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<mods:title id="9B909E7837A8DC8BDF6A6D43CC636675">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 id="DE18AA18980DB46F392767422EE55050">Sands, Arthur F</mods:namePart>
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<mods:namePart id="F57EB9D04E2FCBB4395D7D5B2674C094">Gloeer, Peter</mods:namePart>
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<treatment id="CFE8F3CDA150576C94E3DB2C5E9C9034" ID-GBIF-Taxon="161939042" LSID="urn:lsid:plazi:treatment:CFE8F3CDA150576C94E3DB2C5E9C9034" httpUri="http://treatment.plazi.org/id/CFE8F3CDA150576C94E3DB2C5E9C9034" lastPageNumber="25" pageId="0" pageNumber="25" scope_class="Gastropoda" scope_family="Neritidae" scope_subFamily="Neritininae" scope_superFamily="Neritoidea">
<subSubSection id="70DCBCDB9577C147C58A15DEE03B18AE" pageId="0" pageNumber="25" type="nomenclature">
<paragraph id="215F636B52D39A4C9ACD64BB58F991B4" pageId="0" pageNumber="25">
<taxonomicName id="B77135F6C866EF1043F4702A39DE702D" ID-CoL="7C7NM" LSID="CFE8F3CD-A150-576C-94E3-DB2C5E9C9034" authority="V. Anistratenko in O. Anistratenko et al., 1999" authorityName="V. Anistratenko in O. Anistratenko et al." authorityYear="1999" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999</taxonomicName>
<figureCitation id="76361D216B23B05B71807ABFE297A594" captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">Figure 22A-M</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="48548D7B069A6F68CCF5E4D7D505B634" pageId="0" pageNumber="25" type="reference_group">
<paragraph id="6E6A4242333553CC446760345D82AD99" pageId="0" pageNumber="25">
<taxonomicName id="2E7C57144865FB6D8C23C59396F829B0" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">Theodoxus velox</taxonomicName>
V. Anistratenko in O.
<bibRefCitation id="8A8DA33A6F2C6E50F0C61514B2DEAA70" author="Anistratenko, OY" journalOrPublisher="Vestnik Zoologii" pageId="0" pageNumber="25" pagination="11 - 19" refId="B14" refString="Anistratenko, OY, Starobogatov, YI, Anistratenko, VV, 1999. Mollyuski roda Theodoxus (Gastropoda, Pectinibranchia, Neritidae) Azovo-Chernomorskogo basseyna. Vestnik Zoologii 33 (3): 11 - 19" title="Mollyuski roda Theodoxus (Gastropoda, Pectinibranchia, Neritidae) Azovo-Chernomorskogo basseyna." volume="33" year="1999">Anistratenko et al. 1999</bibRefCitation>
: 17-18, fig. 4.7;
<bibRefCitation id="F3D8CE6DD18EE327868DD847F28315DC" DOI="https://doi.org/10.3897/zookeys.827.31365" author="Wesselingh, FP" journalOrPublisher="ZooKeys" pageId="0" pageNumber="25" pagination="31 - 124" refId="B159" refString="Wesselingh, FP, Neubauer, TA, Anistratenko, VV, Vinarski, MV, Yanina, T, ter Poorten, JJ, Kijashko, P, Albrecht, C, Anistratenko, OY, D'Hont, A, Frolov, P, Gandara, AM, Gittenberger, A, Gogaladze, A, Karpinsky, M, Lattuada, M, Popa, L, Sands, AF, van de Velde, S, Vandendorpe, J, Wilke, T, 2019. Mollusc species from the Pontocaspian region - an expert opinion list. ZooKeys 827: 31 - 124, DOI: https://doi.org/10.3897/zookeys.827.31365" title="Mollusc species from the Pontocaspian region - an expert opinion list." url="https://doi.org/10.3897/zookeys.827.31365" volume="827" year="2019">Wesselingh et al. 2019</bibRefCitation>
: 66.
</paragraph>
</subSubSection>
<subSubSection id="D184483D169AD66B8C60119AA01988F2" pageId="0" pageNumber="25" type="type locality">
<paragraph id="2D3CC08874F919E51670279E848C595A" pageId="0" pageNumber="25">
<typeStatus id="7E778344D730B50E89E78CD4D211C882">Type</typeStatus>
locality.
</paragraph>
<paragraph id="FEEC2E27C731596F764433878ABBA7D4" pageId="0" pageNumber="25">Dnieper Delta, Zburievkiy Liman, Kherson Region, Ukraine.</paragraph>
</subSubSection>
<subSubSection id="9F1331DE4677F1821107F24C28B06A0E" pageId="0" pageNumber="25" type="type material">
<paragraph id="93B33783DB661800DCA2EADB88E7296C" pageId="0" pageNumber="25">
<typeStatus id="6A5FA261C28BC51CFB314BFBB5283CE6">Type</typeStatus>
material.
</paragraph>
<paragraph id="4260B3EF416381DB033F46967A7470E4" pageId="0" pageNumber="25">
<materialsCitation id="839505B508357DE9C3C83A26E8F7D08B" collectionCode="IZAN" country="Turkey" typeStatus="Holotype">
<typeStatus id="6C57036C05B364CF659A2EDB1DE444CF">Holotype</typeStatus>
and five
<typeStatus id="42BEB79FEAE1F262DC5F54376091FF9D">paratypes</typeStatus>
are stored in
<collectionCode id="5E5035730F48805D91674ABE9DB3EF14" collectionName="IZAN">IZAN</collectionCode>
(without coll. no.)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="2E1265794FB1A96FCD0E94736930317A" pageId="0" pageNumber="25" type="remarks">
<paragraph id="CBB1AC0E5976BAA458449278E1551A07" pageId="0" pageNumber="25">Remarks.</paragraph>
<paragraph id="A48EE526BC2EF342D7CA24746C22FC51" pageId="0" pageNumber="25">
This species was recently discussed by
<bibRefCitation id="723E372F6259F6F8090DD225E56CDEAB" DOI="https://doi.org/10.3897/zookeys.827.31365" author="Wesselingh, FP" journalOrPublisher="ZooKeys" pageId="0" pageNumber="25" pagination="31 - 124" refId="B159" refString="Wesselingh, FP, Neubauer, TA, Anistratenko, VV, Vinarski, MV, Yanina, T, ter Poorten, JJ, Kijashko, P, Albrecht, C, Anistratenko, OY, D'Hont, A, Frolov, P, Gandara, AM, Gittenberger, A, Gogaladze, A, Karpinsky, M, Lattuada, M, Popa, L, Sands, AF, van de Velde, S, Vandendorpe, J, Wilke, T, 2019. Mollusc species from the Pontocaspian region - an expert opinion list. ZooKeys 827: 31 - 124, DOI: https://doi.org/10.3897/zookeys.827.31365" title="Mollusc species from the Pontocaspian region - an expert opinion list." url="https://doi.org/10.3897/zookeys.827.31365" volume="827" year="2019">Wesselingh et al. (2019)</bibRefCitation>
. While it was previously considered a junior synonym of
<taxonomicName id="FE0FE2713FD9D6BD7F26AA8532F69DF2" lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
<emphasis id="1B7A5F25365B300E51D2822CC440468D" italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
</taxonomicName>
(
<bibRefCitation id="0389E2B08860B29CAADA6D24AC3B176B" author="Vinarski, MV" journalOrPublisher="Severtsov Institute of Ecology and Evolution of RAS, Moscow" pageId="0" pageNumber="25" refId="B157" refString="Vinarski, MV, Kantor, YI, 2016. Analytical catalogue of fresh and brackish water molluscs of Russia and adjacent countries. A.N. Severtsov Institute of Ecology and Evolution of RAS, Moscow" title="Analytical catalogue of fresh and brackish water molluscs of Russia and adjacent countries. A. N." year="2016">Vinarski and Kantor 2016</bibRefCitation>
), molecular data shows strong monophyletic support for the independence of
<taxonomicName id="AB15F5A3B4146F1E407A9BF607EF6A3F" lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
<emphasis id="87990A993E6B815C54137EFC3562ED64" italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
</taxonomicName>
(
<bibRefCitation id="6BF66C7B9EF3EDCD1748C41B8D5A7F4F" 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 id="E1B8C4BED72E75BC19A97BF38D17B5C2" 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>
). Nevertheless,
<taxonomicName id="0860EC812558DA4A5836C211C8FF5941" lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
<emphasis id="AA86B14CD34A5F203D268B3A2F8849A3" italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
</taxonomicName>
and
<taxonomicName id="5FF7BDFBDE8E58D4A4B2BB6AEB5B526D" lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
<emphasis id="3B009F7EF3CE3DD1F6B1F6D79313CD4D" italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
</taxonomicName>
still hold a sister relationship and likely diverged from one another during the early Pleistocene (
<bibRefCitation id="7DDC39D824AA44B38B86962C6350A892" 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 id="FEB706CFEB28055996E6051A358CD63F" 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>
).
<taxonomicName id="0BEA363A4D2BF8292F01C1D95D68F95D" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">
<emphasis id="B9476BFDD89CFA5C4541D8D9F85C6BA6" italics="true" pageId="0" pageNumber="25">Theodoxus velox</emphasis>
</taxonomicName>
is challenging to distinguish from
<taxonomicName id="8B7FFE79984C3800ACA1AA9C9967139C" lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
<emphasis id="A8289C4C8F58E9FCFDE7F16BA7E2B7C1" italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
</taxonomicName>
given the overlap in geographic range and similarity of conchological features (Figs
<figureCitation id="1199201EAA8E263454C72DA453E36CF5" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Occurrence map for Theodoxus spp. in Asia. All points conform to specimens used in the phylogenetic analyses (Suppl. material 1: Table S 1), except for T. gloeri for which no molecular data are available and only the type and paratype localities are indicated. Species are partitioned across four identical maps (A - D) to limit overlap of points as far as possible. Barring T. gloeri, closer related species are grouped together (Fig. 2)." figureDoi="10.3897/zse.96.48312.figure3" httpUri="https://binary.pensoft.net/fig/376452" pageId="0" pageNumber="25">3</figureCitation>
,
<figureCitation id="A2CE6B6B4901F2D98778679967F62ED0" 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 id="C7F331ED519F566B958C48E11653C304" 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 id="97AAD19478FDCEF481E56AA278198113" captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22</figureCitation>
).
<taxonomicName id="9631D8FDA3B90C01B44BCBE6384AB57A" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">
<emphasis id="2BCD35D4AD9F341531DB9744DB0A15FD" italics="true" pageId="0" pageNumber="25">Theodoxus velox</emphasis>
</taxonomicName>
has less phenotypic variability compared to
<taxonomicName id="22D7E0DE1CF5EE1526F3CB3C3C8935D6" lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
<emphasis id="3F94AA3A537AAAE537321E27B65948CA" italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
</taxonomicName>
(Figs
<figureCitation id="958EC2D0CC1830730F419D334CD8CE80" 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 id="202F840E050EF89BD8E8721BCFA6CF17" 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 id="1F6A6FB63D140A8C460A20FD6CDFC693" captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22</figureCitation>
). Moreover,
<taxonomicName id="E4571517E304AA764C359C3FC02C506C" lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
<emphasis id="71DB5B6A8AE2F292D39CE40BCFE3631C" italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
</taxonomicName>
display more expansive spiral whorls and, in some instances, a more transparent operculum where the conchioline lamella extends deeper into the calcareous base and the callus is less pronounced (Figs
<figureCitation id="3FA24EB69FBC28549FCCDBBEC6E7D4AB" 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 id="BFBFCAD48ED924E358454EFDB24009FF" 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 id="FDA8F91D73F2B24D7B38406BECA9B04C" captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22</figureCitation>
).
</paragraph>
<caption id="7456872F7AF7369DB26434F50A6B1085" doi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25" start="Figure 22" startId="F22">
<paragraph id="5D03B5CFDC0336C8FF0622B3389DB92F" pageId="0" pageNumber="25">
<emphasis id="88FAD36369C5ED49F70550E583E4FFCD" bold="true" pageId="0" pageNumber="25">Figure 22.</emphasis>
<taxonomicName id="8A160A731195D1A52D4C5E2F9CC67ACD" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus velox" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="velox">
<emphasis id="8AF5FD5F6E6C698263D4CC4CEB7BE886" italics="true" pageId="0" pageNumber="25">Theodoxus velox</emphasis>
</taxonomicName>
V. Anistratenko in O. Anistratenko et al., 1999.
<emphasis id="0D2FAFCBD5A78773E063C3EBDDC677A7" bold="true" pageId="0" pageNumber="25">A</emphasis>
-
<emphasis id="6747A0F101BB20946164274B62EE0CA8" bold="true" pageId="0" pageNumber="25">D</emphasis>
. Specimen from the Dnieper River, Kiev, Ukraine (
<collectionCode id="3D6FDBD46B98C5B0229BA6CD391B9E22" collectionName="UGSB">UGSB</collectionCode>
24186);
<emphasis id="8C47045029DFAAD971DD855487E2F536" bold="true" pageId="0" pageNumber="25">E</emphasis>
-
<emphasis id="94D318F3798CFD55B895496CCA658CCE" bold="true" pageId="0" pageNumber="25">H</emphasis>
. Specimen collected in the Danube delta, Vylkove, Ukraine (
<collectionCode id="037495AA55B43CF7DD8E7D4FD250761F" collectionName="UGSB">UGSB</collectionCode>
24184) used in the phylogeny (Fig.
<figureCitation id="D7BF4BFF10863F401C1A71324DCD471D" 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>
);
<emphasis id="8BAD10850ADFDF9DD69DEA5C28398A74" bold="true" pageId="0" pageNumber="25">I</emphasis>
,
<emphasis id="BE32541941C3F7F26E4CEFDD735489B7" bold="true" pageId="0" pageNumber="25">J</emphasis>
.
<typeStatus id="DECF9654427699BF638EC0847BF0041F">Holotype</typeStatus>
from Dnieper Delta, Zbur'ivka Liman, Ukraine (
<collectionCode id="8E1130896DABB4D0BFE6C16D943F1330" collectionName="IZAN">IZAN</collectionCode>
);
<emphasis id="1C8CC21F0F3804439B3A6F128D104F88" bold="true" pageId="0" pageNumber="25">K</emphasis>
-
<emphasis id="D5FD7F27A54F438F743B28D25EF0E9AB" bold="true" pageId="0" pageNumber="25">M</emphasis>
. Specimen from Lake Sapanca, Yeni
<normalizedToken id="A69503BD8229691FF5CAC5BBE2AC42E3" originalValue="Eşme">Esme</normalizedToken>
, Turkey (
<collectionCode id="74F664A32A612F792ECF9F690794B625" collectionName="UGSB">UGSB</collectionCode>
20691).
<taxonomicName id="E3498E698BFC6311C95E8EFC2608F650" baseAuthorityName="Lindholm" baseAuthorityYear="1901" class="Gastropoda" family="Neritidae" genus="Theodoxus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Theodoxus sarmaticus" order="Cycloneritida" pageId="0" pageNumber="25" phylum="Mollusca" rank="species" species="sarmaticus">
<emphasis id="F45224E157A0657E54231590F4C5C8D1" italics="true" pageId="0" pageNumber="25">Theodoxus sarmaticus</emphasis>
</taxonomicName>
(Lindholm, 1901).
<emphasis id="68E7A9F41FE01D82EACD975A4AD052BE" bold="true" pageId="0" pageNumber="25">N</emphasis>
-
<emphasis id="77ED2E4EBBDD30D32D2F998AF2FF60BC" bold="true" pageId="0" pageNumber="25">P</emphasis>
;
<emphasis id="E45B442DB372E3A96DA5268B3A831B7D" bold="true" pageId="0" pageNumber="25">Q</emphasis>
-
<emphasis id="662DBEF09C2B52804B7A2FBA85DE8811" bold="true" pageId="0" pageNumber="25">S</emphasis>
;
<emphasis id="216FE4BA92BAF65D03F2D9D76EDAAC1C" bold="true" pageId="0" pageNumber="25">T</emphasis>
-
<emphasis id="AB6CEC1997E2EE3F23B37F70116C6DFC" bold="true" pageId="0" pageNumber="25">U</emphasis>
. Three
<typeStatus id="9FF34D467D06267D5DED6C987032E1D2">syntypes</typeStatus>
from the Oskol River, near Golubino, Russia stored in
<collectionCode id="5A1D4858CB0CA65C5F55B12A72A1939A" collectionName="Russia, St. Petersburg, Russian Academy of Sciences, Zoological Institute" country="Russia" httpUri="http://grbio.org/cool/z2y8-2jfr" name="Russian Academy of Sciences, Zoological Institute, Zoological Museum" type="Museum">ZIN</collectionCode>
(coll. no. 6099/24). Scale bars:
<quantity id="E1E28BDE50427962FB9AFB94E492900D" metricMagnitude="-3" metricUnit="m" metricValue="1.0" unit="mm" value="1.0">1 mm</quantity>
.
</paragraph>
</caption>
<paragraph id="5C29A676C2FBD87F760C00B6EC90D455" pageId="0" pageNumber="25">
The geographic distribution range of
<taxonomicName id="2CFB38FFE98C4607413ABB3D68C55922" lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
<emphasis id="B776C77995842E773E864BBD98E52317" italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
</taxonomicName>
overlaps with that of
<taxonomicName id="19527C2E23F6A293AD7103423DF52ADA" lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
<emphasis id="70FFA8D699D923CAE60062E2EE47E906" italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
</taxonomicName>
(Lindholm, 1901), which has been considered a junior synonym of
<taxonomicName id="5BAC751BECDD300BBE55D585E4785097" lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
<emphasis id="88738FFF41B6CFC368FFF32E542C32A6" italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
</taxonomicName>
(e.g.
<bibRefCitation id="AD18F9B4CC96CDCE31188B6A26274E6E" author="Vinarski, MV" journalOrPublisher="Severtsov Institute of Ecology and Evolution of RAS, Moscow" pageId="0" pageNumber="25" refId="B157" refString="Vinarski, MV, Kantor, YI, 2016. Analytical catalogue of fresh and brackish water molluscs of Russia and adjacent countries. A.N. Severtsov Institute of Ecology and Evolution of RAS, Moscow" title="Analytical catalogue of fresh and brackish water molluscs of Russia and adjacent countries. A. N." year="2016">Vinarski and Kantor 2016</bibRefCitation>
). While
<taxonomicName id="D715DB2BA3B394E43AE96CD6B5CF9E43" lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
<emphasis id="FA5D08C9286BE34AA76DA0FB6F24AE75" italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
</taxonomicName>
and
<taxonomicName id="16B6744C8BFBA73A2C415F99BEFF9DD7" lsidName="T. fluviatilis" pageId="0" pageNumber="25" rank="species" species="fluviatilis">
<emphasis id="F78B9437AB1339D57DDB0C59F2B8F8CE" italics="true" pageId="0" pageNumber="25">T. fluviatilis</emphasis>
</taxonomicName>
may on occasion share phenotypic similarity, the morphotypes of
<taxonomicName id="A93E2C0315E9C387F2FE7693304A7589" lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
<emphasis id="FCDDF32A9993E6B5AD285F7BA40114B8" italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
</taxonomicName>
closely resemble
<taxonomicName id="0CE18C95E9419879D2100FAE54F4018F" lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
<emphasis id="C832B3F7F1F4F4533D1DE9D0AEFA3D79" italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
</taxonomicName>
(Fig.
<figureCitation id="D94DF84BBD8FDB506891E5C346EF6F08" captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22A-M</figureCitation>
versus Fig.
<figureCitation id="664E71DC46FC91FA88D0CB33337E8F35" captionStart="Figure 22" captionStartId="F22" captionText="Figure 22. Theodoxus velox V. Anistratenko in O. Anistratenko et al., 1999. A - D. Specimen from the Dnieper River, Kiev, Ukraine (UGSB 24186); E - H. Specimen collected in the Danube delta, Vylkove, Ukraine (UGSB 24184) used in the phylogeny (Fig. 2); I, J. Holotype from Dnieper Delta, Zbur'ivka Liman, Ukraine (IZAN); K - M. Specimen from Lake Sapanca, Yeni Esme, Turkey (UGSB 20691). Theodoxus sarmaticus (Lindholm, 1901). N - P; Q - S; T - U. Three syntypes from the Oskol River, near Golubino, Russia stored in ZIN (coll. no. 6099 / 24). Scale bars: 1 mm." figureDoi="10.3897/zse.96.48312.figure22" httpUri="https://binary.pensoft.net/fig/376471" pageId="0" pageNumber="25">22N-U</figureCitation>
). The similarity with
<taxonomicName id="512972C8367B03FF9896CCBA13FFB69E" lsidName="T. velox" pageId="0" pageNumber="25" rank="species" species="velox">
<emphasis id="36B095C152D4FE5D50E708568CF4D7DF" italics="true" pageId="0" pageNumber="25">T. velox</emphasis>
</taxonomicName>
could suggest
<taxonomicName id="999BA0D55A08EB971309EE21497AA417" lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
<emphasis id="91F59C3B73753F07856B18339FF4F33F" italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
</taxonomicName>
is rather conspecific with that species as suggested by
<bibRefCitation id="CEA250D22EC6230F9811DDA3B1E35D2B" author="Gloeer, P" journalOrPublisher="Peter Gloeer, Hetlingen, Germany" pageId="0" pageNumber="25" refId="B62" refString="Gloeer, P, 2019. The freshwater gastropods of the West-Palaearctis. Vol. 1. Fresh- and brackish waters except spring and subterranean snails. Peter Gloeer, Hetlingen, Germany" title="The freshwater gastropods of the West-Palaearctis. Vol. 1. Fresh- and brackish waters except spring and subterranean snails." year="2019">
<normalizedToken id="53C9539027481B3CE93B56A0D541A86B" originalValue="Glöer">Gloeer</normalizedToken>
(2019)
</bibRefCitation>
. Unfortunately, no opercula or soft tissues were preserved among the
<typeStatus id="4717D0D140B84B51108C3867925DEC86">syntypes</typeStatus>
of
<taxonomicName id="650A2A0B70F07FFD1566E40F5A34A25B" lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
<emphasis id="0273EFAD8CA2917744ECBBF543954935" italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
</taxonomicName>
to corroborate this hypothesis. Molecular analyses of topotypic material is required to address this uncertainty. However, should this be confirmed at a later point, the name
<taxonomicName id="A841E3E1174BD6F94F1DB9F266058303" lsidName="T. sarmaticus" pageId="0" pageNumber="25" rank="species" species="sarmaticus">
<emphasis id="E3D239B1D47B33C5402FB8F704ECC613" italics="true" pageId="0" pageNumber="25">T. sarmaticus</emphasis>
</taxonomicName>
would have priority.
</paragraph>
</subSubSection>
<subSubSection id="357563DE498436315D5FC72623722484" pageId="0" pageNumber="25" type="distribution">
<paragraph id="91EE757E1577F5970D1E2FDB37AD0F24" pageId="0" pageNumber="25">Distribution.</paragraph>
<paragraph id="BD175F03A131AA3A69405AD45668A010" pageId="0" pageNumber="25">
This species was indicated to be restricted to drainage systems of the northern Black Sea coast (
<bibRefCitation id="30E4F974CB9CC7CB1EAE86F6E4378CFC" author="Anistratenko, OY" journalOrPublisher="Vestnik Zoologii" pageId="0" pageNumber="25" pagination="11 - 19" refId="B14" refString="Anistratenko, OY, Starobogatov, YI, Anistratenko, VV, 1999. Mollyuski roda Theodoxus (Gastropoda, Pectinibranchia, Neritidae) Azovo-Chernomorskogo basseyna. Vestnik Zoologii 33 (3): 11 - 19" title="Mollyuski roda Theodoxus (Gastropoda, Pectinibranchia, Neritidae) Azovo-Chernomorskogo basseyna." volume="33" year="1999">Anistratenko et al. 1999</bibRefCitation>
;
<bibRefCitation id="8626255F07325D92F8359DE5D45B7350" author="Kantor, YI" journalOrPublisher="KMK Scientific Press, Moscow" pageId="0" pageNumber="25" refId="B78" refString="Kantor, YI, Sysoev, AV, 2006. Morskiye i solonovatovodnyye bryukhonogiye mollyuski Rossii i sopredel'nykh stran: illyustrirovannyy katalog. KMK Scientific Press, Moscow" title="Morskiye i solonovatovodnyye bryukhonogiye mollyuski Rossii i sopredel'nykh stran: illyustrirovannyy katalog." year="2006">Kantor and Sysoev 2006</bibRefCitation>
). Recent molecular data suggest it is distributed as far North as the eastern part of the Baltic Sea and as far South as Anatolia (
<bibRefCitation id="A4D6BD3BCACD61AED3D201D5F9279ACC" 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>
): the only record there derives from Lake Sapanca (
<bibRefCitation id="D7B932AEB246C5DA1ECD2362777E948D" 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 id="304DA5B8720CA4C8E2A3550832C49D41" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Occurrence map for Theodoxus spp. in Asia. All points conform to specimens used in the phylogenetic analyses (Suppl. material 1: Table S 1), except for T. gloeri for which no molecular data are available and only the type and paratype localities are indicated. Species are partitioned across four identical maps (A - D) to limit overlap of points as far as possible. Barring T. gloeri, closer related species are grouped together (Fig. 2)." figureDoi="10.3897/zse.96.48312.figure3" httpUri="https://binary.pensoft.net/fig/376452" pageId="0" pageNumber="25">3C</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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