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<document id="FF623749BDF692628C45A2BB9EE5EA2E" ID-CLB-Dataset="133178" ID-DOI="10.1093/zoolinnean/zlab121" ID-GBIF-Dataset="a2770ec2-41f4-4418-ba75-7ef5a99b88e4" ID-ISSN="0024-4082" ID-Zenodo-Dep="7035584" ID-ZooBank="89452E3C-3AFE-449C-93C6-24EEE19AE45D" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1661767267254" checkinUser="plazi" docAuthor="Delicado, Diana &amp; Hauffe, Torsten" docDate="2022" docId="039D87B34C071B71FC31F9B1FA7675FC" docLanguage="en" docName="zlab121.pdf" docOrigin="Zoological Journal of the Linnean Society 196" docSource="https://doi.org/10.1093/zoolinnean/zlab121" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-2023.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-2023.journal_article" docStyleVersion="7" docTitle="Radomaniola variabilis Delicado &amp; Hauffe 2022, SP. NOV." docType="treatment" docVersion="9" lastPageNumber="430" masterDocId="FFA4FFCB4C251B54FFBAFF85FFD2705A" masterDocTitle="Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years" masterLastPageNumber="441" masterPageNumber="393" pageNumber="427" updateTime="1699373995385" updateUser="ExternalLinkService">
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<mods:title id="13AF103B30E6F678F4DBD37D3A9EC0FC">Shell features and anatomy of the springsnail genus Radomaniola (Caenogastropoda: Hydrobiidae) show a different pace and mode of evolution over five million years</mods:title>
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<mods:namePart id="C5A4451CB4EA6EC85D0FBC2E7139095B">Delicado, Diana</mods:namePart>
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<mods:namePart id="1220467398C596916BB6F254B4081C66">Hauffe, Torsten</mods:namePart>
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<mods:date id="6ADDC02007DA53C1E7429BA78C71714A">2022</mods:date>
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<treatment id="039D87B34C071B71FC31F9B1FA7675FC" ID-DOI="http://doi.org/10.5281/zenodo.7036891" ID-GBIF-Taxon="199898769" ID-Zenodo-Dep="7036891" LSID="urn:lsid:plazi:treatment:039D87B34C071B71FC31F9B1FA7675FC" httpUri="http://treatment.plazi.org/id/039D87B34C071B71FC31F9B1FA7675FC" lastPageId="37" lastPageNumber="430" pageId="34" pageNumber="427">
<subSubSection id="C32E652E4C071B76FC31F9B1FAFC7616" box="[907,1326,1588,1612]" pageId="34" pageNumber="427" type="nomenclature">
<paragraph id="8B8B36A54C071B76FC31F9B1FAFC7616" blockId="34.[907,1326,1588,1612]" box="[907,1326,1588,1612]" pageId="34" pageNumber="427">
<heading id="D0C381C94C071B76FC31F9B1FAFC7616" box="[907,1326,1588,1612]" centered="true" fontSize="9" level="2" pageId="34" pageNumber="427" reason="2">
<taxonomicName id="4C344D264C071B76FC31F9B1FB167610" ID-CoL="9CCKH" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[907,1220,1588,1611]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="variabilis" status="sp. nov.">RADOMANIOLA VARIABILIS</taxonomicName>
<emphasis id="B940EAB74C071B76FB71F9B2FAFC7616" bold="true" box="[1227,1326,1588,1612]" pageId="34" pageNumber="427">
<taxonomicNameLabel id="A27357CC4C071B76FB71F9B2FAFC7616" box="[1227,1326,1588,1612]" pageId="34" pageNumber="427" rank="species">SP. NOV.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C32E652E4C071B71FBA9F9DFFA7675FC" lastPageId="37" lastPageNumber="430" pageId="34" pageNumber="427" type="description">
<paragraph id="8B8B36A54C071B76FBA9F9DFFB747629" blockId="34.[1043,1190,1626,1651]" box="[1043,1190,1626,1651]" pageId="34" pageNumber="427">
(
<figureCitation id="130F2A204C071B76FBA6F9DFFBA67629" box="[1052,1140,1626,1651]" captionStart="Figure 22" captionStartId="35.[163,243,1708,1730]" captionTargetBox="[163,1443,195,1669]" captionTargetId="figure-26@35.[163,1443,195,1669]" captionTargetPageId="35" captionText="Figure 22. Shells, operculum and radulae of Radomaniola variabilis sp. nov. A, B, holotype (MNCN 15.05/200169). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975)." figureDoi="http://doi.org/10.5281/zenodo.7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="34" pageNumber="427">FIGS 22</figureCitation>
,
<figureCitation id="130F2A204C071B76FB38F9DFFB4C7629" box="[1154,1182,1626,1651]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="34" pageNumber="427">23</figureCitation>
)
</paragraph>
<paragraph id="8B8B36A54C071B76FC93F901FA5B76ED" blockId="34.[809,1426,1667,1719]" pageId="34" pageNumber="427">
<emphasis id="B940EAB74C071B76FC93F901FB0776C3" box="[809,1237,1667,1689]" italics="true" pageId="34" pageNumber="427">Z o o b a n k r e g i s t r a t i o n:</emphasis>
z o o b a n k. org:act:
<uuid id="FF920C704C071B76FCCFF924FA5B76ED" box="[885,1417,1697,1719]" pageId="34" pageNumber="427">BACA04A4-0420-4FB3-B15B-A3CD6789A303</uuid>
</paragraph>
<paragraph id="8B8B36A54C071B76FC93F965FA98774F" blockId="34.[809,1425,1760,1813]" pageId="34" pageNumber="427">
<emphasis id="B940EAB74C071B76FC93F965FC7576AF" box="[809,935,1760,1781]" italics="true" pageId="34" pageNumber="427">Etymology:</emphasis>
From Latin
<taxonomicName id="4C344D264C071B76FBFCF965FB6176AF" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[1094,1203,1760,1781]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="variabilis">
<emphasis id="B940EAB74C071B76FBFCF965FB6176AF" box="[1094,1203,1760,1781]" italics="true" pageId="34" pageNumber="427">variabilis</emphasis>
</taxonomicName>
, variable, referring to its high intraspecific variation in shell shape.
</paragraph>
<paragraph id="8B8B36A54C071B70FC93F8BBFF34763C" blockId="34.[809,1426,1853,1906]" lastBlockId="36.[145,761,1556,1638]" lastPageId="36" lastPageNumber="429" pageId="34" pageNumber="427">
<emphasis id="B940EAB74C071B76FC93F8BBFC0F7708" box="[809,989,1853,1875]" italics="true" pageId="34" pageNumber="427">Type material:</emphasis>
<materialsCitation id="3B5C3CF84C071B70FC4FF8B8FF34763C" ID-GBIF-Occurrence="3897142323" collectionCode="MNCN, UGSB" country="Greece" lastPageId="36" lastPageNumber="429" location="Vellas Spring" municipality="Kalpaki" pageId="34" pageNumber="427" specimenCode="MNCN 15.05, UGSB 23975" specimenCount="2" typeStatus="holotype">
<typeStatus id="548F88074C071B76FC4FF8B8FBB17709" box="[1013,1123,1853,1875]" pageId="34" pageNumber="427" type="holotype">Holotype</typeStatus>
(
<emphasis id="B940EAB74C071B76FBCCF8BBFA527708" box="[1142,1408,1853,1875]" italics="true" pageId="34" pageNumber="427">
<specimenCode id="DB929EDE4C071B76FBCCF8BBFACB7708" box="[1142,1305,1853,1875]" collectionCode="MNCN" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="34" pageNumber="427" type="Museum">MNCN 15.05</specimenCode>
/200169
</emphasis>
),
<specimenCount id="9D32FD2C4C071B76FC93F8D9FC327728" box="[809,992,1884,1906]" pageId="34" pageNumber="427" type="generic" typeStatus="paratypes">three paratypes</specimenCount>
(
<specimenCode id="DB929EDE4C071B76FC55F8D9FB547728" box="[1007,1158,1884,1906]" collectionCode="MNCN" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="34" pageNumber="427" type="Museum">MNCN 15.05</specimenCode>
/200170) in the
<collectionCode id="ED25AE604C071B76FA84F8D9FA42772B" box="[1342,1424,1884,1905]" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="34" pageNumber="427" type="Museum">MNCN</collectionCode>
collection and
<specimenCount id="9D32FD2C4C011B70FE87F991FE327673" box="[317,480,1556,1577]" pageId="36" pageNumber="429" type="generic" typeStatus="paratypes">ten paratypes</specimenCount>
in the
<collectionCode id="ED25AE604C011B70FD8DF991FD537673" box="[567,641,1556,1577]" pageId="36" pageNumber="429">UGSB</collectionCode>
collection (
<specimenCode id="DB929EDE4C011B70FF22F9B6FEE67612" box="[152,308,1586,1608]" collectionCode="UGSB" pageId="36" pageNumber="429">UGSB 23975</specimenCode>
). Type locality:
<location id="8EEB607E4C011B70FE51F9B7FD5B7612" LSID="urn:lsid:plazi:treatment:039D87B34C071B71FC31F9B1FA7675FC:8EEB607E4C011B70FE51F9B7FD5B7612" box="[491,649,1586,1608]" country="Greece" municipality="Kalpaki" name="Vellas Spring" pageId="36" pageNumber="429">Vellas Spring</location>
,
<collectingMunicipality id="6BEFACDF4C011B70FD2FF9B7FD267612" box="[661,756,1586,1608]" pageId="36" pageNumber="429">Kalpaki</collectingMunicipality>
,
<collectingCountry id="F32376354C011B70FF2BF9D4FF30763C" box="[145,226,1617,1638]" name="Greece" pageId="36" pageNumber="429">Greece</collectingCountry>
.
</materialsCitation>
</paragraph>
<caption id="DF4B662D4C061B77FF19F929FA767741" ID-DOI="http://doi.org/10.5281/zenodo.7035636" ID-Zenodo-Dep="7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="35" pageNumber="428" startId="35.[163,243,1708,1730]" targetBox="[163,1443,195,1669]" targetPageId="35">
<paragraph id="8B8B36A54C061B77FF19F929FA767741" blockId="35.[163,1444,1708,1819]" pageId="35" pageNumber="428">
<emphasis id="B940EAB74C061B77FF19F929FEC97698" bold="true" box="[163,283,1708,1730]" pageId="35" pageNumber="428">Figure 22.</emphasis>
Shells, operculum and radulae of
<taxonomicName id="4C344D264C061B77FD2BF928FC427699" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[657,912,1709,1731]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="35" pageNumber="428" phylum="Mollusca" rank="species" species="variabilis" status="sp. nov.">
<emphasis id="B940EAB74C061B77FD2BF928FC427699" box="[657,912,1709,1731]" italics="true" pageId="35" pageNumber="428">Radomaniola variabilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27357CC4C061B77FC22F928FC347699" box="[920,998,1709,1731]" pageId="35" pageNumber="428" rank="species">sp. nov.</taxonomicNameLabel>
A, B, holotype (
<emphasis id="B940EAB74C061B77FB2AF928FAAA7699" box="[1168,1400,1709,1731]" italics="true" pageId="35" pageNumber="428">MNCN 15.05/200169</emphasis>
). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975).
</paragraph>
</caption>
<caption id="DF4B662D4C011B70FF2BFAEAFCA7759A" ID-DOI="http://doi.org/10.5281/zenodo.7035642" ID-Zenodo-Dep="7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="36" pageNumber="429" startId="36.[145,223,1391,1413]" targetBox="[145,1425,195,1352]" targetPageId="36">
<paragraph id="8B8B36A54C011B70FF2BFAEAFCA7759A" blockId="36.[145,1426,1391,1472]" pageId="36" pageNumber="429">
<emphasis id="B940EAB74C011B70FF2BFAEAFED775DF" bold="true" box="[145,261,1391,1413]" pageId="36" pageNumber="429">Figure 23.</emphasis>
Anatomy of
<taxonomicName id="4C344D264C011B70FE31FAF5FD5175DC" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[395,643,1392,1414]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="36" pageNumber="429" phylum="Mollusca" rank="species" species="variabilis" status="sp. nov.">
<emphasis id="B940EAB74C011B70FE31FAF5FD5175DC" box="[395,643,1392,1414]" italics="true" pageId="36" pageNumber="429">Radomaniola variabilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27357CC4C011B70FD33FAF5FD0175DC" box="[649,723,1392,1414]" pageId="36" pageNumber="429" rank="species">sp. nov.</taxonomicNameLabel>
AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis.
</paragraph>
</caption>
<paragraph id="8B8B36A54C011B70FF2BF915FB88763D" blockId="36.[145,761,1680,1886]" lastBlockId="36.[809,1425,1556,1639]" pageId="36" pageNumber="429">
<emphasis id="B940EAB74C011B70FF2BF915FEB776FF" box="[145,357,1680,1701]" italics="true" pageId="36" pageNumber="429">Material studied:</emphasis>
<materialsCitation id="3B5C3CF84C011B70FEC1F915FD2A76B9" ID-GBIF-Occurrence="3897142314" collectingDate="2003-06" collectingDateMax="2005-05" collectingDateMin="2003-06" collectionCode="T" country="Greece" county="Muzine" latitude="39.92" location="Qafe Murres" longLatPrecision="700" longitude="20.18" municipality="Kalter Lake" pageId="36" pageNumber="429" specimenCode="MNCN 15.05, UGSB 23975, UGSB 14440, UGSB 14441, UGSB 2294" specimenCount="1">
<collectorName id="26C153734C011B70FEC1F915FDF376FC" box="[379,545,1680,1702]" pageId="36" pageNumber="429">Vellas Spring</collectorName>
,
<collectorName id="26C153734C011B70FD95F915FD4376FC" box="[559,657,1680,1702]" pageId="36" pageNumber="429">Kalpaki</collectorName>
,
<collectingCountry id="F32376354C011B70FD1AF915FD2676FF" box="[672,756,1680,1701]" name="Greece" pageId="36" pageNumber="429">Greece</collectingCountry>
,
<geoCoordinate id="EE0050624C011B70FF2BF92BFEC6769E" box="[145,276,1710,1732]" degrees="39.8660" direction="north" orientation="latitude" pageId="36" pageNumber="429" precision="5" value="39.866">39.8660°N</geoCoordinate>
,
<geoCoordinate id="EE0050624C011B70FE9EF92BFE75769E" box="[292,423,1710,1732]" degrees="20.6243" direction="east" orientation="longitude" pageId="36" pageNumber="429" precision="5" value="20.6243">20.6243°E</geoCoordinate>
,
<collectionCode id="ED25AE604C011B70FE0CF92AFE15769E" box="[438,455,1711,1732]" name="Tavera, Department of Geology and Geophysics" pageId="36" pageNumber="429">T</collectionCode>
.W. and
<collectorName id="26C153734C011B70FD95F92AFDB7769E" box="[559,613,1711,1732]" name="Cardoso A" pageId="36" pageNumber="429">C.A.</collectorName>
,
<date id="FF8A10654C011B70FDCCF92AFD21769E" box="[630,755,1710,1732]" pageId="36" pageNumber="429" value="2005-05">
<collectingDate id="EFCEE98D4C011B70FDCCF92AFD21769E" box="[630,755,1710,1732]" pageId="36" pageNumber="429" value="2005-05">May 2005</collectingDate>
</date>
,
<specimenCode id="DB929EDE4C011B70FF2BF948FEF676B9" box="[145,292,1741,1763]" collectionCode="MNCN" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="36" pageNumber="429" type="Museum">MNCN 15.05</specimenCode>
/200170 and
<specimenCode id="DB929EDE4C011B70FE03F948FD9C76B9" box="[441,590,1741,1763]" collectionCode="UGSB" pageId="36" pageNumber="429">UGSB 23975</specimenCode>
(80% ethanol);
</materialsCitation>
<materialsCitation id="3B5C3CF84C011B70FF2BF969FF297765" ID-GBIF-Occurrence="3897142320" collectingDate="2003-06" collectingDateMax="2005-05" collectingDateMin="2003-06" collectionCode="T" country="Greece" county="Muzine" latitude="39.92" location="Qafe Murres" longLatPrecision="700" longitude="20.18" municipality="Kalter Lake" pageId="36" pageNumber="429" specimenCode="MNCN 15.05, UGSB 23975, UGSB 14440, UGSB 14441, UGSB 2294" specimenCount="1">
unnamed spring,
<quantity id="4CCC9B404C011B70FEE6F969FE457758" box="[348,407,1772,1794]" metricMagnitude="3" metricUnit="m" metricValue="3.0" pageId="36" pageNumber="429" unit="km" value="3.0">3 km</quantity>
west of
<location id="8EEB607E4C011B70FE42F969FD5F775B" LSID="urn:lsid:plazi:treatment:039D87B34C071B71FC31F9B1FA7675FC:8EEB607E4C011B70FE42F969FD5F775B" box="[504,653,1772,1794]" country="Greece" county="Muzine" latitude="39.92" longLatPrecision="700" longitude="20.18" municipality="Kalter Lake" name="Qafe Murres" pageId="36" pageNumber="429">Qafë Murrës</location>
,
<collectingCountry id="F32376354C011B70FD22F969FD277758" box="[664,757,1772,1794]" name="Albania" pageId="36" pageNumber="429">Albania</collectingCountry>
,
<geoCoordinate id="EE0050624C011B70FF2BF88FFF22777A" box="[145,240,1802,1824]" degrees="41.63" direction="north" orientation="latitude" pageId="36" pageNumber="429" precision="555" value="41.63">41.63°N</geoCoordinate>
,
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, Z.F.,
<date id="FF8A10654C011B70FE2EF88EFDC2777A" box="[404,528,1803,1824]" pageId="36" pageNumber="429" value="2003-06">
<collectingDate id="EFCEE98D4C011B70FE2EF88EFDC2777A" box="[404,528,1803,1824]" pageId="36" pageNumber="429" value="2003-06">June 2003</collectingDate>
</date>
,
<specimenCode id="DB929EDE4C011B70FDA7F88EFD64777A" box="[541,694,1803,1824]" collectionCode="UGSB" pageId="36" pageNumber="429">UGSB 14440</specimenCode>
(80% ethanol);
</materialsCitation>
<materialsCitation id="3B5C3CF84C011B70FEBEF8ACFBCB7670" ID-GBIF-Occurrence="3897142302" collectingDate="2003-06" collectingDateMax="2005-05" collectingDateMin="2003-06" collectionCode="T" country="Greece" county="Muzine" latitude="39.92" location="Qafe Murres" longLatPrecision="700" longitude="20.18" municipality="Kalter Lake" pageId="36" pageNumber="429" specimenCode="MNCN 15.05, UGSB 23975, UGSB 14440, UGSB 14441, UGSB 2294" specimenCount="1">
unnamed spring,
<quantity id="4CCC9B404C011B70FE62F8AFFDC77765" box="[472,533,1833,1855]" metricMagnitude="3" metricUnit="m" metricValue="7.0" pageId="36" pageNumber="429" unit="km" value="7.0">7 km</quantity>
south of
<location id="8EEB607E4C011B70FD32F8ACFD267765" LSID="urn:lsid:plazi:treatment:039D87B34C071B71FC31F9B1FA7675FC:8EEB607E4C011B70FD32F8ACFD267765" box="[648,756,1833,1855]" country="Greece" county="Muzine" latitude="39.92" longLatPrecision="700" longitude="20.18" municipality="Kalter Lake" name="Tepelene" pageId="36" pageNumber="429">Tepelenë</location>
,
<collectingCountry id="F32376354C011B70FF2BF8CDFF3C7704" box="[145,238,1864,1886]" name="Albania" pageId="36" pageNumber="429">Albania</collectingCountry>
,
<geoCoordinate id="EE0050624C011B70FF40F8CDFE8B7707" box="[250,345,1864,1885]" degrees="40.27" direction="north" orientation="latitude" pageId="36" pageNumber="429" precision="555" value="40.27">40.27°N</geoCoordinate>
,
<geoCoordinate id="EE0050624C011B70FEDEF8CDFE107707" box="[356,450,1864,1885]" degrees="20.28" direction="east" orientation="longitude" pageId="36" pageNumber="429" precision="555" value="20.28">20.28°E</geoCoordinate>
, Z.F.,
<date id="FF8A10654C011B70FDBDF8CDFD767707" box="[519,676,1864,1886]" pageId="36" pageNumber="429" value="2004-10">
<collectingDate id="EFCEE98D4C011B70FDBDF8CDFD767707" box="[519,676,1864,1886]" pageId="36" pageNumber="429" value="2004-10">October 2004</collectingDate>
</date>
,
<specimenCode id="DB929EDE4C011B70FD0AF8CDFCBC7673" box="[688,878,1556,1885]" collectionCode="UGSB" pageId="36" pageNumber="429">UGSB 14441</specimenCode>
(80% ethanol);
</materialsCitation>
<materialsCitation id="3B5C3CF84C011B70FB9BF991FB85763D" ID-GBIF-Occurrence="3897142310" collectingDate="2003-06" collectingDateMax="2005-05" collectingDateMin="2003-06" collectionCode="T" country="Greece" county="Muzine" latitude="39.92" location="Qafe Murres" longLatPrecision="700" longitude="20.18" municipality="Kalter Lake" pageId="36" pageNumber="429" specimenCode="MNCN 15.05, UGSB 23975, UGSB 14440, UGSB 14441, UGSB 2294" specimenCount="1">
<collectorName id="26C153734C011B70FB9BF991FB837670" box="[1057,1105,1556,1578]" pageId="36" pageNumber="429">Syri</collectorName>
i
<collectingMunicipality id="6BEFACDF4C011B70FBDDF991FB217670" box="[1127,1267,1556,1578]" pageId="36" pageNumber="429">Kaltër Lake</collectingMunicipality>
,
<quantity id="4CCC9B404C011B70FB44F991FAEB7670" box="[1278,1337,1556,1578]" metricMagnitude="3" metricUnit="m" metricValue="7.0" pageId="36" pageNumber="429" unit="km" value="7.0">7 km</quantity>
west of
<collectingCounty id="62EA4E294C011B70FC93F9B7FC527612" box="[809,896,1586,1608]" pageId="36" pageNumber="429">Muzinë</collectingCounty>
,
<collectingCountry id="F32376354C011B70FC30F9B7FC377612" box="[906,997,1586,1608]" name="Albania" pageId="36" pageNumber="429">Albania</collectingCountry>
,
<geoCoordinate id="EE0050624C011B70FC4AF9B6FB9E7612" box="[1008,1100,1587,1608]" degrees="39.92" direction="north" orientation="latitude" pageId="36" pageNumber="429" precision="555" value="39.92">39.92°N</geoCoordinate>
,
<geoCoordinate id="EE0050624C011B70FBEDF9B6FB617612" box="[1111,1203,1587,1608]" degrees="20.18" direction="east" orientation="longitude" pageId="36" pageNumber="429" precision="555" value="20.18">20.18°E</geoCoordinate>
, Z.F.,
<date id="FF8A10654C011B70FB4EF9B7FA5F7612" box="[1268,1421,1586,1608]" pageId="36" pageNumber="429" value="2004-10">
<collectingDate id="EFCEE98D4C011B70FB4EF9B7FA5F7612" box="[1268,1421,1586,1608]" pageId="36" pageNumber="429" value="2004-10">October 2004</collectingDate>
</date>
,
<specimenCode id="DB929EDE4C011B70FC93F9D4FC7D763C" box="[809,943,1617,1638]" collectionCode="UGSB" pageId="36" pageNumber="429">UGSB 2294</specimenCode>
(80% ethanol)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8B36A54C011B71FC93F915FEEB70A3" blockId="36.[809,1424,1680,1886]" lastBlockId="37.[163,778,197,250]" lastPageId="37" lastPageNumber="430" pageId="36" pageNumber="429">
<emphasis id="B940EAB74C011B70FC93F915FC6376FF" box="[809,945,1680,1701]" italics="true" pageId="36" pageNumber="429">Diagnosis:</emphasis>
Protoconch microsculpture pitted; central radular tooth formula 7-C-7/1-1; bursa copulatrix ovoid, duct slightly longer than bursal length; SR1 pyriform, duct short; SR2 pyriform, duct short, slightly larger than SR1; penis unpigmented, gradually tapering, base narrow, approximately as long as head length; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.47).
</paragraph>
<paragraph id="8B8B36A54C001B71FF19FEA6FEED7324" blockId="37.[161,780,290,1907]" pageId="37" pageNumber="430">
<emphasis id="B940EAB74C001B71FF19FEA6FEFE7162" box="[163,300,291,312]" italics="true" pageId="37" pageNumber="430">Description:</emphasis>
Shell ovate-conic, 4.04.5 whorls, height 2.0
<quantity id="4CCC9B404C001B71FF6DFEC4FEE0710D" box="[215,306,321,343]" metricMagnitude="-3" metricUnit="m" metricValue="2.6" pageId="37" pageNumber="430" unit="mm" value="2.6">2.6 mm</quantity>
(
<figureCitation id="130F2A204C001B71FEFCFEC4FE1F710C" box="[326,461,321,343]" captionStart="Figure 22" captionStartId="35.[163,243,1708,1730]" captionTargetBox="[163,1443,195,1669]" captionTargetId="figure-26@35.[163,1443,195,1669]" captionTargetPageId="35" captionText="Figure 22. Shells, operculum and radulae of Radomaniola variabilis sp. nov. A, B, holotype (MNCN 15.05/200169). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975)." figureDoi="http://doi.org/10.5281/zenodo.7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="37" pageNumber="430">Fig. 22AF</figureCitation>
; Supporting Information, Table S6). Periostracum yellowish, sometimes whitish. Protoconch ~425 µm wide, 1.5 whorls; nucleus ~175 µm wide; protoconch microsculpture pitted (
<figureCitation id="130F2A204C001B71FD5AFE18FF1B718B" captionStart="Figure 22" captionStartId="35.[163,243,1708,1730]" captionTargetBox="[163,1443,195,1669]" captionTargetId="figure-26@35.[163,1443,195,1669]" captionTargetPageId="35" captionText="Figure 22. Shells, operculum and radulae of Radomaniola variabilis sp. nov. A, B, holotype (MNCN 15.05/200169). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975)." figureDoi="http://doi.org/10.5281/zenodo.7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="37" pageNumber="430">Fig. 22I</figureCitation>
). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about two-thirds of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (
<figureCitation id="130F2A204C001B71FD5AFD9DFF387216" captionStart="Figure 22" captionStartId="35.[163,243,1708,1730]" captionTargetBox="[163,1443,195,1669]" captionTargetId="figure-26@35.[163,1443,195,1669]" captionTargetPageId="35" captionText="Figure 22. Shells, operculum and radulae of Radomaniola variabilis sp. nov. A, B, holotype (MNCN 15.05/200169). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975)." figureDoi="http://doi.org/10.5281/zenodo.7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="37" pageNumber="430">Fig. 22B, F</figureCitation>
). Umbilicus narrow, not covered by the inner lip. Operculum oval, brownish, about two whorls; muscle attachment area oval and located near the nucleus (
<figureCitation id="130F2A204C001B71FF11FD17FEE272F2" box="[171,304,658,680]" captionStart="Figure 22" captionStartId="35.[163,243,1708,1730]" captionTargetBox="[163,1443,195,1669]" captionTargetId="figure-26@35.[163,1443,195,1669]" captionTargetPageId="35" captionText="Figure 22. Shells, operculum and radulae of Radomaniola variabilis sp. nov. A, B, holotype (MNCN 15.05/200169). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975)." figureDoi="http://doi.org/10.5281/zenodo.7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="37" pageNumber="430">Fig. 22G, H</figureCitation>
). Radular length intermediate, ~600 µm (~25% of total shell length), with ~60 rows of teeth. Central tooth formula 7-C-7/1-1 (
<figureCitation id="130F2A204C001B71FD8EFD55FD6172BF" box="[564,691,720,742]" captionStart="Figure 22" captionStartId="35.[163,243,1708,1730]" captionTargetBox="[163,1443,195,1669]" captionTargetId="figure-26@35.[163,1443,195,1669]" captionTargetPageId="35" captionText="Figure 22. Shells, operculum and radulae of Radomaniola variabilis sp. nov. A, B, holotype (MNCN 15.05/200169). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975)." figureDoi="http://doi.org/10.5281/zenodo.7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="37" pageNumber="430">Fig. 22JL</figureCitation>
); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula (3)4-C-4(3). Inner marginal teeth having 2532 tapered cusps, shortening toward the base. Outer marginal teeth with 2734 sharp cusps (
<figureCitation id="130F2A204C001B71FF11FCECFEE27325" box="[171,304,873,895]" captionStart="Figure 22" captionStartId="35.[163,243,1708,1730]" captionTargetBox="[163,1443,195,1669]" captionTargetId="figure-26@35.[163,1443,195,1669]" captionTargetPageId="35" captionText="Figure 22. Shells, operculum and radulae of Radomaniola variabilis sp. nov. A, B, holotype (MNCN 15.05/200169). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975)." figureDoi="http://doi.org/10.5281/zenodo.7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="37" pageNumber="430">Fig. 22M, N</figureCitation>
).
</paragraph>
<paragraph id="8B8B36A54C001B71FF01FC0DFD0774B4" blockId="37.[161,780,290,1907]" pageId="37" pageNumber="430">
Animal darkly pigmented except for neck and tentacles (
<figureCitation id="130F2A204C001B71FEA0FC23FE4E73E6" box="[282,412,934,956]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="37" pageNumber="430">Fig. 23G, H</figureCitation>
). Ctenidium with 12 or 13 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (
<figureCitation id="130F2A204C001B71FF11FBA4FED5746C" box="[171,263,1057,1079]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="37" pageNumber="430">Fig. 23A</figureCitation>
). Stomach as long as wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (
<figureCitation id="130F2A204C001B71FD1AFBDBFCD6742E" box="[672,772,1118,1140]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="37" pageNumber="430">Fig. 23B</figureCitation>
; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio= 0.47); cerebral ganglia approximately equal in size, presenting small black granules (
<figureCitation id="130F2A204C001B71FDD0FB5CFD1574B4" box="[618,711,1241,1263]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="37" pageNumber="430">Fig. 23C</figureCitation>
).
</paragraph>
<paragraph id="8B8B36A54C001B71FF01FB7DFDE7767B" blockId="37.[161,780,290,1907]" pageId="37" pageNumber="430">
Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland.
<collectingRegion id="49F0F8474C001B71FE2EFAB3FE097511" box="[404,475,1334,1355]" country="Turkey" name="Bursa" pageId="37" pageNumber="430">Bursa</collectingRegion>
copulatrix ovoid, slightly longer than wide. Bursal duct slightly longer than bursal length. Renal oviduct unpigmented, coiled. SR1 pyriform, duct short, joining renal oviduct slightly above the insertion point with bursal duct. SR2 slightly larger than SR1, pyriform, with a short duct, located on renal oviduct near loop (
<figureCitation id="130F2A204C001B71FD38FA68FCD77659" box="[642,773,1517,1539]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="37" pageNumber="430">Fig. 23D, E</figureCitation>
; Supporting Information, Table S8).
</paragraph>
<paragraph id="8B8B36A54C001B71FF01F9AFFED47729" blockId="37.[161,780,290,1907]" pageId="37" pageNumber="430">
Male genitalia with a prostate gland approximately two times longer than wide, bean shaped; seminal duct entering the middle-posterior region; pallial vas deferens emerging close to its anterior edge (
<figureCitation id="130F2A204C001B71FF16F920FEDD76E0" box="[172,271,1701,1723]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="37" pageNumber="430">Fig. 23F</figureCitation>
). Penis unpigmented, gradually tapering, approximately as long as head length, base narrow, weakly folded along inner edge and with one medial outgrowth on its left side (
<figureCitation id="130F2A204C001B71FE50F884FDA6774C" box="[490,628,1793,1815]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="37" pageNumber="430">Fig. 23GK</figureCitation>
; Supporting Information, Table S9); penis attached well behind the right eye; penial duct narrow, near outer edge, almost straight.
</paragraph>
<paragraph id="8B8B36A54C001B71FC81FF40FAAE710C" blockId="37.[827,1443,197,342]" pageId="37" pageNumber="430">
<emphasis id="B940EAB74C001B71FC81FF40FB177080" box="[827,1221,197,218]" italics="true" pageId="37" pageNumber="430">
Habitat:
<taxonomicName id="4C344D264C001B71FC0AFF40FB177080" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[944,1221,197,218]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="variabilis">Radomaniola variabilis</taxonomicName>
</emphasis>
lives in cold water springs and lacustrine habitats; it has been found on stones and aquatic vegetation. It co-occurs with
<taxonomicName id="4C344D264C001B71FC81FEA4FB11716C" authority="Muller, 1774" authorityName="Muller" authorityYear="1774" box="[827,1219,289,311]" class="Gastropoda" family="Planorbidae" genus="Ancylus" kingdom="Animalia" order="Siphonariida" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="fluviatilis">
<emphasis id="B940EAB74C001B71FC81FEA4FBCE716C" box="[827,1052,289,310]" italics="true" pageId="37" pageNumber="430">Ancylus fluviatilis</emphasis>
Müller, 1774
</taxonomicName>
,
<taxonomicName id="4C344D264C001B71FB68FEA4FC08710F" authority="(Muller, 1774)" baseAuthorityName="Muller" baseAuthorityYear="1774" class="Gastropoda" family="Lymnaeidae" genus="Radix" isUncertain="true" kingdom="Animalia" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="peregra">
<emphasis id="B940EAB74C001B71FB68FEA4FAC8716C" box="[1234,1306,289,310]" italics="true" pageId="37" pageNumber="430">Radix</emphasis>
cf.
<emphasis id="B940EAB74C001B71FAF2FEA7FA71716D" box="[1352,1443,290,311]" italics="true" pageId="37" pageNumber="430">peregra</emphasis>
(Müller, 1774)
</taxonomicName>
and
<taxonomicName id="4C344D264C001B71FBA8FEC5FB4E710F" box="[1042,1180,320,341]" class="Gastropoda" family="Bithyniidae" genus="Bithynia" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B940EAB74C001B71FBA8FEC5FBA4710F" box="[1042,1142,320,341]" italics="true" pageId="37" pageNumber="430">Bithynia</emphasis>
sp.
</taxonomicName>
in the
<typeStatus id="548F88074C001B71FB56FEC5FACF710F" box="[1260,1309,320,341]" pageId="37" pageNumber="430">type</typeStatus>
locality.
</paragraph>
<paragraph id="8B8B36A54C001B71FC81FEFBFAA7748A" blockId="37.[824,1444,382,1447]" pageId="37" pageNumber="430">
<emphasis id="B940EAB74C001B71FC81FEFBFC7771C9" box="[827,933,382,403]" italics="true" pageId="37" pageNumber="430">Remarks:</emphasis>
Populations of
<taxonomicName id="4C344D264C001B71FBE5FEFAFB3871C9" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[1119,1258,382,404]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="variabilis">
<emphasis id="B940EAB74C001B71FBE5FEFAFB3871C9" box="[1119,1258,382,404]" italics="true" pageId="37" pageNumber="430">R. variabilis</emphasis>
</taxonomicName>
are particularly variable in the number of cusps on the lateral radular teeth, intensity of body pigmentation, penis width (
<figureCitation id="130F2A204C001B71FCF9FE5FFC4871AA" box="[835,922,474,496]" captionStart="Figure 22" captionStartId="35.[163,243,1708,1730]" captionTargetBox="[163,1443,195,1669]" captionTargetId="figure-26@35.[163,1443,195,1669]" captionTargetPageId="35" captionText="Figure 22. Shells, operculum and radulae of Radomaniola variabilis sp. nov. A, B, holotype (MNCN 15.05/200169). C, paratype (UGSB 23975). D, shell (UGSB 14440). E, F, shell (UGSB 2294). GJ, N, paratypes (UGSB 23975). G, H, operculum (G, inner side; H, outer side). I, protoconch. J, portion of radula ribbon (UGSB 23975). K, portion of radula ribbon (UGSB 2294). L, central radular teeth (UGSB 14440). M, outer marginal teeth (UGSB 2294). N, outer marginal teeth (UGSB 23975)." figureDoi="http://doi.org/10.5281/zenodo.7035636" httpUri="https://zenodo.org/record/7035636/files/figure.png" pageId="37" pageNumber="430">Figs 22</figureCitation>
,
<figureCitation id="130F2A204C001B71FC12FE5EFC1671AA" box="[936,964,475,496]" captionStart="Figure 23" captionStartId="36.[145,223,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-161@36.[145,1425,195,1352]" captionTargetPageId="36" captionText="Figure 23. Anatomy of Radomaniola variabilis sp. nov.AG, I, paratypes (UGSB 23975). H, J, UGSB 14440. K, UGSB 2294. A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, prostate gland. G, H, head of male and penis. IK, penis." figureDoi="http://doi.org/10.5281/zenodo.7035642" httpUri="https://zenodo.org/record/7035642/files/figure.png" pageId="37" pageNumber="430">23</figureCitation>
) and shell morphology (e.g. shells of the Vella Spring population in
<collectingCountry id="F32376354C001B71FBCFFE7CFB167254" box="[1141,1220,505,526]" name="Greece" pageId="37" pageNumber="430">Greece</collectingCountry>
are slightly larger, more elongate than those of the Albanian populations and present a smaller aperture with a thin inner lip). Female genitalia could not be examined in detail because most of the dissected females from the
<typeStatus id="548F88074C001B71FACAFDF1FA7072D3" box="[1392,1442,628,649]" pageId="37" pageNumber="430">type</typeStatus>
locality were parasitized and had stunted organs. Also, we found no females among the few specimens collected at the localities in
<collectingCountry id="F32376354C001B71FBA2FD55FBA472BC" box="[1048,1142,720,742]" name="Albania" pageId="37" pageNumber="430">Albania</collectingCountry>
. Although the ABGD and GMYC methods subdivided this species into three entities, we consider this taxon as a single species based on the geographical proximity and phylogenetic clustering of the studied populations as a monophyletic group, a small intraspecific divergence &lt;1.6% for
<emphasis id="B940EAB74C001B71FACEFCEFFA717325" box="[1396,1443,874,895]" italics="true" pageId="37" pageNumber="430">COI</emphasis>
and similar shape of the shell and penis.
<bibRefCitation id="EFA54B544C001B71FAB8FC0DFC5773E6" author="Feher Z &amp; Eross ZP" pageId="37" pageNumber="430" pagination="3 - 21" refId="ref26787" refString="Feher Z, Eross ZP. 2009. Contribution to the Mollusca fauna of Albania. Results of the field trips of the Hungarian Natural History Museum between 1992 and 2007. Schriften zur Malakozoologie 25: 3 - 21." type="journal article" year="2009">Fehér &amp; Erőss (2009)</bibRefCitation>
assigned the three populations from
<collectingCountry id="F32376354C001B71FAFCFC23FA7073E6" box="[1350,1442,934,956]" name="Albania" pageId="37" pageNumber="430">Albania</collectingCountry>
studied herein to
<taxonomicName id="4C344D264C001B71FBB3FC43FBB67381" authorityName="SUBSP. MAXIMA" authorityYear="2022" box="[1033,1124,966,987]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="curta">
<emphasis id="B940EAB74C001B71FBB3FC43FBB67381" box="[1033,1124,966,987]" italics="true" pageId="37" pageNumber="430">R. curta</emphasis>
</taxonomicName>
based on shell morphology. However, penis features and genetic data suggest that these populations are members of
<taxonomicName id="4C344D264C001B71FB10FB86FAE1744D" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[1194,1331,1026,1048]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="variabilis">
<emphasis id="B940EAB74C001B71FB10FB86FAE1744D" box="[1194,1331,1026,1048]" italics="true" pageId="37" pageNumber="430">R. variabilis</emphasis>
</taxonomicName>
and not of
<taxonomicName id="4C344D264C001B71FC82FBA7FC46746D" authorityName="SUBSP. MAXIMA" authorityYear="2022" box="[824,916,1058,1079]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="curta">
<emphasis id="B940EAB74C001B71FC82FBA7FC46746D" box="[824,916,1058,1079]" italics="true" pageId="37" pageNumber="430">R. curta</emphasis>
</taxonomicName>
. The authors also reported the species
<taxonomicName id="4C344D264C001B71FAF3FBA7FA71746D" authorityName="SUBSP. MAXIMA" authorityYear="2022" box="[1353,1443,1058,1079]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="curta">
<emphasis id="B940EAB74C001B71FAF3FBA7FA71746D" box="[1353,1443,1058,1079]" italics="true" pageId="37" pageNumber="430">R. curta</emphasis>
</taxonomicName>
and
<taxonomicName id="4C344D264C001B71FCD1FBC4FC38740F" baseAuthorityName="Feher &amp; Eross" baseAuthorityYear="2009" box="[875,1002,1088,1110]" class="Gastropoda" family="Hydrobiidae" genus="Orientalia" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="albanica">
<emphasis id="B940EAB74C001B71FCD1FBC4FC38740F" box="[875,1002,1088,1110]" italics="true" pageId="37" pageNumber="430">R. albanica</emphasis>
</taxonomicName>
co-occurring in the Syri i Kaltër Spring (
<bibRefCitation id="EFA54B544C001B71FCF9FBDBFBEA742E" author="Feher Z &amp; Eross ZP" box="[835,1080,1118,1140]" pageId="37" pageNumber="430" pagination="3 - 21" refId="ref26787" refString="Feher Z, Eross ZP. 2009. Contribution to the Mollusca fauna of Albania. Results of the field trips of the Hungarian Natural History Museum between 1992 and 2007. Schriften zur Malakozoologie 25: 3 - 21." type="journal article" year="2009">Fehér &amp; Erőss, 2009</bibRefCitation>
). Based on morphological and molecular results, we conclude that the snails from the same locality examined in the present study are likely to be conspecific with
<taxonomicName id="4C344D264C001B71FB94FB3EFB657495" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[1070,1207,1210,1232]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="variabilis">
<emphasis id="B940EAB74C001B71FB94FB3EFB657495" box="[1070,1207,1210,1232]" italics="true" pageId="37" pageNumber="430">R. variabilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C344D264C001B71FB55FB3EFABC7495" baseAuthorityName="Feher &amp; Eross" baseAuthorityYear="2009" box="[1263,1390,1210,1232]" class="Gastropoda" family="Hydrobiidae" genus="Orientalia" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="albanica">
<emphasis id="B940EAB74C001B71FB55FB3EFABC7495" box="[1263,1390,1210,1232]" italics="true" pageId="37" pageNumber="430">R. albanica</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B8B36A54C001B71FCE9FB5FFA7675FC" blockId="37.[824,1444,382,1447]" pageId="37" pageNumber="430">
<taxonomicName id="4C344D264C001B71FCE9FB5FFB4574B5" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[851,1175,1241,1263]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="variabilis">
<emphasis id="B940EAB74C001B71FCE9FB5FFB4574B5" box="[851,1175,1241,1263]" italics="true" pageId="37" pageNumber="430">Radomaniola variabilis</emphasis>
</taxonomicName>
is morphologically distinguishable from the geographically and phylogenetically proximate species
<taxonomicName id="4C344D264C001B71FAA5FA92FA497571" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[1311,1435,1302,1324]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="dolens">
<emphasis id="B940EAB74C001B71FAA5FA92FA497571" box="[1311,1435,1302,1324]" italics="true" pageId="37" pageNumber="430">R. dolens</emphasis>
</taxonomicName>
,
<taxonomicName id="4C344D264C001B71FC81FAB3FC167510" baseAuthorityName="Feher &amp; Eross" baseAuthorityYear="2009" box="[827,964,1333,1355]" class="Gastropoda" family="Hydrobiidae" genus="Orientalia" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="albanica">
<emphasis id="B940EAB74C001B71FC81FAB3FC167510" box="[827,964,1333,1355]" italics="true" pageId="37" pageNumber="430">R. albanica</emphasis>
</taxonomicName>
and
<taxonomicName id="4C344D264C001B71FBB9FAB3FB477510" baseAuthorityName="Westerlund" baseAuthorityYear="1881" box="[1027,1173,1333,1355]" class="Gastropoda" family="Hydrobiidae" genus="Belgrandiella" kingdom="Animalia" order="Littorinimorpha" pageId="37" pageNumber="430" phylum="Mollusca" rank="species" species="haesitans">
<emphasis id="B940EAB74C001B71FBB9FAB3FB477510" box="[1027,1173,1333,1355]" italics="true" pageId="37" pageNumber="430">R. haesitans</emphasis>
</taxonomicName>
according to its wider shell, wider shell umbilicus, longer penis, shorter bursal duct and SR1 and sequence divergence from these three species (56% average divergence for
<emphasis id="B940EAB74C001B71FADFFA14FA4675FC" box="[1381,1428,1425,1446]" italics="true" pageId="37" pageNumber="430">COI</emphasis>
).
</paragraph>
</subSubSection>
</treatment>
</document>