<documentid="404319F98A0212FA3BCFEC58F3417856"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 & Hauffe, Torsten"docDate="2025"docId="536F827BFFA353122AE1FB7AF3B2F5FA"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 araujoi Delicado, Khalloufi & Hauffe 2025, sp. nov."docType="treatment"docVersion="3"lastPageNumber="16"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="13"updateTime="1738263585638"updateUser="ExternalLinkService">
<mods:titleid="77532D38D04566A8DEC98EE593270C74">A holistic perspective on species delimitation outperforms all methods based on single data types in freshwater gastropods (Caenogastropoda: Hydrobiidae: Pseudamnicola)</mods:title>
<mods:affiliationid="3EB143372F3CF0B8282F5E8990AB7B31">Muséum d’Histoire Naturelle de Marrakech, Cadi Ayyad University, Prince Moulay Abdellah Boulevard, B. P. 2390, 40000 Marrakesh, Morocco</mods:affiliation>
<mods:affiliationid="53BDDD426A3352563E3A2432CC9B324A">Environmental Biomonitoring Laboratory (LR 01 ES 14), Faculty of Sciences of Bizerte, University of Carthage, 7021 Jarzouna, Tunisia</mods:affiliation>
<mods:affiliationid="607E0306576AFCD8BDC3AFFFE33AA647">Department of Biology, University of Fribourg, Chemin du Musée 10, CH- 1700 Fribourg, Switzerland & Swiss Institute of Bioinformatics, Chemin du Musée 10, CH- 1700 Fribourg, Switzerland</mods:affiliation>
<taxonomicNameid="1CC648EEFFA353112AE1FB7AF540F202"authority="Delicado, Khalloufi & Hauffe"authorityName="Delicado, Khalloufi & Hauffe"authorityYear="2025"class="Gastropoda"family="Hydrobiidae"genus="Pseudamnicola"kingdom="Animalia"order="Littorinimorpha"pageId="12"pageNumber="13"phylum="Mollusca"rank="species"species="araujoi"status="sp. nov.">
<figureCitationid="43FD2FE8FFA353112DFDFAA4F592F268"box="[1178,1192,1332,1358]"captionStart="Figure 6"captionStartId="14.[129,194,1387,1411]"captionTargetBox="[129,1473,144,1359]"captionTargetId="figure-304@14.[129,1473,144,1359]"captionTargetPageId="14"captionText="Figure 6. Anatomy of Pseudamnicola araujoi. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach.C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, G, head of male and penis. H, prostate gland."figureDoi="http://doi.org/10.5281/zenodo.14774887"httpUri="https://zenodo.org/record/14774887/files/figure.png"pageId="12"pageNumber="13">6</figureCitation>
Named after Dr Rafael Araujo, curator of Malacology at Museo Nacional de Ciencias Naturales (MNCN-CSIC) (2008–2021), who contributed greatly to the knowledge of freshwater molluscs, including those of
<specimenCodeid="8B609B16FFA353112ADDF93EF569F1E0"box="[954,1107,1710,1734]"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="12"pageNumber="13"type="Museum">MNCN 15.05</specimenCode>
<specimenCodeid="8B609B16FFA353112A26F95DF2E6F1C3"box="[833,988,1741,1765]"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="12"pageNumber="13"type="Museum">MNCN 15.05</specimenCode>
/200505P) in the
<collectionCodeid="BDD7ABA8FFA353112DC0F95DF5C3F1C3"box="[1191,1273,1741,1765]"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="12"pageNumber="13"type="Museum">MNCN</collectionCode>
<specimenCodeid="8B609B16FFA353112AABF819F55BF087"box="[972,1121,1929,1953]"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="12"pageNumber="13"type="Museum">MNCN 15.05</specimenCode>
<taxonomicNameid="1CC648EEFFA253102B6CFAD6F3D1F278"authorityName="Delicado, Khalloufi & Hauffe"authorityYear="2025"box="[523,747,1350,1374]"class="Gastropoda"family="Hydrobiidae"genus="Pseudamnicola"kingdom="Animalia"order="Littorinimorpha"pageId="13"pageNumber="14"phylum="Mollusca"rank="species"species="araujoi">
<paragraphid="DB79336DFFA253102916FA7DF054F1E6"blockId="13.[113,764,1517,1729]"pageId="13"pageNumber="14">Shell small, ovate-conic; protoconch microsculpture pitted; central radular tooth formula 4-C-4/1-1; bursa copulatrix large, pyriform, with a duct longer than bursal length; SR1 long, 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 darkly pigmented, elongate (mean RPG ratio =.60).</paragraph>
). 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 (
); basal tongue V-shaped, length about equal to lateral margin. Lateral tooth formula 3-C-3. Inner marginal teeth having ~20 tapered cusps, shortening towards the base. Outer marginal teeth with ~25 sharp cusps (
<figureCitationid="43FD2FE8FFA253102CEAF8CEF25CF0B3"captionStart="Figure 6"captionStartId="14.[129,194,1387,1411]"captionTargetBox="[129,1473,144,1359]"captionTargetId="figure-304@14.[129,1473,144,1359]"captionTargetPageId="14"captionText="Figure 6. Anatomy of Pseudamnicola araujoi. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach.C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, G, head of male and penis. H, prostate gland."figureDoi="http://doi.org/10.5281/zenodo.14774887"httpUri="https://zenodo.org/record/14774887/files/figure.png"pageId="13"pageNumber="14">Fig. 6F, G</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 (
<figureCitationid="43FD2FE8FFA153132B84F99DF1A4F163"captionStart="Figure 6"captionStartId="14.[129,194,1387,1411]"captionTargetBox="[129,1473,144,1359]"captionTargetId="figure-304@14.[129,1473,144,1359]"captionTargetPageId="14"captionText="Figure 6. Anatomy of Pseudamnicola araujoi. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach.C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, G, head of male and penis. H, prostate gland."figureDoi="http://doi.org/10.5281/zenodo.14774887"httpUri="https://zenodo.org/record/14774887/files/figure.png"pageId="14"pageNumber="15">Fig. 6A</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 (
<figureCitationid="43FD2FE8FFA1531329ECF91BF1ECF185"box="[139,214,1675,1699]"captionStart="Figure 6"captionStartId="14.[129,194,1387,1411]"captionTargetBox="[129,1473,144,1359]"captionTargetId="figure-304@14.[129,1473,144,1359]"captionTargetPageId="14"captionText="Figure 6. Anatomy of Pseudamnicola araujoi. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach.C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, G, head of male and penis. H, prostate gland."figureDoi="http://doi.org/10.5281/zenodo.14774887"httpUri="https://zenodo.org/record/14774887/files/figure.png"pageId="14"pageNumber="15">Fig. 6B</figureCitation>
; Supporting Information, Table S4). Nervous system elongate (mean RPG ratio =.60), darkly pigmented, darker on ganglia than on connectives and commissures; cerebral ganglia approximately equal in size (
<figureCitationid="43FD2FE8FFA1531328C4F979F0D7F027"box="[419,493,1769,1793]"captionStart="Figure 6"captionStartId="14.[129,194,1387,1411]"captionTargetBox="[129,1473,144,1359]"captionTargetId="figure-304@14.[129,1473,144,1359]"captionTargetPageId="14"captionText="Figure 6. Anatomy of Pseudamnicola araujoi. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach.C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, G, head of male and penis. H, prostate gland."figureDoi="http://doi.org/10.5281/zenodo.14774887"httpUri="https://zenodo.org/record/14774887/files/figure.png"pageId="14"pageNumber="15">Fig. 6C</figureCitation>
<taxonomicNameid="1CC648EEFFA153132836FAFAF30BF2A5"authorityName="Delicado, Khalloufi & Hauffe"authorityYear="2025"box="[337,561,1386,1411]"class="Gastropoda"family="Hydrobiidae"genus="Pseudamnicola"kingdom="Animalia"order="Littorinimorpha"pageId="14"pageNumber="15"phylum="Mollusca"rank="species"species="araujoi">
. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacle. F, G, head of male and penis. H, prostate gland.
copulatrix large, pyriform, almost as long as wide. Bursal duct longer than bursal length. Renal oviduct black-pigmented until the insertion of seminal receptacle, making one or two loops. Seminal receptacle elongate, with a short duct, joining renal oviduct at the insertion point with bursal duct (
<figureCitationid="43FD2FE8FFA153132C2CFA5FF480F2C0"box="[1355,1466,1487,1511]"captionStart="Figure 6"captionStartId="14.[129,194,1387,1411]"captionTargetBox="[129,1473,144,1359]"captionTargetId="figure-304@14.[129,1473,144,1359]"captionTargetPageId="14"captionText="Figure 6. Anatomy of Pseudamnicola araujoi. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach.C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, G, head of male and penis. H, prostate gland."figureDoi="http://doi.org/10.5281/zenodo.14774887"httpUri="https://zenodo.org/record/14774887/files/figure.png"pageId="14"pageNumber="15">Fig. 6D, E</figureCitation>
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 (
<figureCitationid="43FD2FE8FFA153132A1CF9FBF2F0F1A5"box="[891,970,1643,1667]"captionStart="Figure 6"captionStartId="14.[129,194,1387,1411]"captionTargetBox="[129,1473,144,1359]"captionTargetId="figure-304@14.[129,1473,144,1359]"captionTargetPageId="14"captionText="Figure 6. Anatomy of Pseudamnicola araujoi. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach.C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, G, head of male and penis. H, prostate gland."figureDoi="http://doi.org/10.5281/zenodo.14774887"httpUri="https://zenodo.org/record/14774887/files/figure.png"pageId="14"pageNumber="15">Fig. 6H</figureCitation>
<figureCitationid="43FD2FE8FFA153132D37F93AF58CF1E4"box="[1104,1206,1706,1730]"captionStart="Figure 6"captionStartId="14.[129,194,1387,1411]"captionTargetBox="[129,1473,144,1359]"captionTargetId="figure-304@14.[129,1473,144,1359]"captionTargetPageId="14"captionText="Figure 6. Anatomy of Pseudamnicola araujoi. A–H, paratypes (UGSB 17299). A, ctenidium and osphradium. B, stomach.C, partial nervous system.D, pallial oviduct. E, bursa copulatrix and seminal receptacle.F, G, head of male and penis. H, prostate gland."figureDoi="http://doi.org/10.5281/zenodo.14774887"httpUri="https://zenodo.org/record/14774887/files/figure.png"pageId="14"pageNumber="15">Fig. 6F, G</figureCitation>
Endemic to a small spring, in which the species lives in high abundance. The substrate is composed of silt and gravel. The average water temperature varies between 14°C in winter and 25°C in summer. The conductivity varies between 0.53 and 0.78 mS. Co-occurring mollusc species are
<taxonomicNameid="1CC648EEFFA0531228A2FF00F044F7E1"authority="Khalloufi, Bejaoui & Delicado, 2017"authorityName="Khalloufi, Bejaoui & Delicado"authorityYear="2017"class="Gastropoda"family="Hydrobiidae"genus="Bullaregia"kingdom="Animalia"order="Littorinimorpha"pageId="15"pageNumber="16"phylum="Mollusca"rank="species"species="tunisiensis">
<taxonomicNameid="1CC648EEFFA053122B77FF3FF039F7C1"authority="(O. F. Muller, 1774)"baseAuthorityName="O. F. Muller"baseAuthorityYear="1774"class="Gastropoda"family="Lymnaeidae"genus="Galba"kingdom="Animalia"pageId="15"pageNumber="16"phylum="Mollusca"rank="species"species="truncatula">
<figureCitationid="43FD2FE8FFA053122BB7FE3AF1E8F6C7"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 S7F–L, S</figureCitation>
<figureCitationid="43FD2FE8FFA0531229B5FE59F02FF6C7"box="[210,277,457,481]"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">8O–Q</figureCitation>
average sequence divergence of 7.4%. The shell and aperture shapes of
<taxonomicNameid="1CC648EEFFA0531228AAFD99F311F506"authorityName="Delicado, Khalloufi & Hauffe"authorityYear="2025"box="[461,555,521,544]"class="Gastropoda"family="Hydrobiidae"genus="Pseudamnicola"kingdom="Animalia"order="Littorinimorpha"pageId="15"pageNumber="16"phylum="Mollusca"rank="species"species="araujoi">
<taxonomicNameid="1CC648EEFFA0531229ABFDB8F071F519"authorityName="Gloer, Bouzid & Boeters"authorityYear="2010"box="[204,331,551,575]"class="Gastropoda"family="Hydrobiidae"genus="Pseudamnicola"kingdom="Animalia"order="Littorinimorpha"pageId="15"pageNumber="16"phylum="Mollusca"rank="species"species="calamensis">
<taxonomicNameid="1CC648EEFFA053122819FDB8F0FEF519"authorityName="Gloer, Bouzid & Boeters"authorityYear="2010"box="[382,452,551,575]"class="Gastropoda"family="Hydrobiidae"genus="Pseudamnicola"kingdom="Animalia"order="Littorinimorpha"pageId="15"pageNumber="16"phylum="Mollusca"rank="species"species="linae">
<taxonomicNameid="1CC648EEFFA053122914FDD7F1D4F579"authorityName="Gloer, Bouzid & Boeters"authorityYear="2010"box="[115,238,583,607]"class="Gastropoda"family="Hydrobiidae"genus="Pseudamnicola"kingdom="Animalia"order="Littorinimorpha"pageId="15"pageNumber="16"phylum="Mollusca"rank="species"species="calamensis">
: figs 6–20), which are mostly known from their shells. The morphology and morphometry of its female genitalia are, however, different from the other two species (
<bibRefCitationid="BF574E9CFFA05312291BFD54F019F5FA"author="Gloer P & Bouzid S & Boeters HD"box="[124,291,708,732]"pageId="15"pageNumber="16"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">
<figureCitationid="43FD2FE8FFA053122B58FD54F34DF5FA"box="[575,631,708,732]"captionStart="Figure 2"captionStartId="4.[129,194,1781,1805]"captionTargetBox="[129,1473,144,1753]"captionTargetPageId="4"captionText="Figure 2. Morphological variation among Pseudamnicola species. Plots in the left and right columns display the results for the unsupervised model-based clustering and canonical variate analysis (CVA), respectively, based on four different datasets: A, principal components of shell shape; B, shell measurements; C, D, female and male genitalia measurements. The number of analyzed individuals (N) is indicated on the left side. Symbols and numbers on the right represent species assignments and correspond to those used in Figure 1. Abbreviations shown in images are explained in the Materials and methods section."figureDoi="http://doi.org/10.5281/zenodo.14774875"httpUri="https://zenodo.org/record/14774875/files/figure.png"pageId="15"pageNumber="16">Fig. 2</figureCitation>
