treatments-xml/data/55/E9/1A/55E91AC28D0857A69BA932BD100648AE.xml

<document ID-DOI="http://dx.doi.org/10.3897/zse.96.48312" ID-GBIF-Dataset="7de3ea60-ba35-4831-ae1d-048dc362f8b0" ID-GBIF-Taxon="161939051" ID-Pensoft-Pub="1860-0743-1-25" ID-Pensoft-UUID="4EDA9B5B777551B7BBB0152DD188E5EF" ID-ZooBank="F2C8585A1268443693348B64AE20F6EE" LSID="urn:lsid:zoobank.org:act:97AF034D-EF6D-4AB2-AC17-0C5B037C8DAD" ModsDocID="1860-0743-1-25" checkinTime="1580744411276" checkinUser="pensoft" docAuthor="Sands, Arthur F, Gloeer, Peter, Guerlek, Mustafa E, Albrecht, Christian &amp; Neubauer, Thomas A" docDate="2020" docId="55E91AC28D0857A69BA932BD100648AE" 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 wesselinghi Sands &amp; Gloeer 2020, sp. nov." docType="treatment" docUuid="97AF034D-EF6D-4AB2-AC17-0C5B037C8DAD" docUuidSource="ZooBank" 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: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:namePart>Sands, Arthur F</mods:namePart>
<mods:affiliation>Justus Liebig Univrsity Giessen, Giessen, Germany</mods:affiliation>
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<mods:namePart>Gloeer, Peter</mods:namePart>
<mods:affiliation>Biodiversity Research Laboratory, Hetlingen, Germany</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Guerlek, Mustafa E</mods:namePart>
<mods:affiliation>Mehmet Akif Ersoy niversity, Burdur, Turkey</mods:affiliation>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Albrecht, Christian</mods:namePart>
<mods:affiliation>Justus Liebig University Giessen, Giessen, Germany</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Neubauer, Thomas A</mods:namePart>
<mods:affiliation>Justus Liebig University Giessen, Giessen, Germany &amp; Naturalis Biodiversity Centre, Leiden, Netherlands</mods:affiliation>
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<taxonomicName LSID="55E91AC2-8D08-57A6-9BA9-32BD100648AE" authority="Sands &amp; Glöer" authorityName="Sands &amp; Gloeer" authorityYear="2020" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus wesselinghi" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="wesselinghi" status="sp. nov.">
Theodoxus wesselinghi Sands &amp; 
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</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="25">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">Figures 23A-H</figureCitation>
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">, 24A-M</figureCitation>
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Radula of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. paratypes. A. Portion of the radula showing full sets of teeth (UGSB 20688); B. Magnified view of the central teeth (UGSB 20688); C. Magnified view of the lateral and marginal teeth (UGSB 20686); D. Magnified view of the first and second rows of marginal teeth (UGSB 20688); E. Magnified view of the first row of marginal teeth (UGSB 20686); F. Magnified view of the faces of inner marginal teeth belonging to the first row (UGSB 20685). Scale bars: 100 μm (A-E), 20 μm (F)." figureDoi="10.3897/zse.96.48312.figure25" httpUri="https://binary.pensoft.net/fig/376474" pageId="0" pageNumber="25">, 25A-F</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="25" type="reference_group">
<paragraph pageId="0" pageNumber="25">
<taxonomicName class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus fluviatilis" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="fluviatilis">Theodoxus fluviatilis</taxonomicName>
: 
<bibRefCitation author="Odabasi, DA" journalOrPublisher="Ecologica Montenegrina" pageId="0" pageNumber="25" pagination="327 - 333" refId="B115" refString="Odabasi, DA, Arslan, N, 2015. Description of a new subterranean nerite: Theodoxus gloeri n. sp. with some data on the freshwater gastropod fauna of Balikdami Wetland (Sakarya River, Turkey). Ecologica Montenegrina 2 (4): 327 - 333" title="Description of a new subterranean nerite: Theodoxus gloeri n. sp. with some data on the freshwater gastropod fauna of Balikdami Wetland (Sakarya River, Turkey)." volume="2" year="2015">
<normalizedToken originalValue="Odabaşı">Odabasi</normalizedToken>
and Arslan 2015
</bibRefCitation>
: 330-331 (
<emphasis bold="true" pageId="0" pageNumber="25">non</emphasis>
<taxonomicName authorityName="Linnaeus" authorityYear="1758" class="Gastropoda" family="Neritidae" genus="Nerita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nerita fluviatilis" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="fluviatilis">Nerita fluviatilis</taxonomicName>
Linnaeus, 1758).
</paragraph>
<paragraph pageId="0" pageNumber="25">
<taxonomicName baseAuthorityName="Recluz" baseAuthorityYear="1841" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus anatolicus" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="anatolicus">Theodoxus anatolicus</taxonomicName>
: 
<bibRefCitation author="Yildirim, MZ" journalOrPublisher="Ecologica Montenegrina" pageId="0" pageNumber="25" pagination="115 - 119" refId="B163" refString="Yildirim, MZ, Koca, SB, Guerlek, ME, Gloeer, P, 2018. A new genus and a species from Turkey, Isparta felei n. gen. n. sp.(Gastropoda: Truncatelloidea: Hydrobiidae). Ecologica Montenegrina 18: 115 - 119" title="A new genus and a species from Turkey, Isparta felei n. gen. n. sp. (Gastropoda: Truncatelloidea: Hydrobiidae)." volume="18" year="2018">
<normalizedToken originalValue="Yıldırım">Yildirim</normalizedToken>
et al. 2018
</bibRefCitation>
: 118 (
<emphasis bold="true" pageId="0" pageNumber="25">non</emphasis>
<taxonomicName authorityName="Recluz" authorityYear="1841" class="Gastropoda" family="Neritidae" genus="Nerita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nerita anatolica" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="anatolica">Nerita anatolica</taxonomicName>
<normalizedToken originalValue="Récluz">Recluz</normalizedToken>
, 1841).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="25" type="type locality">
<paragraph pageId="0" pageNumber="25">
<typeStatus>Type</typeStatus>
locality.
</paragraph>
<paragraph pageId="0" pageNumber="25">
Sakarya River, 
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, Bilecik, Turkey; 
<geoCoordinate degrees="40.0439" direction="north" orientation="latitude" precision="5" value="40.0439">40.0439°N</geoCoordinate>
, 
<geoCoordinate degrees="30.452" direction="east" orientation="longitude" precision="55" value="30.452">30.452°E</geoCoordinate>
(Figs 
<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">3D</figureCitation>
, 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23A, B</figureCitation>
).
</paragraph>
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<typeStatus>Holotype</typeStatus>
.
</emphasis>
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.
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.342200 (Sakarya River, 
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, 
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) stored in 
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: Shell height 6.0 mm, width 6.0 mm (Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24A-D</figureCitation>
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.
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<materialsCitation collectionCode="A" country="Turkey" latitude="40.0439" longitude="30.452" typeStatus="Paratypes">
<emphasis bold="true" pageId="0" pageNumber="25">
<typeStatus>Paratypes</typeStatus>
.
</emphasis>
Twenty-four specimens from Sakarya River, 
<normalizedToken originalValue="Çayköy">Caykoey</normalizedToken>
, Bilecik, 
<collectingCountry name="Turkey" value="Turkey">Turkey</collectingCountry>
; 
<geoCoordinate degrees="40.0439" direction="north" orientation="latitude" precision="5" value="40.0439">40.0439°N</geoCoordinate>
, 
<geoCoordinate degrees="30.452" direction="east" orientation="longitude" precision="55" value="30.452">30.452°E</geoCoordinate>
(Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23A, B</figureCitation>
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(
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
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.342201, 
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.342202; Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24E-G</figureCitation>
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20688, 
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20743, 
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20744; Fig. 
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Radula of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. paratypes. A. Portion of the radula showing full sets of teeth (UGSB 20688); B. Magnified view of the central teeth (UGSB 20688); C. Magnified view of the lateral and marginal teeth (UGSB 20686); D. Magnified view of the first and second rows of marginal teeth (UGSB 20688); E. Magnified view of the first row of marginal teeth (UGSB 20686); F. Magnified view of the faces of inner marginal teeth belonging to the first row (UGSB 20685). Scale bars: 100 μm (A-E), 20 μm (F)." figureDoi="10.3897/zse.96.48312.figure25" httpUri="https://binary.pensoft.net/fig/376474" pageId="0" pageNumber="25">25A, B, D</figureCitation>
)
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. 
<materialsCitation collectionCode="C" country="Turkey" latitude="38.00358" longitude="31.47217">
Twenty-four specimens from an unnamed roadside spring in Fele, 
<normalizedToken originalValue="İsparta">Isparta</normalizedToken>
province, 
<collectingCountry name="Turkey" value="Turkey">Turkey</collectingCountry>
; 
<geoCoordinate degrees="38.00358" direction="north" orientation="latitude" precision="1" value="38.00358">38.00358°N</geoCoordinate>
, 
<geoCoordinate degrees="31.47217" direction="east" orientation="longitude" precision="1" value="31.47217">31.47217°E</geoCoordinate>
(Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23C, D</figureCitation>
): 
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(
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342203, 
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342204; Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24H-J</figureCitation>
) and 
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20684, 
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20735, 
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20736)
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. 
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Twenty-five specimens from Eflatun 
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; 
<geoCoordinate degrees="37.8256" direction="north" orientation="latitude" precision="5" value="37.8256">37.8256°N</geoCoordinate>
, 
<geoCoordinate degrees="31.6748" direction="east" orientation="longitude" precision="5" value="31.6748">31.6748°E</geoCoordinate>
(Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23E, F</figureCitation>
): 
<quantity metricMagnitude="-1" metricUnit="m" metricValue="2.794" unit="in" value="11.0">11 in</quantity>
<collectionCode collectionName="NMNL" country="Netherlands" httpUri="http://grbio.org/cool/1yj9-ymxi" name="Natuurmuseum Nijmegen e.o." type="Herbarium">NMNL</collectionCode>
(
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342205, 
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342206; Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24K-M</figureCitation>
) and 
<quantity metricMagnitude="-1" metricUnit="m" metricValue="3.556" unit="in" value="14.0">14 in</quantity>
<collectionCode collectionName="UGSB">UGSB</collectionCode>
(
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20685, 
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20737, 
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20738; Fig. 
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Radula of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. paratypes. A. Portion of the radula showing full sets of teeth (UGSB 20688); B. Magnified view of the central teeth (UGSB 20688); C. Magnified view of the lateral and marginal teeth (UGSB 20686); D. Magnified view of the first and second rows of marginal teeth (UGSB 20688); E. Magnified view of the first row of marginal teeth (UGSB 20686); F. Magnified view of the faces of inner marginal teeth belonging to the first row (UGSB 20685). Scale bars: 100 μm (A-E), 20 μm (F)." figureDoi="10.3897/zse.96.48312.figure25" httpUri="https://binary.pensoft.net/fig/376474" pageId="0" pageNumber="25">25F</figureCitation>
)
</materialsCitation>
. 
<materialsCitation collectionCode="G" country="Turkey" latitude="39.15277" longitude="31.61562">
Thirty specimens from 
<normalizedToken originalValue="Balıkdamı">Balikdami</normalizedToken>
Wetland spring, 
<normalizedToken originalValue="Eskişehir">Eskisehir</normalizedToken>
, 
<collectingCountry name="Turkey" value="Turkey">Turkey</collectingCountry>
; 
<geoCoordinate degrees="39.15277" direction="north" orientation="latitude" precision="1" value="39.15277">39.15277°N</geoCoordinate>
, 
<geoCoordinate degrees="31.61562" direction="east" orientation="longitude" precision="1" value="31.61562">31.61562°E</geoCoordinate>
(Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23G, H</figureCitation>
): 
<quantity metricMagnitude="-1" metricUnit="m" metricValue="3.302" unit="in" value="13.0">13 in</quantity>
<collectionCode collectionName="NMNL" country="Netherlands" httpUri="http://grbio.org/cool/1yj9-ymxi" name="Natuurmuseum Nijmegen e.o." type="Herbarium">NMNL</collectionCode>
(
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342207) and 
<quantity metricMagnitude="-1" metricUnit="m" metricValue="4.318" unit="in" value="17.0">17 in</quantity>
<collectionCode collectionName="UGSB">UGSB</collectionCode>
(
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20686, 
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20739, 
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20740; Fig. 
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Radula of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. paratypes. A. Portion of the radula showing full sets of teeth (UGSB 20688); B. Magnified view of the central teeth (UGSB 20688); C. Magnified view of the lateral and marginal teeth (UGSB 20686); D. Magnified view of the first and second rows of marginal teeth (UGSB 20688); E. Magnified view of the first row of marginal teeth (UGSB 20686); F. Magnified view of the faces of inner marginal teeth belonging to the first row (UGSB 20685). Scale bars: 100 μm (A-E), 20 μm (F)." figureDoi="10.3897/zse.96.48312.figure25" httpUri="https://binary.pensoft.net/fig/376474" pageId="0" pageNumber="25">25C, E</figureCitation>
)
</materialsCitation>
.
</paragraph>
<caption doi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25" start="Figure 23" startId="F23">
<paragraph pageId="0" pageNumber="25">
<emphasis bold="true" pageId="0" pageNumber="25">Figure 23.</emphasis>
<typeStatus>Type</typeStatus>
locality of 
<taxonomicName authorityName="Sands &amp; Gloeer" authorityYear="2020" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus wesselinghi" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus wesselinghi</emphasis>
</taxonomicName>
Sands &amp; 
<normalizedToken originalValue="Glöer">Gloeer</normalizedToken>
sp. nov. 
<emphasis bold="true" pageId="0" pageNumber="25">A</emphasis>
, 
<emphasis bold="true" pageId="0" pageNumber="25">B</emphasis>
. Sakarya River, 
<normalizedToken originalValue="Çayköy">Caykoey</normalizedToken>
, Bilecik, Turkey, 
<geoCoordinate degrees="40.0439" direction="north" orientation="latitude" precision="5" value="40.0439">40.0439°N</geoCoordinate>
, 
<geoCoordinate degrees="30.452" direction="east" orientation="longitude" precision="55" value="30.452">30.452°E</geoCoordinate>
. Further 
<typeStatus>paratype</typeStatus>
localities: 
<emphasis bold="true" pageId="0" pageNumber="25">C</emphasis>
, 
<emphasis bold="true" pageId="0" pageNumber="25">D</emphasis>
. Unnamed roadside spring in Fele, 
<taxonomicName authorityName="Yildirim, Koca, Gurlek &amp; Gloer" authorityYear="2018" class="Gastropoda" family="Hydrobiidae" genus="Isparta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Isparta" order="Littorinimorpha" pageId="0" pageNumber="25" phylum="Mollusca" rank="genus">Isparta</taxonomicName>
province, Turkey, 
<geoCoordinate degrees="38.00358" direction="north" orientation="latitude" precision="1" value="38.00358">38.00358°N</geoCoordinate>
, 
<geoCoordinate degrees="31.47217" direction="east" orientation="longitude" precision="1" value="31.47217">31.47217°E</geoCoordinate>
; 
<emphasis bold="true" pageId="0" pageNumber="25">E</emphasis>
, 
<emphasis bold="true" pageId="0" pageNumber="25">F</emphasis>
. Eflatun 
<normalizedToken originalValue="Pınarı">Pinari</normalizedToken>
, near 
<normalizedToken originalValue="Sadıkhacı">Sadikhaci</normalizedToken>
, Konya, Turkey, 
<geoCoordinate degrees="37.8256" direction="north" orientation="latitude" precision="5" value="37.8256">37.8256°N</geoCoordinate>
, 
<geoCoordinate degrees="31.6748" direction="east" orientation="longitude" precision="5" value="31.6748">31.6748°E</geoCoordinate>
; 
<emphasis bold="true" pageId="0" pageNumber="25">G</emphasis>
, 
<emphasis bold="true" pageId="0" pageNumber="25">H</emphasis>
. 
<normalizedToken originalValue="Balıkdamı">Balikdami</normalizedToken>
Wetland spring, 
<normalizedToken originalValue="Eskişehir">Eskisehir</normalizedToken>
, Turkey, 
<geoCoordinate degrees="39.15277" direction="north" orientation="latitude" precision="1" value="39.15277">39.15277°N</geoCoordinate>
, 
<geoCoordinate degrees="31.61562" direction="east" orientation="longitude" precision="1" value="31.61562">31.61562°E</geoCoordinate>
.
</paragraph>
</caption>
<caption doi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25" start="Figure 24" startId="F24">
<paragraph pageId="0" pageNumber="25">
<emphasis bold="true" pageId="0" pageNumber="25">Figure 24.</emphasis>
<taxonomicName authorityName="Sands &amp; Gloeer" authorityYear="2020" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus wesselinghi" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus wesselinghi</emphasis>
</taxonomicName>
Sands &amp; 
<normalizedToken originalValue="Glöer">Gloeer</normalizedToken>
sp. nov. 
<emphasis bold="true" pageId="0" pageNumber="25">A</emphasis>
- 
<emphasis bold="true" pageId="0" pageNumber="25">D</emphasis>
. 
<typeStatus>Holotype</typeStatus>
collected in the Sakarya River, 
<normalizedToken originalValue="Çayköy">Caykoey</normalizedToken>
, Bilecik, Turkey (
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342200); 
<emphasis bold="true" pageId="0" pageNumber="25">E</emphasis>
- 
<emphasis bold="true" pageId="0" pageNumber="25">G</emphasis>
. 
<typeStatus>Paratype</typeStatus>
from the same location as the 
<typeStatus>holotype</typeStatus>
(
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342202); 
<emphasis bold="true" pageId="0" pageNumber="25">H</emphasis>
- 
<emphasis bold="true" pageId="0" pageNumber="25">J</emphasis>
. 
<typeStatus>Paratype</typeStatus>
from Fele, 
<taxonomicName authorityName="Yildirim, Koca, Gurlek &amp; Gloer" authorityYear="2018" class="Gastropoda" family="Hydrobiidae" genus="Isparta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Isparta" order="Littorinimorpha" pageId="0" pageNumber="25" phylum="Mollusca" rank="genus">Isparta</taxonomicName>
province, Turkey (
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342204); 
<emphasis bold="true" pageId="0" pageNumber="25">K</emphasis>
- 
<emphasis bold="true" pageId="0" pageNumber="25">M</emphasis>
. 
<typeStatus>Paratype</typeStatus>
from Eflatun 
<normalizedToken originalValue="Pınarı">Pinari</normalizedToken>
, Konya province, Turkey (
<collectionCode collectionName="Netherlands, Leiden, Nationaal Natuurhistorische Museum (&quot;Naturalis&quot;) [formerly Rijksmuseum van Natuurlijke Historie]" country="Netherlands" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34992" name="National Museum of Natural History, Naturalis" type="Museum">RMNH</collectionCode>
.
<collectionCode collectionName="MOL" country="Peru" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15530" name="Universidad Nacional Agraria La Molina" type="Herbarium">MOL</collectionCode>
.342206). All photographed material (A-M) is stored in 
<collectionCode collectionName="NMNL" country="Netherlands" httpUri="http://grbio.org/cool/1yj9-ymxi" name="Natuurmuseum Nijmegen e.o." type="Herbarium">NMNL</collectionCode>
. Scale bars: 
<quantity metricMagnitude="-3" metricUnit="m" metricValue="1.0" unit="mm" value="1.0">1 mm</quantity>
.
</paragraph>
</caption>
<caption doi="10.3897/zse.96.48312.figure25" httpUri="https://binary.pensoft.net/fig/376474" pageId="0" pageNumber="25" start="Figure 25" startId="F25">
<paragraph pageId="0" pageNumber="25">
<emphasis bold="true" pageId="0" pageNumber="25">Figure 25.</emphasis>
Radula of 
<taxonomicName authorityName="Sands &amp; Gloeer" authorityYear="2020" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus wesselinghi" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus wesselinghi</emphasis>
</taxonomicName>
Sands &amp; 
<normalizedToken originalValue="Glöer">Gloeer</normalizedToken>
sp. nov. 
<typeStatus>paratypes</typeStatus>
. 
<emphasis bold="true" pageId="0" pageNumber="25">A</emphasis>
. Portion of the radula showing full sets of teeth (
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20688); 
<emphasis bold="true" pageId="0" pageNumber="25">B</emphasis>
. Magnified view of the central teeth (
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20688); 
<emphasis bold="true" pageId="0" pageNumber="25">C</emphasis>
. Magnified view of the lateral and marginal teeth (
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20686); 
<emphasis bold="true" pageId="0" pageNumber="25">D</emphasis>
. Magnified view of the first and second rows of marginal teeth (
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20688); 
<emphasis bold="true" pageId="0" pageNumber="25">E</emphasis>
. Magnified view of the first row of marginal teeth (
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20686); 
<emphasis bold="true" pageId="0" pageNumber="25">F</emphasis>
. Magnified view of the faces of inner marginal teeth belonging to the first row (
<collectionCode collectionName="UGSB">UGSB</collectionCode>
20685). Scale bars: 100 
<normalizedToken originalValue="μm">μm</normalizedToken>
(A-E), 20 
<normalizedToken originalValue="μm">μm</normalizedToken>
(F).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="25" type="etymology">
<paragraph pageId="0" pageNumber="25">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="25">The species is named in honour of the molluscan palaeontologist Frank P. Wesselingh (Naturalis Biodiversity Center, Leiden, The Netherlands) for his contributions to malacology.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="25" type="description">
<paragraph pageId="0" pageNumber="25">Description.</paragraph>
<paragraph pageId="0" pageNumber="25">
Shell (Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24A-C, E, F, H-M</figureCitation>
): Hemispherical, transversely elongate, consisting of typically three whorls that rapidly grow. Spire well defined, moderate height for 
<taxonomicName authorityName="Montfort" authorityYear="1810" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus</emphasis>
</taxonomicName>
; often corroded along with other parts of shell. Shell height ranges from 4.0-
<quantity metricMagnitude="-2" metricUnit="m" metricValue="6.8" unit="mm" value="68.0">6.8 mm</quantity>
, width from 
<quantity metricMagnitude="-2" metricUnit="m" metricValue="5.45" metricValueMax="7.1" metricValueMin="3.8" unit="mm" value="54.5" valueMax="71.0" valueMin="38.0">3.8-7.1 mm</quantity>
. Juveniles appear more globular. Periostracum is uniformly ivory or solid black, intermediate forms with broad brown-black smudged diagonal stripes also exist; surface glossy, finely striated with growth lines. Aperture semicircular, no serrations on inner lip. Columellar plate smooth, flat to slightly concave, inclined towards aperture; colouration blue-grey in darker shelled individuals to white in lighter forms.
</paragraph>
<paragraph pageId="0" pageNumber="25">
Operculum (Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24D, G</figureCitation>
): Operculum plate made of two parts, calcareous base and conchioline lamella; operculum base light, mostly ivory to white, white lamella with distinct orange edge on border with operculum base. Operculum base left adductor can be blunt and rounded, weak callus on top right edge. Apophysis follows same colour scheme as operculum calcareous base, broader at top and narrower and attenuated at bottom. Very narrow rib-shield, deep rib-pouch present on operculum. Operculum lacks a pseudo-apophysis.
</paragraph>
<paragraph pageId="0" pageNumber="25">
Radula (Fig. 
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Radula of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. paratypes. A. Portion of the radula showing full sets of teeth (UGSB 20688); B. Magnified view of the central teeth (UGSB 20688); C. Magnified view of the lateral and marginal teeth (UGSB 20686); D. Magnified view of the first and second rows of marginal teeth (UGSB 20688); E. Magnified view of the first row of marginal teeth (UGSB 20686); F. Magnified view of the faces of inner marginal teeth belonging to the first row (UGSB 20685). Scale bars: 100 μm (A-E), 20 μm (F)." figureDoi="10.3897/zse.96.48312.figure25" httpUri="https://binary.pensoft.net/fig/376474" pageId="0" pageNumber="25">25A-F</figureCitation>
): R-central tooth flanked by A-central, B-central, C-central, E-lateral on each side. Additionally, two interconnected layers of marginal teeth encase central and lateral teeth. R-central shows some variation among populations, slightly spherical or more squared face with slightly concave anterior edge. A-central large and flat with thin ridge that becomes broad and folded right at end of cusp. B-central diminished, forms irregular 
<normalizedToken originalValue="“S”">&quot;S&quot;</normalizedToken>
shape. C-central equally diminished, hidden below lower edge of E-lateral. E-lateral simple with smooth upper edge. First layer of marginal teeth consists of 37-44 teeth that decrease in size away from E-lateral but increase in size and bear serrations on edges of small faces; semidetached from second layer, which is fused and forms outer wall.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="25" type="differentiating features">
<paragraph pageId="0" pageNumber="25">Differentiating features.</paragraph>
<paragraph pageId="0" pageNumber="25">
Based on conchological features of periostracum colouration and patterning and shell shape, it is difficult to differentiate 
<taxonomicName genus="T." lsidName="T. wesselinghi" pageId="0" pageNumber="25" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">T. wesselinghi</emphasis>
</taxonomicName>
sp. nov. from most Asian 
<taxonomicName authorityName="Montfort" authorityYear="1810" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus</emphasis>
</taxonomicName>
spp. given the variety in colour and patterns among the 
<typeStatus>type</typeStatus>
material (Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24A-M</figureCitation>
). However, in some instances it can be distinguished from Anatolian morphotypes of 
<taxonomicName genus="T." lsidName="T. baeticus" pageId="0" pageNumber="25" rank="species" species="baeticus">
<emphasis italics="true" pageId="0" pageNumber="25">T. baeticus</emphasis>
</taxonomicName>
, which typically displays ivory blotches on a brown background (Fig. 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Theodoxus baeticus (Lamarck, 1822). A - D. Specimen (T. callosus - morphoype) collected in Akyaka, Turkey (UGSB 19118); E - H. Specimen (T. callosus - morphotype) collected in Kato Tragana, Greece (UGSB 2488); I - L. Specimen (T. varius - morphotype) collected at the Blue Eye Nature Monument, Albania (UGSB 24170); M - P. Specimen (T. meridionalis - morphotype) collected at the Ketana Oasis, Gabes, Tunisia (UGSB 18111). All specimens, except A-H, were used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure7" httpUri="https://binary.pensoft.net/fig/376456" pageId="0" pageNumber="25">7A-D</figureCitation>
); 
<taxonomicName genus="T." lsidName="T. altenai" pageId="0" pageNumber="25" rank="species" species="altenai">
<emphasis italics="true" pageId="0" pageNumber="25">T. altenai</emphasis>
</taxonomicName>
with clear ivory checks on a dark brown-black background (Fig. 
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Theodoxus altenai Schuett, 1965. A - C. Holotype from Lake Kirkgoez, Doesemealti, Antalya, Turkey (RNL V. 56 / 1, NMNL). D - G. Topotype (UGSB 24167) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure4" httpUri="https://binary.pensoft.net/fig/376453" pageId="0" pageNumber="25">4A-G</figureCitation>
) and 
<taxonomicName genus="T." lsidName="T. gloeri" pageId="0" pageNumber="25" rank="species" species="gloeri">
<emphasis italics="true" pageId="0" pageNumber="25">T. gloeri</emphasis>
</taxonomicName>
lacking shell pigmentation and bearing strong axial ribs on the shell (Fig. 
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Theodoxus gloeri Odabasi &amp; Arslan, 2015. A - C. Paratype from Balikdami Wetland-Sakarya River, Eskisehir, Turkey (coll. Gloeer). Scale bar: 1 mm." figureDoi="10.3897/zse.96.48312.figure11" httpUri="https://binary.pensoft.net/fig/376460" pageId="0" pageNumber="25">11A-C</figureCitation>
). The light ivory-coloured operculum calcareous base makes this species distinct from 
<taxonomicName genus="T." lsidName="T. gurur" pageId="0" pageNumber="25" rank="species" species="gurur">
<emphasis italics="true" pageId="0" pageNumber="25">T. gurur</emphasis>
</taxonomicName>
sp. nov. (light to dark brown; Figs 
<figureCitation captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Theodoxus gurur Sands &amp; Gloeer sp. nov. A - D. Holotype from the Suyunbasi spring complex, Ayrancilar, Izmir, Turkey stored in NMNL (RMNH. MOL. 342197); E - G. Paratype from same locality and storage facility (RMNH. MOL. 342199). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure13" httpUri="https://binary.pensoft.net/fig/376462" pageId="0" pageNumber="25">13D</figureCitation>
, 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24D, G</figureCitation>
) and 
<taxonomicName genus="T." lsidName="T. wilkei" pageId="0" pageNumber="25" rank="species" species="wilkei">
<emphasis italics="true" pageId="0" pageNumber="25">T. wilkei</emphasis>
</taxonomicName>
sp. nov. (bright orange; Figs 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24D, G</figureCitation>
, 
<figureCitation captionStart="Figure 27" captionStartId="F27" captionText="Figure 27. Theodoxus wilkei Sands &amp; Gloeer sp. nov. A - D. Holotype (RMNH. MOL. 342208); E - G. Paratype (RMNH. MOL. 342210); H - J. Paratype (RMNH. MOL. 342211). All photographed material is from the Cifteler Spring, Cifteler, Eskisehir, Turkey and stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure27" httpUri="https://binary.pensoft.net/fig/376476" pageId="0" pageNumber="25">27D</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="25">
More differentiating features occur in the operculum structure (Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24D, G</figureCitation>
). The presence of an attenuated apophysis distinguishes 
<taxonomicName genus="T." lsidName="T. wesselinghi" pageId="0" pageNumber="25" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">T. wesselinghi</emphasis>
</taxonomicName>
sp. nov. from 
<taxonomicName genus="T." lsidName="T. altenai" pageId="0" pageNumber="25" rank="species" species="altenai">
<emphasis italics="true" pageId="0" pageNumber="25">T. altenai</emphasis>
</taxonomicName>
and 
<taxonomicName genus="T." lsidName="T. jordani" pageId="0" pageNumber="25" rank="species" species="jordani">
<emphasis italics="true" pageId="0" pageNumber="25">T. jordani</emphasis>
</taxonomicName>
, which have non-attenuated apophyses (Figs 
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Theodoxus altenai Schuett, 1965. A - C. Holotype from Lake Kirkgoez, Doesemealti, Antalya, Turkey (RNL V. 56 / 1, NMNL). D - G. Topotype (UGSB 24167) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure4" httpUri="https://binary.pensoft.net/fig/376453" pageId="0" pageNumber="25">4</figureCitation>
, 
<figureCitation captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Theodoxus jordani (G. B. Sowerby I, 1836). A - D. Specimen from the Avakas Gorge, Peyia, Cyprus (UGSB 18321) incorporated into the phylogeny (Fig. 2); E - H. Specimen collected in the Ceyhan River, Eski Misis, Adana, Turkey (UGSB 20777); I - L. Specimen from the Orontes River, Antakya, Hatay, Turkey (UGSB 24175) used in the phylogeny (Fig. 2); M - O. Specimen (T. euphraticus - morphotype) from the Seyed Hosein Park spring, near the Bishapur, Fars, Iran (UGSB 22233); P - R. Topotype of N. jordani from the River Jordan, Israel (UGSB 24177) incorporated into the phylogeny (Fig. 2); S, T. Specimen (T. euphraticus - morphotype) from the Karun River, Ahvaz, Iran (UGSB 21681); U. Specimen (T. niloticus - morphotype) collected in the Nile River, Egypt (UGSB 24178) used in the phylogeny (Fig. 2); V. Specimen (T. euphraticus - morphotype) from the Karun River, Ahvaz, Iran (UGSB 21682). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure15" httpUri="https://binary.pensoft.net/fig/376464" pageId="0" pageNumber="25">15</figureCitation>
, 
<figureCitation captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Theodoxus jordani (G. B. Sowerby I, 1836). A - C. Syntype of N. meridionalis var. mesopotamica from near Diyarbakir, Turkey (ZMZ 528914); D, E. Specimen (T. mesopotamicus - morphotype) collected in Sanliurfa, Turkey (UGSB 23426) and used in the phylogeny (Fig. 2); F - H; K - M. Two syntypes of N. cinctella collected at the source of River Chabur near Ras al-Ayn, Syria (ZMB / Moll 21732); I, J. Topotype of N. cinctella from the same locality (UGSB 18762). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure17" httpUri="https://binary.pensoft.net/fig/376466" pageId="0" pageNumber="25">17</figureCitation>
). A rib-pouch and rib-shield are either totally lacking or extremely diminished in 
<taxonomicName genus="T." lsidName="T. altenai" pageId="0" pageNumber="25" rank="species" species="altenai">
<emphasis italics="true" pageId="0" pageNumber="25">T. altenai</emphasis>
</taxonomicName>
, 
<taxonomicName genus="T." lsidName="T. anatolicus" pageId="0" pageNumber="25" rank="species" species="anatolicus">
<emphasis italics="true" pageId="0" pageNumber="25">T. anatolicus</emphasis>
</taxonomicName>
, 
<taxonomicName genus="T." lsidName="T. jordani" pageId="0" pageNumber="25" rank="species" species="jordani">
<emphasis italics="true" pageId="0" pageNumber="25">T. jordani</emphasis>
</taxonomicName>
, and 
<taxonomicName genus="T." lsidName="T. macri" pageId="0" pageNumber="25" rank="species" species="macri">
<emphasis italics="true" pageId="0" pageNumber="25">T. macri</emphasis>
</taxonomicName>
(Figs 
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Theodoxus altenai Schuett, 1965. A - C. Holotype from Lake Kirkgoez, Doesemealti, Antalya, Turkey (RNL V. 56 / 1, NMNL). D - G. Topotype (UGSB 24167) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure4" httpUri="https://binary.pensoft.net/fig/376453" pageId="0" pageNumber="25">4</figureCitation>
, 
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Theodoxus anatolicus (Recluz, 1841). A - D. Specimen collected at Isikli, Denizli, Turkey (UGSB 24168) incorporated into the phylogeny (Fig. 2); E - G. Lectotype of N. anatolica from Izmir, Turkey (MHNG-MOLL- 15028). H, I. Paralectotype of N. anatolica from Izmir, Turkey (MHNG-MOLL- 15028); J, K. Paralectotype of N. anatolica from Chios, Greece (MNHN-IM- 2000 - 32519); L, M. Syntype of N. belladonna from Izmir, Turkey (ZMZ 528908). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure5" httpUri="https://binary.pensoft.net/fig/376454" pageId="0" pageNumber="25">5</figureCitation>
, 
<figureCitation captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Theodoxus jordani (G. B. Sowerby I, 1836). A - D. Specimen from the Avakas Gorge, Peyia, Cyprus (UGSB 18321) incorporated into the phylogeny (Fig. 2); E - H. Specimen collected in the Ceyhan River, Eski Misis, Adana, Turkey (UGSB 20777); I - L. Specimen from the Orontes River, Antakya, Hatay, Turkey (UGSB 24175) used in the phylogeny (Fig. 2); M - O. Specimen (T. euphraticus - morphotype) from the Seyed Hosein Park spring, near the Bishapur, Fars, Iran (UGSB 22233); P - R. Topotype of N. jordani from the River Jordan, Israel (UGSB 24177) incorporated into the phylogeny (Fig. 2); S, T. Specimen (T. euphraticus - morphotype) from the Karun River, Ahvaz, Iran (UGSB 21681); U. Specimen (T. niloticus - morphotype) collected in the Nile River, Egypt (UGSB 24178) used in the phylogeny (Fig. 2); V. Specimen (T. euphraticus - morphotype) from the Karun River, Ahvaz, Iran (UGSB 21682). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure15" httpUri="https://binary.pensoft.net/fig/376464" pageId="0" pageNumber="25">15</figureCitation>
, 
<figureCitation captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Theodoxus jordani (G. B. Sowerby I, 1836). A - C. Syntype of N. meridionalis var. mesopotamica from near Diyarbakir, Turkey (ZMZ 528914); D, E. Specimen (T. mesopotamicus - morphotype) collected in Sanliurfa, Turkey (UGSB 23426) and used in the phylogeny (Fig. 2); F - H; K - M. Two syntypes of N. cinctella collected at the source of River Chabur near Ras al-Ayn, Syria (ZMB / Moll 21732); I, J. Topotype of N. cinctella from the same locality (UGSB 18762). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure17" httpUri="https://binary.pensoft.net/fig/376466" pageId="0" pageNumber="25">17</figureCitation>
, 
<figureCitation captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Theodoxus macri (G. B. Sowerby II, 1849). A - D. Specimen collected close to Harbiye Falls, Harbiye, Antakya, Turkey (UGSB 24179); E - H. Specimen from Lake Yenisehir, Reyhanli, Turkey (UGSB 24180); I, J. Specimen from Lake Balik, Adalar, Turkey (UGSB 24181). All specimens (A-J) were used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure18" httpUri="https://binary.pensoft.net/fig/376467" pageId="0" pageNumber="25">18</figureCitation>
), while they are more pronounced in 
<taxonomicName genus="T." lsidName="T. wesselinghi" pageId="0" pageNumber="25" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">T. wesselinghi</emphasis>
</taxonomicName>
sp. nov. (Fig. 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24</figureCitation>
). The rib-shield in 
<taxonomicName genus="T." lsidName="T. wesselinghi" pageId="0" pageNumber="25" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">T. wesselinghi</emphasis>
</taxonomicName>
sp. nov. is however less broad than that typically observed in 
<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>
and 
<taxonomicName genus="T." lsidName="T. baeticus" pageId="0" pageNumber="25" rank="species" species="baeticus">
<emphasis italics="true" pageId="0" pageNumber="25">T. baeticus</emphasis>
</taxonomicName>
(Figs 
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Theodoxus baeticus (Lamarck, 1822). A - C. Paralectotype of N. baetica from Andalusia, Spain (MHNG-MOLL- 51319). D) Operculum of a paralectotype of N. baetica from Andalusia (MHNG-MOLL- 51319); E - G. Lectotype of N. baetica from Andalusia (MHNG-MOLL- 51319); H-J; K; L; M; N. Five syntypes of N. meridionalis from Sicily (ZMB, without coll. no.); O - R. Topotype of N. guadianensis collected in the Guadiana River near Mertola, Portugal incorporated into the phylogeny (UGSB 22084); S - V. Specimen from Soller, Balearic Islands used in the phylogeny (UGSB 19162). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure6" httpUri="https://binary.pensoft.net/fig/376455" pageId="0" pageNumber="25">6</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>
). Furthermore, the lack of a pseudo-apophysis differentiates the new species from 
<taxonomicName genus="T." lsidName="T. altenai" pageId="0" pageNumber="25" rank="species" species="altenai">
<emphasis italics="true" pageId="0" pageNumber="25">T. altenai</emphasis>
</taxonomicName>
, 
<taxonomicName genus="T." lsidName="T. anatolicus" pageId="0" pageNumber="25" rank="species" species="anatolicus">
<emphasis italics="true" pageId="0" pageNumber="25">T. anatolicus</emphasis>
</taxonomicName>
, 
<taxonomicName genus="T." lsidName="T. baeticus" pageId="0" pageNumber="25" rank="species" species="baeticus">
<emphasis italics="true" pageId="0" pageNumber="25">T. baeticus</emphasis>
</taxonomicName>
, 
<taxonomicName genus="T." lsidName="T. gurur" pageId="0" pageNumber="25" rank="species" species="gurur">
<emphasis italics="true" pageId="0" pageNumber="25">T. gurur</emphasis>
</taxonomicName>
sp. nov., 
<taxonomicName genus="T." lsidName="T. jordani" pageId="0" pageNumber="25" rank="species" species="jordani">
<emphasis italics="true" pageId="0" pageNumber="25">T. jordani</emphasis>
</taxonomicName>
, and 
<taxonomicName genus="T." lsidName="T. macri" pageId="0" pageNumber="25" rank="species" species="macri">
<emphasis italics="true" pageId="0" pageNumber="25">T. macri</emphasis>
</taxonomicName>
(Figs 
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Theodoxus altenai Schuett, 1965. A - C. Holotype from Lake Kirkgoez, Doesemealti, Antalya, Turkey (RNL V. 56 / 1, NMNL). D - G. Topotype (UGSB 24167) incorporated into the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure4" httpUri="https://binary.pensoft.net/fig/376453" pageId="0" pageNumber="25">4</figureCitation>
- 
<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Theodoxus baeticus (Lamarck, 1822). A - D. Specimen from Cuquillo, Granada, Andalusia, Spain (UGSB 22089); E - H. Specimen (T. meridionalis - morphotype) collected in the Palancia River, Navajas, Spain (UGSB 22090); I - L. Specimen (T. elongatulus - morphotype) from Arrancada, Pombal, Portugal (UGSB 22080); M - P. Specimen (T. velascoi - morphotype) from Montanejos, Castellon, Spain (UGSB 22082). Theodoxus cf. valentinus (Graells, 1846). Q - S. Specimen conforming to T. valentinus from the Verd River, Masalaves, Spain (UGSB 21787); T. Operculum of a specimen conforming to T. valentinus from the same locality as Q-S (UGSB 22087). All specimens, except Q-S, were used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure8" httpUri="https://binary.pensoft.net/fig/376457" pageId="0" pageNumber="25">8</figureCitation>
, 
<figureCitation captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Theodoxus gurur Sands &amp; Gloeer sp. nov. A - D. Holotype from the Suyunbasi spring complex, Ayrancilar, Izmir, Turkey stored in NMNL (RMNH. MOL. 342197); E - G. Paratype from same locality and storage facility (RMNH. MOL. 342199). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure13" httpUri="https://binary.pensoft.net/fig/376462" pageId="0" pageNumber="25">13</figureCitation>
, 
<figureCitation captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Theodoxus jordani (G. B. Sowerby I, 1836). A - D. Specimen from the Avakas Gorge, Peyia, Cyprus (UGSB 18321) incorporated into the phylogeny (Fig. 2); E - H. Specimen collected in the Ceyhan River, Eski Misis, Adana, Turkey (UGSB 20777); I - L. Specimen from the Orontes River, Antakya, Hatay, Turkey (UGSB 24175) used in the phylogeny (Fig. 2); M - O. Specimen (T. euphraticus - morphotype) from the Seyed Hosein Park spring, near the Bishapur, Fars, Iran (UGSB 22233); P - R. Topotype of N. jordani from the River Jordan, Israel (UGSB 24177) incorporated into the phylogeny (Fig. 2); S, T. Specimen (T. euphraticus - morphotype) from the Karun River, Ahvaz, Iran (UGSB 21681); U. Specimen (T. niloticus - morphotype) collected in the Nile River, Egypt (UGSB 24178) used in the phylogeny (Fig. 2); V. Specimen (T. euphraticus - morphotype) from the Karun River, Ahvaz, Iran (UGSB 21682). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure15" httpUri="https://binary.pensoft.net/fig/376464" pageId="0" pageNumber="25">15</figureCitation>
, 
<figureCitation captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Theodoxus jordani (G. B. Sowerby I, 1836). A - C. Syntype of N. meridionalis var. mesopotamica from near Diyarbakir, Turkey (ZMZ 528914); D, E. Specimen (T. mesopotamicus - morphotype) collected in Sanliurfa, Turkey (UGSB 23426) and used in the phylogeny (Fig. 2); F - H; K - M. Two syntypes of N. cinctella collected at the source of River Chabur near Ras al-Ayn, Syria (ZMB / Moll 21732); I, J. Topotype of N. cinctella from the same locality (UGSB 18762). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure17" httpUri="https://binary.pensoft.net/fig/376466" pageId="0" pageNumber="25">17</figureCitation>
, 
<figureCitation captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Theodoxus macri (G. B. Sowerby II, 1849). A - D. Specimen collected close to Harbiye Falls, Harbiye, Antakya, Turkey (UGSB 24179); E - H. Specimen from Lake Yenisehir, Reyhanli, Turkey (UGSB 24180); I, J. Specimen from Lake Balik, Adalar, Turkey (UGSB 24181). All specimens (A-J) were used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure18" httpUri="https://binary.pensoft.net/fig/376467" pageId="0" pageNumber="25">18</figureCitation>
). Additionally, the presence of a weak callus on the top right edge of the operculum base in 
<taxonomicName genus="T." lsidName="T. wesselinghi" pageId="0" pageNumber="25" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">T. wesselinghi</emphasis>
</taxonomicName>
sp. nov. helps to differentiate this species from 
<taxonomicName genus="T." lsidName="T. gurur" pageId="0" pageNumber="25" rank="species" species="gurur">
<emphasis italics="true" pageId="0" pageNumber="25">T. gurur</emphasis>
</taxonomicName>
sp. nov. and 
<taxonomicName genus="T." lsidName="T. jordani" pageId="0" pageNumber="25" rank="species" species="jordani">
<emphasis italics="true" pageId="0" pageNumber="25">T. jordani</emphasis>
</taxonomicName>
, which lack a callus (Figs 
<figureCitation captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Theodoxus gurur Sands &amp; Gloeer sp. nov. A - D. Holotype from the Suyunbasi spring complex, Ayrancilar, Izmir, Turkey stored in NMNL (RMNH. MOL. 342197); E - G. Paratype from same locality and storage facility (RMNH. MOL. 342199). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure13" httpUri="https://binary.pensoft.net/fig/376462" pageId="0" pageNumber="25">13</figureCitation>
, 
<figureCitation captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Theodoxus jordani (G. B. Sowerby I, 1836). A - D. Specimen from the Avakas Gorge, Peyia, Cyprus (UGSB 18321) incorporated into the phylogeny (Fig. 2); E - H. Specimen collected in the Ceyhan River, Eski Misis, Adana, Turkey (UGSB 20777); I - L. Specimen from the Orontes River, Antakya, Hatay, Turkey (UGSB 24175) used in the phylogeny (Fig. 2); M - O. Specimen (T. euphraticus - morphotype) from the Seyed Hosein Park spring, near the Bishapur, Fars, Iran (UGSB 22233); P - R. Topotype of N. jordani from the River Jordan, Israel (UGSB 24177) incorporated into the phylogeny (Fig. 2); S, T. Specimen (T. euphraticus - morphotype) from the Karun River, Ahvaz, Iran (UGSB 21681); U. Specimen (T. niloticus - morphotype) collected in the Nile River, Egypt (UGSB 24178) used in the phylogeny (Fig. 2); V. Specimen (T. euphraticus - morphotype) from the Karun River, Ahvaz, Iran (UGSB 21682). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure15" httpUri="https://binary.pensoft.net/fig/376464" pageId="0" pageNumber="25">15</figureCitation>
, 
<figureCitation captionStart="Figure 17" captionStartId="F17" captionText="Figure 17. Theodoxus jordani (G. B. Sowerby I, 1836). A - C. Syntype of N. meridionalis var. mesopotamica from near Diyarbakir, Turkey (ZMZ 528914); D, E. Specimen (T. mesopotamicus - morphotype) collected in Sanliurfa, Turkey (UGSB 23426) and used in the phylogeny (Fig. 2); F - H; K - M. Two syntypes of N. cinctella collected at the source of River Chabur near Ras al-Ayn, Syria (ZMB / Moll 21732); I, J. Topotype of N. cinctella from the same locality (UGSB 18762). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure17" httpUri="https://binary.pensoft.net/fig/376466" pageId="0" pageNumber="25">17</figureCitation>
, 
<figureCitation captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. Theodoxus macri (G. B. Sowerby II, 1849). A - D. Specimen collected close to Harbiye Falls, Harbiye, Antakya, Turkey (UGSB 24179); E - H. Specimen from Lake Yenisehir, Reyhanli, Turkey (UGSB 24180); I, J. Specimen from Lake Balik, Adalar, Turkey (UGSB 24181). All specimens (A-J) were used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure18" httpUri="https://binary.pensoft.net/fig/376467" pageId="0" pageNumber="25">18</figureCitation>
, 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24</figureCitation>
), as well as from 
<taxonomicName genus="T." lsidName="T. anatolicus" pageId="0" pageNumber="25" rank="species" species="anatolicus">
<emphasis italics="true" pageId="0" pageNumber="25">T. anatolicus</emphasis>
</taxonomicName>
, 
<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>
, 
<taxonomicName genus="T." lsidName="T. major" pageId="0" pageNumber="25" rank="species" species="major">
<emphasis italics="true" pageId="0" pageNumber="25">T. major</emphasis>
</taxonomicName>
, 
<taxonomicName genus="T." lsidName="T. pallidus" pageId="0" pageNumber="25" rank="species" species="pallidus">
<emphasis italics="true" pageId="0" pageNumber="25">T. pallidus</emphasis>
</taxonomicName>
, and 
<taxonomicName genus="T." lsidName="T. wilkei" pageId="0" pageNumber="25" rank="species" species="wilkei">
<emphasis italics="true" pageId="0" pageNumber="25">T. wilkei</emphasis>
</taxonomicName>
sp. nov., which have stronger calluses (Figs 
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Theodoxus anatolicus (Recluz, 1841). A - D. Specimen collected at Isikli, Denizli, Turkey (UGSB 24168) incorporated into the phylogeny (Fig. 2); E - G. Lectotype of N. anatolica from Izmir, Turkey (MHNG-MOLL- 15028). H, I. Paralectotype of N. anatolica from Izmir, Turkey (MHNG-MOLL- 15028); J, K. Paralectotype of N. anatolica from Chios, Greece (MNHN-IM- 2000 - 32519); L, M. Syntype of N. belladonna from Izmir, Turkey (ZMZ 528908). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure5" httpUri="https://binary.pensoft.net/fig/376454" pageId="0" pageNumber="25">5</figureCitation>
, 
<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 19" captionStartId="F19" captionText="Figure 19. Theodoxus major Issel, 1865. A, B. Specimen (T. pallasi - morphotype) collected in the Caspian Sea, offshore from Aktau, Kazakhstan (UGSB 20712); C, D. Specimen (T. pallasi - morphotype) collected in the Aral Sea, Kazakhstan in 1979 (UGSB 19129); E - H. Specimen (T. pallasi - morphotype) from the Shahpol River, Aliabad Askar Khan, Iran (UGSB 18091); I - K. Specimen of T. major collected at the outflow of Lake Akna, Aknalich, Armenia (UGSB 20482); L - N. Specimen (T. astrachanicus - morphotype) collected in Utlyukskij Liman, Ukraine (UGSB 18130); O, P. Specimen of T. major collected in the Hrazdan River at the inflow of the Yerevan Reservoir, Armenia (UGSB 20496); Q - T. Specimen of T. major from Zoeram, North Khorasan, Iran (UGSB 21661); U, V. Specimen of T. schultzii collected in the Caspian Sea, offshore from Aktau, Kazakhstan (UGSB 20714); W - Y. Lectotype of T. pallasi from Caspian Sea, Dagestan, Russia (ZIN 54547 / 63); Za, Zb. Syntype of N. schultzii from the Caspian Sea (ZIN, no. 5 in systematic catalogue). Specimens E-K and O-T used in the phylogeny (Fig. 2). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure19" httpUri="https://binary.pensoft.net/fig/376468" pageId="0" pageNumber="25">19</figureCitation>
, 
<figureCitation captionStart="Figure 20" captionStartId="F20" captionText="Figure 20. Theodoxus pallidus (Dunker, 1861). A - D. Specimen from Haji Abad, Fars, Iran (UGSB 22227); E - H. Syntype of N. pallida from southern Iran (ZMB 108.792); I, J; K, L. Two syntypes of N. pallida from southern Iran (ZMB 108.793); M - P. Specimen (T. doriae - morphotype) from a spring near Harat, Yazd, Iran (UGSB 21706); Q, R; S, T; U, V; W - Y. Four syntypes of N. doriae from the thermal springs of Kerman, Iran (MSNG, without coll. no.). Specimens A-D and M-P were used in the phylogeny. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure20" httpUri="https://binary.pensoft.net/fig/376469" pageId="0" pageNumber="25">20</figureCitation>
, 
<figureCitation captionStart="Figure 24" captionStartId="F24" captionText="Figure 24. Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A - D. Holotype collected in the Sakarya River, Caykoey, Bilecik, Turkey (RMNH. MOL. 342200); E - G. Paratype from the same location as the holotype (RMNH. MOL. 342202); H - J. Paratype from Fele, Isparta province, Turkey (RMNH. MOL. 342204); K - M. Paratype from Eflatun Pinari, Konya province, Turkey (RMNH. MOL. 342206). All photographed material (A-M) is stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure24" httpUri="https://binary.pensoft.net/fig/376473" pageId="0" pageNumber="25">24</figureCitation>
, 
<figureCitation captionStart="Figure 27" captionStartId="F27" captionText="Figure 27. Theodoxus wilkei Sands &amp; Gloeer sp. nov. A - D. Holotype (RMNH. MOL. 342208); E - G. Paratype (RMNH. MOL. 342210); H - J. Paratype (RMNH. MOL. 342211). All photographed material is from the Cifteler Spring, Cifteler, Eskisehir, Turkey and stored in NMNL. Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure27" httpUri="https://binary.pensoft.net/fig/376476" pageId="0" pageNumber="25">27</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="25">
Based on the available data for 
<taxonomicName authorityName="Montfort" authorityYear="1810" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus</emphasis>
</taxonomicName>
ralulae, 
<taxonomicName genus="T." lsidName="T. wesselinghi" pageId="0" pageNumber="25" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">T. wesselinghi</emphasis>
</taxonomicName>
sp. nov. can be distinguished by a more globular R-central face from 
<taxonomicName genus="T." lsidName="T. gurur" pageId="0" pageNumber="25" rank="species" species="gurur">
<emphasis italics="true" pageId="0" pageNumber="25">T. gurur</emphasis>
</taxonomicName>
sp. nov., 
<taxonomicName genus="T." lsidName="T. wilkei" pageId="0" pageNumber="25" rank="species" species="wilkei">
<emphasis italics="true" pageId="0" pageNumber="25">T. wilkei</emphasis>
</taxonomicName>
sp. nov., 
<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>
, and 
<taxonomicName genus="T." lsidName="T. jordani" pageId="0" pageNumber="25" rank="species" species="jordani">
<emphasis italics="true" pageId="0" pageNumber="25">T. jordani</emphasis>
</taxonomicName>
, where it is more rectangular or triangulate (see 
<bibRefCitation DOI="https://doi.org/10.2307/4063879" author="Baker, HB" journalOrPublisher="Proceedings of the Academy of Natural Sciences of Philadelphia" pageId="0" pageNumber="25" pagination="117 - 178" refId="B18" refString="Baker, HB, 1923. Notes on the radula of the Neritidae. Proceedings of the Academy of Natural Sciences of Philadelphia 75: 117 - 178, DOI: https://doi.org/10.2307/4063879" title="Notes on the radula of the Neritidae." url="https://doi.org/10.2307/4063879" volume="75" year="1923">Baker 1923</bibRefCitation>
; 
<bibRefCitation author="Zettler, ML" journalOrPublisher="Mollusca" pageId="0" pageNumber="25" pagination="13 - 72" refId="B166" refString="Zettler, ML, 2008. Zur Taxonomie und Verbreitung der Gattung Theodoxus Montfort, 1810 in Deutschland. Darstellung historischer und rezenter Daten einschliesslich einer Bibliografie. Mollusca 26 (1): 13 - 72" title="Zur Taxonomie und Verbreitung der Gattung Theodoxus Montfort, 1810 in Deutschland. Darstellung historischer und rezenter Daten einschliesslich einer Bibliografie." volume="26" year="2008">Zettler 2008</bibRefCitation>
; Figs 
<figureCitation captionStart="Figure 14" captionStartId="F14" captionText="Figure 14. Radula of Theodoxus gurur Sands &amp; Gloeer sp. nov. paratypes (UGSB 20689). A. Portion of the radula showing full sets of teeth; B. Magnified view of the central and lateral teeth; C. Magnified view of the lateral and marginal teeth; D. Magnified view of the first row of marginal teeth; E. Merging point between the first and second rows of marginal teeth; F. Magnified view of the faces of inner marginal teeth belonging to the first row. Scale bars: 100 μm (A-D), 20 μm (E-F)." figureDoi="10.3897/zse.96.48312.figure14" httpUri="https://binary.pensoft.net/fig/376463" pageId="0" pageNumber="25">14B</figureCitation>
, 
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Radula of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. paratypes. A. Portion of the radula showing full sets of teeth (UGSB 20688); B. Magnified view of the central teeth (UGSB 20688); C. Magnified view of the lateral and marginal teeth (UGSB 20686); D. Magnified view of the first and second rows of marginal teeth (UGSB 20688); E. Magnified view of the first row of marginal teeth (UGSB 20686); F. Magnified view of the faces of inner marginal teeth belonging to the first row (UGSB 20685). Scale bars: 100 μm (A-E), 20 μm (F)." figureDoi="10.3897/zse.96.48312.figure25" httpUri="https://binary.pensoft.net/fig/376474" pageId="0" pageNumber="25">25B</figureCitation>
, 
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Radula of Theodoxus wilkei Sands &amp; Gloeer sp. nov. paratypes (UGSB 20687). A. Portion of the radula showing full sets of teeth; B. Magnified view of the central and lateral teeth; C. Magnified view of the lateral and first and second rows of marginal teeth; D. Magnified view of the lateral teeth; E. The first row of marginal teeth; F. Magnified view of the faces of inner marginal teeth belonging to the first row. Scale bars: 100 μm (A-D), 20 μm (E-F)." figureDoi="10.3897/zse.96.48312.figure28" httpUri="https://binary.pensoft.net/fig/376477" pageId="0" pageNumber="25">28B</figureCitation>
). Furthermore, the smooth upper edge of the E-lateral can be used to distinguish this species from 
<taxonomicName genus="T." lsidName="T. wilkei" pageId="0" pageNumber="25" rank="species" species="wilkei">
<emphasis italics="true" pageId="0" pageNumber="25">T. wilkei</emphasis>
</taxonomicName>
sp. nov. and 
<taxonomicName genus="T." lsidName="T. major" pageId="0" pageNumber="25" rank="species" species="major">
<emphasis italics="true" pageId="0" pageNumber="25">T. major</emphasis>
</taxonomicName>
, which generally have E-laterals with serrated edges (see 
<bibRefCitation DOI="https://doi.org/10.1127/arch.moll/146/213-226" author="Anistratenko, VV" journalOrPublisher="Archiv fuer Molluskenkunde" pageId="0" pageNumber="25" pagination="213 - 226" refId="B17" refString="Anistratenko, VV, Zettler, ML, Anistratenko, OY, 2017. On the taxonomic relationship between Theodoxus pallasi and T. astrachanicus (Gastropoda: Neritidae) from the Ponto-Caspian region. Archiv fuer Molluskenkunde 146 (2): 213 - 226, DOI: https://doi.org/10.1127/arch.moll/146/213-226" title="On the taxonomic relationship between Theodoxus pallasi and T. astrachanicus (Gastropoda: Neritidae) from the Ponto-Caspian region." url="https://doi.org/10.1127/arch.moll/146/213-226" volume="146" year="2017">Anistratenko et al. 2017</bibRefCitation>
; Figs 
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Radula of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. paratypes. A. Portion of the radula showing full sets of teeth (UGSB 20688); B. Magnified view of the central teeth (UGSB 20688); C. Magnified view of the lateral and marginal teeth (UGSB 20686); D. Magnified view of the first and second rows of marginal teeth (UGSB 20688); E. Magnified view of the first row of marginal teeth (UGSB 20686); F. Magnified view of the faces of inner marginal teeth belonging to the first row (UGSB 20685). Scale bars: 100 μm (A-E), 20 μm (F)." figureDoi="10.3897/zse.96.48312.figure25" httpUri="https://binary.pensoft.net/fig/376474" pageId="0" pageNumber="25">25C</figureCitation>
, 
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Radula of Theodoxus wilkei Sands &amp; Gloeer sp. nov. paratypes (UGSB 20687). A. Portion of the radula showing full sets of teeth; B. Magnified view of the central and lateral teeth; C. Magnified view of the lateral and first and second rows of marginal teeth; D. Magnified view of the lateral teeth; E. The first row of marginal teeth; F. Magnified view of the faces of inner marginal teeth belonging to the first row. Scale bars: 100 μm (A-D), 20 μm (E-F)." figureDoi="10.3897/zse.96.48312.figure28" httpUri="https://binary.pensoft.net/fig/376477" pageId="0" pageNumber="25">28C, D</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="25" type="remarks">
<paragraph pageId="0" pageNumber="25">Remarks.</paragraph>
<paragraph pageId="0" pageNumber="25">
<taxonomicName authorityName="Sands &amp; Gloeer" authorityYear="2020" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus wesselinghi" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus wesselinghi</emphasis>
</taxonomicName>
sp. nov. forms part of a larger clade that includes 
<taxonomicName genus="T." lsidName="T. syriacus" pageId="0" pageNumber="25" rank="species" species="syriacus">
<emphasis italics="true" pageId="0" pageNumber="25">T. syriacus</emphasis>
</taxonomicName>
and 
<taxonomicName genus="T." lsidName="T. wilkei" pageId="0" pageNumber="25" rank="species" species="wilkei">
<emphasis italics="true" pageId="0" pageNumber="25">T. wilkei</emphasis>
</taxonomicName>
sp. nov., where it shares a closer sister-species relationship with 
<taxonomicName genus="T." lsidName="T. wilkei" pageId="0" pageNumber="25" rank="species" species="wilkei">
<emphasis italics="true" pageId="0" pageNumber="25">T. wilkei</emphasis>
</taxonomicName>
sp. nov. (
<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>
; 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 &quot; Systematic checklist &quot; for more information). Small green (PP ≥ 0.95) and red (PP &lt;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>
). The three species likely diverged from one another in quick succession over the Pliocene-Pleistocene transition (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 &quot; Systematic checklist &quot; for more information). Small green (PP ≥ 0.95) and red (PP &lt;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>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="25" type="distribution">
<paragraph pageId="0" pageNumber="25">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="25">
Known so far only from the four localities in central-west Anatolia (Figs 
<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">3D</figureCitation>
, 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23A-H</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="25" type="ecology">
<paragraph pageId="0" pageNumber="25">Ecology.</paragraph>
<paragraph pageId="0" pageNumber="25">
<taxonomicName authorityName="Sands &amp; Gloeer" authorityYear="2020" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus wesselinghi" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus wesselinghi</emphasis>
</taxonomicName>
sp. nov. can be found in both springs (
<normalizedToken originalValue="Balıkdamı">Balikdami</normalizedToken>
Wetland, Eflatun 
<normalizedToken originalValue="Pınarı">Pinari</normalizedToken>
, Fele; Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23C-H</figureCitation>
) and streams (Sakarya River, 
<normalizedToken originalValue="Çayköy">Caykoey</normalizedToken>
; Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23A, B</figureCitation>
) with clear water. The occurrence of macrophytes appears negligible to the species as it occurs in localities both with (Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23E-H</figureCitation>
) and without (Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23A-D</figureCitation>
) aquatic plant growth. The floors of all localities are made up of coarse grained sand and large and small rocks and stones that 
<taxonomicName genus="T." lsidName="T. wesselinghi" pageId="0" pageNumber="25" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">T. wesselinghi</emphasis>
</taxonomicName>
sp. nov. is often attached to (personal observation M.E.G.; Fig. 
<figureCitation captionStart="Figure 23" captionStartId="F23" captionText="Figure 23. Type locality of Theodoxus wesselinghi Sands &amp; Gloeer sp. nov. A, B. Sakarya River, Caykoey, Bilecik, Turkey, 40.0439 ° N, 30.452 ° E. Further paratype localities: C, D. Unnamed roadside spring in Fele, Isparta province, Turkey, 38.00358 ° N, 31.47217 ° E; E, F. Eflatun Pinari, near Sadikhaci, Konya, Turkey, 37.8256 ° N, 31.6748 ° E; G, H. Balikdami Wetland spring, Eskisehir, Turkey, 39.15277 ° N, 31.61562 ° E." figureDoi="10.3897/zse.96.48312.figure23" httpUri="https://binary.pensoft.net/fig/376472" pageId="0" pageNumber="25">23H</figureCitation>
). 
<taxonomicName authorityName="Sands &amp; Gloeer" authorityYear="2020" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus wesselinghi" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="wesselinghi">
<emphasis italics="true" pageId="0" pageNumber="25">Theodoxus wesselinghi</emphasis>
</taxonomicName>
sp. nov. co-occurs with 
<taxonomicName baseAuthorityName="Gray" baseAuthorityYear="1843" class="Gastropoda" family="Tateidae" genus="Potamopyrgus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Potamopyrgus antipodarum" order="Littorinimorpha" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="antipodarum">
<emphasis italics="true" pageId="0" pageNumber="25">Potamopyrgus antipodarum</emphasis>
</taxonomicName>
(Gray, 1843) and 
<taxonomicName class="Gastropoda" family="Hydrobiidae" genus="Isparta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Isparta felei" order="Littorinimorpha" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="felei">
<emphasis italics="true" pageId="0" pageNumber="25">Isparta felei</emphasis>
</taxonomicName>
<normalizedToken originalValue="Yıldırım">Yildirim</normalizedToken>
, Koca, 
<normalizedToken originalValue="Gürlek">Guerlek</normalizedToken>
&amp; 
<normalizedToken originalValue="Glöer">Gloeer</normalizedToken>
, 
<quantity metricMagnitude="1" metricUnit="m" metricValue="5.125719999999999" unit="in" value="2018.0">2018 in</quantity>
Fele spring, 
<taxonomicName authorityName="Radoman" authorityYear="1973" class="Gastropoda" family="Hydrobiidae" genus="Falsipyrgula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Falsipyrgula" order="Littorinimorpha" pageId="0" pageNumber="25" phylum="Mollusca" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="25">Falsipyrgula</emphasis>
</taxonomicName>
sp. in Eflatun 
<normalizedToken originalValue="Pınarı">Pinari</normalizedToken>
, and 
<taxonomicName class="Gastropoda" family="Hydrobiidae" genus="Pseudamnicola" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudamnicola natolica" order="Littorinimorpha" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="natolica">
<emphasis italics="true" pageId="0" pageNumber="25">Pseudamnicola natolica</emphasis>
</taxonomicName>
(
<normalizedToken originalValue="Küster">Kuester</normalizedToken>
, 1852) (and possibly also 
<taxonomicName genus="T." lsidName="T. gloeri" pageId="0" pageNumber="25" rank="species" species="gloeri">
<emphasis italics="true" pageId="0" pageNumber="25">T. gloeri</emphasis>
</taxonomicName>
; 
<bibRefCitation author="Odabasi, DA" journalOrPublisher="Ecologica Montenegrina" pageId="0" pageNumber="25" pagination="327 - 333" refId="B115" refString="Odabasi, DA, Arslan, N, 2015. Description of a new subterranean nerite: Theodoxus gloeri n. sp. with some data on the freshwater gastropod fauna of Balikdami Wetland (Sakarya River, Turkey). Ecologica Montenegrina 2 (4): 327 - 333" title="Description of a new subterranean nerite: Theodoxus gloeri n. sp. with some data on the freshwater gastropod fauna of Balikdami Wetland (Sakarya River, Turkey)." volume="2" year="2015">
<normalizedToken originalValue="Odabaşı">Odabasi</normalizedToken>
and Arslan 2015
</bibRefCitation>
) in the 
<normalizedToken originalValue="Balıkdamı">Balikdami</normalizedToken>
Wetland spring (personal observation M.E.G.).
</paragraph>
</subSubSection>
</treatment>
</document>