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<document id="A74DB8EAA19B8AE4B79EBF0AEE4DE32C" 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="039D87B34C3F1B4AFCC9F9D6FF1C7211" 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 nachtigallae Delicado &amp; Hauffe 2022, SP. NOV." docType="treatment" docUuid="0139AB61-FB6A-42C4-814A-78E132AAEC1A" docUuidSource="ZooBank" docVersion="9" lastPageNumber="422" 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="419" updateTime="1699373995385" updateUser="ExternalLinkService">
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<mods:title id="4D4FDBD92135594969BC6491D74FA8A4">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="4425B488E54B0F17EC184E465C2C78D5">Delicado, Diana</mods:namePart>
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<mods:namePart id="95534C7308EE3FEFF2CAB95C0C5C9BBC">Hauffe, Torsten</mods:namePart>
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<mods:title id="369B7B5A5FFDEF1653DEEDFBE24AB370">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="03DA84D7570930B697843FCF2A06915A">2022</mods:date>
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<mods:number id="7042905B2DBC328EF329EC60AC71139E">2022-02-23</mods:number>
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<mods:number id="DE32CE95501187735E32DAB7707ED808">196</mods:number>
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<mods:classification id="31568611833217C56C4D4D9DEA4A1B57">journal article</mods:classification>
<mods:identifier id="1F2019761782561D25802124F9CB6011" type="CLB-Dataset">133178</mods:identifier>
<mods:identifier id="CB73CC3F493295397449A9E0D68F209D" type="DOI">10.1093/zoolinnean/zlab121</mods:identifier>
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<treatment id="039D87B34C3F1B4AFCC9F9D6FF1C7211" ID-DOI="http://doi.org/10.5281/zenodo.7036883" ID-GBIF-Taxon="199898763" ID-Zenodo-Dep="7036883" LSID="urn:lsid:zoobank.org:act:0139AB61-FB6A-42C4-814A-78E132AAEC1A" httpUri="http://treatment.plazi.org/id/039D87B34C3F1B4AFCC9F9D6FF1C7211" lastPageId="30" lastPageNumber="422" pageId="26" pageNumber="419">
<subSubSection id="C32E652E4C3F1B4EFCC9F9D6FA957631" box="[883,1351,1619,1643]" pageId="26" pageNumber="419" type="nomenclature">
<paragraph id="8B8B36A54C3F1B4EFCC9F9D6FA957631" blockId="26.[883,1351,1619,1643]" box="[883,1351,1619,1643]" pageId="26" pageNumber="419">
<heading id="D0C381C94C3F1B4EFCC9F9D6FA957631" box="[883,1351,1619,1643]" centered="true" fontSize="9" level="2" pageId="26" pageNumber="419" reason="2">
<taxonomicName id="4C344D264C3F1B4EFCC9F9D6FB0E7630" ID-CoL="9CCJC" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[883,1244,1619,1642]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="26" pageNumber="440" phylum="Mollusca" rank="species" species="nachtigallae" status="sp. nov.">RADOMANIOLA NACHTIGALLAE</taxonomicName>
<emphasis id="B940EAB74C3F1B4EFB5EF9D3FA957631" bold="true" box="[1252,1351,1619,1643]" pageId="26" pageNumber="419">
<taxonomicNameLabel id="A27357CC4C3F1B4EFB5EF9D3FA957631" box="[1252,1351,1619,1643]" pageId="26" pageNumber="419" rank="species">SP. NOV.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C32E652E4C3F1B4AFBA9F9FFFF1C7211" lastPageId="30" lastPageNumber="423" pageId="26" pageNumber="419" type="description">
<paragraph id="8B8B36A54C3F1B4EFBA9F9FFFB7476C9" blockId="26.[1043,1190,1658,1683]" box="[1043,1190,1658,1683]" pageId="26" pageNumber="419">
(
<figureCitation id="130F2A204C3F1B4EFBA6F9FFFBA676C9" box="[1052,1140,1658,1683]" captionStart="Figure 16" captionStartId="28.[145,225,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-230@28.[145,1425,195,1317]" captionTargetPageId="28" captionText="Figure 16. Shells, operculum and radulae of Radomaniola nachtigallae sp. nov. A, B, holotype (MNCN 15.05/200165). C–J, paratypes (UGSB 18857). 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.7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="26" pageNumber="419">FIGS 16</figureCitation>
,
<figureCitation id="130F2A204C3F1B4EFB38F9FFFB4C76C9" box="[1154,1182,1658,1683]" captionStart="Figure 17" captionStartId="29.[164,243,1315,1337]" captionTargetBox="[163,1443,195,1275]" captionTargetId="figure-249@29.[163,1443,195,1275]" captionTargetPageId="29" captionText="Figure 17. Anatomy of Radomaniola nachtigallae sp. nov. A–H, paratypes (UGSB 18857). 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.7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="26" pageNumber="419">17</figureCitation>
)
</paragraph>
<paragraph id="8B8B36A54C3F1B4EFC93F926FA5B768D" blockId="26.[809,1426,1698,1751]" pageId="26" pageNumber="419">
<emphasis id="B940EAB74C3F1B4EFC93F926FB0776E2" box="[809,1237,1698,1720]" italics="true" pageId="26" pageNumber="419">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="FF920C704C3F1B4EFCCCF944FA5B768D" box="[886,1417,1729,1751]" pageId="26" pageNumber="419">0139AB61-FB6A-42C4-814A-78E132AAEC1A</uuid>
</paragraph>
<paragraph id="8B8B36A54C3F1B4EFC93F885FC53772B" blockId="26.[809,1425,1792,1905]" pageId="26" pageNumber="419">
<emphasis id="B940EAB74C3F1B4EFC93F885FC61774F" box="[809,947,1792,1813]" italics="true" pageId="26" pageNumber="419">Etymology:</emphasis>
Named after Silvia Nachtigall, who helped us with much of the preparation of material and the laboratory work conducted for this and other studies.
</paragraph>
<caption id="DF4B662D4C3E1B4FFF19FAA6FDD1752E" ID-DOI="http://doi.org/10.5281/zenodo.7035618" ID-Zenodo-Dep="7035618" httpUri="https://zenodo.org/record/7035618/files/figure.png" pageId="27" pageNumber="420" startId="27.[163,242,1315,1337]" targetBox="[163,1443,195,1275]" targetPageId="27">
<paragraph id="8B8B36A54C3E1B4FFF19FAA6FDD1752E" blockId="27.[163,1444,1315,1396]" pageId="27" pageNumber="420">
<emphasis id="B940EAB74C3E1B4FFF19FAA6FEC87562" bold="true" box="[163,282,1315,1337]" pageId="27" pageNumber="420">Figure 15.</emphasis>
Anatomy of
<taxonomicName id="4C344D264C3E1B4FFE1FFAA6FD6B7563" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[421,697,1315,1337]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="27" pageNumber="420" phylum="Mollusca" rank="species" species="jovanovskae" status="sp. nov.">
<emphasis id="B940EAB74C3E1B4FFE1FFAA6FD6B7563" box="[421,697,1315,1337]" italics="true" pageId="27" pageNumber="420">Radomaniola jovanovskae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27357CC4C3E1B4FFD7AFAA6FCDF7563" box="[704,781,1315,1337]" pageId="27" pageNumber="420" rank="species">sp. nov.</taxonomicNameLabel>
A–H, paratypes (UGSB 19517). 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="8B8B36A54C3E1B4FFF19FA1FFD097670" blockId="27.[163,780,1434,1578]" pageId="27" pageNumber="420">
<emphasis id="B940EAB74C3E1B4FFF19FA1FFE8575F5" box="[163,343,1434,1455]" italics="true" pageId="27" pageNumber="420">Type material:</emphasis>
<materialsCitation id="3B5C3CF84C3E1B4FFED5FA1FFD097670" ID-GBIF-Occurrence="3897142304" collectionCode="MNCN, UGSB" country="Bosnia and Herzegovina" location="Krupa River" municipality="Krupa na Vrbasu" pageId="27" pageNumber="420" specimenCode="MNCN 15.05, UGSB 18857" specimenCount="101" typeStatus="holotype">
<typeStatus id="548F88074C3E1B4FFED5FA1FFE0F75EA" box="[367,477,1434,1456]" pageId="27" pageNumber="420" type="holotype">Holotype</typeStatus>
(
<emphasis id="B940EAB74C3E1B4FFE4AFA1FFD2875F5" box="[496,762,1434,1455]" italics="true" pageId="27" pageNumber="420">
<specimenCode id="DB929EDE4C3E1B4FFE4AFA1FFD4175F5" box="[496,659,1434,1455]" 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="27" pageNumber="420" type="Museum">MNCN 15.05</specimenCode>
/200165
</emphasis>
),
<specimenCount id="9D32FD2C4C3E1B4FFF19FA3CFE9A7594" box="[163,328,1465,1486]" pageId="27" pageNumber="420" type="generic" typeStatus="paratypes">ten paratypes</specimenCount>
(
<specimenCode id="DB929EDE4C3E1B4FFEE3FA3CFE277594" box="[345,501,1464,1486]" 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="27" pageNumber="420" type="Museum">MNCN 15.05</specimenCode>
/200166) in the
<collectionCode id="ED25AE604C3E1B4FFD0DFA3CFCD87594" box="[695,778,1465,1486]" 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="27" pageNumber="420" type="Museum">MNCN</collectionCode>
collection and&gt;
<specimenCount id="9D32FD2C4C3E1B4FFEDCFA52FDDE75B7" box="[358,524,1495,1517]" pageId="27" pageNumber="420" type="generic" typeStatus="paratypes">100 paratypes</specimenCount>
(
<specimenCode id="DB929EDE4C3E1B4FFDA6FA52FD6475B7" box="[540,694,1495,1517]" collectionCode="UGSB" pageId="27" pageNumber="420">UGSB 18857</specimenCode>
) in the
<collectionCode id="ED25AE604C3E1B4FFF1EFA73FF3C7651" box="[164,238,1526,1547]" pageId="27" pageNumber="420">UGSB</collectionCode>
collection. Type locality: source of the
<location id="8EEB607E4C3E1B4FFD7BFA73FF337670" LSID="urn:lsid:plazi:treatment:039D87B34C3F1B4AFCC9F9D6FF1C7211:8EEB607E4C3E1B4FFD7BFA73FF337670" country="Bosnia and Herzegovina" municipality="Krupa na Vrbasu" name="Krupa River" pageId="27" pageNumber="420">Krupa River</location>
,
<collectingMunicipality id="6BEFACDF4C3E1B4FFF51F990FE607670" box="[235,434,1556,1578]" pageId="27" pageNumber="420">Krupa na Vrbasu</collectingMunicipality>
,
<collectingCountry id="F32376354C3E1B4FFE07F991FD057670" box="[445,727,1556,1578]" name="Bosnia and Herzegovina" pageId="27" pageNumber="420">Bosnia and Herzegovina</collectingCountry>
.
</materialsCitation>
</paragraph>
<paragraph id="8B8B36A54C3E1B4FFF19F9D6FD60769F" blockId="27.[163,779,1619,1733]" pageId="27" pageNumber="420">
<emphasis id="B940EAB74C3E1B4FFF19F9D6FEBE7632" box="[163,364,1619,1640]" italics="true" pageId="27" pageNumber="420">Material studied:</emphasis>
<materialsCitation id="3B5C3CF84C3E1B4FFE38F9D6FD7C769F" ID-GBIF-Occurrence="3897142312" collectionCode="T" collectorName="Diana Delicado &amp; Ju" country="Bosnia and Herzegovina" county="Krupa na Vrbasu" latitude="44.6207" location="Source" longLatPrecision="39543" longitude="5.0" municipality="Krupa River" pageId="27" pageNumber="420" specimenCode="UGSB 18857" specimenCount="1">
<location id="8EEB607E4C3E1B4FFE38F9D6FE037632" LSID="urn:lsid:plazi:treatment:039D87B34C3F1B4AFCC9F9D6FF1C7211:8EEB607E4C3E1B4FFE38F9D6FE037632" box="[386,465,1619,1640]" country="Bosnia and Herzegovina" county="Krupa na Vrbasu" latitude="44.6207" longLatPrecision="39543" longitude="5.0" municipality="Krupa River" name="Source" pageId="27" pageNumber="420">Source</location>
of the
<collectingMunicipality id="6BEFACDF4C3E1B4FFD9EF9D1FD677633" box="[548,693,1619,1641]" pageId="27" pageNumber="420">Krupa River</collectingMunicipality>
,
<collectingCounty id="62EA4E294C3E1B4FFD7BF9D1FEFB76D2" pageId="27" pageNumber="420">Krupa na Vrbasu</collectingCounty>
,
<collectingCountry id="F32376354C3E1B4FFE83F9F7FDA176D2" box="[313,627,1650,1672]" name="Bosnia and Herzegovina" pageId="27" pageNumber="420">Bosnia and Herzegovina</collectingCountry>
,
<geoCoordinate id="EE0050624C3E1B4FFD3EF9F7FCD476D2" box="[644,774,1650,1672]" degrees="44.6207" direction="north" orientation="latitude" pageId="27" pageNumber="420" precision="5" value="44.6207">44.6207°N</geoCoordinate>
, 1 7. 1 1 0
<geoCoordinate id="EE0050624C3E1B4FFF41F915FEFB76FC" box="[251,297,1680,1702]" degrees="5" direction="east" orientation="longitude" pageId="27" pageNumber="420" precision="55555" value="5.0">5°E</geoCoordinate>
,
<collectionCode id="ED25AE604C3E1B4FFE85F914FE8076FC" box="[319,338,1681,1702]" name="Tavera, Department of Geology and Geophysics" pageId="27" pageNumber="420">T</collectionCode>
.H. a n d
<collectorName id="26C153734C3E1B4FFE78F914FE2876FC" box="[450,506,1681,1702]" name="Diana Delicado" pageId="27" pageNumber="420">D.D.</collectorName>
,
<collectorName id="26C153734C3E1B4FFDABF914FDFD76FC" box="[529,559,1681,1702]" pageId="27" pageNumber="420">Ju</collectorName>
n e 2 0 1 6, M N C N 15.05/200166 and
<specimenCode id="DB929EDE4C3E1B4FFEC8F92AFDD5769F" box="[370,519,1711,1733]" collectionCode="UGSB" pageId="27" pageNumber="420">UGSB 18857</specimenCode>
(80% ethanol)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8B36A54C3E1B4FFF19F96BFBD87656" blockId="27.[163,779,1774,1888]" lastBlockId="27.[827,1442,1434,1548]" pageId="27" pageNumber="420">
<emphasis id="B940EAB74C3E1B4FFF19F96BFEFB7759" box="[163,297,1774,1795]" italics="true" pageId="27" pageNumber="420">Diagnosis:</emphasis>
Protoconch microsculpture slightly wrinkled; central radular tooth formula 7-C-7/1- 1; bursa copulatrix globular, with a long duct; SR1 elongate, duct short; SR2 elongate, duct short; penis slightly pigmented on the distal section, gradually tapering, shorter than head length; nervous system weakly pigmented, moderately concentrated (mean RPG ratio = 0.46).
</paragraph>
<paragraph id="8B8B36A54C3E1B48FC81F9B0FDC576A5" blockId="27.[827,1444,1588,1886]" lastBlockId="28.[144,762,1494,1884]" lastPageId="28" lastPageNumber="421" pageId="27" pageNumber="420">
<emphasis id="B940EAB74C3E1B4FFC81F9B0FC187610" box="[827,970,1589,1610]" italics="true" pageId="27" pageNumber="420">Description:</emphasis>
Shell ovate-conic, four whorls, height
<quantity id="4CCC9B404C3E1B4FFC86F9D6FC187633" box="[828,970,1619,1641]" metricMagnitude="-3" metricUnit="m" metricValue="2.5" metricValueMax="2.8" metricValueMin="2.2" pageId="27" pageNumber="420" unit="mm" value="2.5" valueMax="2.8" valueMin="2.2">2.2–2.8 mm</quantity>
(
<figureCitation id="130F2A204C3E1B4FFC67F9D6FBB47633" box="[989,1126,1619,1641]" captionStart="Figure 16" captionStartId="28.[145,225,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-230@28.[145,1425,195,1317]" captionTargetPageId="28" captionText="Figure 16. Shells, operculum and radulae of Radomaniola nachtigallae sp. nov. A, B, holotype (MNCN 15.05/200165). C–J, paratypes (UGSB 18857). 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.7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="27" pageNumber="420">Fig. 16A–D</figureCitation>
; Supporting Information, Table S6). Periostracum yellowish. Protoconch ~550 µm wide, 1.5 whorls; nucleus ~200 µm wide; protoconch microsculpture slightly wrinkled (
<figureCitation id="130F2A204C3E1B4FFACFF92AFCB876BE" captionStart="Figure 16" captionStartId="28.[145,225,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-230@28.[145,1425,195,1317]" captionTargetPageId="28" captionText="Figure 16. Shells, operculum and radulae of Radomaniola nachtigallae sp. nov. A, B, holotype (MNCN 15.05/200165). C–J, paratypes (UGSB 18857). 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.7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="27" pageNumber="420">Fig. 16G</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="130F2A204C3E1B4FFC12F8CDFBDA7704" box="[936,1032,1864,1886]" captionStart="Figure 16" captionStartId="28.[145,225,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-230@28.[145,1425,195,1317]" captionTargetPageId="28" captionText="Figure 16. Shells, operculum and radulae of Radomaniola nachtigallae sp. nov. A, B, holotype (MNCN 15.05/200165). C–J, paratypes (UGSB 18857). 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.7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="27" pageNumber="420">Fig. 16B</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="130F2A204C391B48FE9BF996FE487673" box="[289,410,1555,1577]" captionStart="Figure 16" captionStartId="28.[145,225,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-230@28.[145,1425,195,1317]" captionTargetPageId="28" captionText="Figure 16. Shells, operculum and radulae of Radomaniola nachtigallae sp. nov. A, B, holotype (MNCN 15.05/200165). C–J, paratypes (UGSB 18857). 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.7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="28" pageNumber="421">Fig. 16E, F</figureCitation>
). Radular length intermediate, ~650 µm (~25% of total shell length), with ~60 rows of teeth (
<figureCitation id="130F2A204C391B48FF44F9D4FEB6763D" box="[254,356,1617,1639]" captionStart="Figure 16" captionStartId="28.[145,225,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-230@28.[145,1425,195,1317]" captionTargetPageId="28" captionText="Figure 16. Shells, operculum and radulae of Radomaniola nachtigallae sp. nov. A, B, holotype (MNCN 15.05/200165). C–J, paratypes (UGSB 18857). 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.7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="28" pageNumber="421">Fig. 16H</figureCitation>
). Central tooth formula 7-C-7/1-1 (
<figureCitation id="130F2A204C391B48FF23F9F5FF2076DF" box="[153,242,1647,1670]" captionStart="Figure 16" captionStartId="28.[145,225,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-230@28.[145,1425,195,1317]" captionTargetPageId="28" captionText="Figure 16. Shells, operculum and radulae of Radomaniola nachtigallae sp. nov. A, B, holotype (MNCN 15.05/200165). C–J, paratypes (UGSB 18857). 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.7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="28" pageNumber="421">Fig. 16I</figureCitation>
); basal tongue U-shaped, length about equal to lateral margin. Lateral tooth formula (5)4-C-4(5). Inner marginal teeth having 30–32 tapered cusps, shortening toward the base. Outer marginal teeth with 35–37 sharp cusps (
<figureCitation id="130F2A204C391B48FE17F96FFDD5775A" box="[429,519,1770,1792]" captionStart="Figure 16" captionStartId="28.[145,225,1356,1378]" captionTargetBox="[145,1425,195,1317]" captionTargetId="figure-230@28.[145,1425,195,1317]" captionTargetPageId="28" captionText="Figure 16. Shells, operculum and radulae of Radomaniola nachtigallae sp. nov. A, B, holotype (MNCN 15.05/200165). C–J, paratypes (UGSB 18857). 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.7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="28" pageNumber="421">Fig. 16J</figureCitation>
).
</paragraph>
<caption id="DF4B662D4C391B48FF2BFAC9FD0675C7" ID-DOI="http://doi.org/10.5281/zenodo.7035620" ID-Zenodo-Dep="7035620" httpUri="https://zenodo.org/record/7035620/files/figure.png" pageId="28" pageNumber="421" startId="28.[145,225,1356,1378]" targetBox="[145,1425,195,1317]" targetPageId="28">
<paragraph id="8B8B36A54C391B48FF2BFAC9FD0675C7" blockId="28.[145,1425,1356,1437]" pageId="28" pageNumber="421">
<emphasis id="B940EAB74C391B48FF2BFAC9FEDB7538" bold="true" box="[145,265,1356,1378]" pageId="28" pageNumber="421">Figure 16.</emphasis>
Shells, operculum and radulae of
<taxonomicName id="4C344D264C391B48FDC5FAC8FC497539" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[639,923,1357,1379]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="28" pageNumber="421" phylum="Mollusca" rank="species" species="nachtigallae" status="sp. nov.">
<emphasis id="B940EAB74C391B48FDC5FAC8FC497539" box="[639,923,1357,1379]" italics="true" pageId="28" pageNumber="421">Radomaniola nachtigallae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27357CC4C391B48FC19FAC8FC237539" box="[931,1009,1357,1379]" pageId="28" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
A, B, holotype (
<emphasis id="B940EAB74C391B48FB21FAC8FA517539" box="[1179,1411,1357,1379]" italics="true" pageId="28" pageNumber="421">MNCN 15.05/200165</emphasis>
). C–J, paratypes (UGSB 18857). 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="8B8B36A54C391B48FF13F88CFB9F76A5" blockId="28.[144,762,1494,1884]" lastBlockId="28.[809,1425,1494,1884]" pageId="28" pageNumber="421">
Animal darkly pigmented except for neck and tentacles (
<figureCitation id="130F2A204C391B48FEAFF8A2FEAF7767" box="[277,381,1831,1853]" captionStart="Figure 17" captionStartId="29.[164,243,1315,1337]" captionTargetBox="[163,1443,195,1275]" captionTargetId="figure-249@29.[163,1443,195,1275]" captionTargetPageId="29" captionText="Figure 17. Anatomy of Radomaniola nachtigallae sp. nov. A–H, paratypes (UGSB 18857). 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.7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="28" pageNumber="421">Fig. 17F</figureCitation>
). Ctenidium with 13–15 welldeveloped gill filaments, occupying ~50% of pallial cavity length and positioned posteriorly. Osphradium of intermediate width and opposite middle of ctenidium (
<figureCitation id="130F2A204C391B48FC09F996FBCA7673" box="[947,1048,1555,1577]" captionStart="Figure 17" captionStartId="29.[164,243,1315,1337]" captionTargetBox="[163,1443,195,1275]" captionTargetId="figure-249@29.[163,1443,195,1275]" captionTargetPageId="29" captionText="Figure 17. Anatomy of Radomaniola nachtigallae sp. nov. A–H, paratypes (UGSB 18857). 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.7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="28" pageNumber="421">Fig. 17A</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="130F2A204C391B48FC1DF9EAFBDF76DF" box="[935,1037,1647,1669]" captionStart="Figure 17" captionStartId="29.[164,243,1315,1337]" captionTargetBox="[163,1443,195,1275]" captionTargetId="figure-249@29.[163,1443,195,1275]" captionTargetPageId="29" captionText="Figure 17. Anatomy of Radomaniola nachtigallae sp. nov. A–H, paratypes (UGSB 18857). 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.7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="28" pageNumber="421">Fig. 17B</figureCitation>
; Supporting Information, Table S7). Nervous system slightly pigmented, moderately concentrated (mean RPG ratio = 0.46); cerebral ganglia approximately equal in size, presenting small black granules (
<figureCitation id="130F2A204C391B48FC65F96FFBEC76A5" box="[991,1086,1770,1792]" captionStart="Figure 17" captionStartId="29.[164,243,1315,1337]" captionTargetBox="[163,1443,195,1275]" captionTargetId="figure-249@29.[163,1443,195,1275]" captionTargetPageId="29" captionText="Figure 17. Anatomy of Radomaniola nachtigallae sp. nov. A–H, paratypes (UGSB 18857). 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.7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="28" pageNumber="421">Fig. 17C</figureCitation>
).
</paragraph>
<paragraph id="8B8B36A54C391B49FCFBF88CFD117612" blockId="28.[809,1425,1494,1884]" lastBlockId="29.[163,780,1434,1885]" lastPageId="29" lastPageNumber="422" pageId="28" pageNumber="421">
Female glandular oviduct approximately three times longer than wide. Albumen gland shorter than capsule gland.
<collectingRegion id="49F0F8474C391B48FCC4F8C2FC157706" box="[894,967,1863,1884]" country="Turkey" name="Bursa" pageId="28" pageNumber="421">Bursa</collectingRegion>
copulatrix globular, as long as wide. Bursal duct longer than bursal length. Renal oviduct unpigmented, coiled. SR1 elongate, duct short, joining renal oviduct slightly above the insertion point with bursal duct. SR2 slightly shorter than SR1, elongate, with a short duct, located on renal oviduct near loop (
<figureCitation id="130F2A204C381B49FF11F9B6FEFB7613" box="[171,297,1587,1609]" captionStart="Figure 17" captionStartId="29.[164,243,1315,1337]" captionTargetBox="[163,1443,195,1275]" captionTargetId="figure-249@29.[163,1443,195,1275]" captionTargetPageId="29" captionText="Figure 17. Anatomy of Radomaniola nachtigallae sp. nov. A–H, paratypes (UGSB 18857). 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.7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="29" pageNumber="422">Fig. 17D, E</figureCitation>
; Supporting Information, Table S8).
</paragraph>
<caption id="DF4B662D4C381B49FF1EFAA6FDD1752E" ID-DOI="http://doi.org/10.5281/zenodo.7035622" ID-Zenodo-Dep="7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="29" pageNumber="422" startId="29.[164,243,1315,1337]" targetBox="[163,1443,195,1275]" targetPageId="29">
<paragraph id="8B8B36A54C381B49FF1EFAA6FDD1752E" blockId="29.[163,1443,1315,1396]" pageId="29" pageNumber="422">
<emphasis id="B940EAB74C381B49FF1EFAA6FEC87562" bold="true" box="[164,282,1315,1337]" pageId="29" pageNumber="422">Figure 17.</emphasis>
Anatomy of
<taxonomicName id="4C344D264C381B49FE1EFAA6FD697563" authority="Delicado &amp; Hauffe, 2022" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[420,699,1315,1337]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="29" pageNumber="422" phylum="Mollusca" rank="species" species="nachtigallae" status="sp. nov.">
<emphasis id="B940EAB74C381B49FE1EFAA6FD697563" box="[420,699,1315,1337]" italics="true" pageId="29" pageNumber="422">Radomaniola nachtigallae</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27357CC4C381B49FD78FAA6FCDD7563" box="[706,783,1315,1337]" pageId="29" pageNumber="422" rank="species">sp. nov.</taxonomicNameLabel>
A–H, paratypes (UGSB 18857). 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="8B8B36A54C381B49FF01F9D7FBE37594" blockId="29.[163,780,1434,1885]" lastBlockId="29.[827,1442,1434,1486]" pageId="29" pageNumber="422">
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="130F2A204C381B49FD65F92BFF0676BB" captionStart="Figure 17" captionStartId="29.[164,243,1315,1337]" captionTargetBox="[163,1443,195,1275]" captionTargetId="figure-249@29.[163,1443,195,1275]" captionTargetPageId="29" captionText="Figure 17. Anatomy of Radomaniola nachtigallae sp. nov. A–H, paratypes (UGSB 18857). 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.7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="29" pageNumber="422">Fig. 17H</figureCitation>
). Penis slightly pigmented on the distal section, gradually tapering, shorter than head length, base medium wide, weakly folded along inner edge and with one medial outgrowth on its left side (
<figureCitation id="130F2A204C381B49FD31F8ADFCD77767" box="[651,773,1832,1854]" captionStart="Figure 17" captionStartId="29.[164,243,1315,1337]" captionTargetBox="[163,1443,195,1275]" captionTargetId="figure-249@29.[163,1443,195,1275]" captionTargetPageId="29" captionText="Figure 17. Anatomy of Radomaniola nachtigallae sp. nov. A–H, paratypes (UGSB 18857). 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.7035622" httpUri="https://zenodo.org/record/7035622/files/figure.png" pageId="29" pageNumber="422">Fig. 17F, G</figureCitation>
; Supporting Information, Table S9); penis attached well behind the right eye; penial duct narrow, near outer edge, almost straight.
</paragraph>
<paragraph id="8B8B36A54C381B49FC81FA72FBC176FC" blockId="29.[824,1444,1527,1702]" pageId="29" pageNumber="422">
<emphasis id="B940EAB74C381B49FC81FA72FC717656" box="[827,931,1527,1548]" italics="true" pageId="29" pageNumber="422">Habitat:</emphasis>
This species has been found only in a rheocrene spring of clear and shallow waters presenting low temperature (10.5 °C) and conductivity (465 µS/ cm). Aquatic plants are sparse. Snails are abundant on stones. Other co-occurring gastropods are
<taxonomicName id="4C344D264C381B49FA98F9F7FC8876FC" class="Gastropoda" family="Hydrobiidae" genus="Sadleriana" kingdom="Animalia" order="Littorinimorpha" pageId="29" pageNumber="422" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B940EAB74C381B49FA98F9F7FA7176DD" box="[1314,1443,1650,1671]" italics="true" pageId="29" pageNumber="422">Sadleriana</emphasis>
sp.
</taxonomicName>
and
<taxonomicName id="4C344D264C381B49FC28F915FBC176FC" box="[914,1043,1680,1702]" class="Gastropoda" family="Planorbidae" genus="Ancylus" kingdom="Animalia" order="Siphonariida" pageId="29" pageNumber="422" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B940EAB74C381B49FC28F915FC3E76FF" box="[914,1004,1680,1701]" italics="true" pageId="29" pageNumber="422">Ancylus</emphasis>
sp.
</taxonomicName>
</paragraph>
<paragraph id="8B8B36A54C381B4AFC81F94AFF1C7211" blockId="29.[827,1443,1743,1888]" lastBlockId="30.[145,762,197,587]" lastPageId="30" lastPageNumber="423" pageId="29" pageNumber="422">
<emphasis id="B940EAB74C381B49FC81F94AFC6276BE" box="[827,944,1743,1764]" italics="true" pageId="29" pageNumber="422">Remarks:</emphasis>
This species is closely related to the geographically proximate congeners
<taxonomicName id="4C344D264C381B49FB53F96BFABD7759" baseAuthorityName="Radoman" baseAuthorityYear="1973" box="[1257,1391,1774,1795]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="29" pageNumber="422" phylum="Mollusca" rank="species" species="bosniaca">
<emphasis id="B940EAB74C381B49FB53F96BFABD7759" box="[1257,1391,1774,1795]" italics="true" pageId="29" pageNumber="422">R. bosniaca</emphasis>
</taxonomicName>
and
<taxonomicName id="4C344D264C381B49FC86F888FC0F777B" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[828,989,1804,1826]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="29" pageNumber="422" phylum="Mollusca" rank="species" species="szarowskae">
<emphasis id="B940EAB74C381B49FC86F888FC0F777B" box="[828,989,1804,1826]" italics="true" pageId="29" pageNumber="422">R. szarowskae</emphasis>
</taxonomicName>
. The average
<emphasis id="B940EAB74C381B49FB3AF888FB7D7778" box="[1152,1199,1805,1826]" italics="true" pageId="29" pageNumber="422">COI</emphasis>
sequence divergence between
<taxonomicName id="4C344D264C381B49FC1EF8A9FB87771A" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[932,1109,1835,1857]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="29" pageNumber="422" phylum="Mollusca" rank="species" species="nachtigallae">
<emphasis id="B940EAB74C381B49FC1EF8A9FB87771A" box="[932,1109,1835,1857]" italics="true" pageId="29" pageNumber="422">R. nachtigallae</emphasis>
</taxonomicName>
and these two species is 2% and 4%, respectively. This small genetic divergence contrasts with their considerable morphological differentiation in terms of shell size, shell shape and other characters of the reproductive system. Thus,
<taxonomicName id="4C344D264C3B1B4AFF29FEA7FEED716C" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[147,319,289,311]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="30" pageNumber="423" phylum="Mollusca" rank="species" species="nachtigallae">
<emphasis id="B940EAB74C3B1B4AFF29FEA7FEED716C" box="[147,319,289,311]" italics="true" pageId="30" pageNumber="423">R. nachtigallae</emphasis>
</taxonomicName>
differs from
<taxonomicName id="4C344D264C3B1B4AFE6CFEA7FD88716C" baseAuthorityName="Radoman" baseAuthorityYear="1973" box="[470,602,289,311]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="30" pageNumber="423" phylum="Mollusca" rank="species" species="bosniaca">
<emphasis id="B940EAB74C3B1B4AFE6CFEA7FD88716C" box="[470,602,289,311]" italics="true" pageId="30" pageNumber="423">R. bosniaca</emphasis>
</taxonomicName>
in that it has a larger shell with taller spire whorls, wider base of penis, shorter penis compared with head length, more globular bursa copulatrix, longer bursal duct and more elongated SR1; and differs from
<taxonomicName id="4C344D264C3B1B4AFDEEFE18FD2A71EB" authorityName="Delicado &amp; Hauffe" authorityYear="2022" box="[596,760,412,434]" class="Gastropoda" family="Hydrobiidae" genus="Radomaniola" kingdom="Animalia" order="Littorinimorpha" pageId="30" pageNumber="423" phylum="Mollusca" rank="species" species="szarowskae">
<emphasis id="B940EAB74C3B1B4AFDEEFE18FD2A71EB" box="[596,760,412,434]" italics="true" pageId="30" pageNumber="423">R. szarowskae</emphasis>
</taxonomicName>
in its smaller shell, smaller penis and smaller bursa copulatrix (Supporting Information,
<figureCitation id="130F2A204C3B1B4AFD86FE5CFD4171B4" box="[572,659,473,495]" 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="30" pageNumber="423">Figs S2</figureCitation>
, S
<figureCitation id="130F2A204C3B1B4AFD0AFE5CFD6D71B5" box="[688,703,473,495]" 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). E–J, 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="30" pageNumber="423">6</figureCitation>
, S
<figureCitation id="130F2A204C3B1B4AFD66FE5CFD3871B4" box="[732,746,473,494]" 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). C–J, 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="30" pageNumber="423">8</figureCitation>
). These three species have a similar number of gill filaments and cusps on the radular teeth and RPG ratio.
</paragraph>
</subSubSection>
</treatment>
</document>
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