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322 lines
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<document ID-DOI="http://dx.doi.org/10.3897/zse.96.48312" ID-GBIF-Dataset="7de3ea60-ba35-4831-ae1d-048dc362f8b0" ID-GBIF-Taxon="161939042" ID-Pensoft-Pub="1860-0743-1-25" ID-Pensoft-UUID="4EDA9B5B777551B7BBB0152DD188E5EF" ID-ZooBank="F2C8585A1268443693348B64AE20F6EE" ModsDocID="1860-0743-1-25" checkinTime="1580744411276" checkinUser="pensoft" docAuthor="Sands, Arthur F, Gloeer, Peter, Guerlek, Mustafa E, Albrecht, Christian & Neubauer, Thomas A" docDate="2020" docId="CFE8F3CDA150576C94E3DB2C5E9C9034" docLanguage="en" docName="ZoosystEvol 96(1): 25-66" docOrigin="Zoosystematics and Evolution 96 (1)" docSource="http://dx.doi.org/10.3897/zse.96.48312" docTitle="Theodoxus velox V. Anistratenko in O. Anistratenko et al. 1999" docType="treatment" docVersion="6" id="4EDA9B5B777551B7BBB0152DD188E5EF" lastPageNumber="25" masterDocId="4EDA9B5B777551B7BBB0152DD188E5EF" masterDocTitle="A revision of the extant species of Theodoxus (Gastropoda, Neritidae) in Asia, with the description of three new species" masterLastPageNumber="66" masterPageNumber="25" pageId="0" pageNumber="25" updateTime="1635972787372" updateUser="ExternalLinkService">
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<mods:titleInfo>
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<mods:title>A revision of the extant species of Theodoxus (Gastropoda, Neritidae) in Asia, with the description of three new species</mods:title>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Sands, Arthur F</mods:namePart>
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<mods:affiliation>Justus Liebig Univrsity Giessen, Giessen, Germany</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Gloeer, Peter</mods:namePart>
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<mods:affiliation>Biodiversity Research Laboratory, Hetlingen, Germany</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Guerlek, Mustafa E</mods:namePart>
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<mods:affiliation>Mehmet Akif Ersoy niversity, Burdur, Turkey</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Albrecht, Christian</mods:namePart>
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<mods:affiliation>Justus Liebig University Giessen, Giessen, Germany</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Neubauer, Thomas A</mods:namePart>
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<mods:affiliation>Justus Liebig University Giessen, Giessen, Germany & Naturalis Biodiversity Centre, Leiden, Netherlands</mods:affiliation>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>Zoosystematics and Evolution</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2020</mods:date>
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<mods:detail type="volume">
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<mods:number>96</mods:number>
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<mods:detail type="issue">
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<mods:number>1</mods:number>
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<mods:extent unit="page">
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<mods:start>25</mods:start>
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<mods:end>66</mods:end>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zse.96.48312</mods:url>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zse.96.48312</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1860-0743-1-25</mods:identifier>
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<treatment ID-GBIF-Taxon="161939042" LSID="urn:lsid:plazi:treatment:CFE8F3CDA150576C94E3DB2C5E9C9034" httpUri="http://treatment.plazi.org/id/CFE8F3CDA150576C94E3DB2C5E9C9034" lastPageNumber="25" pageId="0" pageNumber="25">
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<subSubSection pageId="0" pageNumber="25" type="nomenclature">
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<paragraph pageId="0" pageNumber="25">
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<taxonomicName LSID="CFE8F3CD-A150-576C-94E3-DB2C5E9C9034" authority="V. Anistratenko in O. Anistratenko et al., 1999" authorityName="V. Anistratenko in O. Anistratenko et al." authorityYear="1999" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999</taxonomicName>
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<figureCitation captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">Figure 22A-M</figureCitation>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="25" type="reference_group">
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<paragraph pageId="0" pageNumber="25">
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<taxonomicName class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">Theodoxus velox</taxonomicName>
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V. Anistratenko in O.
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<bibRefCitation author="Anistratenko, OY" journalOrPublisher="Vestnik Zoologii" pageId="0" pageNumber="25" pagination="11 - 19" refId="B14" refString="Anistratenko, OY, Starobogatov, YI, Anistratenko, VV, 1999. Mollyuski roda Theodoxus (Gastropoda, Pectinibranchia, Neritidae) Azovo-Chernomorskogo basseyna. Vestnik Zoologii 33 (3): 11 - 19" title="Mollyuski roda Theodoxus (Gastropoda, Pectinibranchia, Neritidae) Azovo-Chernomorskogo basseyna." volume="33" year="1999">Anistratenko et al. 1999</bibRefCitation>
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: 17-18, fig. 4.7;
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<bibRefCitation DOI="https://doi.org/10.3897/zookeys.827.31365" author="Wesselingh, FP" journalOrPublisher="ZooKeys" pageId="0" pageNumber="25" pagination="31 - 124" refId="B159" refString="Wesselingh, FP, Neubauer, TA, Anistratenko, VV, Vinarski, MV, Yanina, T, ter Poorten, JJ, Kijashko, P, Albrecht, C, Anistratenko, OY, D'Hont, A, Frolov, P, Gandara, AM, Gittenberger, A, Gogaladze, A, Karpinsky, M, Lattuada, M, Popa, L, Sands, AF, van de Velde, S, Vandendorpe, J, Wilke, T, 2019. Mollusc species from the Pontocaspian region - an expert opinion list. ZooKeys 827: 31 - 124, DOI: https://doi.org/10.3897/zookeys.827.31365" title="Mollusc species from the Pontocaspian region - an expert opinion list." url="https://doi.org/10.3897/zookeys.827.31365" volume="827" year="2019">Wesselingh et al. 2019</bibRefCitation>
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: 66.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="25" type="type locality">
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<paragraph pageId="0" pageNumber="25">
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<typeStatus>Type</typeStatus>
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locality.
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</paragraph>
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<paragraph pageId="0" pageNumber="25">Dnieper Delta, Zburievkiy Liman, Kherson Region, Ukraine.</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="25" type="type material">
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<paragraph pageId="0" pageNumber="25">
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<typeStatus>Type</typeStatus>
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material.
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</paragraph>
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<paragraph pageId="0" pageNumber="25">
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<materialsCitation collectionCode="IZAN" country="Turkey" typeStatus="Holotype">
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<typeStatus>Holotype</typeStatus>
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and five
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<typeStatus>paratypes</typeStatus>
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are stored in
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<collectionCode collectionName="IZAN">IZAN</collectionCode>
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(without coll. no.)
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</materialsCitation>
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.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="25" type="remarks">
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<paragraph pageId="0" pageNumber="25">Remarks.</paragraph>
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<paragraph pageId="0" pageNumber="25">
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This species was recently discussed by
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<bibRefCitation DOI="https://doi.org/10.3897/zookeys.827.31365" author="Wesselingh, FP" journalOrPublisher="ZooKeys" pageId="0" pageNumber="25" pagination="31 - 124" refId="B159" refString="Wesselingh, FP, Neubauer, TA, Anistratenko, VV, Vinarski, MV, Yanina, T, ter Poorten, JJ, Kijashko, P, Albrecht, C, Anistratenko, OY, D'Hont, A, Frolov, P, Gandara, AM, Gittenberger, A, Gogaladze, A, Karpinsky, M, Lattuada, M, Popa, L, Sands, AF, van de Velde, S, Vandendorpe, J, Wilke, T, 2019. Mollusc species from the Pontocaspian region - an expert opinion list. ZooKeys 827: 31 - 124, DOI: https://doi.org/10.3897/zookeys.827.31365" title="Mollusc species from the Pontocaspian region - an expert opinion list." url="https://doi.org/10.3897/zookeys.827.31365" volume="827" year="2019">Wesselingh et al. (2019)</bibRefCitation>
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. While it was previously considered a junior synonym of
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<taxonomicName genus="T." lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
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<emphasis italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
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</taxonomicName>
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(
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<bibRefCitation author="Vinarski, MV" journalOrPublisher="Severtsov Institute of Ecology and Evolution of RAS, Moscow" pageId="0" pageNumber="25" refId="B157" refString="Vinarski, MV, Kantor, YI, 2016. Analytical catalogue of fresh and brackish water molluscs of Russia and adjacent countries. A.N. Severtsov Institute of Ecology and Evolution of RAS, Moscow" title="Analytical catalogue of fresh and brackish water molluscs of Russia and adjacent countries. A. N." year="2016">Vinarski and Kantor 2016</bibRefCitation>
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), molecular data shows strong monophyletic support for the independence of
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<taxonomicName genus="T." lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
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<emphasis italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
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</taxonomicName>
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(
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<bibRefCitation DOI="https://doi.org/10.1111/jbi.13590" author="Sands, AF" journalOrPublisher="Journal of Biogeography" pageId="0" pageNumber="25" pagination="1274 - 1286" refId="B138" refString="Sands, AF, Sereda, SV, Stelbrink, B, Neubauer, TA, Lazarev, S, Wilke, T, Albrecht, C, 2019a. Contributions of biogeographical functions to species accumulation may change over time in refugial regions. Journal of Biogeography 46 (6): 1274 - 1286, DOI: https://doi.org/10.1111/jbi.13590" title="Contributions of biogeographical functions to species accumulation may change over time in refugial regions." url="https://doi.org/10.1111/jbi.13590" volume="46" year="2019 a">Sands et al. 2019a</bibRefCitation>
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; Fig.
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dated phylogeny of the genus Theodoxus constructed with BEAST, based on existing GenBank and newly incorporated data for COI, 16 S and ATPα (Suppl. material 1: Table S 1). Morphospecies where associations are queried are highlighted by a question mark after the species name (see the " Systematic checklist " for more information). Small green (PP ≥ 0.95) and red (PP <0.95) dots on nodes indicate the relative posterior probability support of divergence events. Node labels (1 - 32) correspond to divergence dates and HPD intervals in millions of years ago (Mya) for selected nodes (see Suppl. material 1: Table S 3). Dashed dark blue and light blue lines represent the minimum interspecific and maximum intraspecific variation determined by Sands et al. (2019 a) to aid Theodoxus spp. delimitation (see see Methods, Phylogenetics and species delimitation and that study for more information on the delimitation methods)." figureDoi="10.3897/zse.96.48312.figure2" httpUri="https://binary.pensoft.net/fig/376451" pageId="0" pageNumber="25">2</figureCitation>
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). Nevertheless,
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<taxonomicName genus="T." lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
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<emphasis italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
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</taxonomicName>
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and
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<taxonomicName genus="T." lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
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<emphasis italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
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</taxonomicName>
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still hold a sister relationship and likely diverged from one another during the early Pleistocene (
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<bibRefCitation DOI="https://doi.org/10.1111/jbi.13590" author="Sands, AF" journalOrPublisher="Journal of Biogeography" pageId="0" pageNumber="25" pagination="1274 - 1286" refId="B138" refString="Sands, AF, Sereda, SV, Stelbrink, B, Neubauer, TA, Lazarev, S, Wilke, T, Albrecht, C, 2019a. Contributions of biogeographical functions to species accumulation may change over time in refugial regions. Journal of Biogeography 46 (6): 1274 - 1286, DOI: https://doi.org/10.1111/jbi.13590" title="Contributions of biogeographical functions to species accumulation may change over time in refugial regions." url="https://doi.org/10.1111/jbi.13590" volume="46" year="2019 a">Sands et al. 2019a</bibRefCitation>
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; Fig.
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dated phylogeny of the genus Theodoxus constructed with BEAST, based on existing GenBank and newly incorporated data for COI, 16 S and ATPα (Suppl. material 1: Table S 1). Morphospecies where associations are queried are highlighted by a question mark after the species name (see the " Systematic checklist " for more information). Small green (PP ≥ 0.95) and red (PP <0.95) dots on nodes indicate the relative posterior probability support of divergence events. Node labels (1 - 32) correspond to divergence dates and HPD intervals in millions of years ago (Mya) for selected nodes (see Suppl. material 1: Table S 3). Dashed dark blue and light blue lines represent the minimum interspecific and maximum intraspecific variation determined by Sands et al. (2019 a) to aid Theodoxus spp. delimitation (see see Methods, Phylogenetics and species delimitation and that study for more information on the delimitation methods)." figureDoi="10.3897/zse.96.48312.figure2" httpUri="https://binary.pensoft.net/fig/376451" pageId="0" pageNumber="25">2</figureCitation>
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).
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<taxonomicName class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">
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<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus velox</emphasis>
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</taxonomicName>
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is challenging to distinguish from
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<taxonomicName genus="T." lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
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<emphasis italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
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</taxonomicName>
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given the overlap in geographic range and similarity of conchological features (Figs
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<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Occurrence map for Theodoxus spp. in Asia. All points conform to specimens used in the phylogenetic analyses (Suppl. material 1: Table S 1), except for T. gloeri for which no molecular data are available and only the type and paratype localities are indicated. Species are partitioned across four identical maps (A - D) to limit overlap of points as far as possible. Barring T. gloeri, closer related species are grouped together (Fig. 2)." figureDoi="10.3897/zse.96.48312.figure3" httpUri="https://binary.pensoft.net/fig/376452" pageId="0" pageNumber="25">3</figureCitation>
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,
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<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Theodoxus fluviatilis (Linnaeus, 1758). A - D. Specimen collected close to Oued Laabid, Morocco (UGSB 18106) (Fig. 2); E - H. Specimen from the Danube River, Woerth an der Donau, Germany (UGSB 24173); I - L. Specimen (T. euxinus - morphotype) collected in the Papuc Creek, Kiyikoey, Turkey (UGSB 24172) (Fig. 2); M - P. Specimen (T. euxinus - morphotype) from Ovidiopol, Ukraine (UGSB 18124); Q - S. Specimen (T. heldreichi - morphotype) collected in Lake Egirdir, Turkey (UGSB 20361); T, U. Specimen (T. danasteri - morphotype) collected in Krasna Kosa, Ukraine (UGSB 18417) (Fig. 2); V. Opercula of a T. danasteri - morphotype from Lake Skadar, Montenegro (UGSB 24171). Specimens A-D, I-L and T-U were used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure9" httpUri="https://binary.pensoft.net/fig/376458" pageId="0" pageNumber="25">9</figureCitation>
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,
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<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Theodoxus fluviatilis (Linnaeus, 1758). A - D. Specimen (conforming to T. sarmaticus) from the Dnieper-Bug estuary, Ukraine (UGSB 18328); E - H. Specimen (T. subthermalis - morphotype) from Novy Afon, Abkhazia, Georgia (UGSB 18402); I - K. Syntype of N. saulcyi from near the Penteli Monastery, Greece (MHNG-MOLL- 111736); L. Specimen (T. saulcyi - morphotype) from Stilos, Crete, Greece (UGSB 20519) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure10" httpUri="https://binary.pensoft.net/fig/376459" pageId="0" pageNumber="25">10</figureCitation>
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,
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<figureCitation captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22</figureCitation>
|
|||
|
|
).
|
|||
|
|
<taxonomicName class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus velox</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
has less phenotypic variability compared to
|
|||
|
|
<taxonomicName genus="T." lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(Figs
|
|||
|
|
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Theodoxus fluviatilis (Linnaeus, 1758). A - D. Specimen collected close to Oued Laabid, Morocco (UGSB 18106) (Fig. 2); E - H. Specimen from the Danube River, Woerth an der Donau, Germany (UGSB 24173); I - L. Specimen (T. euxinus - morphotype) collected in the Papuc Creek, Kiyikoey, Turkey (UGSB 24172) (Fig. 2); M - P. Specimen (T. euxinus - morphotype) from Ovidiopol, Ukraine (UGSB 18124); Q - S. Specimen (T. heldreichi - morphotype) collected in Lake Egirdir, Turkey (UGSB 20361); T, U. Specimen (T. danasteri - morphotype) collected in Krasna Kosa, Ukraine (UGSB 18417) (Fig. 2); V. Opercula of a T. danasteri - morphotype from Lake Skadar, Montenegro (UGSB 24171). Specimens A-D, I-L and T-U were used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure9" httpUri="https://binary.pensoft.net/fig/376458" pageId="0" pageNumber="25">9</figureCitation>
|
|||
|
|
,
|
|||
|
|
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Theodoxus fluviatilis (Linnaeus, 1758). A - D. Specimen (conforming to T. sarmaticus) from the Dnieper-Bug estuary, Ukraine (UGSB 18328); E - H. Specimen (T. subthermalis - morphotype) from Novy Afon, Abkhazia, Georgia (UGSB 18402); I - K. Syntype of N. saulcyi from near the Penteli Monastery, Greece (MHNG-MOLL- 111736); L. Specimen (T. saulcyi - morphotype) from Stilos, Crete, Greece (UGSB 20519) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure10" httpUri="https://binary.pensoft.net/fig/376459" pageId="0" pageNumber="25">10</figureCitation>
|
|||
|
|
,
|
|||
|
|
<figureCitation captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22</figureCitation>
|
|||
|
|
). Moreover,
|
|||
|
|
<taxonomicName genus="T." lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
display more expansive spiral whorls and, in some instances, a more transparent operculum where the conchioline lamella extends deeper into the calcareous base and the callus is less pronounced (Figs
|
|||
|
|
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Theodoxus fluviatilis (Linnaeus, 1758). A - D. Specimen collected close to Oued Laabid, Morocco (UGSB 18106) (Fig. 2); E - H. Specimen from the Danube River, Woerth an der Donau, Germany (UGSB 24173); I - L. Specimen (T. euxinus - morphotype) collected in the Papuc Creek, Kiyikoey, Turkey (UGSB 24172) (Fig. 2); M - P. Specimen (T. euxinus - morphotype) from Ovidiopol, Ukraine (UGSB 18124); Q - S. Specimen (T. heldreichi - morphotype) collected in Lake Egirdir, Turkey (UGSB 20361); T, U. Specimen (T. danasteri - morphotype) collected in Krasna Kosa, Ukraine (UGSB 18417) (Fig. 2); V. Opercula of a T. danasteri - morphotype from Lake Skadar, Montenegro (UGSB 24171). Specimens A-D, I-L and T-U were used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure9" httpUri="https://binary.pensoft.net/fig/376458" pageId="0" pageNumber="25">9</figureCitation>
|
|||
|
|
,
|
|||
|
|
<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Theodoxus fluviatilis (Linnaeus, 1758). A - D. Specimen (conforming to T. sarmaticus) from the Dnieper-Bug estuary, Ukraine (UGSB 18328); E - H. Specimen (T. subthermalis - morphotype) from Novy Afon, Abkhazia, Georgia (UGSB 18402); I - K. Syntype of N. saulcyi from near the Penteli Monastery, Greece (MHNG-MOLL- 111736); L. Specimen (T. saulcyi - morphotype) from Stilos, Crete, Greece (UGSB 20519) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure10" httpUri="https://binary.pensoft.net/fig/376459" pageId="0" pageNumber="25">10</figureCitation>
|
|||
|
|
,
|
|||
|
|
<figureCitation captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22</figureCitation>
|
|||
|
|
).
|
|||
|
|
</paragraph>
|
|||
|
|
<caption doi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25" start="Figure 22" startId="F22">
|
|||
|
|
<paragraph pageId="0" pageNumber="25">
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">Figure 22.</emphasis>
|
|||
|
|
<taxonomicName class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus velox</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
V. Anistratenko in O. Anistratenko et al., 1999.
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">A</emphasis>
|
|||
|
|
-
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">D</emphasis>
|
|||
|
|
. Specimen from the Dnieper River, Kiev, Ukraine (
|
|||
|
|
<collectionCode collectionName="UGSB">UGSB</collectionCode>
|
|||
|
|
24186);
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">E</emphasis>
|
|||
|
|
-
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">H</emphasis>
|
|||
|
|
. Specimen collected in the Danube delta, Vylkove, Ukraine (
|
|||
|
|
<collectionCode collectionName="UGSB">UGSB</collectionCode>
|
|||
|
|
24184) used in the phylogeny (Fig.
|
|||
|
|
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Dated phylogeny of the genus Theodoxus constructed with BEAST, based on existing GenBank and newly incorporated data for COI, 16 S and ATPα (Suppl. material 1: Table S 1). Morphospecies where associations are queried are highlighted by a question mark after the species name (see the " Systematic checklist " for more information). Small green (PP ≥ 0.95) and red (PP <0.95) dots on nodes indicate the relative posterior probability support of divergence events. Node labels (1 - 32) correspond to divergence dates and HPD intervals in millions of years ago (Mya) for selected nodes (see Suppl. material 1: Table S 3). Dashed dark blue and light blue lines represent the minimum interspecific and maximum intraspecific variation determined by Sands et al. (2019 a) to aid Theodoxus spp. delimitation (see see Methods, Phylogenetics and species delimitation and that study for more information on the delimitation methods)." figureDoi="10.3897/zse.96.48312.figure2" httpUri="https://binary.pensoft.net/fig/376451" pageId="0" pageNumber="25">2</figureCitation>
|
|||
|
|
);
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">I</emphasis>
|
|||
|
|
,
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">J</emphasis>
|
|||
|
|
.
|
|||
|
|
<typeStatus>Holotype</typeStatus>
|
|||
|
|
from Dnieper Delta, Zbur'ivka Liman, Ukraine (
|
|||
|
|
<collectionCode collectionName="IZAN">IZAN</collectionCode>
|
|||
|
|
);
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">K</emphasis>
|
|||
|
|
-
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">M</emphasis>
|
|||
|
|
. Specimen from Lake Sapanca, Yeni
|
|||
|
|
<normalizedToken originalValue="Eşme">Esme</normalizedToken>
|
|||
|
|
, Turkey (
|
|||
|
|
<collectionCode collectionName="UGSB">UGSB</collectionCode>
|
|||
|
|
20691).
|
|||
|
|
<taxonomicName baseAuthorityName="Lindholm" baseAuthorityYear="1901" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus sarmaticus" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="sarmaticus">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus sarmaticus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(Lindholm, 1901).
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">N</emphasis>
|
|||
|
|
-
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">P</emphasis>
|
|||
|
|
;
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">Q</emphasis>
|
|||
|
|
-
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">S</emphasis>
|
|||
|
|
;
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">T</emphasis>
|
|||
|
|
-
|
|||
|
|
<emphasis bold="true" pageId="0" pageNumber="25">U</emphasis>
|
|||
|
|
. Three
|
|||
|
|
<typeStatus>syntypes</typeStatus>
|
|||
|
|
from the Oskol River, near Golubino, Russia stored in
|
|||
|
|
<collectionCode collectionName="Russia, St. Petersburg, Russian Academy of Sciences, Zoological Institute" country="Russia" httpUri="http://grbio.org/cool/z2y8-2jfr" name="Russian Academy of Sciences, Zoological Institute, Zoological Museum" type="Museum">ZIN</collectionCode>
|
|||
|
|
(coll. no. 6099/24). Scale bars:
|
|||
|
|
<quantity metricMagnitude="-3" metricUnit="m" metricValue="1.0" unit="mm" value="1.0">1 mm</quantity>
|
|||
|
|
.
|
|||
|
|
</paragraph>
|
|||
|
|
</caption>
|
|||
|
|
<paragraph pageId="0" pageNumber="25">
|
|||
|
|
The geographic distribution range of
|
|||
|
|
<taxonomicName genus="T." lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
overlaps with that of
|
|||
|
|
<taxonomicName genus="T." lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(Lindholm, 1901), which has been considered a junior synonym of
|
|||
|
|
<taxonomicName genus="T." lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(e.g.
|
|||
|
|
<bibRefCitation author="Vinarski, MV" journalOrPublisher="Severtsov Institute of Ecology and Evolution of RAS, Moscow" pageId="0" pageNumber="25" refId="B157" refString="Vinarski, MV, Kantor, YI, 2016. Analytical catalogue of fresh and brackish water molluscs of Russia and adjacent countries. A.N. Severtsov Institute of Ecology and Evolution of RAS, Moscow" title="Analytical catalogue of fresh and brackish water molluscs of Russia and adjacent countries. A. N." year="2016">Vinarski and Kantor 2016</bibRefCitation>
|
|||
|
|
). While
|
|||
|
|
<taxonomicName genus="T." lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
and
|
|||
|
|
<taxonomicName genus="T." lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
may on occasion share phenotypic similarity, the morphotypes of
|
|||
|
|
<taxonomicName genus="T." lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
closely resemble
|
|||
|
|
<taxonomicName genus="T." lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(Fig.
|
|||
|
|
<figureCitation captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22A-M</figureCitation>
|
|||
|
|
versus Fig.
|
|||
|
|
<figureCitation captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22N-U</figureCitation>
|
|||
|
|
). The similarity with
|
|||
|
|
<taxonomicName genus="T." lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
could suggest
|
|||
|
|
<taxonomicName genus="T." lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is rather conspecific with that species as suggested by
|
|||
|
|
<bibRefCitation author="Gloeer, P" journalOrPublisher="Peter Gloeer, Hetlingen, Germany" pageId="0" pageNumber="25" refId="B62" refString="Gloeer, P, 2019. The freshwater gastropods of the West-Palaearctis. Vol. 1. Fresh- and brackish waters except spring and subterranean snails. Peter Gloeer, Hetlingen, Germany" title="The freshwater gastropods of the West-Palaearctis. Vol. 1. Fresh- and brackish waters except spring and subterranean snails." year="2019">
|
|||
|
|
<normalizedToken originalValue="Glöer">Gloeer</normalizedToken>
|
|||
|
|
(2019)
|
|||
|
|
</bibRefCitation>
|
|||
|
|
. Unfortunately, no opercula or soft tissues were preserved among the
|
|||
|
|
<typeStatus>syntypes</typeStatus>
|
|||
|
|
of
|
|||
|
|
<taxonomicName genus="T." lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
to corroborate this hypothesis. Molecular analyses of topotypic material is required to address this uncertainty. However, should this be confirmed at a later point, the name
|
|||
|
|
<taxonomicName genus="T." lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
|
|||
|
|
<emphasis italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
would have priority.
|
|||
|
|
</paragraph>
|
|||
|
|
</subSubSection>
|
|||
|
|
<subSubSection pageId="0" pageNumber="25" type="distribution">
|
|||
|
|
<paragraph pageId="0" pageNumber="25">Distribution.</paragraph>
|
|||
|
|
<paragraph pageId="0" pageNumber="25">
|
|||
|
|
This species was indicated to be restricted to drainage systems of the northern Black Sea coast (
|
|||
|
|
<bibRefCitation author="Anistratenko, OY" journalOrPublisher="Vestnik Zoologii" pageId="0" pageNumber="25" pagination="11 - 19" refId="B14" refString="Anistratenko, OY, Starobogatov, YI, Anistratenko, VV, 1999. Mollyuski roda Theodoxus (Gastropoda, Pectinibranchia, Neritidae) Azovo-Chernomorskogo basseyna. Vestnik Zoologii 33 (3): 11 - 19" title="Mollyuski roda Theodoxus (Gastropoda, Pectinibranchia, Neritidae) Azovo-Chernomorskogo basseyna." volume="33" year="1999">Anistratenko et al. 1999</bibRefCitation>
|
|||
|
|
;
|
|||
|
|
<bibRefCitation author="Kantor, YI" journalOrPublisher="KMK Scientific Press, Moscow" pageId="0" pageNumber="25" refId="B78" refString="Kantor, YI, Sysoev, AV, 2006. Morskiye i solonovatovodnyye bryukhonogiye mollyuski Rossii i sopredel'nykh stran: illyustrirovannyy katalog. KMK Scientific Press, Moscow" title="Morskiye i solonovatovodnyye bryukhonogiye mollyuski Rossii i sopredel'nykh stran: illyustrirovannyy katalog." year="2006">Kantor and Sysoev 2006</bibRefCitation>
|
|||
|
|
). Recent molecular data suggest it is distributed as far North as the eastern part of the Baltic Sea and as far South as Anatolia (
|
|||
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<bibRefCitation DOI="https://doi.org/10.1111/jbi.13590" author="Sands, AF" journalOrPublisher="Journal of Biogeography" pageId="0" pageNumber="25" pagination="1274 - 1286" refId="B138" refString="Sands, AF, Sereda, SV, Stelbrink, B, Neubauer, TA, Lazarev, S, Wilke, T, Albrecht, C, 2019a. Contributions of biogeographical functions to species accumulation may change over time in refugial regions. Journal of Biogeography 46 (6): 1274 - 1286, DOI: https://doi.org/10.1111/jbi.13590" title="Contributions of biogeographical functions to species accumulation may change over time in refugial regions." url="https://doi.org/10.1111/jbi.13590" volume="46" year="2019 a">Sands et al. 2019a</bibRefCitation>
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): the only record there derives from Lake Sapanca (
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<bibRefCitation DOI="https://doi.org/10.1111/jbi.13590" author="Sands, AF" journalOrPublisher="Journal of Biogeography" pageId="0" pageNumber="25" pagination="1274 - 1286" refId="B138" refString="Sands, AF, Sereda, SV, Stelbrink, B, Neubauer, TA, Lazarev, S, Wilke, T, Albrecht, C, 2019a. Contributions of biogeographical functions to species accumulation may change over time in refugial regions. Journal of Biogeography 46 (6): 1274 - 1286, DOI: https://doi.org/10.1111/jbi.13590" title="Contributions of biogeographical functions to species accumulation may change over time in refugial regions." url="https://doi.org/10.1111/jbi.13590" volume="46" year="2019 a">Sands et al. 2019a</bibRefCitation>
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; Fig.
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<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Occurrence map for Theodoxus spp. in Asia. All points conform to specimens used in the phylogenetic analyses (Suppl. material 1: Table S 1), except for T. gloeri for which no molecular data are available and only the type and paratype localities are indicated. Species are partitioned across four identical maps (A - D) to limit overlap of points as far as possible. Barring T. gloeri, closer related species are grouped together (Fig. 2)." figureDoi="10.3897/zse.96.48312.figure3" httpUri="https://binary.pensoft.net/fig/376452" pageId="0" pageNumber="25">3C</figureCitation>
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).
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</paragraph>
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</subSubSection>
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</treatment>
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</document>
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