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<document id="B3CB27FB351A59384CE6451ED086F8F2" 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="536F827BFFB6530629F3F88FF1FCF7E1" 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 sardiniensis Delicado, Boulaassafer &amp; Hauffe 2025, sp. nov." docType="treatment" docVersion="3" lastPageNumber="27" 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="26" updateTime="1738263585638" updateUser="ExternalLinkService">
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<mods:title id="5E4632963E9D22B7BDB38332EE699BF6">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:namePart id="75F1656D27BA16A41885E78B1903E7D7">Delicado, Diana</mods:namePart>
<mods:affiliation id="BAEEFC7A978FB0D6C4DCA61B8A21E604">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="6EA1F035C6537200FA510E98DBCAF105" type="email">didelicado@gmail.com</mods:nameIdentifier>
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<mods:namePart id="523557B07046481CB2540D531E7A4D47">Boulaassafer, Khadija</mods:namePart>
<mods:affiliation id="85CE524754804932FF453991ACD6BF5A">Muséum dHistoire 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="79714AF4610C03BFE90CF77A68BA4DFC">Khalloufi, Noureddine</mods:namePart>
<mods:affiliation id="0F069AC3CA3EDA2577AA67AF22F10636">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="CA91F4D65C8DB9B048DABDAE8103540B">Hauffe, Torsten</mods:namePart>
<mods:affiliation id="BBBB871703D330F5637CD3FA8DB72233">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:date id="D78ABC0C26F2DF2DD9DD6F9AE8FC3CF2">2025</mods:date>
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<treatment id="536F827BFFB6530629F3F88FF1FCF7E1" ID-DOI="http://doi.org/10.5281/zenodo.14774877" ID-Zenodo-Dep="14774877" LSID="urn:lsid:plazi:treatment:536F827BFFB6530629F3F88FF1FCF7E1" httpUri="http://treatment.plazi.org/id/536F827BFFB6530629F3F88FF1FCF7E1" lastPageId="27" lastPageNumber="27" pageId="25" pageNumber="26" scope_family="Hydrobiidae" scope_subClass="Caenogastropoda">
<subSubSection id="93DC60E6FFB6530429F3F88FF338F07F" pageId="25" pageNumber="26" type="nomenclature">
<paragraph id="DB79336DFFB6530429F3F88FF338F07F" blockId="25.[148,728,1823,1881]" pageId="25" pageNumber="26">
<emphasis id="E9B2EF7FFFB6530429F3F88FF338F07F" bold="true" pageId="25" pageNumber="26">
<taxonomicName id="1CC648EEFFB6530429F3F88FF088F07E" authority="Delicado, Boulaassafer &amp; Hauffe" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="sardiniensis" status="sp. nov.">
<emphasis id="E9B2EF7FFFB6530429F3F88FF0F3F01C" bold="true" box="[148,457,1823,1850]" italics="true" pageId="25" pageNumber="26">Pseudamnicola sardiniensis</emphasis>
Delicado, Boulaassafer &amp; Hauffe
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<taxonomicNameLabel id="F2815204FFB6530428D1F8AFF338F07F" box="[438,514,1855,1881]" pageId="25" pageNumber="26" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
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<subSubSection id="93DC60E6FFB653042814F8E5F59EF3BA" pageId="25" pageNumber="26" type="description">
<paragraph id="DB79336DFFB653042814F8E5F0C3F0A9" blockId="25.[113,600,1909,2004]" box="[371,505,1909,1936]" pageId="25" pageNumber="26">
<emphasis id="E9B2EF7FFFB653042814F8E5F0C3F0A9" bold="true" box="[371,505,1909,1936]" pageId="25" pageNumber="26">
(
<figureCitation id="43FD2FE8FFB65304281AF8E5F0F3F0B6" box="[381,457,1909,1936]" captionStart="Figure 15" captionStartId="25.[113,178,1025,1049]" captionTargetBox="[114,1458,146,998]" captionTargetId="figure-389@25.[114,1458,146,998]" captionTargetPageId="25" captionText="Figure 15. Shells, operculum, and radulae of Pseudamnicola sardiniensis. A, holotype (MNCN 15.05/200504H). BF, paratypes (UGSB 15501). A, B, shell.C, protoconch.D, portion of radula ribbon.E, central radular teeth. F, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774909" httpUri="https://zenodo.org/record/14774909/files/figure.png" pageId="25" pageNumber="26">Figs 15</figureCitation>
,
<figureCitation id="43FD2FE8FFB6530428B5F8E6F0D4F0B6" box="[466,494,1910,1936]" captionStart="Figure 16" captionStartId="26.[129,194,1022,1046]" captionTargetBox="[129,1473,144,994]" captionTargetId="figure-484@26.[129,1473,144,994]" captionTargetPageId="26" captionText="Figure 16. Anatomy of Pseudamnicola sardiniensis. AF, paratypes (UGSB 15501). A, ctenidium and osphradium.B, stomach. C, partial nervous system. D, head of male and penis. E, penis.F, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774911" httpUri="https://zenodo.org/record/14774911/files/figure.png" pageId="25" pageNumber="26">16</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="DB79336DFFB653042916F80CF323F092" blockId="25.[113,600,1909,2004]" box="[113,537,1948,1972]" pageId="25" pageNumber="26">
<taxonomicName id="1CC648EEFFB653042916F80CF00FF092" box="[113,309,1948,1972]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="undefined-7">
<emphasis id="E9B2EF7FFFB653042916F80CF039F092" box="[113,259,1948,1972]" italics="true" pageId="25" pageNumber="26">Pseudamnicola</emphasis>
sp. 7
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<bibRefCitation id="BF574E9CFFB653042836F80CF323F092" author="Delicado D &amp; Hauffe T &amp; Wilke T" box="[337,537,1948,1972]" pageId="25" pageNumber="26" 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="E9B2EF7FFFB6530428D5F80DF0DBF092" box="[434,481,1948,1972]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2018
</bibRefCitation>
</paragraph>
<paragraph id="DB79336DFFB6530429EAF82CF362F0F2" blockId="25.[113,600,1909,2004]" box="[141,600,1980,2004]" pageId="25" pageNumber="26">
<taxonomicName id="1CC648EEFFB6530429EAF82CF06AF0F5" box="[141,336,1980,2004]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="25" pageNumber="26" phylum="Mollusca" rank="species" species="undefined-7">
<emphasis id="E9B2EF7FFFB6530429EAF82CF025F0F2" box="[141,287,1980,2004]" italics="true" pageId="25" pageNumber="26">Pseudamnicola</emphasis>
sp. 7
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<bibRefCitation id="BF574E9CFFB65304280BF82CF362F0F2" author="Boulaassafer K &amp; Ghamizi M &amp; Machordom A" box="[364,600,1980,2004]" pageId="25" pageNumber="26" 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="E9B2EF7FFFB653042896F82CF31AF0F2" box="[497,544,1980,2004]" italics="true" pageId="25" pageNumber="26">et al.</emphasis>
2020
</bibRefCitation>
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<emphasis id="E9B2EF7FFFB653042A4DFBF5F522F35B" box="[810,1048,1125,1149]" italics="true" pageId="25" pageNumber="26">ZooBank registration:</emphasis>
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urn:lsid:zoobank.org:act:
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<emphasis id="E9B2EF7FFFB653042D5BFB2CF598F3F0" box="[1084,1186,1212,1238]" italics="true" pageId="25" pageNumber="26">Etymology</emphasis>
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<paragraph id="DB79336DFFB653042A4DFB73F2DBF23C" blockId="25.[810,1459,1251,1306]" pageId="25" pageNumber="26">
The species name refers to the occurrence of this species on the island of
<collectingRegion id="1902FD8FFFB653042AE1FA92F2E7F23C" box="[902,989,1282,1306]" country="Italy" name="Sardegna" pageId="25" pageNumber="26">Sardinia</collectingRegion>
.
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<emphasis id="E9B2EF7FFFB653042D49FAAEF595F271" box="[1070,1199,1341,1367]" italics="true" pageId="25" pageNumber="26">Type material</emphasis>
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(
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<specimenCode id="8B609B16FFB653042AF9FAF4F510F25D" box="[926,1066,1379,1403]" 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="25" pageNumber="26" type="Museum">MNCN 15.05</specimenCode>
/200504H
</emphasis>
) and
<specimenCount id="CDC0F8E4FFB653042DB2FAF3F449F25A" box="[1237,1395,1379,1404]" count="3" pageId="25" pageNumber="26" type="generic" typeStatus="paratype">three paratypes</specimenCount>
in the
</materialsCitation>
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<collectionCode id="BDD7ABA8FFB653042A4CFA13F254F2BD" box="[811,878,1411,1435]" pageId="25" pageNumber="26">UGSB</collectionCode>
collection (
<specimenCode id="8B609B16FFB653042A88FA13F5BCF2BC" box="[1007,1158,1410,1435]" collectionCode="UGSB" pageId="25" pageNumber="26">UGSB 15501</specimenCode>
).
<typeStatus id="047D8DCFFFB653042DFBFA13F5EBF2BD" box="[1180,1233,1411,1435]" pageId="25" pageNumber="26">Type</typeStatus>
locality: stream near
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sa
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,
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,
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,
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.
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<emphasis id="E9B2EF7FFFB653042D45FA4DF581F2D1" box="[1058,1211,1501,1527]" italics="true" pageId="25" pageNumber="26">Material studied</emphasis>
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<paragraph id="DB79336DFFB653042A4DF993F5BAF17C" blockId="25.[810,1459,1501,1626]" pageId="25" pageNumber="26">
<materialsCitation id="6BAE3930FFB653042A4DF993F541F17C" collectingDate="2000-09" collectionCode="MNCN, UGSB" collectorName="Colomba MS" country="Italy" county="Nurallao" latitude="39.84" location="Stream" longLatPrecision="6" longitude="9.1052" municipality="Mela" pageId="25" pageNumber="26" specimenCode="MNCN 15.05, UGSB 15501" specimenCount="1" stateProvince="Sardegna">
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near
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sa
<collectingMunicipality id="3B1DA917FFB653042A9BF993F508F13D" box="[1020,1074,1539,1563]" pageId="25" pageNumber="26">Mela</collectingMunicipality>
,
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,
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,
<collectingCountry id="A3D173FDFFB653042C62F993F408F13D" box="[1285,1330,1539,1563]" name="Italy" pageId="25" pageNumber="26">Italy</collectingCountry>
,
<geoCoordinate id="BEF255AAFFB653042C59F993F495F13D" box="[1342,1455,1539,1563]" degrees="39.8400" direction="north" orientation="latitude" pageId="25" pageNumber="26" precision="5" value="39.84">39.8400°N</geoCoordinate>
,
<geoCoordinate id="BEF255AAFFB653042A4DF9B2F2B0F11C" box="[810,906,1570,1594]" degrees="9.1052" direction="east" orientation="longitude" pageId="25" pageNumber="26" precision="5" value="9.1052">9.1052°E</geoCoordinate>
, leg.
<collectorName id="763356BBFFB653042AA1F9B3F2CDF11D" box="[966,1015,1571,1595]" name="Colomba MS" pageId="25" pageNumber="26">M.S.</collectorName>
,
<date id="AF7815ADFFB653042D62F9B3F58FF11D" box="[1029,1205,1571,1595]" pageId="25" pageNumber="26" value="2000-09">
<collectingDate id="BF3CEC45FFB653042D62F9B3F58FF11D" box="[1029,1205,1571,1595]" pageId="25" pageNumber="26" value="2000-09">September 2000</collectingDate>
</date>
,
<specimenCode id="8B609B16FFB653042DA4F9B3F463F11C" box="[1219,1369,1570,1595]" 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="25" pageNumber="26" type="Museum">MNCN 15.05</specimenCode>
/200504 and
<specimenCode id="8B609B16FFB653042A31F9D2F2E5F17C" box="[854,991,1602,1626]" collectionCode="UGSB" pageId="25" pageNumber="26">UGSB 15501</specimenCode>
(80% ethanol)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFB653042D27F9EAF5B8F073" pageId="25" pageNumber="26" type="diagnosis">
<paragraph id="DB79336DFFB653042D27F9EAF5A7F1B5" blockId="25.[1088,1181,1658,1683]" box="[1088,1181,1658,1683]" pageId="25" pageNumber="26">
<emphasis id="E9B2EF7FFFB653042D27F9EAF5A7F1B5" box="[1088,1181,1658,1683]" italics="true" pageId="25" pageNumber="26">Diagnosis</emphasis>
</paragraph>
<paragraph id="DB79336DFFB653042A4DF930F5B8F073" blockId="25.[809,1459,1696,1877]" pageId="25" pageNumber="26">Shell small, ovate-conic, with a short spire; protoconch microsculpture pitted; central radular tooth formula (5)4-C-4(5)/1-1; penis gradually tapering, with many folds over the entire surface, a moderately wide base and a small patch of pigmentation on its distal region; nervous system darkly pigmented, elongate (mean RPG ratio =.64).</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFB653072D5FF8E5F2C4F2F5" lastPageId="26" lastPageNumber="27" pageId="25" pageNumber="26" type="description">
<paragraph id="DB79336DFFB653042D5FF8E5F59EF0A8" blockId="25.[810,1460,1909,2002]" box="[1080,1188,1909,1934]" pageId="25" pageNumber="26">
<emphasis id="E9B2EF7FFFB653042D5FF8E5F59EF0A8" box="[1080,1188,1909,1934]" italics="true" pageId="25" pageNumber="26">Description</emphasis>
</paragraph>
<paragraph id="DB79336DFFB653072A4DF80BF3A6F273" blockId="25.[810,1460,1909,2002]" lastBlockId="26.[127,778,1122,1961]" lastPageId="26" lastPageNumber="27" pageId="25" pageNumber="26">
Shell ovate-conic, whorls 4, height 2.0
<quantity id="1C3E9E88FFB653042DA8F80AF427F094" box="[1231,1309,1946,1970]" metricMagnitude="-3" metricUnit="m" metricValue="2.5" pageId="25" pageNumber="26" unit="mm" value="2.5">2.5 mm</quantity>
(
<figureCitation id="43FD2FE8FFB653042C56F80BF496F094" box="[1329,1452,1946,1971]" captionStart="Figure 15" captionStartId="25.[113,178,1025,1049]" captionTargetBox="[114,1458,146,998]" captionTargetId="figure-389@25.[114,1458,146,998]" captionTargetPageId="25" captionText="Figure 15. Shells, operculum, and radulae of Pseudamnicola sardiniensis. A, holotype (MNCN 15.05/200504H). BF, paratypes (UGSB 15501). A, B, shell.C, protoconch.D, portion of radula ribbon.E, central radular teeth. F, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774909" httpUri="https://zenodo.org/record/14774909/files/figure.png" pageId="25" pageNumber="26">Fig. 15A, B</figureCitation>
; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~400 µm wide, whorls 1.5; nucleus ~150 µm wide; protoconch microsculpture pitted (
<figureCitation id="43FD2FE8FFB553072B6AFB11F34AF3BF" box="[525,624,1153,1177]" captionStart="Figure 15" captionStartId="25.[113,178,1025,1049]" captionTargetBox="[114,1458,146,998]" captionTargetId="figure-389@25.[114,1458,146,998]" captionTargetPageId="25" captionText="Figure 15. Shells, operculum, and radulae of Pseudamnicola sardiniensis. A, holotype (MNCN 15.05/200504H). BF, paratypes (UGSB 15501). A, B, shell.C, protoconch.D, portion of radula ribbon.E, central radular teeth. F, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774909" httpUri="https://zenodo.org/record/14774909/files/figure.png" pageId="26" pageNumber="27">Fig. 15C</figureCitation>
). Teleoconch whorls moderately convex, with deep sutures; body whorl large, occupying about three-quarters of total shell length. Aperture longer than wide, slightly oval; inner lip thicker than outer lip; peristome margin simple, straight. Umbilicus narrow, not covered by the inner lip.
<typeStatus id="047D8DCFFFB553072898FA8EF35CF210" box="[511,614,1310,1334]" pageId="26" pageNumber="27" type="holotype">Holotype</typeStatus>
: SL =
<quantity id="1C3E9E88FFB553072BCEFA8EF23CF213" box="[681,774,1310,1334]" metricMagnitude="-3" metricUnit="m" metricValue="2.29" pageId="26" pageNumber="27" unit="mm" value="2.29">2.29 mm</quantity>
, SW =
<quantity id="1C3E9E88FFB5530729A4FAADF01AF273" box="[195,288,1341,1365]" metricMagnitude="-3" metricUnit="m" metricValue="1.75" pageId="26" pageNumber="27" unit="mm" value="1.75">1.75 mm</quantity>
, AL =
<quantity id="1C3E9E88FFB553072802FAADF0F8F273" box="[357,450,1341,1365]" metricMagnitude="-3" metricUnit="m" metricValue="1.34" pageId="26" pageNumber="27" unit="mm" value="1.34">1.34 mm</quantity>
, and AW =
<quantity id="1C3E9E88FFB553072B5DFAADF3A2F273" box="[570,664,1341,1365]" metricMagnitude="-3" metricUnit="m" metricValue="1.03" pageId="26" pageNumber="27" unit="mm" value="1.03">1.03 mm</quantity>
.
</paragraph>
<caption id="8FB963E5FFB5530729E6FC6EF235F314" ID-DOI="http://doi.org/10.5281/zenodo.14774911" ID-Zenodo-Dep="14774911" httpUri="https://zenodo.org/record/14774911/files/figure.png" pageId="26" pageNumber="27" startId="26.[129,194,1022,1046]" targetBox="[129,1473,144,994]" targetPageId="26" targetType="figure">
<paragraph id="DB79336DFFB5530729E6FC6EF235F314" blockId="26.[129,1424,1022,1075]" pageId="26" pageNumber="27">
<emphasis id="E9B2EF7FFFB5530729E6FC6EF1DFF330" bold="true" box="[129,229,1022,1046]" pageId="26" pageNumber="27">Figure 16.</emphasis>
Anatomy of
<taxonomicName id="1CC648EEFFB55307283AFC6EF352F330" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[349,616,1022,1046]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="26" pageNumber="27" phylum="Mollusca" rank="species" species="sardiniensis">
<emphasis id="E9B2EF7FFFB55307283AFC6EF352F330" bold="true" box="[349,616,1022,1046]" italics="true" pageId="26" pageNumber="27">Pseudamnicola sardiniensis</emphasis>
</taxonomicName>
. AF, paratypes (UGSB 15501). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, head of male and penis. E, penis. F, prostate gland.
</paragraph>
</caption>
<paragraph id="DB79336DFFB5530729FBFACDF3F1F2B2" blockId="26.[127,778,1122,1961]" pageId="26" pageNumber="27">Operculum oval, yellowish, about two and a half whorls; muscle attachment area oval and located near the nucleus.</paragraph>
<paragraph id="DB79336DFFB5530729FBFA0BF23EF176" blockId="26.[127,778,1122,1961]" pageId="26" pageNumber="27">
Radular length intermediate, ~500 µm (~25% of total shell length), with ~45 rows of teeth (
<figureCitation id="43FD2FE8FFB5530728A5FA2BF322F2F4" box="[450,536,1466,1491]" captionStart="Figure 15" captionStartId="25.[113,178,1025,1049]" captionTargetBox="[114,1458,146,998]" captionTargetId="figure-389@25.[114,1458,146,998]" captionTargetPageId="25" captionText="Figure 15. Shells, operculum, and radulae of Pseudamnicola sardiniensis. A, holotype (MNCN 15.05/200504H). BF, paratypes (UGSB 15501). A, B, shell.C, protoconch.D, portion of radula ribbon.E, central radular teeth. F, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774909" httpUri="https://zenodo.org/record/14774909/files/figure.png" pageId="26" pageNumber="27">Fig.15D</figureCitation>
). Central tooth formula (5)4-C-4(5)/1-1 (
<figureCitation id="43FD2FE8FFB55307282EFA4AF09EF2D4" box="[329,420,1498,1522]" captionStart="Figure 15" captionStartId="25.[113,178,1025,1049]" captionTargetBox="[114,1458,146,998]" captionTargetId="figure-389@25.[114,1458,146,998]" captionTargetPageId="25" captionText="Figure 15. Shells, operculum, and radulae of Pseudamnicola sardiniensis. A, holotype (MNCN 15.05/200504H). BF, paratypes (UGSB 15501). A, B, shell.C, protoconch.D, portion of radula ribbon.E, central radular teeth. F, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774909" httpUri="https://zenodo.org/record/14774909/files/figure.png" pageId="26" pageNumber="27">Fig. 15E</figureCitation>
); basal tongue V-shaped, length about equal to lateral margin. Lateral tooth formula 4-C-4. Inner marginal teeth having 2025 tapered cusps, shortening towards the base. Outer marginal teeth with ~25 sharp cusps (
<figureCitation id="43FD2FE8FFB553072BC6F9A8F3CEF176" box="[673,756,1592,1616]" captionStart="Figure 15" captionStartId="25.[113,178,1025,1049]" captionTargetBox="[114,1458,146,998]" captionTargetId="figure-389@25.[114,1458,146,998]" captionTargetPageId="25" captionText="Figure 15. Shells, operculum, and radulae of Pseudamnicola sardiniensis. A, holotype (MNCN 15.05/200504H). BF, paratypes (UGSB 15501). A, B, shell.C, protoconch.D, portion of radula ribbon.E, central radular teeth. F, inner and outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.14774909" httpUri="https://zenodo.org/record/14774909/files/figure.png" pageId="26" pageNumber="27">Fig. 15F</figureCitation>
).
</paragraph>
<paragraph id="DB79336DFFB5530729FBF9C7F5F9F3BF" blockId="26.[127,778,1122,1961]" lastBlockId="26.[825,1476,1122,1491]" pageId="26" pageNumber="27">
Animal darkly pigmented, except for neck and tentacles (
<figureCitation id="43FD2FE8FFB553072B84F9C7F194F1A9" captionStart="Figure 16" captionStartId="26.[129,194,1022,1046]" captionTargetBox="[129,1473,144,994]" captionTargetId="figure-484@26.[129,1473,144,994]" captionTargetPageId="26" captionText="Figure 16. Anatomy of Pseudamnicola sardiniensis. AF, paratypes (UGSB 15501). A, ctenidium and osphradium.B, stomach. C, partial nervous system. D, head of male and penis. E, penis.F, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774911" httpUri="https://zenodo.org/record/14774911/files/figure.png" pageId="26" pageNumber="27">Fig. 16D</figureCitation>
). Snout as long as wide, with medium distal lobation; foot size intermediate, with dorsal pigmentation. Ctenidium with ~15 well-developed gill filaments, occupying ~50% of pallial cavity length and posteriorly positioned. Osphradium of intermediate width and opposite middle of ctenidium (
<figureCitation id="43FD2FE8FFB553072BF8F964F3C3F02A" box="[671,761,1780,1804]" captionStart="Figure 16" captionStartId="26.[129,194,1022,1046]" captionTargetBox="[129,1473,144,994]" captionTargetId="figure-484@26.[129,1473,144,994]" captionTargetPageId="26" captionText="Figure 16. Anatomy of Pseudamnicola sardiniensis. AF, paratypes (UGSB 15501). A, ctenidium and osphradium.B, stomach. C, partial nervous system. D, head of male and penis. E, penis.F, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774911" httpUri="https://zenodo.org/record/14774911/files/figure.png" pageId="26" pageNumber="27">Fig. 16A</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="43FD2FE8FFB5530729ECF8E1F1D8F0AF" box="[139,226,1905,1929]" captionStart="Figure 16" captionStartId="26.[129,194,1022,1046]" captionTargetBox="[129,1473,144,994]" captionTargetId="figure-484@26.[129,1473,144,994]" captionTargetPageId="26" captionText="Figure 16. Anatomy of Pseudamnicola sardiniensis. AF, paratypes (UGSB 15501). A, ctenidium and osphradium.B, stomach. C, partial nervous system. D, head of male and penis. E, penis.F, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774911" httpUri="https://zenodo.org/record/14774911/files/figure.png" pageId="26" pageNumber="27">Fig. 16B</figureCitation>
; Supporting Information, Table S4). Nervous system elongate (mean RPG ratio =.64), darkly pigmented, darker on ganglia than on connectives and commissures; cerebral ganglia approximately equal in size (
<figureCitation id="43FD2FE8FFB553072D3BFB11F589F3BF" box="[1116,1203,1153,1177]" captionStart="Figure 16" captionStartId="26.[129,194,1022,1046]" captionTargetBox="[129,1473,144,994]" captionTargetId="figure-484@26.[129,1473,144,994]" captionTargetPageId="26" captionText="Figure 16. Anatomy of Pseudamnicola sardiniensis. AF, paratypes (UGSB 15501). A, ctenidium and osphradium.B, stomach. C, partial nervous system. D, head of male and penis. E, penis.F, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774911" httpUri="https://zenodo.org/record/14774911/files/figure.png" pageId="26" pageNumber="27">Fig. 16C</figureCitation>
).
</paragraph>
<paragraph id="DB79336DFFB553072A32FB31F55DF39F" blockId="26.[825,1476,1122,1491]" box="[853,1127,1185,1209]" pageId="26" pageNumber="27">Female genitalia unknown.</paragraph>
<paragraph id="DB79336DFFB553072A32FB50F2C4F2F5" blockId="26.[825,1476,1122,1491]" pageId="26" pageNumber="27">
Male genitalia with a prostate gland approximately twice longer than wide, bean-shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (
<figureCitation id="43FD2FE8FFB553072D79FA8EF54DF210" box="[1054,1143,1310,1334]" captionStart="Figure 16" captionStartId="26.[129,194,1022,1046]" captionTargetBox="[129,1473,144,994]" captionTargetId="figure-484@26.[129,1473,144,994]" captionTargetPageId="26" captionText="Figure 16. Anatomy of Pseudamnicola sardiniensis. AF, paratypes (UGSB 15501). A, ctenidium and osphradium.B, stomach. C, partial nervous system. D, head of male and penis. E, penis.F, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774911" httpUri="https://zenodo.org/record/14774911/files/figure.png" pageId="26" pageNumber="27">Fig. 16F</figureCitation>
). Penis 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="43FD2FE8FFB553072C25FACDF480F252" box="[1346,1466,1373,1397]" captionStart="Figure 16" captionStartId="26.[129,194,1022,1046]" captionTargetBox="[129,1473,144,994]" captionTargetId="figure-484@26.[129,1473,144,994]" captionTargetPageId="26" captionText="Figure 16. Anatomy of Pseudamnicola sardiniensis. AF, paratypes (UGSB 15501). A, ctenidium and osphradium.B, stomach. C, partial nervous system. D, head of male and penis. E, penis.F, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.14774911" httpUri="https://zenodo.org/record/14774911/files/figure.png" pageId="26" pageNumber="27">Fig. 16D, E</figureCitation>
; Supporting Information, Table S6); base moderately wide; attachment area central; penial duct narrow, curved, coursing close to outer edge.
</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFB553072D3FFA62F28AF176" pageId="26" pageNumber="27" type="biology_ecology">
<paragraph id="DB79336DFFB553072D3FFA62F599F12A" blockId="26.[1112,1187,1522,1548]" box="[1112,1187,1522,1548]" pageId="26" pageNumber="27">
<emphasis id="E9B2EF7FFFB553072D3FFA62F599F12A" box="[1112,1187,1522,1548]" italics="true" pageId="26" pageNumber="27">Habitat</emphasis>
</paragraph>
<paragraph id="DB79336DFFB553072A5EF989F28AF176" blockId="26.[825,1474,1561,1616]" pageId="26" pageNumber="27">Endemic to a small stream, where the species was found in low abundance.</paragraph>
</subSubSection>
<subSubSection id="93DC60E6FFB553062D34F9FFF1FCF7E1" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27" type="discussion">
<paragraph id="DB79336DFFB553072D34F9FFF59DF1AF" blockId="26.[1107,1191,1647,1673]" box="[1107,1191,1647,1673]" pageId="26" pageNumber="27">
<emphasis id="E9B2EF7FFFB553072D34F9FFF59DF1AF" box="[1107,1191,1647,1673]" italics="true" pageId="26" pageNumber="27">Remarks</emphasis>
</paragraph>
<paragraph id="DB79336DFFB553062A5EF906F1FCF7E1" blockId="26.[825,1475,1686,1961]" lastBlockId="27.[113,762,144,199]" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27">
We dissected three of the
<specimenCount id="CDC0F8E4FFB553072D2EF906F5D5F188" box="[1097,1263,1686,1710]" count="8" pageId="26" pageNumber="27" type="generic">eight specimens</specimenCount>
of
<taxonomicName id="1CC648EEFFB553072C75F907F4A0F188" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[1298,1434,1686,1710]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="26" pageNumber="27" phylum="Mollusca" rank="species" species="sardiniensis">
<emphasis id="E9B2EF7FFFB553072C75F907F4A0F188" box="[1298,1434,1686,1710]" italics="true" pageId="26" pageNumber="27">P. sardiniensis</emphasis>
</taxonomicName>
deposited in the UGSB collection and found no female specimens. The shell of this species has a lower spire than that of the other two newly discovered species in
<collectingRegion id="1902FD8FFFB553072DC6F964F5CFF02A" box="[1185,1269,1780,1804]" country="Italy" name="Sardegna" pageId="26" pageNumber="27">Sardinia</collectingRegion>
(
<taxonomicName id="1CC648EEFFB553072C6DF964F4F9F02D" box="[1290,1475,1780,1804]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="26" pageNumber="27" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="E9B2EF7FFFB553072C6DF964F4A6F02A" box="[1290,1436,1780,1804]" italics="true" pageId="26" pageNumber="27">Pseudamnicola</emphasis>
sp.
</taxonomicName>
10 and 11). Its penis is slightly narrower than in the other two species. There is, however, a large genetic divergence (7.5% for
<emphasis id="E9B2EF7FFFB553072A5CF8C2F25FF04F" box="[827,869,1874,1897]" italics="true" pageId="26" pageNumber="27">COI</emphasis>
) between
<taxonomicName id="1CC648EEFFB553072AB4F8C3F560F04C" authorityName="Delicado, Boulaassafer &amp; Hauffe" authorityYear="2025" box="[979,1114,1874,1898]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="26" pageNumber="27" phylum="Mollusca" rank="species" species="sardiniensis">
<emphasis id="E9B2EF7FFFB553072AB4F8C3F560F04C" box="[979,1114,1874,1898]" italics="true" pageId="26" pageNumber="27">P. sardiniensis</emphasis>
</taxonomicName>
and the other two species endemic to
<collectingRegion id="1902FD8FFFB553072A33F8E1F292F0AF" box="[852,936,1905,1929]" country="Italy" name="Sardegna" pageId="26" pageNumber="27">Sardinia</collectingRegion>
(
<taxonomicName id="1CC648EEFFB553072AD0F8E1F553F0AF" box="[951,1129,1905,1929]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="26" pageNumber="27" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="E9B2EF7FFFB553072AD0F8E1F573F0AF" box="[951,1097,1905,1929]" italics="true" pageId="26" pageNumber="27">Pseudamnicola</emphasis>
sp.
</taxonomicName>
10 and 11). The average sequence divergence with
<taxonomicName id="1CC648EEFFB553072A81F801F564F08E" baseAuthorityName="Falkner and Boeters" baseAuthorityYear="2003" box="[998,1118,1937,1960]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="26" pageNumber="27" phylum="Mollusca" rank="species" species="moussonii">
<emphasis id="E9B2EF7FFFB553072A81F801F564F08E" box="[998,1118,1937,1960]" italics="true" pageId="26" pageNumber="27">P. moussonii</emphasis>
</taxonomicName>
, another species of
<taxonomicName id="1CC648EEFFB553072C57F801F4F8F08F" authorityName="Paulucci" authorityYear="1878" box="[1328,1474,1937,1961]" class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" kingdom="Animalia" order="Littorinimorpha" pageId="26" pageNumber="27" phylum="Mollusca" rank="genus">
<emphasis id="E9B2EF7FFFB553072C57F801F4F8F08F" box="[1328,1474,1937,1961]" italics="true" pageId="26" pageNumber="27">Pseudamnicola</emphasis>
</taxonomicName>
reported from the island (
<bibRefCitation id="BF574E9CFFB45306281DFF00F3A2F78E" author="Giusti F &amp; Castagnolo L" box="[378,664,144,168]" pageId="27" pageNumber="28" pagination="227 - 49" refId="ref21558" refString="Giusti F, Castagnolo L. Notulae Malacologicae, XXX. I molluschi viventi, terrestri e d'acqua dolce, nello studio biogeografico dell'isola di Sardegna. Biogeographia - The Journal of Integrative Biogeography 1983; 8: 227 - 49." type="journal article" year="1983">Giusti and Castagnolo 1983</bibRefCitation>
), was 8% for
<emphasis id="E9B2EF7FFFB4530629F3FF20F1FAF7E1" box="[148,192,176,199]" italics="true" pageId="27" pageNumber="28">COI</emphasis>
.
</paragraph>
</subSubSection>
</treatment>
</document>