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<document id="2E7E32C97830A6F7AF6F1DFFE72BA106" 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="039D87B34C311B42FF22FA67FDDA760F" 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 curta SUBSP. MAXIMA 2022, SUBSP. NOV." docType="treatment" docUuid="D86C5BA4-235B-410B-93B7-531D97B30A4D" docUuidSource="ZooBank" docVersion="9" lastPageNumber="414" 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="413" updateTime="1699373995385" updateUser="ExternalLinkService">
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<mods:title id="4E60855FE22C9EDEA4E8796D411FC198">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="31210535A7B5EC0F075576C451E1C548">Delicado, Diana</mods:namePart>
</mods:name>
<mods:name id="8D63E25F25CA42963B691C0570EB5B78" type="personal">
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<mods:namePart id="BAA98A44345BC65A5BB5755D230DC16D">Hauffe, Torsten</mods:namePart>
</mods:name>
<mods:typeOfResource id="10142977610B653CB3E1785E2AC2BC18">text</mods:typeOfResource>
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<mods:title id="D10A1207A7007068BC8C480BE76C5626">Zoological Journal of the Linnean Society</mods:title>
</mods:titleInfo>
<mods:part id="7FBA0B47DA909B9C86D06AB57821F601">
<mods:date id="55028E118C310D8124CD090953DDEF3B">2022</mods:date>
<mods:detail id="5C89B663278A756E150CD03E19A3942A" type="pubDate">
<mods:number id="7C6E9E8264C9C9E048D56F7DC8DA8904">2022-02-23</mods:number>
</mods:detail>
<mods:detail id="6D0BB983CAC62651AC6AACB034FEFCB1" type="volume">
<mods:number id="C42322B73CAE103686A595469FF27BD5">196</mods:number>
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<mods:start id="1D14011753F206CB782678B8BE344913">393</mods:start>
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<mods:location id="0844ADCFE01A3308E5E8C0539B262487">
<mods:url id="C40822A4AAA0C528255B9605A45503BF">https://doi.org/10.1093/zoolinnean/zlab121</mods:url>
</mods:location>
<mods:classification id="CA551A63E7A1640227CB2091424DBFE8">journal article</mods:classification>
<mods:identifier id="13B86C04764E86B443820F99BB70A646" type="CLB-Dataset">133178</mods:identifier>
<mods:identifier id="E7BCEB1450CD88EF347F10CAF8304D9A" type="DOI">10.1093/zoolinnean/zlab121</mods:identifier>
<mods:identifier id="0197AE02D8E904B10CFDF1CF9D382980" type="GBIF-Dataset">a2770ec2-41f4-4418-ba75-7ef5a99b88e4</mods:identifier>
<mods:identifier id="5AA10F187CB75897E0F1BC276BE1EFA3" type="ISSN">0024-4082</mods:identifier>
<mods:identifier id="3F8075F475068D23DF9E9C6AB5FA7DEC" type="Zenodo-Dep">7035584</mods:identifier>
<mods:identifier id="335918A507A5EEB1109E5BAD9E0A07CE" type="ZooBank">89452E3C-3AFE-449C-93C6-24EEE19AE45D</mods:identifier>
</mods:mods>
<treatment id="039D87B34C311B42FF22FA67FDDA760F" ID-DOI="http://doi.org/10.5281/zenodo.7036871" ID-GBIF-Taxon="199898766" ID-Zenodo-Dep="7036871" LSID="urn:lsid:zoobank.org:act:D86C5BA4-235B-410B-93B7-531D97B30A4D" httpUri="http://treatment.plazi.org/id/039D87B34C311B42FF22FA67FDDA760F" lastPageId="22" lastPageNumber="414" pageId="20" pageNumber="413">
<subSubSection id="C32E652E4C311B40FF22FA67FD2375A0" box="[152,753,1506,1530]" pageId="20" pageNumber="413" type="nomenclature">
<paragraph id="8B8B36A54C311B40FF22FA67FD2375A0" blockId="20.[152,753,1506,1530]" box="[152,753,1506,1530]" pageId="20" pageNumber="413">
<heading id="D0C381C94C311B40FF22FA67FD2375A0" box="[152,753,1506,1530]" centered="true" fontSize="9" level="2" pageId="20" pageNumber="413" reason="2">
<taxonomicName id="4C344D264C311B40FF22FA67FD8A75A3" ID-CoL="4R9KW" authority="SUBSP. MAXIMA" authorityName="SUBSP. MAXIMA" authorityYear="2022" box="[152,600,1506,1530]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="20" pageNumber="413" phylum="Mollusca" rank="species" species="curta" status="subsp. nov.">
<emphasis id="B940EAB74C311B40FF22FA67FE2775A0" bold="true" box="[152,501,1506,1530]" pageId="20" pageNumber="413">
<emphasis id="B940EAB74C311B40FF22FA67FE4E75A3" bold="true" box="[152,412,1506,1529]" italics="true" pageId="20" pageNumber="413">RADOMANIOLA CURTA</emphasis>
SUBSP.
</emphasis>
MAXIMA
</taxonomicName>
<emphasis id="B940EAB74C311B40FDDAFA60FD2375A0" bold="true" box="[608,753,1506,1530]" pageId="20" pageNumber="413">
<taxonomicNameLabel id="A27357CC4C311B40FDDAFA60FD2375A0" box="[608,753,1506,1530]" pageId="20" pageNumber="413" rank="subSpecies">SUBSP. NOV.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C32E652E4C311B42FEC1F98CFDDA760F" lastPageId="22" lastPageNumber="415" pageId="20" pageNumber="413" type="description">
<paragraph id="8B8B36A54C311B40FEC1F98CFDDC7678" blockId="20.[379,526,1545,1570]" box="[379,526,1545,1570]" pageId="20" pageNumber="413">
(
<figureCitation id="130F2A204C311B40FE3EF98CFE0E7678" box="[388,476,1545,1570]" captionStart="Figure 10" captionStartId="21.[162,240,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-266@21.[163,1443,195,1317]" captionTargetPageId="21" captionText="Figure 10. Shells, operculum and radulae of Radomaniola curta maxima subsp. nov.A, B, holotype (MNCN 15.05/200153). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="20" pageNumber="413">FIGS 10</figureCitation>
,
<figureCitation id="130F2A204C311B40FE50F98CFDD47678" box="[490,518,1545,1570]" captionStart="Figure 11" captionStartId="22.[145,227,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-26@22.[145,1425,195,1352]" captionTargetPageId="22" captionText="Figure 11. Anatomy of Radomaniola curta maxima subsp. nov. A–H, paratypes (UGSB 19540). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, head of male and penis. G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.7035607" httpUri="https://zenodo.org/record/7035607/files/figure.png" pageId="20" pageNumber="413">11</figureCitation>
)
</paragraph>
<paragraph id="8B8B36A54C311B40FF2BF9B7FD3C763C" blockId="20.[145,762,1585,1638]" pageId="20" pageNumber="413">
<emphasis id="B940EAB74C311B40FF2BF9B7FDEF761D" box="[145,573,1585,1607]" italics="true" pageId="20" pageNumber="413">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="FF920C704C311B40FF65F9D5FD3C763C" box="[223,750,1616,1638]" pageId="20" pageNumber="413">D86C5BA4-235B-410B-93B7-531D97B30A4D</uuid>
</paragraph>
<paragraph id="8B8B36A54C311B40FF2BF90BFE537699" blockId="20.[145,759,1678,1731]" pageId="20" pageNumber="413">
<emphasis id="B940EAB74C311B40FF2BF90BFEDD76F9" box="[145,271,1678,1699]" italics="true" pageId="20" pageNumber="413">Etymology:</emphasis>
From Latin
<emphasis id="B940EAB74C311B40FE19F90AFDDE76FE" box="[419,524,1679,1700]" italics="true" pageId="20" pageNumber="413">maximus</emphasis>
, very large, referring to its large body size.
</paragraph>
<paragraph id="8B8B36A54C311B40FF2BF967FEF2772B" blockId="20.[145,762,1761,1905]" pageId="20" pageNumber="413">
<emphasis id="B940EAB74C311B40FF2BF967FE9776AC" box="[145,325,1761,1783]" italics="true" pageId="20" pageNumber="413">Type material:</emphasis>
<materialsCitation id="3B5C3CF84C311B40FEE7F964FEF2772B" ID-GBIF-Occurrence="3897142309" collectionCode="MNCN, UGSB" country="Serbia" location="Crnojevica River" pageId="20" pageNumber="413" specimenCode="MNCN 15.05, UGSB 19540" specimenCount="520" typeStatus="holotype">
<typeStatus id="548F88074C311B40FEE7F964FE1976AD" box="[349,459,1761,1783]" pageId="20" pageNumber="413" type="holotype">Holotype</typeStatus>
(
<emphasis id="B940EAB74C311B40FE64F967FD3A76AC" box="[478,744,1761,1783]" italics="true" pageId="20" pageNumber="413">
<specimenCode id="DB929EDE4C311B40FE64F967FD5376AC" box="[478,641,1761,1783]" 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="20" pageNumber="413" type="Museum">MNCN 15.05</specimenCode>
/200153
</emphasis>
),
<specimenCount id="9D32FD2C4C311B40FF2BF885FEFD774F" box="[145,303,1792,1813]" pageId="20" pageNumber="413" type="generic" typeStatus="paratypes">20 paratypes</specimenCount>
(
<specimenCode id="DB929EDE4C311B40FEFAF885FE0D774C" box="[320,479,1792,1814]" 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="20" pageNumber="413" type="Museum">MNCN 15.05</specimenCode>
/200154) in the
<collectionCode id="ED25AE604C311B40FD1FF885FD2A774F" box="[677,760,1792,1813]" 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="20" pageNumber="413" type="Museum">MNCN</collectionCode>
collection and&gt;
<specimenCount id="9D32FD2C4C311B40FEEEF89BFE28776E" box="[340,506,1822,1844]" pageId="20" pageNumber="413" type="generic" typeStatus="paratypes">500 paratypes</specimenCount>
(
<specimenCode id="DB929EDE4C311B40FDB0F89AFD76776E" box="[522,676,1822,1844]" collectionCode="UGSB" pageId="20" pageNumber="413">UGSB 19540</specimenCode>
) in the
<collectionCode id="ED25AE604C311B40FF28F8B8FF0C7708" box="[146,222,1853,1874]" pageId="20" pageNumber="413">UGSB</collectionCode>
collection. Type locality:
<location id="8EEB607E4C311B40FD99F8B8FD217709" LSID="urn:lsid:plazi:treatment:039D87B34C311B42FF22FA67FDDA760F:8EEB607E4C311B40FD99F8B8FD217709" box="[547,755,1853,1875]" country="Serbia" name="Crnojevica River" pageId="20" pageNumber="413">Crnojevića River</location>
,
<collectingCountry id="F32376354C311B40FF2BF8D9FECF772B" box="[145,285,1884,1905]" name="Serbia" pageId="20" pageNumber="413">Montenegro</collectingCountry>
.
</materialsCitation>
</paragraph>
<paragraph id="8B8B36A54C311B40FC93FF43FBBA7162" blockId="20.[809,1424,198,312]" pageId="20" pageNumber="413">
<emphasis id="B940EAB74C311B40FC93FF43FC207081" box="[809,1010,198,219]" italics="true" pageId="20" pageNumber="413">Material studied:</emphasis>
<materialsCitation id="3B5C3CF84C311B40FBB2FF43FBB67162" ID-GBIF-Occurrence="3897142311" collectingDate="2009-09" collectionCode="T" collectorName="Diehl E" country="Serbia" latitude="42.355" location="Crnojevica River" longLatPrecision="6" longitude="19.0284" pageId="20" pageNumber="413" specimenCode="MNCN 15.05, UGSB 19540" specimenCount="1">
<location id="8EEB607E4C311B40FBB2FF43FB197086" LSID="urn:lsid:plazi:treatment:039D87B34C311B42FF22FA67FDDA760F:8EEB607E4C311B40FBB2FF43FB197086" box="[1032,1227,198,220]" country="Serbia" latitude="42.355" longLatPrecision="6" longitude="19.0284" name="Crnojevica River" pageId="20" pageNumber="413">Crnojevića River</location>
, near the bridge,
<collectingCountry id="F32376354C311B40FC90FF60FC6E70A0" box="[810,956,229,250]" name="Serbia" pageId="20" pageNumber="413">Montenegro</collectingCountry>
,
<geoCoordinate id="EE0050624C311B40FC70FF60FB9B70A1" box="[970,1097,229,251]" degrees="42.3550" direction="north" orientation="latitude" pageId="20" pageNumber="413" precision="5" value="42.355">42.3550°N</geoCoordinate>
,
<geoCoordinate id="EE0050624C311B40FBEDFF60FB0770A0" box="[1111,1237,229,250]" degrees="19.0284" direction="east" orientation="longitude" pageId="20" pageNumber="413" precision="5" value="19.0284">19.0284°E</geoCoordinate>
,
<collectionCode id="ED25AE604C311B40FB59FF60FB2570A0" box="[1251,1271,229,250]" name="Tavera, Department of Geology and Geophysics" pageId="20" pageNumber="413">T</collectionCode>
.H., C.W. and
<collectorName id="26C153734C311B40FC93FE81FCB27143" box="[809,864,260,281]" name="Diehl E" pageId="20" pageNumber="413">D.E.</collectorName>
,
<date id="FF8A10654C311B40FCC8FE81FB937143" box="[882,1089,260,282]" pageId="20" pageNumber="413" value="2009-09">
<collectingDate id="EFCEE98D4C311B40FCC8FE81FB937143" box="[882,1089,260,282]" pageId="20" pageNumber="413" value="2009-09">September 2009</collectingDate>
</date>
,
<specimenCode id="DB929EDE4C311B40FBEBFE81FB217143" box="[1105,1267,259,281]" 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="20" pageNumber="413" type="Museum">MNCN 15.05</specimenCode>
/200154 and
<specimenCode id="DB929EDE4C311B40FC93FEA7FC6F7162" box="[809,957,290,312]" collectionCode="UGSB" pageId="20" pageNumber="413">UGSB 19540</specimenCode>
(80% ethanol)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8B36A54C311B40FC93FEE7FA837275" blockId="20.[809,1426,353,559]" pageId="20" pageNumber="413">
<emphasis id="B940EAB74C311B40FC93FEE7FC73712D" box="[809,929,354,375]" italics="true" pageId="20" pageNumber="413">Diagnosis:</emphasis>
Protoconch microsculpture pitted; central radular tooth formula 5-C-5/1-1; bursa copulatrix ovoid, with a long duct; SR1 elongate, duct short; SR2 small, pyriform, duct short; penis unpigmented, gradually tapering, shorter than head length, sometimes with one medial outgrowth on its left side; nervous system unpigmented, elongate (mean RPG ratio = 0.52).
</paragraph>
<paragraph id="8B8B36A54C311B40FC93FDDDFBBF74CF" blockId="20.[807,1426,599,1909]" pageId="20" pageNumber="413">
<emphasis id="B940EAB74C311B40FC93FDDDFC607237" box="[809,946,600,621]" italics="true" pageId="20" pageNumber="413">Description:</emphasis>
Shell ovate-conic, 4.0–4.5 whorls, height
<quantity id="4CCC9B404C311B40FC93FDF3FC6A72D6" box="[809,952,630,652]" metricMagnitude="-3" metricUnit="m" metricValue="3.0" metricValueMax="3.5" metricValueMin="2.5" pageId="20" pageNumber="413" unit="mm" value="3.0" valueMax="3.5" valueMin="2.5">2.5–3.5 mm</quantity>
(
<figureCitation id="130F2A204C311B40FC71FDF3FB8672D1" box="[971,1108,630,652]" captionStart="Figure 10" captionStartId="21.[162,240,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-266@21.[163,1443,195,1317]" captionTargetPageId="21" captionText="Figure 10. Shells, operculum and radulae of Radomaniola curta maxima subsp. nov.A, B, holotype (MNCN 15.05/200153). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="20" pageNumber="413">Fig. 10A–D</figureCitation>
; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~400 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture pitted (
<figureCitation id="130F2A204C311B40FB86FD57FB7272BD" box="[1084,1184,722,744]" captionStart="Figure 10" captionStartId="21.[162,240,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-266@21.[163,1443,195,1317]" captionTargetPageId="21" captionText="Figure 10. Shells, operculum and radulae of Radomaniola curta maxima subsp. nov.A, B, holotype (MNCN 15.05/200153). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="20" pageNumber="413">Fig. 10G</figureCitation>
). Teleoconch whorls slightly convex, with deep sutures; body whorl large, occupying about three-quarters of total shell length. Aperture oval; inner lip thicker than outer lip; peristome margin simple, straight (
<figureCitation id="130F2A204C311B40FBF2FCC8FB7A7338" box="[1096,1192,845,867]" captionStart="Figure 10" captionStartId="21.[162,240,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-266@21.[163,1443,195,1317]" captionTargetPageId="21" captionText="Figure 10. Shells, operculum and radulae of Radomaniola curta maxima subsp. nov.A, B, holotype (MNCN 15.05/200153). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="20" pageNumber="413">Fig. 10B</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="130F2A204C311B40FBEAFC2CFB1F73E4" box="[1104,1229,937,959]" captionStart="Figure 10" captionStartId="21.[162,240,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-266@21.[163,1443,195,1317]" captionTargetPageId="21" captionText="Figure 10. Shells, operculum and radulae of Radomaniola curta maxima subsp. nov.A, B, holotype (MNCN 15.05/200153). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="20" pageNumber="413">Fig. 10E, F</figureCitation>
). Radular length intermediate, ~700 µm (~25% of total shell length), with ~50 rows of teeth (
<figureCitation id="130F2A204C311B40FC4FFC63FB8673A1" box="[1013,1108,998,1020]" captionStart="Figure 10" captionStartId="21.[162,240,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-266@21.[163,1443,195,1317]" captionTargetPageId="21" captionText="Figure 10. Shells, operculum and radulae of Radomaniola curta maxima subsp. nov.A, B, holotype (MNCN 15.05/200153). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="20" pageNumber="413">Fig. 10H</figureCitation>
). Central tooth formula 5-C- 5/1-1 (
<figureCitation id="130F2A204C311B40FCC9FB80FC1C7440" box="[883,974,1029,1051]" captionStart="Figure 10" captionStartId="21.[162,240,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-266@21.[163,1443,195,1317]" captionTargetPageId="21" captionText="Figure 10. Shells, operculum and radulae of Radomaniola curta maxima subsp. nov.A, B, holotype (MNCN 15.05/200153). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="20" pageNumber="413">Fig. 10I</figureCitation>
); basal tongue V-shaped, length about equal to lateral margin. Lateral tooth formula (4)3-C- 3(4). Inner marginal teeth bearing 18–21 tapered cusps, shortening toward the base. Outer marginal teeth with 22–23 sharp cusps (
<figureCitation id="130F2A204C311B40FBBCFB05FB8C74CF" box="[1030,1118,1152,1174]" captionStart="Figure 10" captionStartId="21.[162,240,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-266@21.[163,1443,195,1317]" captionTargetPageId="21" captionText="Figure 10. Shells, operculum and radulae of Radomaniola curta maxima subsp. nov.A, B, holotype (MNCN 15.05/200153). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth." figureDoi="http://doi.org/10.5281/zenodo.7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="20" pageNumber="413">Fig. 10J</figureCitation>
).
</paragraph>
<paragraph id="8B8B36A54C311B40FCFBFB1BFC00765F" blockId="20.[807,1426,599,1909]" pageId="20" pageNumber="413">
Animal darkly pigmented except for neck and tentacles (
<figureCitation id="130F2A204C311B40FC17FB38FBC77488" box="[941,1045,1213,1235]" captionStart="Figure 11" captionStartId="22.[145,227,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-26@22.[145,1425,195,1352]" captionTargetPageId="22" captionText="Figure 11. Anatomy of Radomaniola curta maxima subsp. nov. A–H, paratypes (UGSB 19540). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, head of male and penis. G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.7035607" httpUri="https://zenodo.org/record/7035607/files/figure.png" pageId="20" pageNumber="413">Fig. 11F</figureCitation>
). Ctenidium with 13–16 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (
<figureCitation id="130F2A204C311B40FC17FAB2FBDF7517" box="[941,1037,1335,1357]" captionStart="Figure 11" captionStartId="22.[145,227,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-26@22.[145,1425,195,1352]" captionTargetPageId="22" captionText="Figure 11. Anatomy of Radomaniola curta maxima subsp. nov. A–H, paratypes (UGSB 19540). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, head of male and penis. G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.7035607" httpUri="https://zenodo.org/record/7035607/files/figure.png" pageId="20" pageNumber="413">Fig. 11A</figureCitation>
). Stomach approximately as long as wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (
<figureCitation id="130F2A204C311B40FC26FA16FC2875F3" box="[924,1018,1427,1449]" captionStart="Figure 11" captionStartId="22.[145,227,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-26@22.[145,1425,195,1352]" captionTargetPageId="22" captionText="Figure 11. Anatomy of Radomaniola curta maxima subsp. nov. A–H, paratypes (UGSB 19540). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, head of male and penis. G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.7035607" httpUri="https://zenodo.org/record/7035607/files/figure.png" pageId="20" pageNumber="413">Fig. 11B</figureCitation>
; Supporting Information, Table S7). Nervous system unpigmented, elongate (mean RPG ratio = 0.52); cerebral ganglia approximately equal in size (
<figureCitation id="130F2A204C311B40FCDEFA75FC11765F" box="[868,963,1520,1542]" captionStart="Figure 11" captionStartId="22.[145,227,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-26@22.[145,1425,195,1352]" captionTargetPageId="22" captionText="Figure 11. Anatomy of Radomaniola curta maxima subsp. nov. A–H, paratypes (UGSB 19540). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, head of male and penis. G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.7035607" httpUri="https://zenodo.org/record/7035607/files/figure.png" pageId="20" pageNumber="413">Fig. 11C</figureCitation>
).
</paragraph>
<paragraph id="8B8B36A54C311B40FCFBF98BFA9B7743" blockId="20.[807,1426,599,1909]" pageId="20" pageNumber="413">
Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland.
<collectingRegion id="49F0F8474C311B40FCC2F9C9FC6C763B" box="[888,958,1612,1633]" country="Turkey" name="Bursa" pageId="20" pageNumber="413">Bursa</collectingRegion>
copulatrix ovoid, about as long as wide. Bursal duct longer than bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, with a short duct, joining renal oviduct slightly above the insertion point with bursal duct. SR2 shorter than SR1, pyriform, with a short duct, located on renal oviduct near loop (
<figureCitation id="130F2A204C311B40FC8BF881FC7D7743" box="[817,943,1796,1818]" captionStart="Figure 11" captionStartId="22.[145,227,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-26@22.[145,1425,195,1352]" captionTargetPageId="22" captionText="Figure 11. Anatomy of Radomaniola curta maxima subsp. nov. A–H, paratypes (UGSB 19540). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, head of male and penis. G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.7035607" httpUri="https://zenodo.org/record/7035607/files/figure.png" pageId="20" pageNumber="413">Fig. 11D, E</figureCitation>
; Supporting Information, Table S8).
</paragraph>
<paragraph id="8B8B36A54C311B41FCFBF8A7FD7E76CB" blockId="20.[807,1426,599,1909]" lastBlockId="21.[163,780,1476,1681]" lastPageId="21" lastPageNumber="414" pageId="20" pageNumber="413">
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="130F2A204C301B41FD26FA41FD2E7583" box="[668,764,1476,1498]" captionStart="Figure 11" captionStartId="22.[145,227,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-26@22.[145,1425,195,1352]" captionTargetPageId="22" captionText="Figure 11. Anatomy of Radomaniola curta maxima subsp. nov. A–H, paratypes (UGSB 19540). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, head of male and penis. G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.7035607" httpUri="https://zenodo.org/record/7035607/files/figure.png" pageId="21" pageNumber="414">Fig. 11H</figureCitation>
). Penis unpigmented, gradually tapering, shorter than head length, weakly folded along inner edge and with one medial outgrowth (sometimes absent) on its left side (
<figureCitation id="130F2A204C301B41FF58F9BBFEB37609" box="[226,353,1598,1620]" captionStart="Figure 11" captionStartId="22.[145,227,1391,1413]" captionTargetBox="[145,1425,195,1352]" captionTargetId="figure-26@22.[145,1425,195,1352]" captionTargetPageId="22" captionText="Figure 11. Anatomy of Radomaniola curta maxima subsp. nov. A–H, paratypes (UGSB 19540). A, ctenidium and osphradium. B, stomach. C, partial nervous system. D, pallial oviduct. E, bursa copulatrix and seminal receptacles. F, head of male and penis. G, penis. H, prostate gland." figureDoi="http://doi.org/10.5281/zenodo.7035607" httpUri="https://zenodo.org/record/7035607/files/figure.png" pageId="21" pageNumber="414">Fig. 11F, G</figureCitation>
; Supporting Information, Table S9); penis attached to the area close to the right eye; penial duct narrow, near outer edge, almost straight.
</paragraph>
<caption id="DF4B662D4C301B41FF18FAC9FD3475C7" ID-DOI="http://doi.org/10.5281/zenodo.7035605" ID-Zenodo-Dep="7035605" httpUri="https://zenodo.org/record/7035605/files/figure.png" pageId="21" pageNumber="414" startId="21.[162,240,1356,1378]" targetBox="[163,1443,195,1317]" targetPageId="21">
<paragraph id="8B8B36A54C301B41FF18FAC9FD3475C7" blockId="21.[162,1443,1356,1437]" pageId="21" pageNumber="414">
<emphasis id="B940EAB74C301B41FF18FAC9FEC47538" bold="true" box="[162,278,1356,1378]" pageId="21" pageNumber="414">Figure 10.</emphasis>
Shells, operculum and radulae of
<taxonomicName id="4C344D264C301B41FDC1FAC8FC4D7539" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[635,927,1357,1379]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="414" phylum="Mollusca" rank="subSpecies" species="curta" status="subsp. nov." subSpecies="maxima">
<emphasis id="B940EAB74C301B41FDC1FAC8FC4D7539" box="[635,927,1357,1379]" italics="true" pageId="21" pageNumber="414">Radomaniola curta maxima</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27357CC4C301B41FC1FFAC8FBC77539" box="[933,1045,1357,1379]" pageId="21" pageNumber="414" rank="subSpecies">subsp. nov.</taxonomicNameLabel>
A, B, holotype (
<emphasis id="B940EAB74C301B41FB0EFAC8FA447539" box="[1204,1430,1357,1379]" italics="true" pageId="21" pageNumber="414">MNCN 15.05/200153</emphasis>
). C–J, paratypes (UGSB 19540). C, D, shells. E, F, operculum (E, inner side; F, outer side). G, protoconch. H, portion of radula ribbon. I, central radular teeth. J, outer marginal teeth.
</paragraph>
</caption>
<paragraph id="8B8B36A54C301B41FF19F932FDF8773D" blockId="21.[163,780,1719,1895]" pageId="21" pageNumber="414">
<emphasis id="B940EAB74C301B41FF19F932FED17696" box="[163,259,1719,1740]" italics="true" pageId="21" pageNumber="414">Habitat:</emphasis>
This is a riverine subspecies collected from the margins of the Crnojevića River when the water level was low. Snails were found in high abundance on stones and small rocks. Co-occurring species are
<taxonomicName id="4C344D264C301B41FF6FF8B7FEB47712" box="[213,358,1842,1864]" class="Gastropoda" family="Planorbidae" genus="Ancylus" kingdom="Animalia" order="Siphonariida" pageId="21" pageNumber="414" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B940EAB74C301B41FF6FF8B7FEE4771D" box="[213,310,1842,1863]" italics="true" pageId="21" pageNumber="414">Ancylus</emphasis>
sp.
</taxonomicName>
,
<taxonomicName id="4C344D264C301B41FECDF8B7FDFA7712" box="[375,552,1842,1864]" class="Gastropoda" family="Neritidae" genus="Theodoxus" kingdom="Animalia" order="Cycloneritida" pageId="21" pageNumber="414" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B940EAB74C301B41FECDF8B7FE2B771D" box="[375,505,1842,1863]" italics="true" pageId="21" pageNumber="414">Theodoxus</emphasis>
sp.
</taxonomicName>
,
<taxonomicName id="4C344D264C301B41FD83F8B7FD007712" box="[569,722,1842,1864]" class="Gastropoda" family="Bithyniidae" genus="Bithynia" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="414" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B940EAB74C301B41FD83F8B7FD77771D" box="[569,677,1842,1863]" italics="true" pageId="21" pageNumber="414">Bithynia</emphasis>
sp.
</taxonomicName>
and
<taxonomicName id="4C344D264C301B41FF19F8D4FDF7773D" authority="(Bole, 1961)" baseAuthorityName="Bole" baseAuthorityYear="1961" box="[163,549,1873,1895]" class="Gastropoda" family="Hydrobiidae" genus="Anagastina" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="414" phylum="Mollusca" rank="species" species="matjasici">
<emphasis id="B940EAB74C301B41FF19F8D4FE44773C" box="[163,406,1873,1894]" italics="true" pageId="21" pageNumber="414">Anagastina matjasici</emphasis>
(Bole, 1961)
</taxonomicName>
.
</paragraph>
<paragraph id="8B8B36A54C301B42FC81FA46FDDA760F" blockId="21.[827,1443,1475,1896]" lastBlockId="22.[145,761,1508,1622]" lastPageId="22" lastPageNumber="415" pageId="21" pageNumber="414">
<emphasis id="B940EAB74C301B41FC81FA46FC787582" box="[827,938,1475,1496]" italics="true" pageId="21" pageNumber="414">Remarks:</emphasis>
Although specimens of
<emphasis id="B940EAB74C301B41FB60FA41FAA67583" box="[1242,1396,1476,1497]" italics="true" pageId="21" pageNumber="414">R. c. maxima</emphasis>
are larger than those of other conspecific populations and present a more ovate shell shape, they are included in
<taxonomicName id="4C344D264C301B41FCE1F9A5FC6E766F" authorityName="SUBSP. MAXIMA" authorityYear="2022" box="[859,956,1568,1589]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="414" phylum="Mollusca" rank="species" species="curta">
<emphasis id="B940EAB74C301B41FCE1F9A5FC6E766F" box="[859,956,1568,1589]" italics="true" pageId="21" pageNumber="414">R. curta</emphasis>
</taxonomicName>
by the presence of a long SR1, an ovoid bursa copulatrix with a long duct, a gradually tapering penis with a medial outgrowth on the left edge and an RPG ratio from moderately concentrated to elongate. All these features have been described for
<emphasis id="B940EAB74C301B41FA94F91EFA7176EA" box="[1326,1443,1691,1712]" italics="true" pageId="21" pageNumber="414">
R. c.
<taxonomicName id="4C344D264C301B41FADCF91EFA7176EA" authorityName="SUBSP. MAXIMA" authorityYear="2022" box="[1382,1443,1691,1712]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="414" phylum="Mollusca" rank="species" species="curta">curta</taxonomicName>
</emphasis>
by
<bibRefCitation id="EFA54B544C301B41FCDBF93CFBFB7694" author="Radoman P" box="[865,1065,1721,1743]" pageId="21" pageNumber="414" pagination="1 - 256" refId="ref28653" refString="Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Srpska Akademija Nauka i Umetnosti, Posebna Izdanja (Odeljenje PrirodnoMatematitschkich Nauka) 57: i - ii + 1 - 256." type="journal article" year="1983">Radoman (1983)</bibRefCitation>
. In this monograph,
<bibRefCitation id="EFA54B544C301B41FA8AF93CFC5176B7" author="Radoman P" pageId="21" pageNumber="414" pagination="1 - 256" refId="ref28653" refString="Radoman P. 1983. Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Srpska Akademija Nauka i Umetnosti, Posebna Izdanja (Odeljenje PrirodnoMatematitschkich Nauka) 57: i - ii + 1 - 256." type="journal article" year="1983">Radoman (1983)</bibRefCitation>
stated that
<taxonomicName id="4C344D264C301B41FBAAF952FB4476B6" box="[1040,1174,1751,1772]" class="Gastropoda" family="Hydrobiidae" genus="Orientalina" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="414" phylum="Mollusca" rank="genus">
<emphasis id="B940EAB74C301B41FBAAF952FB4476B6" box="[1040,1174,1751,1772]" italics="true" pageId="21" pageNumber="414">Orientalina</emphasis>
</taxonomicName>
(=
<taxonomicName id="4C344D264C301B41FB06F95DFA8476B7" authorityName="Szarowska" authorityYear="2006" box="[1212,1366,1752,1773]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="414" phylum="Mollusca" rank="genus">
<emphasis id="B940EAB74C301B41FB06F95DFA8476B7" box="[1212,1366,1752,1773]" italics="true" pageId="21" pageNumber="414">Radomaniola</emphasis>
</taxonomicName>
)
<taxonomicName id="4C344D264C301B41FADCF95DFA7076B7" authorityName="SUBSP. MAXIMA" authorityYear="2022" box="[1382,1442,1752,1773]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="21" pageNumber="414" phylum="Mollusca" rank="species" species="curta">
<emphasis id="B940EAB74C301B41FADCF95DFA7076B7" box="[1382,1442,1752,1773]" italics="true" pageId="21" pageNumber="414">curta</emphasis>
</taxonomicName>
subspecies differ from each other mainly in shell rather than in anatomical features, which applies to
<emphasis id="B940EAB74C301B41FC86F8B1FC1C7713" box="[828,974,1844,1865]" italics="true" pageId="21" pageNumber="414">R. c. maxima</emphasis>
. Other than by shell shape and size, this subspecies is diagnosable by its small number of cusps on the radular teeth, a high RPG ratio and a wide base of the penis. Our phylogenetic inferences also place the studied specimens of
<emphasis id="B940EAB74C331B42FE0AF9A7FD91766D" box="[432,579,1570,1591]" italics="true" pageId="22" pageNumber="415">R. c. maxima</emphasis>
unambiguously within the
<taxonomicName id="4C344D264C331B42FEB4F9C4FEBB760C" authorityName="SUBSP. MAXIMA" authorityYear="2022" box="[270,361,1601,1622]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="22" pageNumber="415" phylum="Mollusca" rank="species" species="curta">
<emphasis id="B940EAB74C331B42FEB4F9C4FEBB760C" box="[270,361,1601,1622]" italics="true" pageId="22" pageNumber="415">R. curta</emphasis>
</taxonomicName>
clade (
<figureCitation id="130F2A204C331B42FE03F9C5FE2B760F" box="[441,505,1600,1622]" captionStart="Figure 2" captionStartId="7.[163,242,1198,1220]" captionTargetBox="[165,1434,223,1142]" captionTargetId="figure-295@7.[163,1443,195,1158]" captionTargetPageId="7" captionText="Figure 2. Majority-rule consensus tree from a Bayesian analysis based on the multilocus dataset (COI, 16S and 28S) of Radomaniola. Branch support values (BPP, Bayesian posterior probability; BS, bootstrap support) are provided on nodes. Tip labels correspond to the sequence codes given in the Supporting Information (Table S1). Vertical bars on the right represent entities identified by the molecular species delimitation methods automatic barcode gap discovery (ABGD), Bayesian generalized mixed Yule-coalescent method (bGMYC) and Poisson tree processes species delimitation method in a Bayesian framework (bPTP) and the multi-rate version (mPTP) with Bayesian support values. Scale bar below topology: substitutions per site." figureDoi="http://doi.org/10.5281/zenodo.7035589" httpUri="https://zenodo.org/record/7035589/files/figure.png" pageId="22" pageNumber="415">Fig. 2</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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