mauritianum</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B5">Baker 1945</xref>: 90; Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: 180) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part></tp:taxon-name></italic> spec. 2 (<xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>: 255, fig. 11b). However there seems to be some variation, especially in the structure of the penial caecum between these species: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gracile">gracile</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mauritianum">mauritianum</tp:taxon-name-part></tp:taxon-name></italic> have a robust penial caecum, large at the base and progressively tapering towards the tip (for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gracile">gracile</tp:taxon-name-part></tp:taxon-name></italic>, see Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: fig. 84.10; <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>; for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mauritianum">mauritianum</tp:taxon-name-part></tp:taxon-name></italic>, see Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: fig. 84.3), whereas <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> has a long slender penial caecum almost uniform in diameter, thin along its entire length (Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: fig. 84.4; <xref ref-type="bibr" rid="B87">Marcus and Marcus 1968</xref>: fig. 9). Based on penial caecum structure, our specimens apparently do not match those assigned to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> but are more similar to those assigned to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mauritianum">mauritianum</tp:taxon-name-part></tp:taxon-name></italic>. As already rightly observed by <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen (2013)</xref>, it is not clear whether these differences are due to individual variation or to phylogenetic divergence. The relationships between these taxa are also uncertain, as is whether these names have been used consistently in the literature. Although they are currently regarded as synonyms (<xref ref-type="bibr" rid="B96">MolluscaBase eds 2024b</xref>), their status and relationships are still not clear and may only be defined after the designation of neotypes and study of an appropriate number of populations using an integrative approach with conchological, anatomical and molecular characters. For now we consider this species according to its current concept (e.g., <xref ref-type="bibr" rid="B66">Horsák et al. 2020</xref>).</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulinum">clavulinum</tp:taxon-name-part></tp:taxon-name></italic> has been found in the tropical greenhouse of <named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>, where it is the commonest and most abundant achatinid species, and in the Biodiversity Garden (Botanical Garden of Padua), where it is rather uncommon. This is the first report from Italy.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Stylommatophora</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Subulinidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">ECB092BC-C5B8-515D-A42A-8B0AF2D7A34A</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>(Rang, 1831)</tp:taxon-authority>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Helix">Helix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannensis">hannensis</tp:taxon-name-part></tp:taxon-name>
                <comment>Rang, 1831: 41–42, pl. 3, fig. 8. Type locality: Senegal, Cape Verde Peninsula, Hann village (“village de Hann sur la presqu’île du Cap-Verd”). Type material: unknown.</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="material" id="SECID0EORAG">
            <title>Material examined.</title>
            <p>Italy • 47 shells and 1 spirit specimen; Trento, Tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>); <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[11.113911,46.062544]}" id="NCID0E2RAG">46°03'45.16"N, 11°06'50.08"E</named-content></named-content>; 4 Jan. 2019, 10 Feb. 2019, 04 May 2019; D. Barbato, G. Bolzonella leg.; GMC 51196 • 17 shells; same locality; 01 Feb. 2022; D. Barbato, A. Benocci leg.; GMC 51187 • 1 shell; same locality; 02 Feb. 2023; F. Rossi leg.; GMC 57345 • 6 shells; same locality; 09 Feb. 2023; D. Barbato, A. Benocci leg.; GMC 57352.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EASAG">
            <title>Description.</title>
            <p><bold><italic>Shell</italic></bold> (Figs <xref ref-type="fig" rid="F5">17–20</xref>). Dextral, very small in size, very minutely perforate, elongate, very slender, conical, rather robust, pearly off-white, glossy or waxy and sub-transparent when fresh, with 5–6 slightly convex whorls separated by moderately deep sutures. Apex obtuse, rounded, and smooth. Last whorl ~ ½ of shell height and less convex than preceding ones. Aperture small, ~ 1/3 of shell height, obliquely pyriform, slightly prosocline. Peristome interrupted, not thickened, only slightly reflected on columella, sometimes with subtle callous rim on parietum; columella straight; outer margin sinuous in lateral view (approximately inverse S-shaped). Protoconch smooth; teleoconch with weak irregular collabral growth lines. Shell dimensions: <abbrev xlink:title="shell height" id="ABBRID0EPSAG">SH</abbrev> 4.0–4.8 mm; <abbrev xlink:title="shell diameter" id="ABBRID0ETSAG">SD</abbrev> 1.6–1.8 mm; <abbrev xlink:title="aperture height" id="ABBRID0EXSAG">AH</abbrev> 1.6–1.7 mm; <abbrev xlink:title="aperture width" id="ABBRID0E2SAG">AW</abbrev> 0.9–1.0 mm.</p>
            <fig id="F5" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures17-20</object-id>
              <object-id content-type="arpha">070A9A95-5F33-5E2B-B827-52EF611438E9</object-id>
              <label>Figures 17–20.</label>
              <caption>
                <p>Shells of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 May 2019. Shell material of this species belonged mainly to juveniles; the shells depicted in the figures were the best available, although some are from specimens that were not fully grown.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g005.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103571.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103571</uri>
              </graphic>
            </fig>
            <p><bold><italic>Female distal genitalia</italic></bold> (Fig. <xref ref-type="fig" rid="F3">14</xref>). Free oviduct very short and wide. Bursa copulatrix sac-like, oval with short slender duct (as long as bursa copulatrix), initially not flared. Vagina very short and wide (as long as free oviduct).</p>
            <p><bold><italic>Male distal genitalia</italic></bold> (Fig. <xref ref-type="fig" rid="F3">14</xref>). Vas deferens almost uniform in diameter (very thin to thin along its entire length), entering penial complex at its proximal end. Penial complex apparently consisting only of penis. Penis short, almost uniformly cylindrical with thin short penial sheath enveloping its distal tract. Penial retractor muscle inserted on proximal end of penis.</p>
            <p><bold><italic>Genital atrium</italic></bold> (Fig. <xref ref-type="fig" rid="F3">14</xref>). Rather long.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="remarks" id="SECID0E3UAG">
            <title>Remarks.</title>
            <p>The species was first named <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Helix">Helix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="clavulus">clavulus</tp:taxon-name-part></tp:taxon-name></italic> by Férussac (1821: 52) based on specimens from Guadeloupe and then <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Helix">Helix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="goodalli">goodalli</tp:taxon-name-part></tp:taxon-name></italic> by <xref ref-type="bibr" rid="B90">Miller (1822</xref>: 381) on specimens from near Bristol, England. Unfortunately Férussac’s name was not accompanied by a description, a definition, or an indication and so it is not available, whereas Miller’s name, extensively used until the early 20<sup>th</sup> century (cf. <xref ref-type="bibr" rid="B115">Pilsbry 1906b</xref>), turned out to be a junior homonym and was replaced by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Bulimus">Bulimus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pumilus">pumilus</tp:taxon-name-part></tp:taxon-name></italic> established by <xref ref-type="bibr" rid="B109">Pfeiffer (1840)</xref> on specimens from Cuba (cf. <xref ref-type="bibr" rid="B118">Pilsbry 1910</xref>). <xref ref-type="bibr" rid="B114">Pilsbry (1906a</xref>: 141–142) also discussed the hypothesis of <xref ref-type="bibr" rid="B177">Wollaston (1878</xref>: 510) that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Helix">Helix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="goodalli">goodalli</tp:taxon-name-part></tp:taxon-name></italic> was a junior synonym of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Helix">Helix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic> established by <xref ref-type="bibr" rid="B127">Rang (1831)</xref> on specimens from the Cape Verde Peninsula, Senegal, observing: “whether this course was well-founded is a question which must remain unsettled until specimens from Rang’s original locality can be compared.” Consequently he never adopted Rang’s name for this species (cf. <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: 181–182). The synonymy of the two species was reproposed by <xref ref-type="bibr" rid="B58">Groh (1983)</xref> based on study of the original descriptions and the literature, and has subsequently been adopted by most recent authors (e.g., <xref ref-type="bibr" rid="B171">von Proschwitz 1994</xref>; <xref ref-type="bibr" rid="B27">Cowie 1997</xref>, <xref ref-type="bibr" rid="B28">2000</xref>; <xref ref-type="bibr" rid="B23">Chase and Robinson 2001</xref>; <xref ref-type="bibr" rid="B6">Bank and Menkhorst 2008</xref>; <xref ref-type="bibr" rid="B50">Gerber and Clark 2015</xref>; <xref ref-type="bibr" rid="B66">Horsák et al. 2020</xref>). Perplexity persists about the real identity of the species described by Rang. Only the dimensions, which are consistent with those of an <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part></tp:taxon-name></italic> species, support Groh’s interpretation. Otherwise the situation remains as described by Pilsbry more than a hundred years ago: type material of Rang’s species is unknown; no one has reported or studied material from the type locality, which when Rang visited it, was a small village, today englobed in the city of Dakar (where a green area, the Parc forestier et zoologique de Hann, still survives in Hann); finally Rang’s description and illustration are completely inadequate to establish the identity of the species he treated; his figure depicts a snail with shell having all the whorls quite round, whereas this species has the last whorl almost flat (incidentally <xref ref-type="bibr" rid="B131">Robinson (1999</xref>: table 1) considered <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic> to be absent from Africa).</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic> is regarded as native to tropical America (<xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>; <xref ref-type="bibr" rid="B78">Kerney and Cameron 1979</xref>; <xref ref-type="bibr" rid="B38">Deisler and Abbott 1984</xref>; <xref ref-type="bibr" rid="B27">Cowie 1997</xref>; <xref ref-type="bibr" rid="B30">Cowie et al. 2008</xref>; <xref ref-type="bibr" rid="B19">Brook et al. 2010</xref>; <xref ref-type="bibr" rid="B92">Miquel and Herrera 2014</xref>) where it is widespread in Central America and the West Indies. On the contrary <xref ref-type="bibr" rid="B131">Robinson (1999</xref>: Table <xref ref-type="table" rid="T1">1</xref>) regarded it as native to East Asia. It has been introduced into South America, Atlantic islands, East Africa, West Indian Ocean islands, South-east Asia, and Pacific islands. It has also been reported from the mid temperate latitudes of the northern hemisphere where it only occurs in greenhouses and hothouses (see Table <xref ref-type="table" rid="T2">2</xref> for details and references). Since these reports are only based on shell identifications, it is not possible to exclude that some are misidentifications (e.g., <xref ref-type="bibr" rid="B99">Muratov 2010</xref>: fig. 28).</p>
            <table-wrap id="T2" position="float" orientation="portrait">
              <label>Table 2.</label>
              <caption>
                <p>Geographical distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic>. Asterisks indicate countries / islands where the species has been recorded only in greenhouses or very disturbed anthropogenic habitats.</p>
              </caption>
              <table id="TID0EHZDI" rules="all">
                <tbody>
                  <tr>
                    <th rowspan="1" colspan="1">Regions</th>
                    <th rowspan="1" colspan="1">Countries / Islands</th>
                    <th rowspan="1" colspan="1">References</th>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>North America</bold>
                    </td>
                    <td rowspan="1" colspan="1">United States*</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B40">Dundee (1974)</xref>, <xref ref-type="bibr" rid="B101">Nekola (2014)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Central America</bold>
                    </td>
                    <td rowspan="1" colspan="1">Guatemala, Mexico, Nicaragua, and Panama</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B158">Thompson (2011)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>West Indies</bold>
                    </td>
                    <td rowspan="1" colspan="1">Bahamas, Cuba, Hispaniola, Jamaica, Lesser Antilles (Barbados, Curaçao, Guadeloupe, Martinique, and Saint Martin)</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B38">Deisler and Abbott (1984)</xref>, <xref ref-type="bibr" rid="B23">Chase and Robinson (2001)</xref>, <xref ref-type="bibr" rid="B133">Rosenberg and Muratov (2006)</xref>, <xref ref-type="bibr" rid="B43">Espinosa and Ortea (2009)</xref>, <xref ref-type="bibr" rid="B85">Maceira et al. (2013)</xref>, <xref ref-type="bibr" rid="B22">Charles (2015)</xref>, <xref ref-type="bibr" rid="B39">Delannoye et al. (2015)</xref>, <xref ref-type="bibr" rid="B68">Hovestadt and van Leeuwen (2017)</xref>, <xref ref-type="bibr" rid="B67">Hovestadt and Neckheim (2020)</xref>, <xref ref-type="bibr" rid="B44">Espinosa and Robinson (2021)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South America</bold>
                    </td>
                    <td rowspan="1" colspan="1">Argentina, Brazil, Ecuador, Suriname, and Uruguay</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B163">van Regteren Altena (1975)</xref>, <xref ref-type="bibr" rid="B145">Simone (2006)</xref>, <xref ref-type="bibr" rid="B136">Rumi et al. (2010)</xref>, <xref ref-type="bibr" rid="B169">Virgillito and Miquel (2013)</xref>, <xref ref-type="bibr" rid="B93">Miquel and Jaime (2018)</xref>, <xref ref-type="bibr" rid="B15">Breure et al. (2022a)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Atlantic Ocean</bold>
                    </td>
                    <td rowspan="1" colspan="1">Bermuda, Capo Verde, Saint Helena, and São Tomé</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B34">Crowley and Pain (1977)</xref>, <xref ref-type="bibr" rid="B58">Groh (1983)</xref>, <xref ref-type="bibr" rid="B9">Bieler and Slapcinsky (2000)</xref>, <xref ref-type="bibr" rid="B64">Holyoak et al. (2020)</xref>, <xref ref-type="bibr" rid="B80">Key et al. (2021)</xref>, <xref ref-type="bibr" rid="B123">Preece et al. (2022)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Europe</bold>
                    </td>
                    <td rowspan="1" colspan="1">Austria*, Czech Republic*, Denmark*, France*, Germany*, Great Britain*, Ireland*, Italy*, Netherlands* and Sweden*</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B78">Kerney and Cameron (1979)</xref>, <xref ref-type="bibr" rid="B171">von Proschwitz (1994)</xref>, <xref ref-type="bibr" rid="B83">Leiss and Reischütz (1996)</xref>, <xref ref-type="bibr" rid="B65">Horsák et al. (2004</xref>, <xref ref-type="bibr" rid="B66">2020</xref>), <xref ref-type="bibr" rid="B128">Reischütz et al. (2018)</xref>, <xref ref-type="bibr" rid="B82">Kwitt et al. (2019)</xref>, <xref ref-type="bibr" rid="B1">Anderson and Rowson (2020)</xref>, this paper</td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Africa</bold>
                    </td>
                    <td rowspan="1" colspan="1">Mozambique</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B99">Muratov (2010)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Indian Ocean</bold>
                    </td>
                    <td rowspan="1" colspan="1">Madagascar and Seychelles</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B51">Gerlach (2006)</xref>, <xref ref-type="bibr" rid="B6">Bank and Menkhorst (2008)</xref>, <xref ref-type="bibr" rid="B42">Emberton et al. (2010)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South-East Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Singapore</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B63">Ho (1995)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Oceania</bold>
                    </td>
                    <td rowspan="1" colspan="1">Pacific Islands (American Samoa, Belau/Palau, Cook Islands, Federated States of Micronesia, Fiji, French Polynesia, Galapagos Islands, Guam, Hawaii, Pitcairn Islands, Samoa, Solomon Islands, Tonga, and Vanuatu)</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B151">Solem (1989)</xref>, <xref ref-type="bibr" rid="B28">Cowie (2000</xref>, <xref ref-type="bibr" rid="B29">2001</xref>), <xref ref-type="bibr" rid="B19">Brook et al. (2010)</xref>, <xref ref-type="bibr" rid="B18">Brook (2014)</xref>, <xref ref-type="bibr" rid="B92">Miquel and Herrera (2014)</xref>, <xref ref-type="bibr" rid="B31">Cowie et al. (2017)</xref></td>
                  </tr>
                </tbody>
              </table>
            </table-wrap>
            <p>The genital anatomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part></tp:taxon-name></italic> species was investigated by <xref ref-type="bibr" rid="B5">Baker (1945)</xref>, Baker in <xref ref-type="bibr" rid="B119">Pilsbry (1946)</xref>, and <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen (2013)</xref>. Only three <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part></tp:taxon-name></italic> species have been studied: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic> (see <xref ref-type="bibr" rid="B5">Baker 1945</xref>: 86, as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pumilum">pumilum</tp:taxon-name-part></tp:taxon-name></italic>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pyrgula">pyrgula</tp:taxon-name-part></tp:taxon-name></italic> (Schmacker &amp; Boettger, 1891) (see <xref ref-type="bibr" rid="B5">Baker 1945</xref>: 87; Baker in <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: 183–184, figs 88 2, 3) and an unidentified species from Misali islet, Zanzibar (see <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen 2013</xref>: 251, fig. 9), but only the anatomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pyrgula">pyrgula</tp:taxon-name-part></tp:taxon-name></italic> is adequately described. The distal genitalia of the only adult specimen that we have been able to study (Fig. <xref ref-type="fig" rid="F3">14</xref>) agrees with the general scheme described for these species and in particular with the features reported by <xref ref-type="bibr" rid="B5">Baker (1945)</xref> and Baker in <xref ref-type="bibr" rid="B119">Pilsbry (1946)</xref> for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pumilum">pumilum</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pyrgula">pyrgula</tp:taxon-name-part></tp:taxon-name></italic> and with the description of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pumilum">pumilum</tp:taxon-name-part></tp:taxon-name></italic> given by <xref ref-type="bibr" rid="B5">Baker (1945)</xref>. The major difference between the two species consists in the swelling between the base of the duct of the bursa copulatrix and the proximal vagina: well developed in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pyrgula">pyrgula</tp:taxon-name-part></tp:taxon-name></italic> and much less enlarged in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="pumilum">pumilum</tp:taxon-name-part></tp:taxon-name></italic>.</p>
            <p>Thus, little continues to be known about the genital anatomy of this genus. We need to ascertain whether the proximal complex of the penis is really undivided, to understand the relationships between the vas deferens and the penial sheath and whether the different structure of the female distal genitalia is due to individual variation or to phylogenetic divergence.</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Opeas">Opeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hannense">hannense</tp:taxon-name-part></tp:taxon-name></italic> has only been found in the tropical greenhouse of <named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>, where it is uncommon. This is the first report from Italy.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Stylommatophora</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Subulinidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">63A046FF-D7BC-53FD-BB21-A07CD63234F0</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceum">achatinaceum</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>(Pfeiffer, 1846)</tp:taxon-authority>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Bulimus">Bulimus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceus">achatinaceus</tp:taxon-name-part></tp:taxon-name>
                <comment>Pfeiffer, 1846: 82. Type locality: “Java”. Type material: lectotype no. ZMB Moll. 65746, Zoological Museum, Berlin (<xref ref-type="bibr" rid="B100">Naggs 1994</xref>: fig. 1).</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="material" id="SECID0ENNBG">
            <title>Material examined.</title>
            <p>Italy • 173 shells and 21 spirit specimens; Padua, Biodiversity Garden (Botanical Garden of Padua); <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[11.880658,45.397942]}" id="NCID0EWNBG">45°23'52.59"N, 11°52'50.37"E</named-content></named-content>; 06 Mar. 2019; D. Barbato leg.; GMC 57373.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0E2NBG">
            <title>Description.</title>
            <p><bold><italic>Shell</italic></bold> (Figs <xref ref-type="fig" rid="F6">21–24</xref>). Dextral, medium in size, minutely perforate to imperforate, elongate, slender, conical, rather robust, pearly off-white, opaque, with 7–9 slightly convex whorls, separated by rather deep and in places irregularly crenulate sutures. Apex obtuse, rounded, and smooth. Last whorl ~ 1/3 of shell height. Aperture small, ~ 1/4 of shell height, obliquely pyriform, slightly prosocline. Peristome interrupted, slightly thickened along outer margin, slightly reflected on columella, with callous rim on parietum and columella; columella straight; outer margin sinuous in lateral view (approximately inverse S-shaped). Protoconch smooth; teleoconch with evident irregular collabral striae. Shell dimensions: <abbrev xlink:title="shell height" id="ABBRID0EKOBG">SH</abbrev> 9.4–13.1 mm; <abbrev xlink:title="shell diameter" id="ABBRID0EOOBG">SD</abbrev> 2.9–3.8 mm; <abbrev xlink:title="aperture height" id="ABBRID0ESOBG">AH</abbrev> 3.0–3.5 mm; <abbrev xlink:title="aperture width" id="ABBRID0EWOBG">AW</abbrev> 1.8–2.2 mm.</p>
            <fig id="F6" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures21-24</object-id>
              <object-id content-type="arpha">06F0F8D5-E651-582F-959E-9C5340EB47A1</object-id>
              <label>Figures 21–24.</label>
              <caption>
                <p>Shells of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceum">achatinaceum</tp:taxon-name-part></tp:taxon-name></italic> from the Biodiversity Garden (Botanical Garden of Padua), D. Barbato leg. 06 Mar. 2019.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g006.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103572.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103572</uri>
              </graphic>
            </fig>
            <p><bold><italic>Female distal genitalia</italic></bold> (Figs <xref ref-type="fig" rid="F7">25</xref>, <xref ref-type="fig" rid="F7">27</xref>). Free oviduct long and wide. Bursa copulatrix sac-like, oval with long slender duct (slightly longer than bursa copulatrix), initially slightly flared, and medially convoluted around free oviduct. Vagina long and wide (longer than free oviduct) with huge proximal lateral bulge containing large ligula.</p>
            <fig id="F7" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures25-27</object-id>
              <object-id content-type="arpha">A6727875-FD82-5952-9853-6BCD9F6797AC</object-id>
              <label>Figures 25–27.</label>
              <caption>
                <p>Genital anatomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceum">achatinaceum</tp:taxon-name-part></tp:taxon-name></italic> from the Biodiversity Garden (Botanical Garden of Padua), D. Barbato leg. 06 Mar. 2019: <bold>25</bold> genitalia (hermaphrodite gonad excluded) <bold>26</bold> detail of proximal penial complex <bold>27</bold> internal structure of vagina.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g007.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103573.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103573</uri>
              </graphic>
            </fig>
            <p><bold><italic>Male distal genitalia</italic></bold> (Figs <xref ref-type="fig" rid="F7">25–27</xref>). Vas deferens of variable diameter (proximal tract narrow, medial tract slightly wider and final tract very narrow), entering penial complex at its proximal end. Penial complex consisting of epiphallus, penial caecum and penis. Epiphallus very short. Penial caecum very short (as long as epiphallus). Penis very long, distinctly divided into proximal and distal parts by difference in calibre; proximal part longer (almost twice distal penis), slender and thin walled; distal penis shorter (half proximal penis), thick, muscular walled and enveloped by penial sheath. Penial retractor muscle bifid, one branch inserted on proximal end of penis, the other branch on tip of penial caecum.</p>
            <p><bold><italic>Genital atrium</italic></bold> (Figs <xref ref-type="fig" rid="F7">25</xref>, <xref ref-type="fig" rid="F7">27</xref>). Short.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="remarks" id="SECID0EZRBG">
            <title>Remarks.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceum">achatinaceum</tp:taxon-name-part></tp:taxon-name></italic> is one of the best known subulinids thanks to the excellent anatomical study and the careful taxonomic revision by <xref ref-type="bibr" rid="B100">Naggs (1994)</xref>. Our anatomical study (Figs <xref ref-type="fig" rid="F7">25–27</xref>) fully agrees with that of <xref ref-type="bibr" rid="B100">Naggs (1994)</xref>.</p>
            <p>This species is regarded as native to tropical Asia, where it occurs from Nepal and Sri Lanka to South East Asia. Outside this area it is found in Australia and Pacific islands, West Indian Ocean islands, Europe, and the West Asia (see Table <xref ref-type="table" rid="T3">3</xref> for details and references).</p>
            <table-wrap id="T3" position="float" orientation="portrait">
              <label>Table 3.</label>
              <caption>
                <p>Geographical distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="achatinaceum">achatinaceum</tp:taxon-name-part></tp:taxon-name></italic>. Asterisks indicate countries / islands where the species has been recorded only in greenhouses or very disturbed anthropogenic habitats; hash symbol denotes one record based on a specimen of unknown origin recovered from sandy detritus collected on a beach.</p>
              </caption>
              <table id="TID0ESBAK" rules="all">
                <tbody>
                  <tr>
                    <th rowspan="1" colspan="1">Regions</th>
                    <th rowspan="1" colspan="1">Countries / Islands</th>
                    <th rowspan="1" colspan="1">References</th>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Europe</bold>
                    </td>
                    <td rowspan="1" colspan="1">Austria*, Italy*, and Malta<bold><sup>#</sup></bold></td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B66">Horsák et al. (2020)</xref>, <xref ref-type="bibr" rid="B24">Cilia et al. (2022)</xref>, this paper</td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>West Indian Ocean</bold>
                    </td>
                    <td rowspan="1" colspan="1">Mascarene and Seychelles</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B100">Naggs (1994)</xref>, <xref ref-type="bibr" rid="B57">Griffiths and Florens (2006)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>West Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Iraq</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B72">Hussein et al. (2018)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Nepal and Sri Lanka</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B100">Naggs (1994)</xref>, <xref ref-type="bibr" rid="B21">Budha et al. (2015)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South-East Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Borneo, Indonesia, and Singapore</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B100">Naggs (1994)</xref>, <xref ref-type="bibr" rid="B141">Schilthuizen and Rutjes (2001)</xref>, <xref ref-type="bibr" rid="B156">Tan et al. (2015)</xref>, <xref ref-type="bibr" rid="B105">Nurinsiyaha et al. (2016)</xref>, <xref ref-type="bibr" rid="B113">Phung et al. (2017)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>East Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Dongsha Island and Hong Kong</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B100">Naggs (1994)</xref>, <xref ref-type="bibr" rid="B178">Wu et al. (2007)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Oceania</bold>
                    </td>
                    <td rowspan="1" colspan="1">Australia and Pacific islands (American Samoa, Cook Islands, Fiji, French Polynesia, Guam, Hawaii, Northern Mariana Islands, Samoa, and Tonga)</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B100">Naggs (1994)</xref>, <xref ref-type="bibr" rid="B28">Cowie (2000</xref>, <xref ref-type="bibr" rid="B29">2001</xref>), <xref ref-type="bibr" rid="B144">Shea (2007)</xref>, <xref ref-type="bibr" rid="B19">Brook et al. (2010)</xref>, <xref ref-type="bibr" rid="B153">Stanisic et al. (2010)</xref>, <xref ref-type="bibr" rid="B79">Kerr and Bauman (2013)</xref>, <xref ref-type="bibr" rid="B18">Brook (2014)</xref>, <xref ref-type="bibr" rid="B31">Cowie et al. (2017)</xref></td>
                  </tr>
                </tbody>
              </table>
            </table-wrap>
            <p>The species was found in the Biodiversity Garden (Botanical Garden of Padua), where it forms a well-established population, as in Vienna Zoo, Austria (<xref ref-type="bibr" rid="B66">Horsák et al. 2020</xref>). In contrast, the report from Malta is only based on a specimen of unknown origin recovered from sandy detritus collected on a beach (<xref ref-type="bibr" rid="B24">Cilia et al. 2022</xref>). This is the first report from Italy and the third from Europe.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Stylommatophora</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Subulinidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">C3F6AA0A-9993-54F7-8563-FC134A3EC4D8</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>(Bruguière, 1789)</tp:taxon-authority>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Bulimus">Bulimus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octonus">octonus</tp:taxon-name-part></tp:taxon-name>
                <comment>Bruguière, 1789: 325. Type locality: “… dans les îles Antilles. M. de Badier l’a trouvé abondamment à l’île de la Guadeloupe, &amp; j’en ai vu chez M. d’Antic qui lui ont été envoyés de l’île de Saint-Domingue”. Type material: presumed lost (<xref ref-type="bibr" rid="B149">Smith 1992</xref>).</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="material" id="SECID0E12BG">
            <title>Material examined.</title>
            <p>Italy • 112 shells and 2 spirit specimens; Trento, Tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>); <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[11.113911,46.062544]}" id="NCID0EH3BG">46°03'45.16"N, 11°06'50.08"E</named-content></named-content>; 04 Jan. 2019; 10 Feb. 2019; 04 May 2019; D. Barbato, G. Bolzonella leg.; GMC 51197 • 29 shells; same locality; 01 Feb. 2022; D. Barbato, A. Benocci leg.; GMC 51188 • 1 shell; same locality; 09 Feb. 2023; D. Barbato, A. Benocci leg.; GMC 57353 • 1 shell; Padua, Biodiversity Garden (Botanical Garden of Padua); <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[11.880658,45.397942]}" id="NCID0EP3BG">45°23'52.59"N, 11°52'50.37"E</named-content></named-content>; 06 Mar. 2019; D. Barbato leg.; GMC 57374.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EU3BG">
            <title>Description.</title>
            <p><bold><italic>Shell</italic></bold> (Figs <xref ref-type="fig" rid="F1">7</xref>, <xref ref-type="fig" rid="F1">8</xref>, <xref ref-type="fig" rid="F8">28–30</xref>). Shell dextral, medium in size, imperforate, elongate, slender, conical, rather robust, pearly off-white, glossy or waxy and sub-transparent when fresh, with 7–9 slightly convex whorls, separated by rather deep and in places irregularly crenulate sutures. Apex obtuse, rounded, and smooth; last whorl ~ 1/3 of shell height. Aperture small, ~ 1/5 of shell height, ovate, slightly prosocline. Peristome interrupted, not thickened or reflected, with callous rim on parietum and columella; columella concave and obliquely truncate at base; outer margin straight in lateral view. Protoconch smooth; teleoconch with fine, wrinkled, irregular collabral growth lines. Shell dimensions: <abbrev xlink:title="shell height" id="ABBRID0EL4BG">SH</abbrev> 12.1–16.0 mm; <abbrev xlink:title="shell diameter" id="ABBRID0EP4BG">SD</abbrev> 3.3–4.1 mm; <abbrev xlink:title="aperture height" id="ABBRID0ET4BG">AH</abbrev> 2.5–3.2 mm; <abbrev xlink:title="aperture width" id="ABBRID0EX4BG">AW</abbrev> 1.9–2.4 mm.</p>
            <fig id="F8" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures28-30</object-id>
              <object-id content-type="arpha">4D1CB999-17E6-57AB-A902-663EE6296B10</object-id>
              <label>Figures 28–30.</label>
              <caption>
                <p>Shells of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 Jan. 2019.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g008.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103574.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103574</uri>
              </graphic>
            </fig>
            <p><bold><italic>Female distal genitalia</italic></bold> (Fig. <xref ref-type="fig" rid="F9">31</xref>). Free oviduct very short and wide. Bursa copulatrix sac-like, oval with short slender duct (as long as bursa copulatrix), initially not flared. Vagina very long and slender.</p>
            <fig id="F9" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures31-32</object-id>
              <object-id content-type="arpha">CD9DB10B-FE5B-59EB-AE5B-365EBB016EFC</object-id>
              <label>Figures 31, 32.</label>
              <caption>
                <p>Genital anatomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; G. Bolzonella leg. 04 Jan. 2019: <bold>31</bold> genitalia (hermaphrodite gonad excluded) <bold>32</bold> detail of proximal penial complex.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g009.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103575.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103575</uri>
              </graphic>
            </fig>
            <p><bold><italic>Male distal genitalia</italic></bold> (Figs <xref ref-type="fig" rid="F9">31</xref>, <xref ref-type="fig" rid="F9">32</xref>). Vas deferens of varying diameter (proximal tract narrow, medial tract slightly wider and final tract very narrow), entering penial complex near proximal end. Penial complex consisting of epiphallus, penial caecum and penis. Epiphallus very short. Penial caecum very short (as long as epiphallus). Penis very long, divided distinctly into proximal and distal parts by difference in calibre, without penial sheath; proximal part longer (twice distal penis), slender and thin walled; distal penis shorter (half proximal penis) and thick, muscular walled. Penial retractor muscle bifid, one branch inserted on tip of penial flagellum, the other branch on tip of penial caecum.</p>
            <p><bold><italic>Genital atrium</italic></bold> (Fig. <xref ref-type="fig" rid="F9">31</xref>). Very short.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="remarks" id="SECID0E4AAI">
            <title>Remarks.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> is a well-known travelling snail that occurs worldwide (<xref ref-type="bibr" rid="B131">Robinson 1999</xref>) but its native range is uncertain. It was described from Guadeloupe and Hispaniola and has been reported as native to tropical America (<xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>), the Caribbean (<xref ref-type="bibr" rid="B38">Deisler and Abbott 1984</xref>) and Latin America (<xref ref-type="bibr" rid="B131">Robinson 1999</xref>) but this was disputed on the grounds that the other species of the genus occur in Africa (<xref ref-type="bibr" rid="B9">Bieler and Slapcinsky 2000</xref>). <xref ref-type="bibr" rid="B51">Gerlach (2006)</xref> regarded it as native to the Seychelles, based on a subfossil record from Aldabra reported by <xref ref-type="bibr" rid="B52">Gerlach and van Bruggen (1999)</xref>. Others suggested that it was originally from tropical America (<xref ref-type="bibr" rid="B57">Griffiths and Florens 2006</xref>), south-east Asia (<xref ref-type="bibr" rid="B136">Rumi et al. 2010</xref>) or Africa (<xref ref-type="bibr" rid="B14">Breure et al. 2016</xref>; <xref ref-type="bibr" rid="B68">Hovestadt and van Leeuwen 2017</xref>). Multi-gene phylogenetic analysis of the achatinoid snails by <xref ref-type="bibr" rid="B48">Fontanilla et al. (2017)</xref> found that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="striatella">striatella</tp:taxon-name-part></tp:taxon-name></italic> (Rang, 1831) formed a monophyletic group belonging to an unsupported clade including only Old World species, but unfortunately the study included a small selection of subulinine genera, only one of which was from neotropical America.</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> is now distributed widely in humid tropical and subtropical lowlands across the world. It occurs in Central America, the West Indies, South America, sub-Saharan Africa, West Indian Ocean islands, South, South-East, and East Asia, New Guinea, Australia, and Pacific islands. In the mid temperate latitudes of the northern hemisphere, it only occurs in greenhouses and hothouses (see Table <xref ref-type="table" rid="T4">4</xref> for details and references).</p>
            <table-wrap id="T4" position="float" orientation="portrait">
              <label>Table 4.</label>
              <caption>
                <p>Geographical distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic>. Asterisks indicate countries / islands where the species has been recorded only in greenhouses or very disturbed anthropogenic habitats.</p>
              </caption>
              <table id="TID0ELIAK" rules="all">
                <tbody>
                  <tr>
                    <th rowspan="1" colspan="1">Regions</th>
                    <th rowspan="1" colspan="1">Countries / Islands</th>
                    <th rowspan="1" colspan="1">References</th>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>North America</bold>
                    </td>
                    <td rowspan="1" colspan="1">United States*</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B40">Dundee (1974)</xref>, <xref ref-type="bibr" rid="B101">Nekola (2014)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Central America</bold>
                    </td>
                    <td rowspan="1" colspan="1">Belize, Costa Rica, Guatemala, Honduras, Mexico, Nicaragua, Panama, and Salvador</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B158">Thompson (2011)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>West Indies</bold>
                    </td>
                    <td rowspan="1" colspan="1">Bahamas, Cuba, Hispaniola, Jamaica, and Lesser Antilles (Barbados, Curaçao, Guadeloupe, Martinique, Saint Barthélemy, Saint Martin, and Trinidad)</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B38">Deisler and Abbott (1984)</xref>, <xref ref-type="bibr" rid="B23">Chase and Robinson (2001)</xref>, <xref ref-type="bibr" rid="B133">Rosenberg and Muratov (2006)</xref>, <xref ref-type="bibr" rid="B43">Espinosa and Ortea (2009)</xref>, <xref ref-type="bibr" rid="B132">Robinson et al. (2009)</xref>, <xref ref-type="bibr" rid="B138">Rutherford (2011)</xref>, <xref ref-type="bibr" rid="B85">Maceira et al. (2013)</xref>, <xref ref-type="bibr" rid="B22">Charles (2015)</xref>, <xref ref-type="bibr" rid="B39">Delannoye et al. (2015)</xref>, <xref ref-type="bibr" rid="B14">Breure et al. (2016)</xref>, <xref ref-type="bibr" rid="B61">Herrera-Uria (2016)</xref>, <xref ref-type="bibr" rid="B68">Hovestadt and van Leeuwen (2017)</xref>, <xref ref-type="bibr" rid="B67">Hovestadt and Neckheim (2020)</xref>, <xref ref-type="bibr" rid="B44">Espinosa and Robinson (2021)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South America</bold>
                    </td>
                    <td rowspan="1" colspan="1">Brazil, Columbia, Ecuador, Peru, Suriname, and Venezuela</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B4">Baker (1927)</xref>, <xref ref-type="bibr" rid="B87">Marcus and Marcus (1968)</xref>, <xref ref-type="bibr" rid="B163">van Regteren Altena (1975)</xref>, <xref ref-type="bibr" rid="B145">Simone (2006)</xref>, <xref ref-type="bibr" rid="B136">Rumi et al. (2010)</xref>, <xref ref-type="bibr" rid="B15">Breure et al. (2022a)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Atlantic Ocean</bold>
                    </td>
                    <td rowspan="1" colspan="1">Bermuda</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B9">Bieler and Slapcinsky (2000)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Europe</bold>
                    </td>
                    <td rowspan="1" colspan="1">Austria*, Czech Republic*, Denmark*, Great Britain*, Ireland*, Italy*, Netherlands*, and Sweden*</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B78">Kerney and Cameron (1979)</xref>, <xref ref-type="bibr" rid="B171">von Proschwitz (1994</xref>, <xref ref-type="bibr" rid="B173">2016</xref>), <xref ref-type="bibr" rid="B75">Juricková (2006)</xref>, <xref ref-type="bibr" rid="B37">Da Sois (2015)</xref>, <xref ref-type="bibr" rid="B128">Reischütz et al. (2018)</xref>, <xref ref-type="bibr" rid="B1">Anderson and Rowson (2020)</xref>, <xref ref-type="bibr" rid="B66">Horsák et al. (2020)</xref>, this paper</td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Africa</bold>
                    </td>
                    <td rowspan="1" colspan="1">South Africa, Tanzania, and Zimbabwe</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B160">van Bruggen (1981)</xref>, <xref ref-type="bibr" rid="B134">Rowson (2007)</xref>, <xref ref-type="bibr" rid="B60">Herbert (2010)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Indian Ocean</bold>
                    </td>
                    <td rowspan="1" colspan="1">Aldabra, Madagascar, Maldives, Mascarene, Pemba, and Seychelles</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B124">Probst (2001)</xref>, <xref ref-type="bibr" rid="B51">Gerlach (2006)</xref>, <xref ref-type="bibr" rid="B57">Griffiths and Florens (2006)</xref>, <xref ref-type="bibr" rid="B134">Rowson (2007)</xref>, <xref ref-type="bibr" rid="B41">Emberton and Griffiths (2009)</xref>, <xref ref-type="bibr" rid="B135">Rowson et al. (2010)</xref>, <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen (2013)</xref>, <xref ref-type="bibr" rid="B55">Gittenberger et al. (2019)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">India and Sri Lanka</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B126">Ranawana (2006)</xref>, <xref ref-type="bibr" rid="B125">Raheem et al. (2014)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South-East Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Indonesia, Peninsular Malaysia, Sabah in Malaysian Borneo, Singapore, and Vietnam</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B140">Schileyko (2011)</xref>, <xref ref-type="bibr" rid="B156">Tan et al. (2015)</xref>, <xref ref-type="bibr" rid="B49">Foon et al. (2017)</xref>, <xref ref-type="bibr" rid="B113">Phung et al. (2017)</xref>, <xref ref-type="bibr" rid="B104">Nurinsiyaha and Hausdorf (2019)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>East Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Dongsha Islands and Japan*</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B91">Minato (1988)</xref>, <xref ref-type="bibr" rid="B178">Wu et al. (2007)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Oceania</bold>
                    </td>
                    <td rowspan="1" colspan="1">Australia, New Guinea, and Pacific islands (American Samoa, Belau/Palau, Cook Islands, Federated States of Micronesia, Fiji, French Polynesia, Galapagos Islands, Guam, Hawaii, Marshall Islands, New Caledonia, Northern Mariana Islands, Pitcairn Islands, Samoa, Solomon Islands, Tonga, and Vanuatu)</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B28">Cowie (2000</xref>, <xref ref-type="bibr" rid="B29">2001</xref>), <xref ref-type="bibr" rid="B176">Wiktor (2003)</xref>, <xref ref-type="bibr" rid="B144">Shea (2007)</xref>, <xref ref-type="bibr" rid="B19">Brook et al. (2010)</xref>, <xref ref-type="bibr" rid="B137">Rundell (2010)</xref>, <xref ref-type="bibr" rid="B153">Stanisic et al. (2010)</xref>, <xref ref-type="bibr" rid="B17">Brodie and Barker (2012)</xref>, <xref ref-type="bibr" rid="B79">Kerr and Bauman (2013)</xref>, <xref ref-type="bibr" rid="B18">Brook (2014)</xref>, <xref ref-type="bibr" rid="B92">Miquel and Herrera (2014)</xref>, <xref ref-type="bibr" rid="B31">Cowie et al. (2017)</xref></td>
                  </tr>
                </tbody>
              </table>
            </table-wrap>
            <p>The distal genitalia of <named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content> specimens show a general scheme identical to that described by <xref ref-type="bibr" rid="B4">Baker (1927</xref>: 3–4, pl. 20, fig. 99), <xref ref-type="bibr" rid="B87">Marcus and Marcus (1968</xref>: 189–190) and <xref ref-type="bibr" rid="B3">Araújo and Bessa (1993</xref>: 493–495, figs 7–11), but differing in the size of the penial complex from that described by <xref ref-type="bibr" rid="B175">Wiegmann (1894</xref>: 214–216, pl. 16, fig. 3; reproduced by <xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>: 173, fig. 83h), <xref ref-type="bibr" rid="B139">Schileyko (1999</xref>: fig. 662 <abbrev xlink:title="bursa copulatrix (gametolytic gland)" id="ABBRID0EETAI">BC</abbrev>) and <xref ref-type="bibr" rid="B54">Gittenberger and van Bruggen (2013</xref>: 250). Instead of a very short penial complex, in agreement with the first description by <xref ref-type="bibr" rid="B4">Baker (1927)</xref>, <named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content> specimens have a long slender penial complex consisting of a short proximal portion (the epiphallus and the penial caecum closely juxtaposed to each other), a long slender medial portion (the proximal penis) and a short swollen distal portion (the distal penis). In particular, based on the male genitalia of a specimen from Dunoon (Guyana) mounted in glycerine jelly and viewed by transmitted light, <xref ref-type="bibr" rid="B4">Baker (1927)</xref> described the proximal portion as consisting of a flagellar appendix [the penial caecum] and a very short, thick walled tract [the epiphallus]; the vas deferens entering the penial complex at the proximal end of the epiphallus; the penial retractor joining at the tip of penial caecum; a rather elongate papilla present at the internal opening of the epiphallus into the proximal penis. Baker also described a penial sheath enveloping the distal penis joined by a muscular branch originating from the right lower tentacle retractor; finally he interpreted the structure that <xref ref-type="bibr" rid="B175">Wiegmann (1894)</xref> described and figured as the penis as being only the distal penis surrounded by a heavy muscular sheath.</p>
            <p>The small size and the very fine structure of the sections of the proximal portion of the penial complex make dissection difficult and differentiation of its components elusive. Our results substantially agree with Baker’s description. The differences are: the vas deferens enters the penial complex near the base and not at the tip of the epiphallus; the penial retractor consists of two branches, one joined to the epiphallus, the other to the penial caecum; the small size of the proximal penis makes the penial papilla impossible to detect by stereomicroscope; a classical penial sheath (such as that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allopeas">Allopeas</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paropeas">Paropeas</tp:taxon-name-part></tp:taxon-name></italic>) is absent, although a muscular branch from the right lower tentacle retractor joins the distal penis directly on the penial wall. Unfortunately, the scarcity of material prevented a more careful anatomical examination. Some uncertainties about its real organisation and the meaning of such differences among the various anatomical reports therefore remain.</p>
            <p>Multi-gene phylogenetic analysis of the achatinoid snails by <xref ref-type="bibr" rid="B48">Fontanilla et al. (2017)</xref> found that the genus, as currently conceived, is polyphyletic because the three <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part></tp:taxon-name></italic> species examined did not cluster together. The meaning of variation in penis size in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> is also uncertain. <xref ref-type="bibr" rid="B175">Wiegmann (1894)</xref> ventured that the extreme reduction of the male distal genitalia was related to “unisexuality” or sequential hermaphroditism (... dass die betreffenden Thiere durch Verkümmerung des männlichen Theils der Genitalien eingeschlechtig geworden sind, oder aber dass die weibliche Geschlechtsreife der männlichen vorausgeht). Others, such as <xref ref-type="bibr" rid="B4">Baker (1927)</xref>, supposed that the species was a sequential hermaphrodite, but whereas Wiegmann saw it as protogynous, Baker viewed it as protandrous. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> is actually a facultatively self-fertilising egg-retaining species showing no evidence of sequential hermaphroditism (<xref ref-type="bibr" rid="B8">Bessa and Araújo 1995</xref>; <xref ref-type="bibr" rid="B36">D’Ávila et al. 2018</xref>). Thus the reduced penis may be related to loss of biparental reproduction as already supposed by Wiegmann. However this may be because <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> is a complex of species. More research is needed to address these questions.</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Subulina">Subulina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="octona">octona</tp:taxon-name-part></tp:taxon-name></italic> is common in the tropical greenhouse of <named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>, and has been found in the Biodiversity Garden (Botanical Garden of Padua), where only one specimen was collected. This is the first report from Italy.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Stylommatophora</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Subulinidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">11E65CC4-D8CB-53D8-ADEE-277338C7C64A</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>(Swainson, 1840)</tp:taxon-authority>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Macrospira">Macrospira</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name>
                <comment>Swainson, 1840: 335, fig. 97e, f. Type locality: no locality given; according to <xref ref-type="bibr" rid="B146">Smith (1892</xref>: 269), Saint Vincent, Lesser Antilles, West Indies. Type material: unknown. Note: Swainson attributed the new species to the reverend L. Guilding from St. Vincent, from whom he received the material used for the description. Probably this material was accompanied by a manuscript name which Swainson adopted for denoting the species.</comment>
              </tp:nomenclature-citation>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Achatina">Achatina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gundlachi">gundlachi</tp:taxon-name-part></tp:taxon-name>
                <comment>Pfeiffer, 1850: 80. Type locality: Cuba. Type material: unknown.</comment>
              </tp:nomenclature-citation>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="caledonica">caledonica</tp:taxon-name-part></tp:taxon-name>
                <comment>Crosse, 1867: 186–187, Pl. 7, fig. 4. Type locality: Nouméa, New Caledonia. Type material: 1 syntype in Crosse collection (MNHN-IM-2000-4720) (<xref ref-type="bibr" rid="B16">Breure et al. 2022b</xref>).</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="material" id="SECID0EOZAI">
            <title>Material examined.</title>
            <p>Italy • 2 shells; Trento, Tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>); <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[11.113911,46.062544]}" id="NCID0E2ZAI">46°03'45.16"N, 11°06'50.08"E</named-content></named-content>; 01 Feb. 2022; D. Barbato, A. Benocci leg.; GMC 51189.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EA1AI">
            <title>Description.</title>
            <p><bold><italic>Shell</italic></bold> (Figs <xref ref-type="fig" rid="F10">33</xref>, <xref ref-type="fig" rid="F10">34</xref>). Dextral, very small in size, imperforate, elongate, very slender, cylindro-conical, thin and fragile, pearly off-white, colourless, glossy and transparent when fresh, ~ 4 slightly convex whorls separated by rather deep sutures. Apex obtuse, rounded, and smooth. Last whorl ~ 2/3 of shell height. Aperture small, ~ 1/3 of shell height, ovate to pyriform, basally flared, slightly prosocline. Peristome interrupted, not thickened or reflected, with callous rim on parietum and columella; columella straight or slightly concave; outer margin slightly arched forward in the middle in lateral view. Protoconch smooth; teleoconch with very fine collabral lines and very fine spiral grooves particularly evident on last whorl. Shell dimensions: <abbrev xlink:title="shell height" id="ABBRID0ET1AI">SH</abbrev> 2.8 mm; <abbrev xlink:title="shell diameter" id="ABBRID0EX1AI">SD</abbrev> 1.0 mm; <abbrev xlink:title="aperture height" id="ABBRID0E21AI">AH</abbrev> 1.1 mm; <abbrev xlink:title="aperture width" id="ABBRID0E61AI">AW</abbrev> 0.6 mm.</p>
            <fig id="F10" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.1208.119147.figures33-34</object-id>
              <object-id content-type="arpha">D408B20A-A88F-526C-A7DE-966769CFF8C4</object-id>
              <label>Figures 33, 34.</label>
              <caption>
                <p>Shells of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> from the tropical greenhouse of the Science Museum of Trento (<named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content>), D. Barbato &amp; A. Benocci leg. 01 Feb. 2022.</p>
              </caption>
              <graphic xlink:href="zookeys-1208-099_article-119147__-g010.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1103576.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1103576</uri>
              </graphic>
            </fig>
            <p><bold><italic>Body and anatomy</italic>.</bold> Unknown.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="remarks" id="SECID0EG3AI">
            <title>Remarks.</title>
            <p>The early taxonomy of this land snail revolves around three named species: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Macrospira">Macrospira</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> Swainson, 1840, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Achatina">Achatina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gundlachi">gundlachi</tp:taxon-name-part></tp:taxon-name></italic> Pfeiffer, 1850 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="caledonica">caledonica</tp:taxon-name-part></tp:taxon-name></italic> Crosse, 1867.</p>
            <p>Commenting on the land mollusc species introduced to Saint Helena, Edgar Smith maintained that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Achatina">Achatina</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gundlachi">gundlachi</tp:taxon-name-part></tp:taxon-name></italic> was a junior synonym of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Macrospira">Macrospira</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> based on examination of Guilding’s material from St. Vincent, West Indies, deposited in the British Museum (<xref ref-type="bibr" rid="B146">Smith 1892</xref>) and that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="caledonica">caledonica</tp:taxon-name-part></tp:taxon-name></italic> was also a junior synonym of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaspira">Megaspira</tp:taxon-name-part></tp:taxon-name></italic> [sic] <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Megaspira"/><tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> (Smith, 1895).</p>
            <p><xref ref-type="bibr" rid="B117">Pilsbry (1908)</xref>, in his exhaustive revision of orthurethrous snails included in the second edition of the Manual of Conchology, partly rejected Smith’s conclusions, regarding <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Macrospira">Macrospira</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> as a species inadequately described and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic> as a section of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic>. He used <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gundlachi">gundlachi</tp:taxon-name-part></tp:taxon-name></italic> as the valid name for the species. However many years later, dealing with this group again, he adopted <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> as the valid name (<xref ref-type="bibr" rid="B119">Pilsbry 1946</xref>).</p>
            <p>No subsequent authors made any significant contribution for a better taxonomic framework of the species. They repeated what <xref ref-type="bibr" rid="B119">Pilsbry (1946)</xref> proposed, believing that a species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic>, sometimes considered a subgenus or a synonym of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic>, could be found almost everywhere in the world, having spread from the Mesoamerican area (e.g., <xref ref-type="bibr" rid="B131">Robinson 1999</xref>). Indeed many consider it native to the West Indies, Neotropics or Caribbean basin (<xref ref-type="bibr" rid="B40">Dundee 1974</xref>; <xref ref-type="bibr" rid="B27">Cowie 1997</xref>; <xref ref-type="bibr" rid="B131">Robinson 1999</xref>; <xref ref-type="bibr" rid="B23">Chase and Robinson 2001</xref>; <xref ref-type="bibr" rid="B133">Rosenberg and Muratov 2006</xref>; <xref ref-type="bibr" rid="B158">Thompson 2011</xref>; <xref ref-type="bibr" rid="B92">Miquel and Herrera 2014</xref>; <xref ref-type="bibr" rid="B105">Nurinsiyaha et al. 2016</xref>; <xref ref-type="bibr" rid="B104">Nurinsiyaha and Hausdorf 2019</xref>). Others report it to be native to southern Europe (<xref ref-type="bibr" rid="B26">Cotton 1954</xref>) or North America north of Mexico (<xref ref-type="bibr" rid="B101">Nekola 2014</xref>).</p>
            <p>A species, sometimes named <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> (or <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="caledonica">caledonica</tp:taxon-name-part></tp:taxon-name></italic> (or <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="caledonica">caledonica</tp:taxon-name-part></tp:taxon-name></italic>) or <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gundlachi">gundlachi</tp:taxon-name-part></tp:taxon-name></italic> (or <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gundlachi">gundlachi</tp:taxon-name-part></tp:taxon-name></italic>), is currently reported from North, Central and South America, the Caribbean and the Indo-Pacific Region from South-East Asia to Hawaii and Cook Islands (see Table <xref ref-type="table" rid="T5">5</xref> for details and references). It has also been reported from Saint Helena (<xref ref-type="bibr" rid="B146">Smith 1892</xref>) based on a misidentification of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acicula">acicula</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B34">Crowley and Pain 1977</xref>), and from Barbados, Curaçao and Galapagos, where it was no longer found in recent field surveys (<xref ref-type="bibr" rid="B23">Chase and Robinson 2001</xref>; <xref ref-type="bibr" rid="B92">Miquel and Herrera 2014</xref>; <xref ref-type="bibr" rid="B68">Hovestadt and van Leeuwen 2017</xref>).</p>
            <table-wrap id="T5" position="float" orientation="portrait">
              <label>Table 5.</label>
              <caption>
                <p>Geographical distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic>. Asterisks indicate countries / islands where the species has been recorded only in greenhouses or very disturbed anthropogenic habitats.</p>
              </caption>
              <table id="TID0E3QAK" rules="all">
                <tbody>
                  <tr>
                    <th rowspan="1" colspan="1">Regions</th>
                    <th rowspan="1" colspan="1">Countries / Islands</th>
                    <th rowspan="1" colspan="1">References</th>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>North America</bold>
                    </td>
                    <td rowspan="1" colspan="1">United States*</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B119">Pilsbry (1946)</xref>, <xref ref-type="bibr" rid="B40">Dundee (1974)</xref>, <xref ref-type="bibr" rid="B101">Nekola (2014)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Central America</bold>
                    </td>
                    <td rowspan="1" colspan="1">Nicaragua</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B158">Thompson (2011)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>West Indies</bold>
                    </td>
                    <td rowspan="1" colspan="1">Cuba, Hispaniola, Jamaica, Lesser Antilles (Guadeloupe, Martinique, Saint Martin, and Saint Vincent)</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B146">Smith (1892)</xref>, <xref ref-type="bibr" rid="B133">Rosenberg and Muratov (2006)</xref>, <xref ref-type="bibr" rid="B85">Maceira et al. (2013)</xref>, <xref ref-type="bibr" rid="B22">Charles (2015)</xref>, <xref ref-type="bibr" rid="B67">Hovestadt and Neckheim (2020)</xref>, <xref ref-type="bibr" rid="B44">Espinosa and Robinson (2021)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South America</bold>
                    </td>
                    <td rowspan="1" colspan="1">Brazil</td>
                    <td rowspan="1" colspan="1">
                      <xref ref-type="bibr" rid="B145">Simone (2006)</xref>
                    </td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>South-East Asia</bold>
                    </td>
                    <td rowspan="1" colspan="1">Indonesia, Peninsular Malaysia, and Philippines</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B168">Vermeulen and Whitten (1998)</xref>, <xref ref-type="bibr" rid="B84">Maassen (2001)</xref>, <xref ref-type="bibr" rid="B59">Groh (2015)</xref>, <xref ref-type="bibr" rid="B105">Nurinsiyaha et al. (2016)</xref>, <xref ref-type="bibr" rid="B49">Foon et al. (2017)</xref>, <xref ref-type="bibr" rid="B104">Nurinsiyaha and Hausdorf (2019)</xref></td>
                  </tr>
                  <tr>
                    <td rowspan="1" colspan="1">
                      <bold>Oceania</bold>
                    </td>
                    <td rowspan="1" colspan="1">Australia, New Guinea, and Pacific Islands (Cook Islands, Guam, Hawaii, and New Caledonia)</td>
                    <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B150">Solem (1964)</xref>, <xref ref-type="bibr" rid="B159">van Benthem Jutting (1964)</xref>, <xref ref-type="bibr" rid="B28">Cowie (2000</xref>, <xref ref-type="bibr" rid="B29">2001</xref>), <xref ref-type="bibr" rid="B144">Shea (2007)</xref>, <xref ref-type="bibr" rid="B153">Stanisic et al. (2010)</xref></td>
                  </tr>
                </tbody>
              </table>
            </table-wrap>
            <p>It is difficult to say anything new about this group of species, since there is no anatomical data. We can rely on shell characters, the most interesting of which are the spiral sculpture particularly evident in the last whorl, the non-truncated or slightly truncated columella and the basally flared aperture. Based on a close resemblance to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic>, the species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic> have been placed in the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name> but this similarity could also be due to convergence: true <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> have no microsculpture on the teleoconch, have a truncated columella and have no basally flared aperture. In their phylogenetic analysis of the achatinoideans <xref ref-type="bibr" rid="B48">Fontanilla et al. (2017)</xref> examined a <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> species – <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gokweana">gokweana</tp:taxon-name-part></tp:taxon-name></italic> (Boettger, 1870) – which may actually be <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic>, if the material investigated, collected by DG Herbert, matches the description of this species given by him (cf. <xref ref-type="bibr" rid="B60">Herbert 2010</xref>: 127 for description and figure). If this is confirmed, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic> does not belong to the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name> but to a distinct group of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name>. This of course does not resolve the relationships between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic>: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> may really belong to the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Ferussaciidae</tp:taxon-name-part></tp:taxon-name> or to the same or a different group of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Achatinidae</tp:taxon-name-part></tp:taxon-name>, which may include <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic>.</p>
            <p>MolluscaBase lists eight <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic> species (<xref ref-type="bibr" rid="B97">MolluscaBase eds 2024c</xref>), but except for the widespread <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aperta">aperta</tp:taxon-name-part></tp:taxon-name></italic> and the south American <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="blandiana">blandiana</tp:taxon-name-part></tp:taxon-name></italic> Crosse, 1880, all the others occur from Madagascar to Southeast Asia. There is great uncertainty about species-level taxonomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic>. Some of the species listed by MolluscaBase may prove to be true <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> based on the apparent absence of spiral microsculpture, truncated columella and not basally flared aperture: this could be true of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="philippinica">philippinica</tp:taxon-name-part></tp:taxon-name></italic> von Möllendorff, 1890 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sheilae">sheilae</tp:taxon-name-part></tp:taxon-name></italic> Groh, 2015 (see <xref ref-type="bibr" rid="B59">Groh 2015</xref>). On the other hand, some species reported as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part></tp:taxon-name></italic> by MolluscaBase, such as the East African <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="callipeplum">callipeplum</tp:taxon-name-part></tp:taxon-name></italic> (Connolly, 1923), for which clear spiral sculpture is reported by <xref ref-type="bibr" rid="B165">Verdcourt (1986)</xref>, van Bruggen and van Goethem (2001), and <xref ref-type="bibr" rid="B161">van Bruggen (2008)</xref>, or the South African <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cecilioides">Cecilioides</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gokweana">gokweana</tp:taxon-name-part></tp:taxon-name></italic>, for which clear spiral sculpture is described by <xref ref-type="bibr" rid="B60">Herbert (2010)</xref>, may prove to be true <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geostilbia">Geostilbia</tp:taxon-name-part></tp:taxon-name></italic>.</p>
            <p>We assign two shells found in the litter of the tropical greenhouse of <named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content> to this species (unfortunately no living specimen was found during our collecting). This is the first report of the species from Europe.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
    </sec>
  </body>
  <back>
    <ack>
      <title>﻿Acknowledgements</title>
      <p>We thank the staff of <named-content content-type="dwc:institutional_code" xlink:title="Science Museum of Trentoe" xlink:href="http://grbio.org/institution/museo-delle-scienze-0">MUSE</named-content> (Trento, Italy), in particular Francesca Rossi for permission and assistance during sampling and Andrea Bianchi for information on the flora of the Afromontane greenhouse. We also thank the staff of the Botanical Garden of Padua, especially Barbara Baldan for authorisation to visit the greenhouses and Roberto Tacchetto for assistance during field sampling. Special thanks also to Gregorio Bolzonella (Siena, Italy) for help during collecting, Sigrid Hof (Frankfurt am Main, Germany) for loan of specimens from Senckenberg Forschungsinstitut, Giovanni Cappelli (Siena, Italy) for photographs of the shells, and Helen Ampt (Siena, Italy) for English language revision.</p>
    </ack>
    <sec sec-type="﻿Additional information" id="SECID0EVUBI">
      <title>﻿Additional information</title>
      <sec sec-type="Conflict of interest" id="SECID0EZUBI">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="SECID0E5UBI">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Funding" id="SECID0EDVBI">
        <title>Funding</title>
        <p>D. Barbato and G. Manganelli were funded under the National Recovery and Resilience Plan (NRRP), Mission 4 Component 2 Investment 1.4 - Call for tender No. 3138 of 16 December 2021, rectified by Decree n.3175 of 18 December 2021 of Italian Ministry of University and Research funded by the European Union – NextGenerationEU; Award Number: Project code CN_00000033, Concession Decree No. 1034 of 17 June 2022 adopted by the Italian Ministry of University and Research, CUP B63C22000650007 Project title “National Biodiversity Future Center - NBFC”.</p>
      </sec>
      <sec sec-type="Author contributions" id="SECID0EIVBI">
        <title>Author contributions</title>
        <p>Conceptualisation: DB and GM; Methodology, Formal analysis, Investigation, Data Curation: AB, DB, FG, and GM; Writing - Original draft &amp; Writing - Review and Editing: AB, DB, FG, and GM; Supervision: GM; Funding Acquisition: GM.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="SECID0ENVBI">
        <title>Author ORCIDs</title>
        <p>Giuseppe Manganelli <ext-link xlink:href="https://orcid.org/0000-0002-8453-280X" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-8453-280X</ext-link></p>
        <p>Andrea Benocci <ext-link xlink:href="https://orcid.org/0000-0003-3145-215X" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0003-3145-215X</ext-link></p>
        <p>Debora Barbato <ext-link xlink:href="https://orcid.org/0000-0003-1105-1711" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0003-1105-1711</ext-link></p>
        <p>Folco Giusti <ext-link xlink:href="https://orcid.org/0000-0001-8722-4653" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0001-8722-4653</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="SECID0EJWBI">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text.</p>
      </sec>
    </sec>
    <ref-list>
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