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<document id="CA6D340983A7A8B71B4FAA7F020934F7" ID-CLB-Dataset="307941" ID-DOI="10.1093/zoolinnean/zlae010" ID-GBIF-Dataset="5265b49c-637e-4b38-97cb-e0f7c234cf88" ID-ISSN="0024-4082" ID-Zenodo-Dep="14774861" ID-ZooBank="D1C88C6-B4B9-46D0-94AD-BA7439E6DF22" IM.bibliography_approvedBy="valdenar" IM.illustrations_approvedBy="valdenar" IM.materialsCitations_approvedBy="valdenar" IM.metadata_approvedBy="valdenar" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="valdenar" checkinTime="1738153874994" checkinUser="plazi" docAuthor="Delicado, Diana, Boulaassafer, Khadija, Khalloufi, Noureddine &amp; Hauffe, Torsten" docDate="2025" docId="536F827BFFA0530C29E0FD6BF054F176" docLanguage="en" docName="zlae010.pdf" docOrigin="Zoological Journal of the Linnean Society 203 (1)" docSource="https://doi.org/10.1093/zoolinnean/zlae010" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.10:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="10" docTitle="Pseudamnicola argentariensis Delicado, Boulaassafer &amp; Hauffe 2025, sp. nov." docType="treatment" docVersion="3" lastPageNumber="18" masterDocId="AF56FA03FFAF531D2967FF90F13AF726" masterDocTitle="A holistic perspective on species delimitation outperforms all methods based on single data types in freshwater gastropods (Caenogastropoda: Hydrobiidae: Pseudamnicola)" masterLastPageNumber="31" masterPageNumber="1" pageNumber="16" updateTime="1738263585638" updateUser="ExternalLinkService">
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<mods:title id="C37D1B6BE7145059F52E01162BAE34ED">A holistic perspective on species delimitation outperforms all methods based on single data types in freshwater gastropods (Caenogastropoda: Hydrobiidae: Pseudamnicola)</mods:title>
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<mods:role id="C32ABD404DAEF38126A77DBE8B37B538">
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<mods:namePart id="9265C5E5BB6CCA9A5DCA8D5E41B6164A">Delicado, Diana</mods:namePart>
<mods:affiliation id="C6946CC472C87BA75CC55559AFBC6C56">Department of Animal Ecology &amp; Systematics, Justus Liebig University Giessen, Heinrich-Buff-Ring 26 - 32 IFZ, D- 35392 Giessen, Germany</mods:affiliation>
<mods:nameIdentifier id="1067A346B91EEAA0378F00419703D113" type="email">didelicado@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name id="7A5FF85C1C5EC1438D107646E45C7582" type="personal">
<mods:role id="C4A687494A45FC3B881BEDAAC675D968">
<mods:roleTerm id="2E6AED09EF98BFC40D311798A50731FE">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="E5039F0B47D3E0EBCB4628E8721E9822">Boulaassafer, Khadija</mods:namePart>
<mods:affiliation id="8663765302376392A4425A9A268BC7FB">Muséum d’Histoire Naturelle de Marrakech, Cadi Ayyad University, Prince Moulay Abdellah Boulevard, B. P. 2390, 40000 Marrakesh, Morocco</mods:affiliation>
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<mods:namePart id="34F883C9AC8CD5B6DF2A25AEABA57035">Khalloufi, Noureddine</mods:namePart>
<mods:affiliation id="213227AF09D1A1EE0E09726A3D98D553">Environmental Biomonitoring Laboratory (LR 01 ES 14), Faculty of Sciences of Bizerte, University of Carthage, 7021 Jarzouna, Tunisia</mods:affiliation>
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<mods:namePart id="5F9870E70B849736AC26CAA06BB2914F">Hauffe, Torsten</mods:namePart>
<mods:affiliation id="9459CFA890E1B40ED1CAAD895F0421D4">Department of Biology, University of Fribourg, Chemin du Musée 10, CH- 1700 Fribourg, Switzerland &amp; Swiss Institute of Bioinformatics, Chemin du Musée 10, CH- 1700 Fribourg, Switzerland</mods:affiliation>
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<mods:title id="1DAD86D3AB034AF9973D3E9537802EEF">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="9F06408EDBF13FA1E784584832FFFBDE">2025</mods:date>
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<mods:number id="59BAD5B9E7065835B7514E9E0EBA2DF1">203</mods:number>
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<mods:identifier id="3C81DE49940379476007F80E05E6F1E6" type="CLB-Dataset">307941</mods:identifier>
<mods:identifier id="45F96D3FD0DB4B52B6466543E974B279" type="DOI">10.1093/zoolinnean/zlae010</mods:identifier>
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<treatment id="536F827BFFA0530C29E0FD6BF054F176" ID-DOI="http://doi.org/10.5281/zenodo.14774865" ID-Zenodo-Dep="14774865" LSID="urn:lsid:plazi:treatment:536F827BFFA0530C29E0FD6BF054F176" httpUri="http://treatment.plazi.org/id/536F827BFFA0530C29E0FD6BF054F176" lastPageId="17" lastPageNumber="18" pageId="15" pageNumber="16" scope_family="Hydrobiidae" scope_subClass="Caenogastropoda">
<subSubSection id="93DC60E6FFA0531229E0FD6BF338F413" pageId="15" pageNumber="16" type="nomenclature">
<paragraph id="DB79336DFFA0531229E0FD6BF338F413" blockId="15.[135,741,763,821]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA0531229E0FD6BF338F413" bold="true" pageId="15" pageNumber="16">
<taxonomicName id="1CC648EEFFA0531229E0FD6BF088F413" authority="Delicado, Boulaassafer &amp; Hauffe" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="15" pageNumber="16" phylum="Mollusca" rank="species" species="argentariensis" status="sp. nov.">
<emphasis id="E9B2EF7FFFA0531229E0FD6BF0ECF433" bold="true" box="[135,470,763,790]" italics="true" pageId="15" pageNumber="16">Pseudamnicola argentariensis</emphasis>
Delicado, Boulaassafer &amp; Hauffe
</taxonomicName>
<taxonomicNameLabel id="F2815204FFA0531228D1FC8BF338F413" box="[438,514,795,821]" pageId="15" pageNumber="16" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
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<subSubSection id="93DC60E6FFA0531228E6FCC9F0D1F455" box="[385,491,857,884]" pageId="15" pageNumber="16" type="description">
<paragraph id="DB79336DFFA0531228E6FCC9F0D1F455" blockId="15.[113,600,857,951]" box="[385,491,857,884]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA0531228E6FCC9F0D1F455" bold="true" box="[385,491,857,884]" pageId="15" pageNumber="16">
(
<figureCitation id="43FD2FE8FFA0531228ECFCC9F0F3F452" box="[395,457,857,884]" captionStart="Figure 7" captionStartId="16.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1322]" captionTargetId="figure-321@16.[129,1473,144,1322]" captionTargetPageId="16" captionText="Figure 7. Shells, operculum, and radulae of Pseudamnicola argentariensis. A, holotype (MNCN 15.05/200500H). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="15" pageNumber="16">Figs 7</figureCitation>
,
<figureCitation id="43FD2FE8FFA0531228B5FCC9F0DAF455" box="[466,480,857,883]" captionStart="Figure 8" captionStartId="17.[113,178,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-159@17.[114,1458,144,1358]" captionTargetPageId="17" captionText="Figure 8. Anatomy of Pseudamnicola argentariensis. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis.H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="15" pageNumber="16">8</figureCitation>
)
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFA053122916FC10F0D9F336" pageId="15" pageNumber="16" type="reference_group">
<paragraph id="DB79336DFFA053122916FC10F362F491" blockId="15.[113,600,857,951]" pageId="15" pageNumber="16">
<taxonomicName id="1CC648EEFFA053122916FC10F008F4BE" box="[113,306,896,920]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="15" pageNumber="16" phylum="Mollusca" rank="species" species="undefined-1">
<emphasis id="E9B2EF7FFFA053122916FC10F039F4BE" box="[113,259,896,920]" italics="true" pageId="15" pageNumber="16">Pseudamnicola</emphasis>
sp. 1
</taxonomicName>
—
<bibRefCitation id="BF574E9CFFA053122836FC10F323F4BE" author="Delicado D &amp; Hauffe T &amp; Wilke T" box="[337,537,896,920]" pageId="15" pageNumber="16" refId="ref20825" refString="Delicado D, Hauffe T, Wilke T. Ecological opportunity may facilitate diversification in Palearctic freshwater organisms: a case study on hydrobiid gastropods. BMC Evolutionary Biology 2018; 18: 55. https: // doi. org / 10.1186 / s 12862 - 018 - 1169 - 2" type="journal volume" year="2018">
Delicado
<emphasis id="E9B2EF7FFFA0531228D5FC11F0DBF4BE" box="[434,481,896,920]" italics="true" pageId="15" pageNumber="16">et al.</emphasis>
2018
</bibRefCitation>
<taxonomicName id="1CC648EEFFA0531229EAFC0FF077F491" box="[141,333,927,951]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="15" pageNumber="16" phylum="Mollusca" rank="species" species="undefined-1">
<emphasis id="E9B2EF7FFFA0531229EAFC0FF025F491" box="[141,287,927,951]" italics="true" pageId="15" pageNumber="16">Pseudamnicola</emphasis>
sp. 1
</taxonomicName>
—
<bibRefCitation id="BF574E9CFFA05312280BFC0FF362F491" author="Boulaassafer K &amp; Ghamizi M &amp; Machordom A" box="[364,600,927,951]" pageId="15" pageNumber="16" pagination="396 - 416" refId="ref20276" refString="Boulaassafer K, Ghamizi M, Machordom A et al. Phylogenetic relationships within Pseudamnicola Paulucci, 1878 (Caenogastropoda: Truncatelloidea) indicate two independent dispersal events from different continents to the Balearic Islands. Systematics and Biodiversity 2020; 18: 396 - 416. https: // doi. org / 10.1080 / 14772000.2020.17714 66" type="journal article" year="2020">
Boulaassafer
<emphasis id="E9B2EF7FFFA053122896FC30F31AF491" box="[497,544,927,951]" italics="true" pageId="15" pageNumber="16">et al.</emphasis>
2020
</bibRefCitation>
</paragraph>
<paragraph id="DB79336DFFA053122916FC49F0D9F336" blockId="15.[113,762,984,1040]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA053122916FC49F060F4D6" box="[113,346,985,1009]" italics="true" pageId="15" pageNumber="16">ZooBank registration:</emphasis>
<uri id="AF573F6FFFA053122812FC48F0D9F336" pageId="15" pageNumber="16">
urn:lsid:zoobank.org:act:
<uuid id="AF6009B8FFA053122B14FC48F0D9F336" pageId="15" pageNumber="16">FBFFB25D-0A78-4EAF-A578-60EC1F6A761D</uuid>
</uri>
</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFA0531228E4FBBFF0D3F36F" box="[387,489,1071,1097]" pageId="15" pageNumber="16" type="etymology">
<paragraph id="DB79336DFFA0531228E4FBBFF0D3F36F" blockId="15.[387,489,1071,1097]" box="[387,489,1071,1097]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA0531228E4FBBFF0D3F36F" box="[387,489,1071,1097]" italics="true" pageId="15" pageNumber="16">Etymology</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFA053122916FBC6F069F103" pageId="15" pageNumber="16" type="materials_examined">
<paragraph id="DB79336DFFA053122916FBC6F076F3AB" blockId="15.[113,762,1110,1166]" pageId="15" pageNumber="16">
<materialsCitation id="6BAE3930FFA053122916FBC6F072F3AB" country="Italy" location="Referring" municipality="Monte Argentario" pageId="15" pageNumber="16" specimenCount="1">
<location id="DE1965B6FFA053122916FBC6F1E8F348" LSID="urn:lsid:plazi:treatment:536F827BFFA0530C29E0FD6BF054F176:DE1965B6FFA053122916FBC6F1E8F348" box="[113,210,1110,1134]" country="Italy" municipality="Monte Argentario" name="Referring" pageId="15" pageNumber="16">Referring</location>
to
<collectingMunicipality id="3B1DA917FFA05312299DFBC6F08DF348" box="[250,439,1110,1134]" pageId="15" pageNumber="16">Monte Argentario</collectingMunicipality>
(
<collectingCountry id="A3D173FDFFA0531228AFFBC6F0C6F348" box="[456,508,1110,1134]" name="Italy" pageId="15" pageNumber="16">Italy</collectingCountry>
), a municipality located near the
<typeStatus id="047D8DCFFFA0531229AEFBE6F1CDF3A8" box="[201,247,1142,1166]" pageId="15" pageNumber="16">type</typeStatus>
locality
</materialsCitation>
.
</paragraph>
<paragraph id="DB79336DFFA053122812FB3EF0CDF3E1" blockId="15.[112,762,1197,1385]" box="[373,503,1197,1223]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA053122812FB3EF0CDF3E1" box="[373,503,1197,1223]" italics="true" pageId="15" pageNumber="16">Type material</emphasis>
</paragraph>
<paragraph id="DB79336DFFA053122916FB43F050F24F" blockId="15.[112,762,1197,1385]" pageId="15" pageNumber="16">
<materialsCitation id="6BAE3930FFA053122916FB43F050F24F" collectionCode="MNCN, UGSB" country="Italy" county="Grosseto Province" location="Fontana Carpina" municipality="Monte Argentario" pageId="15" pageNumber="16" specimenCode="MNCN 15.05, UGSB 11999" specimenCount="54" typeStatus="holotype">
<typeStatus id="047D8DCFFFA053122916FB43F1EEF3CD" box="[113,212,1235,1259]" pageId="15" pageNumber="16" type="holotype">Holotype</typeStatus>
(
<emphasis id="E9B2EF7FFFA053122983FB44F0E5F3CD" box="[228,479,1235,1259]" italics="true" pageId="15" pageNumber="16">
<specimenCode id="8B609B16FFA053122983FB44F04BF3CD" box="[228,369,1235,1259]" collectionCode="MNCN" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="15" pageNumber="16" type="Museum">MNCN 15.05</specimenCode>
/200500H
</emphasis>
; dissected animal) and
<specimenCount id="CDC0F8E4FFA053122BA8FB43F1EFF22D" count="4" pageId="15" pageNumber="16" type="generic" typeStatus="paratype">four paratypes</specimenCount>
(
<specimenCode id="8B609B16FFA053122983FB63F046F22C" box="[228,380,1266,1291]" collectionCode="MNCN" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="15" pageNumber="16" type="Museum">MNCN 15.05</specimenCode>
/200500P) in the
<collectionCode id="BDD7ABA8FFA053122B5CFB63F3B7F22D" box="[571,653,1267,1291]" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="15" pageNumber="16" type="Museum">MNCN</collectionCode>
collection and ~
<specimenCount id="CDC0F8E4FFA0531229CEFA82F016F20C" box="[169,300,1298,1322]" count="50" pageId="15" pageNumber="16" type="generic" typeStatus="paratype">50 paratypes</specimenCount>
in the
<collectionCode id="BDD7ABA8FFA05312280BFA82F095F20C" box="[364,431,1298,1322]" pageId="15" pageNumber="16">UGSB</collectionCode>
collection (
<specimenCode id="8B609B16FFA053122B43FA82F38CF20C" box="[548,694,1298,1322]" collectionCode="UGSB" pageId="15" pageNumber="16">UGSB 11999</specimenCode>
). Type locality: a spring below the
<location id="DE1965B6FFA0531228E1FAA2F30EF26F" LSID="urn:lsid:plazi:treatment:536F827BFFA0530C29E0FD6BF054F176:DE1965B6FFA0531228E1FAA2F30EF26F" box="[390,564,1329,1353]" country="Italy" county="Grosseto Province" municipality="Monte Argentario" name="Fontana Carpina" pageId="15" pageNumber="16">Fontana Carpina</location>
,
<collectingMunicipality id="3B1DA917FFA053122B5BFAA2F3CFF26F" box="[572,757,1329,1353]" pageId="15" pageNumber="16">Monte Argentario</collectingMunicipality>
,
<collectingCounty id="32184BE1FFA053122916FAC1F015F24F" box="[113,303,1361,1385]" pageId="15" pageNumber="16">Grosseto Province</collectingCounty>
,
<collectingCountry id="A3D173FDFFA05312285FFAC1F05CF24F" box="[312,358,1361,1385]" name="Italy" pageId="15" pageNumber="16">Italy</collectingCountry>
.
</materialsCitation>
</paragraph>
<paragraph id="DB79336DFFA05312280EFA18F338F284" blockId="15.[361,514,1416,1442]" box="[361,514,1416,1442]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA05312280EFA18F338F284" box="[361,514,1416,1442]" italics="true" pageId="15" pageNumber="16">Material studied</emphasis>
</paragraph>
<paragraph id="DB79336DFFA053122916FA3FF069F103" blockId="15.[113,763,1455,1573]" pageId="15" pageNumber="16">
<materialsCitation id="6BAE3930FFA053122916FA3FF075F103" collectingDate="2012-09" collectionCode="MNCN, UGSB" collectorName="Solis-Lemus C" country="Italy" county="Grosseto Province" latitude="42.4242" location="Fontana Carpina" longLatPrecision="6" longitude="11.0908" municipality="Monte Argentario" pageId="15" pageNumber="16" specimenCode="MNCN 15.05, UGSB 11999" specimenCount="1">
A spring below the
<location id="DE1965B6FFA053122803FA3FF327F2E1" LSID="urn:lsid:plazi:treatment:536F827BFFA0530C29E0FD6BF054F176:DE1965B6FFA053122803FA3FF327F2E1" box="[356,541,1455,1479]" country="Italy" county="Grosseto Province" latitude="42.4242" longLatPrecision="6" longitude="11.0908" municipality="Monte Argentario" name="Fontana Carpina" pageId="15" pageNumber="16">Fontana Carpina</location>
,
<collectingMunicipality id="3B1DA917FFA053122B55FA3FF3CCF2E1" box="[562,758,1455,1479]" pageId="15" pageNumber="16">Monte Argentario</collectingMunicipality>
,
<collectingCounty id="32184BE1FFA053122916FA5EF00EF2C0" box="[113,308,1486,1510]" pageId="15" pageNumber="16">Grosseto Province</collectingCounty>
,
<collectingCountry id="A3D173FDFFA053122826FA5EF054F2C0" box="[321,366,1486,1510]" name="Italy" pageId="15" pageNumber="16">Italy</collectingCountry>
,
<geoCoordinate id="BEF255AAFFA05312281BFA5EF0D7F2C0" box="[380,493,1486,1510]" degrees="42.4242" direction="north" orientation="latitude" pageId="15" pageNumber="16" precision="5" value="42.4242">42.4242°N</geoCoordinate>
,
<geoCoordinate id="BEF255AAFFA05312289DFA5EF35DF2C0" box="[506,615,1486,1510]" degrees="11.0908" direction="east" orientation="longitude" pageId="15" pageNumber="16" precision="5" value="11.0908">11.0908°E</geoCoordinate>
, leg.
<collectorName id="763356BBFFA053122BC5FA5EF3F1F2C0" box="[674,715,1486,1510]" name="Solis-Lemus C" pageId="15" pageNumber="16">S.C.</collectorName>
and M.C.,
<date id="AF7815ADFFA0531229DEFA7DF057F123" box="[185,365,1517,1541]" pageId="15" pageNumber="16" value="2012-09">
<collectingDate id="BF3CEC45FFA0531229DEFA7DF057F123" box="[185,365,1517,1541]" pageId="15" pageNumber="16" value="2012-09">September 2012</collectingDate>
</date>
,
<specimenCode id="8B609B16FFA053122818FA7DF323F123" box="[383,537,1517,1541]" collectionCode="MNCN" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="15" pageNumber="16" type="Museum">MNCN 15.05</specimenCode>
/200500 and
<specimenCode id="8B609B16FFA053122BD0FA7DF188F103" collectionCode="UGSB" pageId="15" pageNumber="16">UGSB 11999</specimenCode>
(80% ethanol)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFA0531228E0F9D5F318F019" pageId="15" pageNumber="16" type="diagnosis">
<paragraph id="DB79336DFFA0531228E0F9D5F0DEF178" blockId="15.[112,764,1605,1855]" box="[391,484,1605,1630]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA0531228E0F9D5F0DEF178" box="[391,484,1605,1630]" italics="true" pageId="15" pageNumber="16">Diagnosis</emphasis>
</paragraph>
<paragraph id="DB79336DFFA053122916F9FBF318F019" blockId="15.[112,764,1605,1855]" pageId="15" pageNumber="16">Shell relatively slender (SL/SW ratio&gt; 1.5); protoconch microsculpture pitted; central radular tooth formula 4-C-4/1-1; bursa copulatrix large, pyriform, with a duct longer than bursal length; SR1 elongate, with a short duct; penis gradually tapering, with many folds over the entire surface and a small patch of pigmentation on its distal region; nervous system slightly pigmented, elongate (mean RPG ratio =.61).</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFA0531228E7F8CFF2A6F16F" pageId="15" pageNumber="16" type="description">
<paragraph id="DB79336DFFA0531228E7F8CFF0D6F05E" blockId="15.[113,763,1887,1980]" box="[384,492,1887,1912]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA0531228E7F8CFF0D6F05E" box="[384,492,1887,1912]" italics="true" pageId="15" pageNumber="16">Description</emphasis>
</paragraph>
<paragraph id="DB79336DFFA053122916F815F2C1F6A5" blockId="15.[113,763,1887,1980]" lastBlockId="15.[808,1461,144,1609]" pageId="15" pageNumber="16">
Shell ovate-conic, whorls 4–5, height
<quantity id="1C3E9E88FFA05312289DF814F34FF0BA" box="[506,629,1924,1948]" metricMagnitude="-3" metricUnit="m" metricValue="3.0" metricValueMax="3.5" metricValueMin="2.5" pageId="15" pageNumber="16" unit="mm" value="3.0" valueMax="3.5" valueMin="2.5">2.5–3.5 mm</quantity>
(
<figureCitation id="43FD2FE8FFA053122BE0F815F3C9F0BB" box="[647,755,1925,1949]" captionStart="Figure 7" captionStartId="16.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1322]" captionTargetId="figure-321@16.[129,1473,144,1322]" captionTargetPageId="16" captionText="Figure 7. Shells, operculum, and radulae of Pseudamnicola argentariensis. A, holotype (MNCN 15.05/200500H). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="15" pageNumber="16">Fig. 7A–C</figureCitation>
; Supporting Information, Table S3). Periostracum yellowish. Protoconch ~450 µm wide, whorls 1.5; nucleus ~125 µm wide; protoconch microsculpture pitted (
<figureCitation id="43FD2FE8FFA053122DB0FF3FF419F7E1" box="[1239,1315,175,199]" captionStart="Figure 7" captionStartId="16.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1322]" captionTargetId="figure-321@16.[129,1473,144,1322]" captionTargetPageId="16" captionText="Figure 7. Shells, operculum, and radulae of Pseudamnicola argentariensis. A, holotype (MNCN 15.05/200500H). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="15" pageNumber="16">Fig. 7G</figureCitation>
). Teleoconch whorls moderately convex, with deep sutures; body whorl large, occupying about two-thirds of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (
<figureCitation id="43FD2FE8FFA053122AEEFEBDF2EFF662" box="[905,981,301,325]" captionStart="Figure 7" captionStartId="16.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1322]" captionTargetId="figure-321@16.[129,1473,144,1322]" captionTargetPageId="16" captionText="Figure 7. Shells, operculum, and radulae of Pseudamnicola argentariensis. A, holotype (MNCN 15.05/200500H). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="15" pageNumber="16">Fig. 7D</figureCitation>
). Umbilicus narrow, not covered by the inner lip.
<typeStatus id="047D8DCFFFA053122A32FEDCF286F642" box="[853,956,332,356]" pageId="15" pageNumber="16" type="holotype">Holotype</typeStatus>
: SL =
<quantity id="1C3E9E88FFA053122D66FEDCF566F645" box="[1025,1116,332,356]" metricMagnitude="-3" metricUnit="m" metricValue="3.46" pageId="15" pageNumber="16" unit="mm" value="3.46">3.46 mm</quantity>
, SW =
<quantity id="1C3E9E88FFA053122DCBFEDCF432F645" box="[1196,1288,332,356]" metricMagnitude="-3" metricUnit="m" metricValue="2.01" pageId="15" pageNumber="16" unit="mm" value="2.01">2.01 mm</quantity>
, AL =
<quantity id="1C3E9E88FFA053122C35FEDCF495F645" box="[1362,1455,332,356]" metricMagnitude="-3" metricUnit="m" metricValue="1.69" pageId="15" pageNumber="16" unit="mm" value="1.69">1.69 mm</quantity>
, and AW =
<quantity id="1C3E9E88FFA053122AFDFEFBF2C2F6A5" box="[922,1016,363,387]" metricMagnitude="-3" metricUnit="m" metricValue="1.24" pageId="15" pageNumber="16" unit="mm" value="1.24">1.24 mm</quantity>
.
</paragraph>
<paragraph id="DB79336DFFA053122A22FE1BF24AF6C7" blockId="15.[808,1461,144,1609]" pageId="15" pageNumber="16">
Operculum oval, brownish, about two and a half whorls; muscle attachment area oval and located near the nucleus (
<figureCitation id="43FD2FE8FFA053122CEAFE3AF265F6C7" captionStart="Figure 7" captionStartId="16.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1322]" captionTargetId="figure-321@16.[129,1473,144,1322]" captionTargetPageId="16" captionText="Figure 7. Shells, operculum, and radulae of Pseudamnicola argentariensis. A, holotype (MNCN 15.05/200500H). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="15" pageNumber="16">Fig. 7E, F</figureCitation>
).
</paragraph>
<paragraph id="DB79336DFFA053122A22FE79F404F5BB" blockId="15.[808,1461,144,1609]" pageId="15" pageNumber="16">
Radular length intermediate, ~675 µm (~25% of total shell length), with ~65 rows of teeth (
<figureCitation id="43FD2FE8FFA053122D15FD98F584F539" box="[1138,1214,520,544]" captionStart="Figure 7" captionStartId="16.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1322]" captionTargetId="figure-321@16.[129,1473,144,1322]" captionTargetPageId="16" captionText="Figure 7. Shells, operculum, and radulae of Pseudamnicola argentariensis. A, holotype (MNCN 15.05/200500H). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="15" pageNumber="16">Fig. 7H</figureCitation>
). Central tooth formula 4-C-4/1-1 (
<figureCitation id="43FD2FE8FFA053122AC1FDB7F2D2F519" box="[934,1000,551,575]" captionStart="Figure 7" captionStartId="16.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1322]" captionTargetId="figure-321@16.[129,1473,144,1322]" captionTargetPageId="16" captionText="Figure 7. Shells, operculum, and radulae of Pseudamnicola argentariensis. A, holotype (MNCN 15.05/200500H). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="15" pageNumber="16">Fig. 7I</figureCitation>
); basal tongue V-shaped, length about equal to lateral margin. Lateral tooth formula 3-C-3. Inner marginal teeth having 20–23 tapered cusps, shortening towards the base. Outer marginal teeth with ~25 sharp cusps (
<figureCitation id="43FD2FE8FFA053122D8AFD15F417F5BB" box="[1261,1325,645,669]" captionStart="Figure 7" captionStartId="16.[129,194,1350,1374]" captionTargetBox="[129,1473,144,1322]" captionTargetId="figure-321@16.[129,1473,144,1322]" captionTargetPageId="16" captionText="Figure 7. Shells, operculum, and radulae of Pseudamnicola argentariensis. A, holotype (MNCN 15.05/200500H). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="15" pageNumber="16">Fig. 7J</figureCitation>
).
</paragraph>
<paragraph id="DB79336DFFA053122A22FD35F59DF313" blockId="15.[808,1461,144,1609]" pageId="15" pageNumber="16">
Animal darkly pigmented, except for neck and tentacles (
<figureCitation id="43FD2FE8FFA053122CEAFD35F27CF5FA" captionStart="Figure 8" captionStartId="17.[113,178,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-159@17.[114,1458,144,1358]" captionTargetPageId="17" captionText="Figure 8. Anatomy of Pseudamnicola argentariensis. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis.H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="15" pageNumber="16">Fig. 8F</figureCitation>
). Snout as long as wide, with medium distal lobation; foot size intermediate, with dorsal pigmentation. Ctenidium with 17–19 well-developed gill filaments, occupying ~50% of pallial cavity length and posteriorly positioned. Osphradium of intermediate width and opposite middle of ctenidium (
<figureCitation id="43FD2FE8FFA053122CEAFCD1F272F45F" captionStart="Figure 8" captionStartId="17.[113,178,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-159@17.[114,1458,144,1358]" captionTargetPageId="17" captionText="Figure 8. Anatomy of Pseudamnicola argentariensis. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis.H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="15" pageNumber="16">Fig. 8A</figureCitation>
). Stomach slightly longer than wide, with two chambers almost equal in size and a medium-sized posterior caecum; style sac longer than wide, surrounded by an unpigmented intestine (
<figureCitation id="43FD2FE8FFA053122A52FC2FF2BAF4F1" box="[821,896,959,983]" captionStart="Figure 8" captionStartId="17.[113,178,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-159@17.[114,1458,144,1358]" captionTargetPageId="17" captionText="Figure 8. Anatomy of Pseudamnicola argentariensis. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis.H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="15" pageNumber="16">Fig. 8B</figureCitation>
; Supporting Information, Table S4). Nervous system elongate (mean RPG ratio =.61), slightly pigmented, darker on ganglia than on connectives and commissures; cerebral ganglia approximately equal in size (
<figureCitation id="43FD2FE8FFA053122D2AFB8DF5ADF313" box="[1101,1175,1053,1077]" captionStart="Figure 8" captionStartId="17.[113,178,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-159@17.[114,1458,144,1358]" captionTargetPageId="17" captionText="Figure 8. Anatomy of Pseudamnicola argentariensis. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis.H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="15" pageNumber="16">Fig. 8C</figureCitation>
).
</paragraph>
<paragraph id="DB79336DFFA053122A22FBACF5A9F209" blockId="15.[808,1461,144,1609]" pageId="15" pageNumber="16">
Female glandular oviduct approximately four times longer than wide. Albumen gland shorter than capsule gland.
<collectingRegion id="1902FD8FFFA053122C1EFBCCF489F355" box="[1401,1459,1116,1139]" country="Turkey" name="Bursa" pageId="15" pageNumber="16">Bursa</collectingRegion>
copulatrix pyriform, slightly longer than wide. Bursal duct longer than bursal length. Renal oviduct black-pigmented until the area above the insertion of seminal receptacle, coiled. Seminal receptacle elongate, with a short duct, joining renal oviduct slightly above the insertion point with bursal duct (
<figureCitation id="43FD2FE8FFA053122C24FB68F491F229" box="[1347,1451,1272,1296]" captionStart="Figure 8" captionStartId="17.[113,178,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-159@17.[114,1458,144,1358]" captionTargetPageId="17" captionText="Figure 8. Anatomy of Pseudamnicola argentariensis. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis.H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="15" pageNumber="16">Fig. 8D, E</figureCitation>
; Supporting Information, Table S5).
</paragraph>
<paragraph id="DB79336DFFA053122A22FAA7F2A6F16F" blockId="15.[808,1461,144,1609]" pageId="15" pageNumber="16">
Male genitalia with a prostate gland approximately twice as long as wide, bean-shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (
<figureCitation id="43FD2FE8FFA053122AC8FA05F2C5F28B" box="[943,1023,1429,1453]" captionStart="Figure 8" captionStartId="17.[113,178,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-159@17.[114,1458,144,1358]" captionTargetPageId="17" captionText="Figure 8. Anatomy of Pseudamnicola argentariensis. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis.H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="15" pageNumber="16">Fig. 8H</figureCitation>
). Penis almost as long as head, gradually tapering, with many folds over the entire surface and a small patch of pigmentation on its distal region (
<figureCitation id="43FD2FE8FFA053122DB4FA43F409F2CD" box="[1235,1331,1491,1515]" captionStart="Figure 8" captionStartId="17.[113,178,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-159@17.[114,1458,144,1358]" captionTargetPageId="17" captionText="Figure 8. Anatomy of Pseudamnicola argentariensis. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis.H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="15" pageNumber="16">Fig. 8F, G</figureCitation>
; Supporting Information, Table S6); base moderately wide; attached well behind the right eye; penial duct narrow, curved, coursing close to outer edge.
</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFA053122D2EF9F4F489F185" pageId="15" pageNumber="16" type="biology_ecology">
<paragraph id="DB79336DFFA053122D2EF9F4F5AEF158" blockId="15.[1097,1172,1636,1662]" box="[1097,1172,1636,1662]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA053122D2EF9F4F5AEF158" box="[1097,1172,1636,1662]" italics="true" pageId="15" pageNumber="16">Habitat</emphasis>
</paragraph>
<paragraph id="DB79336DFFA053122A4DF91BF489F185" blockId="15.[810,1459,1675,1699]" box="[810,1459,1675,1699]" pageId="15" pageNumber="16">Endemic to a coastal spring on the Monte Argentario Peninsula.</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFA0530C2D23F952F054F176" lastPageId="17" lastPageNumber="18" pageId="15" pageNumber="16" type="discussion">
<paragraph id="DB79336DFFA053122D23F952F5A2F1FA" blockId="15.[1092,1176,1730,1756]" box="[1092,1176,1730,1756]" pageId="15" pageNumber="16">
<emphasis id="E9B2EF7FFFA053122D23F952F5A2F1FA" box="[1092,1176,1730,1756]" italics="true" pageId="15" pageNumber="16">Remarks</emphasis>
</paragraph>
<paragraph id="DB79336DFFA0530D2A4DF979F052F070" blockId="15.[810,1459,1769,1981]" lastBlockId="16.[129,778,1478,1972]" lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16">
Two hydrobiid populations occurring in Monte Argentario at the Fonte delle tre
<collectingCountry id="A3D173FDFFA053122A90F898F527F006" box="[1015,1053,1800,1824]" name="Fiji" pageId="15" pageNumber="16">Viti</collectingCountry>
Spring and Romito Spring were identified as
<taxonomicName id="1CC648EEFFA053122AE0F8B7F5C9F019" authority="(Calcara 1841)" baseAuthorityName="Calcara" baseAuthorityYear="1841" box="[903,1267,1831,1855]" class="Gastropoda" family="Amnicolidae" genus="Amnicola" kingdom="Animalia" order="Littorinimorpha" pageId="15" pageNumber="16" phylum="Mollusca" rank="species" species="moussoni">
<emphasis id="E9B2EF7FFFA053122AE0F8B7F573F019" box="[903,1097,1831,1855]" italics="true" pageId="15" pageNumber="16">Amnicola moussoni</emphasis>
(
<bibRefCitation id="BF574E9CFFA053122D3AF8B7F5D2F019" author="Calcara P" box="[1117,1256,1831,1855]" pageId="15" pageNumber="16" pagination="1 - 11" refId="ref20402" refString="Calcara P. Monograf dei generi Spirorbis e Succinea seguite da alcune nuove specie di conchiglie siciliane. Estratto dal Giornale Letterario 1841; 226: 1 - 11." type="journal article" year="1841">Calcara 1841</bibRefCitation>
)
</taxonomicName>
(=
<taxonomicName id="1CC648EEFFA053122C46F8B7F2B0F078" baseAuthorityName="Falkner and Boeters" baseAuthorityYear="2003" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="15" pageNumber="16" phylum="Mollusca" rank="species" species="moussonii">
<emphasis id="E9B2EF7FFFA053122C46F8B7F2B0F078" italics="true" pageId="15" pageNumber="16">Pseudamnicola moussonii</emphasis>
</taxonomicName>
) and
<taxonomicName id="1CC648EEFFA053122AADF8D7F436F078" authority="(Benoit, 1875)" baseAuthorityName="Benoit" baseAuthorityYear="1875" box="[970,1292,1862,1887]" class="Gastropoda" family="Amnicolidae" genus="Amnicola" kingdom="Animalia" order="Littorinimorpha" pageId="15" pageNumber="16" phylum="Mollusca" rank="species" species="vestita">
<emphasis id="E9B2EF7FFFA053122AADF8D7F551F078" box="[970,1131,1863,1887]" italics="true" pageId="15" pageNumber="16">Amnicola vestita</emphasis>
(Benoit, 1875)
</taxonomicName>
, respectively, by
<bibRefCitation id="BF574E9CFFA053122A4CF8F6F2EBF058" author="Paulucci M" box="[811,977,1894,1918]" pageId="15" pageNumber="16" pagination="5 - 64" refId="ref23104" refString="Paulucci M. Fauna Italiana. Comunicazioni malacologiche. Articolo nono. Conchiglie terrestri e d'acqua dolce del Monte Argentare e delle isole circostanti. Bullettino della Societa Malacologica Italiana 1886; 12: 5 - 64." type="journal article" year="1886">Paulucci (1886)</bibRefCitation>
. Our studied population near Fontana Caprina is here considered a distinct species because: (i) the two cited species by
<bibRefCitation id="BF574E9CFFA053122AF3F835F503F09B" author="Paulucci M" box="[916,1081,1957,1981]" pageId="15" pageNumber="16" pagination="5 - 64" refId="ref23104" refString="Paulucci M. Fauna Italiana. Comunicazioni malacologiche. Articolo nono. Conchiglie terrestri e d'acqua dolce del Monte Argentare e delle isole circostanti. Bullettino della Societa Malacologica Italiana 1886; 12: 5 - 64." type="journal article" year="1886">Paulucci (1886)</bibRefCitation>
were originally described from Sicily, and our integrative evidence suggested that the
<taxonomicName id="1CC648EEFFBF530D2B10FA56F1F0F2DB" pageId="16" pageNumber="17">
<emphasis id="E9B2EF7FFFBF530D2B10FA56F233F2F8" box="[631,777,1478,1502]" italics="true" pageId="16" pageNumber="17">Pseudamnicola</emphasis>
species
</taxonomicName>
from Sicily have more restricted ranges than previously thought; (ii) according to
<bibRefCitation id="BF574E9CFFBF530D28ECF994F320F13A" author="Giusti F" box="[395,538,1540,1564]" pageId="16" pageNumber="17" pagination="99 - 355" refId="ref21518" refString="Giusti F. Notulae Malacologicae XXIII. I molluschi terrestri, salmastri e di acqua dolce dell'Elba, Giannutri e scogli minori dell'Arcipelago Toscano. Biogeographia - The Journal of Integrative Biogeography 1976; 5: 99 - 355." type="journal article" year="1976">Giusti (1976)</bibRefCitation>
,
<taxonomicName id="1CC648EEFFBF530D2B42F995F345F13A" baseAuthorityName="Benoit" baseAuthorityYear="1875" box="[549,639,1541,1564]" class="Gastropoda" family="Amnicolidae" genus="Amnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="vestita">
<emphasis id="E9B2EF7FFFBF530D2B42F995F345F13A" box="[549,639,1541,1564]" italics="true" pageId="16" pageNumber="17">A. vestita</emphasis>
</taxonomicName>
is a synonym of
<taxonomicName id="1CC648EEFFBF530D29FDF9B4F02AF11D" baseAuthorityName="Falkner and Boeters" baseAuthorityYear="2003" box="[154,272,1572,1595]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="moussonii">
<emphasis id="E9B2EF7FFFBF530D29FDF9B4F02AF11D" box="[154,272,1572,1595]" italics="true" pageId="16" pageNumber="17">P. moussonii</emphasis>
</taxonomicName>
; and (iii) there are clear genetic (6.2% average sequence divergence for
<emphasis id="E9B2EF7FFFBF530D2801F9D3F0A8F17C" box="[358,402,1603,1626]" italics="true" pageId="16" pageNumber="17">COI</emphasis>
) and morphological differences (e.g. in shell or female genitalia features) between
<taxonomicName id="1CC648EEFFBF530D2B0EF9F3F233F15C" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[617,777,1635,1658]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="argentariensis">
<emphasis id="E9B2EF7FFFBF530D2B0EF9F3F233F15C" box="[617,777,1635,1658]" italics="true" pageId="16" pageNumber="17">P. argentariensis</emphasis>
</taxonomicName>
and
<taxonomicName id="1CC648EEFFBF530D29D3F912F008F1BF" baseAuthorityName="Falkner and Boeters" baseAuthorityYear="2003" box="[180,306,1666,1689]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="moussonii">
<emphasis id="E9B2EF7FFFBF530D29D3F912F008F1BF" box="[180,306,1666,1689]" italics="true" pageId="16" pageNumber="17">P. moussonii</emphasis>
</taxonomicName>
.
<bibRefCitation id="BF574E9CFFBF530D2822F912F36BF1BC" author="Manganelli G &amp; Barbato D &amp; Benocci A" box="[325,593,1666,1690]" pageId="16" pageNumber="17" pagination="103 - 28" refId="ref22589" refString="Manganelli G, Barbato D, Benocci A. I molluschi terrestri e d'acqua dolce del Monte Argentario. Atti della Societa Toscana di Scienze Naturali Residente in Pisa Memorie Serie B 2017; 123: 103 - 28." type="journal article" year="2017">
Manganelli
<emphasis id="E9B2EF7FFFBF530D28A3F912F0C1F1BC" box="[452,507,1666,1690]" italics="true" pageId="16" pageNumber="17">et al.</emphasis>
(2017)
</bibRefCitation>
reported several
<taxonomicName id="1CC648EEFFBF530D29E6F931F029F19F" authorityName="Paulucci" authorityYear="1878" box="[129,275,1697,1721]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="genus">
<emphasis id="E9B2EF7FFFBF530D29E6F931F029F19F" box="[129,275,1697,1721]" italics="true" pageId="16" pageNumber="17">Pseudamnicola</emphasis>
</taxonomicName>
populations in Monte Argentario, one from the same area as our studied snails, and tentatively classified them into
<taxonomicName id="1CC648EEFFBF530D29D6F970F013F1D1" baseAuthorityName="Falkner and Boeters" baseAuthorityYear="2003" box="[177,297,1760,1783]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="moussonii">
<emphasis id="E9B2EF7FFFBF530D29D6F970F013F1D1" box="[177,297,1760,1783]" italics="true" pageId="16" pageNumber="17">P. moussonii</emphasis>
</taxonomicName>
. Based on our species delimitation results, we recommend further genetic verification of this species identification, because
<taxonomicName id="1CC648EEFFBF530D284FF88FF0FCF010" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[296,454,1823,1846]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="argentariensis">
<emphasis id="E9B2EF7FFFBF530D284FF88FF0FCF010" box="[296,454,1823,1846]" italics="true" pageId="16" pageNumber="17">P. argentariensis</emphasis>
</taxonomicName>
can potentially occur in other localities in the region.
</paragraph>
<caption id="8FB963E5FFBF530D29E6FAD6F054F2B0" ID-DOI="http://doi.org/10.5281/zenodo.14774891" ID-Zenodo-Dep="14774891" httpUri="https://zenodo.org/record/14774891/files/figure.png" pageId="16" pageNumber="17" startId="16.[129,194,1350,1374]" targetBox="[129,1473,144,1322]" targetPageId="16" targetType="figure">
<paragraph id="DB79336DFFBF530D29E6FAD6F054F2B0" blockId="16.[129,1454,1349,1430]" pageId="16" pageNumber="17">
<emphasis id="E9B2EF7FFFBF530D29E6FAD6F1E2F278" bold="true" box="[129,216,1350,1374]" pageId="16" pageNumber="17">Figure 7.</emphasis>
Shells, operculum, and radulae of
<taxonomicName id="1CC648EEFFBF530D2B7DFAD5F201F278" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[538,827,1349,1374]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="argentariensis">
<emphasis id="E9B2EF7FFFBF530D2B7DFAD5F201F278" bold="true" box="[538,827,1349,1374]" italics="true" pageId="16" pageNumber="17">Pseudamnicola argentariensis</emphasis>
</taxonomicName>
. A, holotype (
<emphasis id="E9B2EF7FFFBF530D2AA2FAD6F597F278" box="[965,1197,1350,1374]" italics="true" pageId="16" pageNumber="17">MNCN 15.05/200500H</emphasis>
). B–J, paratypes (UGSB 11999). A–D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, inner and outer marginal teeth.
</paragraph>
</caption>
<paragraph id="DB79336DFFBF530D29FBF8CDF5DAF1BC" blockId="16.[129,778,1478,1972]" lastBlockId="16.[825,1475,1478,1972]" pageId="16" pageNumber="17">
<taxonomicName id="1CC648EEFFBF530D29FBF8CDF083F053" ID-CoL="C86QQ" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[156,441,1885,1909]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="argentariensis">
<emphasis id="E9B2EF7FFFBF530D29FBF8CDF083F053" box="[156,441,1885,1909]" italics="true" pageId="16" pageNumber="17">Pseudamnicola argentariensis</emphasis>
</taxonomicName>
is one of the few slender-shelled representatives of the genus. In comparison to other slender-shelled species, it can be distinguished by: its wider spire and larger aperture from
<taxonomicName id="1CC648EEFFBF530D2D6AFA56F5D7F2F8" ID-CoL="4NJX9" baseAuthorityName="Kuster" baseAuthorityYear="1852" box="[1037,1261,1478,1502]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="luteolus">
<emphasis id="E9B2EF7FFFBF530D2D6AFA56F5D7F2F8" box="[1037,1261,1478,1502]" italics="true" pageId="16" pageNumber="17">Pseudamnicola luteolus</emphasis>
</taxonomicName>
(Küster 1852-1853) from
<collectingCountry id="A3D173FDFFBF530D2A15FA75F281F2DB" box="[882,955,1509,1533]" name="Algeria" pageId="16" pageNumber="17">Algeria</collectingCountry>
(
<bibRefCitation id="BF574E9CFFBF530D2AACFA75F54DF2DB" author="Gloer P &amp; Bouzid S &amp; Boeters HD" box="[971,1143,1509,1533]" pageId="16" pageNumber="17" pagination="1 - 22" refId="ref21786" refString="Gloer P, Bouzid S, Boeters HD. Revision of the genera Pseudamnicola Paulucci 1878 and Mercuria Boeters 1971 from Algeria with particular emphasis on museum collections (Gastropoda: Prosobranchia: Hydrobiidae). Archiv fur Molluskenkunde 2010; 139: 1 - 22. https: // doi. org / 10.1127 / arch. moll / 1869 - 0963 / 139 / 001 - 022" type="journal article" year="2010">
Glöer
<emphasis id="E9B2EF7FFFBF530D2D6AFA76F507F2DB" box="[1037,1085,1509,1533]" italics="true" pageId="16" pageNumber="17">et al.</emphasis>
2010
</bibRefCitation>
: fig. 21); its smaller shell, larger aperture, less convex spire whorls, longer bursal duct, and wider penis from
<taxonomicName id="1CC648EEFFBF530D2ACCF9B4F516F11D" ID-CoL="6WCHQ" authorityName="Gloer &amp; Zettler" authorityYear="2007" box="[939,1068,1572,1595]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="granjaensis">
<emphasis id="E9B2EF7FFFBF530D2ACCF9B4F516F11D" box="[939,1068,1572,1595]" italics="true" pageId="16" pageNumber="17">P. granjaensis</emphasis>
</taxonomicName>
from Majorca (
<bibRefCitation id="BF574E9CFFBF530D2DABF9B4F4A9F11A" author="Delicado D &amp; Machordom A &amp; Ramos MA" box="[1228,1427,1572,1596]" pageId="16" pageNumber="17" pagination="38 - 71" refId="ref20668" refString="Delicado D, Machordom A, Ramos MA. Vicariant versus dispersal processes in the settlement of Pseudamnicola (Caenogastropoda, Hydrobiidae) in the Mediterranean Balearic Islands. Zoological Journal of the Linnean Society 2014; 171: 38 - 71. https: // doi. org / 10.1111 / zoj. 12124" type="journal article" year="2014">
Delicado
<emphasis id="E9B2EF7FFFBF530D2C4AF9B4F461F11A" box="[1325,1371,1572,1596]" italics="true" pageId="16" pageNumber="17">et al.</emphasis>
2014
</bibRefCitation>
: figs 4, 8); and its more globous shell, wider spire, and larger aperture from
<taxonomicName id="1CC648EEFFBF530D2A15F9F2F4B3F15D" ID-CoL="78CHT" authority="Gloer &amp; Beckmann, 2007" authorityName="Gloer &amp; Beckmann" authorityYear="2007" box="[882,1417,1634,1659]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="sciaccaensis">
<emphasis id="E9B2EF7FFFBF530D2A15F9F2F540F15C" box="[882,1146,1634,1658]" italics="true" pageId="16" pageNumber="17">Pseudamnicola sciaccaensis</emphasis>
<bibRefCitation id="BF574E9CFFBF530D2DE7F9F2F4B3F15D" author="Gloer P &amp; Beckmann KH" box="[1152,1417,1634,1659]" pageId="16" pageNumber="17" pagination="121 - 3" refId="ref21703" refString="Gloer P, Beckmann KH. Pseudamnicola sciaccaensis n. sp., a new small prosobranch snail from Sicily (Gastropoda: Hydrobiidae). Mollusca 2007; 25: 121 - 3." type="journal article" year="2007">Glöer &amp; Beckmann, 2007</bibRefCitation>
</taxonomicName>
from Sicily (
<bibRefCitation id="BF574E9CFFBF530D2AE7F912F5A8F1BC" author="Gloer P &amp; Beckmann KH" box="[896,1170,1666,1690]" pageId="16" pageNumber="17" pagination="121 - 3" refId="ref21703" refString="Gloer P, Beckmann KH. Pseudamnicola sciaccaensis n. sp., a new small prosobranch snail from Sicily (Gastropoda: Hydrobiidae). Mollusca 2007; 25: 121 - 3." type="journal article" year="2007">Glöer and Beckmann 2007</bibRefCitation>
: fig. 1).
</paragraph>
<paragraph id="DB79336DFFBF530D2A32F931F4F8F095" blockId="16.[825,1475,1478,1972]" pageId="16" pageNumber="17">
Snails were found in high abundance at the
<typeStatus id="047D8DCFFFBF530D2C6AF932F401F19C" box="[1293,1339,1698,1722]" pageId="16" pageNumber="17">type</typeStatus>
locality, with the great majority being juveniles. Adult shells differed from the ones of the closely related
<taxonomicName id="1CC648EEFFBF530D2D33F970F58CF1DE" ID-CoL="4NJX7" baseAuthorityName="Issel" baseAuthorityYear="1866" box="[1108,1206,1760,1784]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="lucensis">
<emphasis id="E9B2EF7FFFBF530D2D33F970F58CF1DE" box="[1108,1206,1760,1784]" italics="true" pageId="16" pageNumber="17">P. lucensis</emphasis>
</taxonomicName>
according to their generally higher spire and larger aperture (Supporting Information,
<figureCitation id="43FD2FE8FFBF530D2A5EF88EF2BFF011" box="[825,901,1822,1847]" captionStart="Figure 9" captionStartId="18.[129,194,1896,1920]" captionTargetBox="[129,1473,690,1868]" captionTargetId="figure-231@18.[129,1473,690,1868]" captionTargetPageId="18" captionText="Figure 9. Shells, operculum, and radulae of Pseudamnicola bodoni. A, holotype (MNCN 15.05/200502H). B, paratype (UGSB 988). C, shell (UGSB 5697). D, shell (UGSB 16628). E, F, operculum (E, inner side; F, outer side) (UGSB 988). G, protoconch (UGSB 988). H, portion of radula ribbon (UGSB 988). I, central radular teeth (UGSB 988). J, inner and outer marginal teeth (UGSB 988)." figureDoi="http://doi.org/10.5281/zenodo.14774895" httpUri="https://zenodo.org/record/14774895/files/figure.png" pageId="16" pageNumber="17">Fig. S9</figureCitation>
).
<taxonomicName id="1CC648EEFFBF530D2AC6F88EF5FEF010" ID-CoL="C86QQ" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[929,1220,1822,1846]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="16" pageNumber="17" phylum="Mollusca" rank="species" species="argentariensis">
<emphasis id="E9B2EF7FFFBF530D2AC6F88EF5FEF010" box="[929,1220,1822,1846]" italics="true" pageId="16" pageNumber="17">Pseudamnicola argentariensis</emphasis>
</taxonomicName>
differs anatomically by having a larger bursa copulatrix, more slender penis, longer radula, and fewer cusps on the lateral radular teeth (
<bibRefCitation id="BF574E9CFFBF530D2C24F8CDF481F053" author="Giusti F" box="[1347,1467,1885,1909]" pageId="16" pageNumber="17" pagination="99 - 355" refId="ref21518" refString="Giusti F. Notulae Malacologicae XXIII. I molluschi terrestri, salmastri e di acqua dolce dell'Elba, Giannutri e scogli minori dell'Arcipelago Toscano. Biogeographia - The Journal of Integrative Biogeography 1976; 5: 99 - 355." type="journal article" year="1976">Giusti 1976</bibRefCitation>
; Supporting Information,
<figureCitation id="43FD2FE8FFBF530D2D22F8ECF5A7F0B3" box="[1093,1181,1916,1941]" captionStart="Figure 10" captionStartId="19.[115,180,1386,1410]" captionTargetBox="[114,1458,144,1358]" captionTargetId="figure-312@19.[114,1458,144,1358]" captionTargetPageId="19" captionText="Figure 10.Anatomy of Pseudamnicola bodoni. A–H, paratypes (UGSB 988). A, ctenidium and osphradium. B, stomach. C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, head of male and penis.G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774897" httpUri="https://zenodo.org/record/14774897/files/figure.png" pageId="16" pageNumber="17">Fig. S10</figureCitation>
). Based on these morphological differences, previous studies identified these snails as a
</paragraph>
<paragraph id="DB79336DFFBE530C2916FFDAF078F779" blockId="17.[113,322,74,96]" box="[113,322,74,96]" pageId="17" pageNumber="18">
<emphasis id="E9B2EF7FFFBE530C2916FFDAF191F746" bold="true" box="[113,171,74,96]" pageId="17" pageNumber="18">18 •</emphasis>
Delicado
<emphasis id="E9B2EF7FFFBE530C287EFFDBF078F779" box="[281,322,74,96]" italics="true" pageId="17" pageNumber="18">et al.</emphasis>
</paragraph>
<caption id="8FB963E5FFBE530C2916FAFAF47EF2B8" ID-DOI="http://doi.org/10.5281/zenodo.14774893" ID-Zenodo-Dep="14774893" httpUri="https://zenodo.org/record/14774893/files/figure.png" pageId="17" pageNumber="18" startId="17.[113,178,1386,1410]" targetBox="[114,1458,144,1358]" targetPageId="17" targetType="figure">
<paragraph id="DB79336DFFBE530C2916FAFAF47EF2B8" blockId="17.[113,1425,1386,1439]" pageId="17" pageNumber="18">
<emphasis id="E9B2EF7FFFBE530C2916FAFAF1F3F2A4" bold="true" box="[113,201,1386,1410]" pageId="17" pageNumber="18">Figure 8.</emphasis>
Anatomy of
<taxonomicName id="1CC648EEFFBE530C2825FAFAF359F2A4" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[322,611,1386,1410]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="17" pageNumber="18" phylum="Mollusca" rank="species" species="argentariensis">
<emphasis id="E9B2EF7FFFBE530C2825FAFAF359F2A4" bold="true" box="[322,611,1386,1410]" italics="true" pageId="17" pageNumber="18">Pseudamnicola argentariensis</emphasis>
</taxonomicName>
. A–H, paratypes (UGSB 11999). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle. F, head of male and penis. G, penis. H, prostate gland.
</paragraph>
</caption>
<paragraph id="DB79336DFFBE530C2916FA69F054F176" blockId="17.[113,763,1529,1616]" pageId="17" pageNumber="18">
different species of
<taxonomicName id="1CC648EEFFBE530C285AFA69F0F5F137" authorityName="Paulucci" authorityYear="1878" box="[317,463,1529,1553]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="17" pageNumber="18" phylum="Mollusca" rank="genus">
<emphasis id="E9B2EF7FFFBE530C285AFA69F0F5F137" box="[317,463,1529,1553]" italics="true" pageId="17" pageNumber="18">Pseudamnicola</emphasis>
</taxonomicName>
, not belonging to
<taxonomicName id="1CC648EEFFBE530C2BF4FA6AF3CFF137" baseAuthorityName="Issel" baseAuthorityYear="1866" box="[659,757,1529,1553]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="17" pageNumber="18" phylum="Mollusca" rank="species" species="lucensis">
<emphasis id="E9B2EF7FFFBE530C2BF4FA6AF3CFF137" box="[659,757,1529,1553]" italics="true" pageId="17" pageNumber="18">P. lucensis</emphasis>
</taxonomicName>
, and referred to as
<taxonomicName id="1CC648EEFFBE530C285DF988F333F116" box="[314,521,1560,1584]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="17" pageNumber="18" phylum="Mollusca" rank="species" species="undefined-1">
<emphasis id="E9B2EF7FFFBE530C285DF988F0F6F116" box="[314,460,1560,1584]" italics="true" pageId="17" pageNumber="18">Pseudamnicola</emphasis>
sp. 1
</taxonomicName>
(
<bibRefCitation id="BF574E9CFFBE530C2B7AF988F3CFF116" author="Delicado D &amp; Hauffe T &amp; Wilke T" box="[541,757,1560,1584]" pageId="17" pageNumber="18" refId="ref20825" refString="Delicado D, Hauffe T, Wilke T. Ecological opportunity may facilitate diversification in Palearctic freshwater organisms: a case study on hydrobiid gastropods. BMC Evolutionary Biology 2018; 18: 55. https: // doi. org / 10.1186 / s 12862 - 018 - 1169 - 2" type="journal volume" year="2018">
Delicado
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</paragraph>
</subSubSection>
</treatment>
</document>
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