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<document id="E61E0CB9C65FD652417B5C2F9D69518C" 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="039D87B34C051B76FF5DF96DFAD475A5" 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 szarowskae Delicado &amp; Hauffe 2022, SP. NOV." docType="treatment" docVersion="9" lastPageNumber="427" 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="425" updateTime="1699373995385" updateUser="ExternalLinkService">
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<mods:title id="9155296FEFDEF128C8314E0725E3FDDC">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="E38CA883DE34A7D20E5B5F4A3243F23D">Delicado, Diana</mods:namePart>
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<mods:namePart id="2432998342B24F1E2799034F3E44290A">Hauffe, Torsten</mods:namePart>
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<mods:title id="4FEDBAD515F2BFE83A949DE3DFEFD954">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="A604E0BA78F72B07FE32424988C5D913">2022</mods:date>
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<treatment id="039D87B34C051B76FF5DF96DFAD475A5" ID-DOI="http://doi.org/10.5281/zenodo.7036889" ID-GBIF-Taxon="199898770" ID-Zenodo-Dep="7036889" LSID="urn:lsid:plazi:treatment:039D87B34C051B76FF5DF96DFAD475A5" httpUri="http://treatment.plazi.org/id/039D87B34C051B76FF5DF96DFAD475A5" lastPageId="34" lastPageNumber="427" pageId="32" pageNumber="425">
<subSubSection id="C32E652E4C051B74FF5DF96DFD7176A5" box="[231,675,1767,1791]" pageId="32" pageNumber="425" type="nomenclature">
<paragraph id="8B8B36A54C051B74FF5DF96DFD7176A5" blockId="32.[231,675,1767,1791]" box="[231,675,1767,1791]" pageId="32" pageNumber="425">
<heading id="D0C381C94C051B74FF5DF96DFD7176A5" box="[231,675,1767,1791]" centered="true" fontSize="9" level="2" pageId="32" pageNumber="425" reason="2">
<taxonomicName id="4C344D264C051B74FF5DF96DFDEB76A4" ID-CoL="9CC4K" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[231,569,1768,1791]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="32" pageNumber="425" phylum="Mollusca" rank="species" species="szarowskae" status="sp. nov.">RADOMANIOLA SZAROWSKAE</taxonomicName>
<emphasis id="B940EAB74C051B74FDFAF96EFD7176A5" bold="true" box="[576,675,1767,1791]" pageId="32" pageNumber="425">
<taxonomicNameLabel id="A27357CC4C051B74FDFAF96EFD7176A5" box="[576,675,1767,1791]" pageId="32" pageNumber="425" rank="species">SP. NOV.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C32E652E4C051B76FEC1F88BFAD475A5" lastPageId="34" lastPageNumber="427" pageId="32" pageNumber="425" type="description">
<paragraph id="8B8B36A54C051B74FEC1F88BFDDC777D" blockId="32.[379,526,1806,1831]" box="[379,526,1806,1831]" pageId="32" pageNumber="425">
(
<figureCitation id="130F2A204C051B74FE3EF88BFE0E777D" box="[388,476,1806,1831]" captionStart="Figure 20" captionStartId="33.[163,241,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-258@33.[163,1443,195,1317]" captionTargetPageId="33" captionText="Figure 20. Shells, operculum and radulae of Radomaniola szarowskae sp. nov. A, B, holotype (MNCN 15.05/200167). CJ, paratypes (UGSB 18556). 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.7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="32" pageNumber="425">FIGS 20</figureCitation>
,
<figureCitation id="130F2A204C051B74FE50F88BFDD4777D" box="[490,518,1806,1831]" captionStart="Figure 21" captionStartId="34.[145,224,1314,1336]" captionTargetBox="[145,1425,195,1275]" captionTargetId="figure-246@34.[145,1425,195,1275]" captionTargetPageId="34" captionText="Figure 21. Anatomy of Radomaniola szarowskae sp. nov. AH, paratypes (UGSB 18566). 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.7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="32" pageNumber="425">21</figureCitation>
)
</paragraph>
<paragraph id="8B8B36A54C051B74FF2BF8B2FD0F7731" blockId="32.[145,762,1846,1899]" pageId="32" pageNumber="425">
<emphasis id="B940EAB74C051B74FF2BF8B2FDEF7716" box="[145,573,1846,1868]" italics="true" pageId="32" pageNumber="425">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="FF920C704C051B74FF64F8D0FD0F7731" box="[222,733,1877,1899]" pageId="32" pageNumber="425">605481A3-A28F-4DC0-B532-9A1534B21887</uuid>
</paragraph>
<paragraph id="8B8B36A54C051B74FC93FF40FADA716D" blockId="32.[809,1426,197,311]" pageId="32" pageNumber="425">
<emphasis id="B940EAB74C051B74FC93FF40FC787080" box="[809,938,197,218]" italics="true" pageId="32" pageNumber="425">Etymology:</emphasis>
Named after Dr Magdalena Szarowska, in recognition of her many contributions to the evolution of Truncatelloidean gastropods of the Balkan Peninsula, including species of this genus.
</paragraph>
<paragraph id="8B8B36A54C051B74FC93FEE4FBFA71AA" blockId="32.[809,1426,352,496]" pageId="32" pageNumber="425">
<emphasis id="B940EAB74C051B74FC93FEE4FC0F712F" box="[809,989,352,374]" italics="true" pageId="32" pageNumber="425">Type material:</emphasis>
<materialsCitation id="3B5C3CF84C051B74FC4FFEE5FBFA71AA" ID-GBIF-Occurrence="3897142315" collectionCode="MNCN, UGSB" country="Croatia" location="Cetina River" municipality="Cetina" pageId="32" pageNumber="425" specimenCode="MNCN 15.05, UGSB 18566" specimenCount="61" typeStatus="holotype">
<typeStatus id="548F88074C051B74FC4FFEE5FBB1712C" box="[1013,1123,352,374]" pageId="32" pageNumber="425" type="holotype">Holotype</typeStatus>
(
<emphasis id="B940EAB74C051B74FBCCFEE5FA52712F" box="[1142,1408,352,373]" italics="true" pageId="32" pageNumber="425">
<specimenCode id="DB929EDE4C051B74FBCCFEE5FACA712F" box="[1142,1304,352,373]" 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="32" pageNumber="425" type="Museum">MNCN 15.05</specimenCode>
/200167
</emphasis>
),
<specimenCount id="9D32FD2C4C051B74FC93FEFAFC1C71CE" box="[809,974,383,404]" pageId="32" pageNumber="425" type="generic" typeStatus="paratypes">ten paratypes</specimenCount>
(
<specimenCode id="DB929EDE4C051B74FC65FEFAFBA971CE" box="[991,1147,382,404]" 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="32" pageNumber="425" type="Museum">MNCN 15.05</specimenCode>
/200168) in the
<collectionCode id="ED25AE604C051B74FA87FEFAFA4271CE" box="[1341,1424,383,404]" 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="32" pageNumber="425" type="Museum">MNCN</collectionCode>
collection and
<specimenCount id="9D32FD2C4C051B74FC5BFE18FB5071E9" box="[993,1154,413,435]" pageId="32" pageNumber="425" type="generic" typeStatus="paratypes">60 paratypes</specimenCount>
(
<specimenCode id="DB929EDE4C051B74FB2EFE18FAE471E9" box="[1172,1334,413,435]" collectionCode="UGSB" pageId="32" pageNumber="425">UGSB 18566</specimenCode>
) in the
<collectionCode id="ED25AE604C051B74FC93FE39FCA1718B" box="[809,883,444,465]" pageId="32" pageNumber="425">UGSB</collectionCode>
collection. Type locality: source of the
<location id="8EEB607E4C051B74FAF8FE39FCB571AA" LSID="urn:lsid:plazi:treatment:039D87B34C051B76FF5DF96DFAD475A5:8EEB607E4C051B74FAF8FE39FCB571AA" country="Croatia" municipality="Cetina" name="Cetina River" pageId="32" pageNumber="425">Cetina River</location>
,
<collectingMunicipality id="6BEFACDF4C051B74FCCBFE5FFC6D71AA" box="[881,959,474,496]" pageId="32" pageNumber="425">Cetina</collectingMunicipality>
,
<collectingCountry id="F32376354C051B74FC70FE5FFBF171AA" box="[970,1059,474,496]" name="Croatia" pageId="32" pageNumber="425">Croatia</collectingCountry>
.
</materialsCitation>
</paragraph>
<paragraph id="8B8B36A54C051B74FC93FDA4FAAF722E" blockId="32.[807,1425,545,628]" pageId="32" pageNumber="425">
<emphasis id="B940EAB74C051B74FC93FDA4FC3C726C" box="[809,1006,545,566]" italics="true" pageId="32" pageNumber="425">Material studied:</emphasis>
<materialsCitation id="3B5C3CF84C051B74FBB9FDA4FAAB722E" ID-GBIF-Occurrence="3897142303" collectingDate="2016-06" collectionCode="T" collectorName="Diana Delicado" country="Croatia" county="Cetina" latitude="43.9766" location="Source" longLatPrecision="6" longitude="16.4301" municipality="Cetina River" pageId="32" pageNumber="425" specimenCode="MNCN 15.05, UGSB 18566" specimenCount="1">
<location id="8EEB607E4C051B74FBB9FDA4FB82726C" LSID="urn:lsid:plazi:treatment:039D87B34C051B76FF5DF96DFAD475A5:8EEB607E4C051B74FBB9FDA4FB82726C" box="[1027,1104,545,566]" country="Croatia" county="Cetina" latitude="43.9766" longLatPrecision="6" longitude="16.4301" municipality="Cetina River" name="Source" pageId="32" pageNumber="425">Source</location>
of the
<collectingMunicipality id="6BEFACDF4C051B74FB1BFDA4FAE3726D" box="[1185,1329,545,567]" pageId="32" pageNumber="425">Cetina River</collectingMunicipality>
,
<collectingCounty id="62EA4E294C051B74FA84FDA4FA5E726D" box="[1342,1420,545,567]" pageId="32" pageNumber="425">Cetina</collectingCounty>
,
<collectingCountry id="F32376354C051B74FC93FDC5FCAC720C" box="[809,894,576,598]" name="Croatia" pageId="32" pageNumber="425">Croatia</collectingCountry>
,
<geoCoordinate id="EE0050624C051B74FC3DFDC5FC29720C" box="[903,1019,576,598]" degrees="43.9766" direction="north" orientation="latitude" pageId="32" pageNumber="425" precision="5" value="43.9766">43.9766°N</geoCoordinate>
,
<geoCoordinate id="EE0050624C051B74FBBEFDC5FBA5720C" box="[1028,1143,576,598]" degrees="16.4301" direction="east" orientation="longitude" pageId="32" pageNumber="425" precision="5" value="16.4301">16.4301°E</geoCoordinate>
,
<collectionCode id="ED25AE604C051B74FBC5FDC5FB43720F" box="[1151,1169,576,597]" name="Tavera, Department of Geology and Geophysics" pageId="32" pageNumber="425">T</collectionCode>
.H. and
<collectorName id="26C153734C051B74FB65FDC5FAC0720F" box="[1247,1298,576,597]" name="Diana Delicado" pageId="32" pageNumber="425">D.D.</collectorName>
,
<date id="FF8A10654C051B74FAA0FDC5FA5F720C" box="[1306,1421,576,598]" pageId="32" pageNumber="425" value="2016-06">
<collectingDate id="EFCEE98D4C051B74FAA0FDC5FA5F720C" box="[1306,1421,576,598]" pageId="32" pageNumber="425" value="2016-06">June 2016</collectingDate>
</date>
,
<specimenCode id="DB929EDE4C051B74FC9DFDDAFC65722E" box="[807,951,606,628]" 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="32" pageNumber="425" type="Museum">MNCN 15.05</specimenCode>
/200168 and
<specimenCode id="DB929EDE4C051B74FBFCFDDAFB05722E" box="[1094,1239,606,628]" collectionCode="UGSB" pageId="32" pageNumber="425">UGSB 18566</specimenCode>
(80% ethanol)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8B36A54C051B74FC93FD23FB9473EA" blockId="32.[809,1426,677,944]" pageId="32" pageNumber="425">
<emphasis id="B940EAB74C051B74FC93FD23FC7472E1" box="[809,934,678,699]" italics="true" pageId="32" pageNumber="425">Diagnosis:</emphasis>
Shell ovate-conic, 3.54.0 whorls (apex often eroded); protoconch microsculpture wrinkled; central radular tooth formula 6-C-6/1-1; bursa copulatrix ovoid, with a duct as long as bursal length; SR1 elongate, duct short; SR2 elongate, same length as SR1, duct short; penis unpigmented, gradually tapering, base wide, shorter than head length; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.47).
</paragraph>
<paragraph id="8B8B36A54C051B74FC93FC5FFAA77559" blockId="32.[808,1426,985,1896]" pageId="32" pageNumber="425">
<emphasis id="B940EAB74C051B74FC93FC5FFC6073B5" box="[809,946,986,1007]" italics="true" pageId="32" pageNumber="425">Description:</emphasis>
Shell ovate-conic, 3.54.0 whorls, height 2.33.0 mm, apex often eroded (
<figureCitation id="130F2A204C051B74FB30FC7DFAD57457" box="[1162,1287,1016,1038]" captionStart="Figure 20" captionStartId="33.[163,241,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-258@33.[163,1443,195,1317]" captionTargetPageId="33" captionText="Figure 20. Shells, operculum and radulae of Radomaniola szarowskae sp. nov. A, B, holotype (MNCN 15.05/200167). CJ, paratypes (UGSB 18556). 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.7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="32" pageNumber="425">Fig. 20AD</figureCitation>
; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~500 µm wide, 1.5 whorls; nucleus ~150 µm wide; protoconch microsculpture pitted (
<figureCitation id="130F2A204C051B74FAA0FBD1FA537433" box="[1306,1409,1108,1130]" captionStart="Figure 20" captionStartId="33.[163,241,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-258@33.[163,1443,195,1317]" captionTargetPageId="33" captionText="Figure 20. Shells, operculum and radulae of Radomaniola szarowskae sp. nov. A, B, holotype (MNCN 15.05/200167). CJ, paratypes (UGSB 18556). 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.7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="32" pageNumber="425">Fig. 20G</figureCitation>
). Teleoconch whorls convex, with deep sutures; body whorl large, occupying about three-quarters of total shell length. Aperture slightly oval; inner lip thicker than outer lip; peristome margin simple, straight (
<figureCitation id="130F2A204C051B74FADCFB4BFC877558" captionStart="Figure 20" captionStartId="33.[163,241,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-258@33.[163,1443,195,1317]" captionTargetPageId="33" captionText="Figure 20. Shells, operculum and radulae of Radomaniola szarowskae sp. nov. A, B, holotype (MNCN 15.05/200167). CJ, paratypes (UGSB 18556). 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.7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="32" pageNumber="425">Fig. 20B</figureCitation>
). Umbilicus narrow, not covered by the inner lip.
</paragraph>
<paragraph id="8B8B36A54C051B74FCFBFA89FC6E7504" blockId="32.[808,1426,985,1896]" pageId="32" pageNumber="425">
Operculum oval, yellowish, about two whorls; muscle attachment area oval and located near the nucleus (
<figureCitation id="130F2A204C051B74FC8BFACCFC7E7505" box="[817,940,1353,1375]" captionStart="Figure 20" captionStartId="33.[163,241,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-258@33.[163,1443,195,1317]" captionTargetPageId="33" captionText="Figure 20. Shells, operculum and radulae of Radomaniola szarowskae sp. nov. A, B, holotype (MNCN 15.05/200167). CJ, paratypes (UGSB 18556). 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.7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="32" pageNumber="425">Fig. 20E, F</figureCitation>
).
</paragraph>
<paragraph id="8B8B36A54C051B74FCFBFAEDFA81766F" blockId="32.[808,1426,985,1896]" pageId="32" pageNumber="425">
Radular length intermediate, ~700 µm (~25% of total shell length), with ~50 rows of teeth (
<figureCitation id="130F2A204C051B74FAA4FA03FA5075C6" box="[1310,1410,1414,1436]" captionStart="Figure 20" captionStartId="33.[163,241,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-258@33.[163,1443,195,1317]" captionTargetPageId="33" captionText="Figure 20. Shells, operculum and radulae of Radomaniola szarowskae sp. nov. A, B, holotype (MNCN 15.05/200167). CJ, paratypes (UGSB 18556). 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.7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="32" pageNumber="425">Fig. 20H</figureCitation>
). Central tooth formula 6-C-6/1-1 (
<figureCitation id="130F2A204C051B74FB23FA20FB3C75E0" box="[1177,1262,1445,1467]" captionStart="Figure 20" captionStartId="33.[163,241,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-258@33.[163,1443,195,1317]" captionTargetPageId="33" captionText="Figure 20. Shells, operculum and radulae of Radomaniola szarowskae sp. nov. A, B, holotype (MNCN 15.05/200167). CJ, paratypes (UGSB 18556). 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.7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="32" pageNumber="425">Fig. 20I</figureCitation>
); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula 4-C-4. Inner marginal teeth having 2025 tapered cusps, shortening toward the base. Outer marginal teeth with 2024 sharp cusps (
<figureCitation id="130F2A204C051B74FB56F9A5FA97766F" box="[1260,1349,1568,1590]" captionStart="Figure 20" captionStartId="33.[163,241,1356,1378]" captionTargetBox="[163,1443,195,1317]" captionTargetId="figure-258@33.[163,1443,195,1317]" captionTargetPageId="33" captionText="Figure 20. Shells, operculum and radulae of Radomaniola szarowskae sp. nov. A, B, holotype (MNCN 15.05/200167). CJ, paratypes (UGSB 18556). 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.7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="32" pageNumber="425">Fig. 20J</figureCitation>
).
</paragraph>
<paragraph id="8B8B36A54C051B75FCFBF9BBFE15764C" blockId="32.[808,1426,985,1896]" lastBlockId="33.[163,780,1475,1896]" lastPageId="33" lastPageNumber="426" pageId="32" pageNumber="425">
Animal darkly pigmented except for neck and tentacles (
<figureCitation id="130F2A204C051B74FC17F9D8FBC77628" box="[941,1045,1629,1651]" captionStart="Figure 21" captionStartId="34.[145,224,1314,1336]" captionTargetBox="[145,1425,195,1275]" captionTargetId="figure-246@34.[145,1425,195,1275]" captionTargetPageId="34" captionText="Figure 21. Anatomy of Radomaniola szarowskae sp. nov. AH, paratypes (UGSB 18566). 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.7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="32" pageNumber="425">Fig. 21F</figureCitation>
). Ctenidium with 1315 welldeveloped gill filaments, occupying ~50% of pallial cavity length and posteriorly positioned. Osphradium of intermediate width and opposite middle of ctenidium (
<figureCitation id="130F2A204C051B74FC09F95DFBCA76B7" box="[947,1048,1752,1774]" captionStart="Figure 21" captionStartId="34.[145,224,1314,1336]" captionTargetBox="[145,1425,195,1275]" captionTargetId="figure-246@34.[145,1425,195,1275]" captionTargetPageId="34" captionText="Figure 21. Anatomy of Radomaniola szarowskae sp. nov. AH, paratypes (UGSB 18566). 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.7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="32" pageNumber="425">Fig. 21A</figureCitation>
). Stomach slightly longer than wide, with two chambers almost equal in size; style sac longer than wide, surrounded by an unpigmented intestine (
<figureCitation id="130F2A204C051B74FC26F8B1FC287713" box="[924,1018,1844,1866]" captionStart="Figure 21" captionStartId="34.[145,224,1314,1336]" captionTargetBox="[145,1425,195,1275]" captionTargetId="figure-246@34.[145,1425,195,1275]" captionTargetPageId="34" captionText="Figure 21. Anatomy of Radomaniola szarowskae sp. nov. AH, paratypes (UGSB 18566). 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.7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="32" pageNumber="425">Fig. 21B</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="130F2A204C041B75FEE3F984FE6A764C" box="[345,440,1537,1559]" captionStart="Figure 21" captionStartId="34.[145,224,1314,1336]" captionTargetBox="[145,1425,195,1275]" captionTargetId="figure-246@34.[145,1425,195,1275]" captionTargetPageId="34" captionText="Figure 21. Anatomy of Radomaniola szarowskae sp. nov. AH, paratypes (UGSB 18566). 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.7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="33" pageNumber="426">Fig. 21C</figureCitation>
).
</paragraph>
<caption id="DF4B662D4C041B75FF19FAC9FD3475C7" ID-DOI="http://doi.org/10.5281/zenodo.7035630" ID-Zenodo-Dep="7035630" httpUri="https://zenodo.org/record/7035630/files/figure.png" pageId="33" pageNumber="426" startId="33.[163,241,1356,1378]" targetBox="[163,1443,195,1317]" targetPageId="33">
<paragraph id="8B8B36A54C041B75FF19FAC9FD3475C7" blockId="33.[163,1443,1356,1437]" pageId="33" pageNumber="426">
<emphasis id="B940EAB74C041B75FF19FAC9FECB753B" bold="true" box="[163,281,1356,1378]" pageId="33" pageNumber="426">Figure 20.</emphasis>
Shells, operculum and radulae of
<taxonomicName id="4C344D264C041B75FD3BFAC8FC587539" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[641,906,1357,1379]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="33" pageNumber="426" phylum="Mollusca" rank="species" species="szarowskae" status="sp. nov.">
<emphasis id="B940EAB74C041B75FD3BFAC8FC587539" box="[641,906,1357,1379]" italics="true" pageId="33" pageNumber="426">Radomaniola szarowskae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27357CC4C041B75FC2BFAC8FC0F7539" box="[913,989,1357,1379]" pageId="33" pageNumber="426" rank="species">sp. nov.</taxonomicNameLabel>
A, B, holotype (
<emphasis id="B940EAB74C041B75FBC5FAC8FAB07539" box="[1151,1378,1357,1379]" italics="true" pageId="33" pageNumber="426">MNCN 15.05/200167</emphasis>
). CJ, paratypes (UGSB 18556). 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="8B8B36A54C041B75FF01F99AFD587770" blockId="33.[163,780,1475,1896]" pageId="33" pageNumber="426">
Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland.
<collectingRegion id="49F0F8474C041B75FEF5F9D8FE477628" box="[335,405,1629,1650]" country="Turkey" name="Bursa" pageId="33" pageNumber="426">Bursa</collectingRegion>
copulatrix ovoid, slightly longer than wide. Bursal duct as long as bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, duct short, joining renal oviduct slightly above the insertion point with bursal duct. SR2 as long as SR1, elongate, with a short duct, located on renal oviduct near loop (
<figureCitation id="130F2A204C041B75FD65F973FF3D7770" captionStart="Figure 21" captionStartId="34.[145,224,1314,1336]" captionTargetBox="[145,1425,195,1275]" captionTargetId="figure-246@34.[145,1425,195,1275]" captionTargetPageId="34" captionText="Figure 21. Anatomy of Radomaniola szarowskae sp. nov. AH, paratypes (UGSB 18566). 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.7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="33" pageNumber="426">Fig. 21D, E</figureCitation>
; Supporting Information, Table S8).
</paragraph>
<paragraph id="8B8B36A54C041B75FF01F8B6FA4A76CB" blockId="33.[163,780,1475,1896]" lastBlockId="33.[827,1444,1475,1681]" pageId="33" pageNumber="426">
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="130F2A204C041B75FACDFA67FCB9764C" captionStart="Figure 21" captionStartId="34.[145,224,1314,1336]" captionTargetBox="[145,1425,195,1275]" captionTargetId="figure-246@34.[145,1425,195,1275]" captionTargetPageId="34" captionText="Figure 21. Anatomy of Radomaniola szarowskae sp. nov. AH, paratypes (UGSB 18566). 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.7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="33" pageNumber="426">Fig. 21H</figureCitation>
). Penis unpigmented, gradually tapering, shorter than head length, base wide, weakly folded along inner edge and with one medial outgrowth on its left side (
<figureCitation id="130F2A204C041B75FCCDF9D8FC227628" box="[887,1008,1629,1651]" captionStart="Figure 21" captionStartId="34.[145,224,1314,1336]" captionTargetBox="[145,1425,195,1275]" captionTargetId="figure-246@34.[145,1425,195,1275]" captionTargetPageId="34" captionText="Figure 21. Anatomy of Radomaniola szarowskae sp. nov. AH, paratypes (UGSB 18566). 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.7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="33" pageNumber="426">Fig. 21F, G</figureCitation>
); penis attached close to the right eye; penial duct narrow, near outer edge, almost straight.
</paragraph>
<paragraph id="8B8B36A54C041B76FC81F93FFD2A75BA" blockId="33.[825,1444,1722,1897]" lastBlockId="34.[145,761,1452,1505]" lastPageId="34" lastPageNumber="427" pageId="33" pageNumber="426">
<emphasis id="B940EAB74C041B75FC81F93FFC4A7695" box="[827,920,1722,1743]" italics="true" pageId="33" pageNumber="426">Habitat:</emphasis>
The source of the Cetina River is a large, deep, karst spring located in the north-western slopes of the Dinara Alps. Snails are distributed on stones submerged at
<quantity id="4CCC9B404C041B75FCE1F893FC4F7776" box="[859,925,1814,1836]" metricMagnitude="-1" metricUnit="m" metricValue="5.0" pageId="33" pageNumber="426" unit="m" value="0.5">0.5 m</quantity>
along the shore of the quiet pool formed by the spring waters. The temperature and conductivity of the water are low (9.6 °C and 350 µS/cm) and aquatic vegetation is sparse. Other co-occurring gastropods are
<taxonomicName id="4C344D264C071B76FF2BFA4EFD9B75BA" authority="(Bourguignat, 1887)" baseAuthorityName="Bourguignat" baseAuthorityYear="1887" box="[145,585,1483,1505]" class="Gastropoda" family="Hydrobiidae" genus="Horatia" kingdom="Animalia" order="Littorinimorpha" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="klecakiana">
<emphasis id="B940EAB74C071B76FF2BFA4EFEB475BA" box="[145,358,1483,1504]" italics="true" pageId="34" pageNumber="427">Horatia klecakiana</emphasis>
(Bourguignat, 1887)
</taxonomicName>
and
<taxonomicName id="4C344D264C071B76FDC1FA4EFD2A75BA" box="[635,760,1483,1504]" class="Gastropoda" family="Planorbidae" genus="Ancylus" kingdom="Animalia" order="Siphonariida" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B940EAB74C071B76FDC1FA4EFD0175BA" box="[635,723,1483,1504]" italics="true" pageId="34" pageNumber="427">Ancylus</emphasis>
sp.
</taxonomicName>
</paragraph>
<caption id="DF4B662D4C071B76FF2BFAA7FE237529" ID-DOI="http://doi.org/10.5281/zenodo.7035634" ID-Zenodo-Dep="7035634" httpUri="https://zenodo.org/record/7035634/files/figure.png" pageId="34" pageNumber="427" startId="34.[145,224,1314,1336]" targetBox="[145,1425,195,1275]" targetPageId="34">
<paragraph id="8B8B36A54C071B76FF2BFAA7FE237529" blockId="34.[145,1426,1314,1395]" pageId="34" pageNumber="427">
<emphasis id="B940EAB74C071B76FF2BFAA7FEDA7562" bold="true" box="[145,264,1314,1336]" pageId="34" pageNumber="427">Figure 21.</emphasis>
Anatomy of
<taxonomicName id="4C344D264C071B76FE2FFAA6FD717563" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[405,675,1315,1337]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="szarowskae" status="sp. nov.">
<emphasis id="B940EAB74C071B76FE2FFAA6FD717563" box="[405,675,1315,1337]" italics="true" pageId="34" pageNumber="427">Radomaniola szarowskae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27357CC4C071B76FD10FAA6FD257563" box="[682,759,1315,1337]" pageId="34" pageNumber="427" rank="species">sp. nov.</taxonomicNameLabel>
AH, paratypes (UGSB 18566). 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.
</paragraph>
</caption>
<paragraph id="8B8B36A54C071B76FF2BF98CFAD475A5" blockId="34.[145,761,1545,1905]" lastBlockId="34.[809,1424,1452,1535]" pageId="34" pageNumber="427">
<emphasis id="B940EAB74C071B76FF2BF98CFF2C7644" box="[145,254,1545,1566]" italics="true" pageId="34" pageNumber="427">Remarks:</emphasis>
This is one of the largest species recorded so far for this genus. Conchologically,
<taxonomicName id="4C344D264C071B76FDEEF9ACFD2B7667" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[596,761,1576,1598]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="szarowskae">
<emphasis id="B940EAB74C071B76FDEEF9ACFD2B7667" box="[596,761,1576,1598]" italics="true" pageId="34" pageNumber="427">R. szarowskae</emphasis>
</taxonomicName>
resembles
<emphasis id="B940EAB74C071B76FEABF9CDFE7F7606" box="[273,429,1607,1629]" italics="true" pageId="34" pageNumber="427">R. c. kicavica</emphasis>
, but it differs anatomically from this subspecies according to its larger penis, larger and more globous bursa copulatrix, less concentrated nervous ring and a larger number of cusps on the lateral radular teeth. Phylogenetically, the two taxa are distantly positioned from one another (
<figureCitation id="130F2A204C071B76FF23F97AFF0E774E" box="[153,220,1791,1813]" 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="34" pageNumber="427">Fig. 2</figureCitation>
), and their
<emphasis id="B940EAB74C071B76FED0F97AFE48774E" box="[362,410,1791,1812]" italics="true" pageId="34" pageNumber="427">COI</emphasis>
average sequence divergence is 5.6%.
<taxonomicName id="4C344D264C071B76FF43F898FDFD7768" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[249,559,1821,1842]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="szarowskae">
<emphasis id="B940EAB74C071B76FF43F898FDFD7768" box="[249,559,1821,1842]" italics="true" pageId="34" pageNumber="427">Radomaniola szarowskae</emphasis>
</taxonomicName>
differs from the phylogenetically and geographically close species
<taxonomicName id="4C344D264C071B76FF2BF8D9FEC5772A" baseAuthorityName="Radoman" baseAuthorityYear="1973" box="[145,279,1883,1905]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="bosniaca">
<emphasis id="B940EAB74C071B76FF2BF8D9FEC5772A" box="[145,279,1883,1905]" italics="true" pageId="34" pageNumber="427">R. bosniaca</emphasis>
</taxonomicName>
and
<taxonomicName id="4C344D264C071B76FEE8F8D9FDD3772A" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[338,513,1883,1905]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="34" pageNumber="427" phylum="Mollusca" rank="species" species="nachtigallae">
<emphasis id="B940EAB74C071B76FEE8F8D9FDD3772A" box="[338,513,1883,1905]" italics="true" pageId="34" pageNumber="427">R. nachtigallae</emphasis>
</taxonomicName>
in that it has a more conic and larger shell, more globous and larger bursa copulatrix and in the shape and size of the penis (Supporting Information,
<figureCitation id="130F2A204C071B76FBF4FA6CFB7175A5" box="[1102,1187,1513,1535]" 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="34" pageNumber="427">Figs S2</figureCitation>
, S
<figureCitation id="130F2A204C071B76FB04FA6CFB1E75A5" box="[1214,1228,1513,1535]" captionStart="Figure 6" captionStartId="16.[145,227,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-231@16.[145,1425,195,1317]" captionTargetPageId="16" captionText="Figure 6. Shells, operculum and radulae of Radomaniola curta montenegrensis subsp. nov. A, B, holotype (MNCN 15.05/200157). C, paratype (UGSB 19515). D, shell (UGSB 19043). EJ, paratypes (UGSB 19515). 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.7035597" httpUri="https://zenodo.org/record/7035597/files/figure.png" pageId="34" pageNumber="427">6</figureCitation>
, S
<figureCitation id="130F2A204C071B76FB53FA6FFB2575A5" box="[1257,1271,1514,1535]" captionStart="Figure 8" captionStartId="18.[142,220,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-210@18.[145,1425,195,1317]" captionTargetPageId="18" captionText="Figure 8. Shells, operculum and radulae of Radomaniola curta omblensis subsp. nov.A, B, holotype (MNCN 15.05/200155). CJ, paratypes (UGSB 18778). 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.7035601" httpUri="https://zenodo.org/record/7035601/files/figure.png" pageId="34" pageNumber="427">8</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>