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<document id="EEF724B0D82D5FBDD8D7C7B9D9DFC827" ID-DOI="10.5852/ejt.2024.926.2469" ID-GBIF-Dataset="4511b4bd-f026-49d3-9d4d-a39875e75cc5" ID-ISSN="2118-9773" ID-Zenodo-Dep="10847806" ID-ZooBank="A00D7669-2151-46AE-A066-9AF4D0F2BEEE" IM.bibliography_approvedBy="guilherme" IM.illustrations_approvedBy="admin" IM.materialsCitations_approvedBy="guilherme" IM.metadata_approvedBy="guilherme" IM.tables_approvedBy="guilherme" IM.taxonomicNames_approvedBy="guilherme" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="guilherme" checkinTime="1711011351298" checkinUser="plazi" docAuthor="Jochum, Adrienne, Michalik, Peter, Inäbnit, Thomas, Kneubühler, Jeannette, Slapnik, Rajko, Vrabec, Marko, Schilthuizen, Menno &amp; Ruthensteiner, Bernhard" docDate="2024" docId="03B0879DFF91FFEDFE8DD11BFB77FDD5" docLanguage="en" docName="EJT.2024.926.1-62.pdf" docOrigin="European Journal of Taxonomy 926" docSource="https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2469/10973" docStyle="DocumentStyle:EF2B578F1D15862ADE45B0C07C620911.14:EJT.2018-.journal_article.type1" docStyleId="EF2B578F1D15862ADE45B0C07C620911" docStyleName="EJT.2018-.journal_article.type1" docStyleVersion="14" docTitle="Zospeum kolbae Jochum, Inabnit, Kneubuhler &amp; Ruthensteiner 2024, sp. nov." docType="treatment" docUuid="02B1232B-1BC5-404A-8F6D-FD189619B3AA" docUuidSource="ZooBank" docVersion="13" lastPageNumber="46" masterDocId="FF89FFE5FFBAFFC0FFFFD700FFD4FFD4" masterDocTitle="3 D X-ray microscopy (Micro-CT) and SEM reveal Zospeum troglobalcanicum Absolon, 1916 and allied species from the Western Balkans (Ellobioidea: Carychiidae)" masterLastPageNumber="62" masterPageNumber="1" pageNumber="44" updateTime="1722640713706" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="01E4EA1DE5D33A0863A23E0876785FAD">3 D X-ray microscopy (Micro-CT) and SEM reveal Zospeum troglobalcanicum Absolon, 1916 and allied species from the Western Balkans (Ellobioidea: Carychiidae)</mods:title>
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<mods:namePart id="3D1724BB89FD1701A9165EA126E8CAD9">Jochum, Adrienne</mods:namePart>
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<mods:namePart id="6EDAEF7B9EE357313CA6A9A0FBF09BA6">Michalik, Peter</mods:namePart>
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<mods:affiliation id="507C7096244D2E2E76156DCC696DC0FD">Naturhistorisches Museum Bern, Bernastrasse 15, CH- 3005 Bern, Switzerland. &amp; Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, CH- 3012 Bern, Switzerland.</mods:affiliation>
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<mods:affiliation id="A2A53FCD10C77166D65E8A2F8FD99111">Department of Geology, Faculty of Natural Sciences and Engineering, Aškerčeva 12, University of Ljubljana, 1000 Ljubljana, Slovenia.</mods:affiliation>
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<emphasis id="B96DEA99FF91FFEBFE8DD11BFDEEF9E1" bold="true" box="[370,570,1562,1589]" italics="true" pageId="43" pageNumber="44">Zospeum kolbae</emphasis>
Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner
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<figureCitation id="13222A0EFF91FFEBFD31D161FCFCF9AF" box="[718,808,1633,1659]" captionStart="Fig" captionStartId="45.[189,232,1787,1813]" captionTargetBox="[220,1366,561,1748]" captionTargetId="figure-212@45.[220,1366,561,1748]" captionTargetPageId="45" captionText="Fig. 16. Light microscopic images of full-bodied Zospeum Bourguignat, 1856 from Njeguši, St John’s cave. A–B. Zospeum kolbae Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner sp. nov. (NMBE 571122– 571123), individuals assessed by DNA sequencing with sigmoid intestine showing through shells (shells were destroyed post imaging for tissue extraction). A. Holotype (NMBE 571122), shell of aliquot, aperture, and aperture facing left view. B. Paratype (NMBE 571123), shell of aliquot, aperture, and aperture facing left view. C. Undescribed Zospeum sp. 1 (NMBE 577052) showing all perspectives. D–E. Undescribed Zospeum sp. 1 (NMBE 577053/2) showing all perspectives." figureDoi="http://doi.org/10.5281/zenodo.10847850" httpUri="https://zenodo.org/record/10847850/files/figure.png" pageId="43" pageNumber="44">Figs 16</figureCitation>
, 
<figureCitation id="13222A0EFF91FFEBFCCBD161FCB1F9AF" box="[820,869,1633,1659]" captionStart="Fig" captionStartId="55.[189,232,1822,1848]" captionTargetBox="[191,1400,633,1782]" captionTargetId="figure-236@55.[191,1400,621,1782]" captionTargetPageId="55" captionText="Fig. 22. Sites of localities and collections. A. Site of type locality of Zospeum njunjicae Jochum, Schilthuizen &amp; Ruthensteiner sp. nov., Golubova pećina, Gornja Seoca, Montenegro. Credit: I. Njunjić. B. Collection site within St John’s cave, type locality of Z. kolbae Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner sp. nov. and Z. njegusiense Jochum &amp; Ruthensteiner sp. nov., Njeguši, Montenegro (42.4307° N, 18.8115° E) with speleologist, P. Kunisch. Credit: Péter Lenkei. C. Entrance to Golubova pećina (42.2093° N, 19.1306° E), Gornja Seoca, Montenegro. Credit: I. Njunjić." figureDoi="http://doi.org/10.5281/zenodo.10847870" httpUri="https://zenodo.org/record/10847870/files/figure.png" pageId="43" pageNumber="44">22C</figureCitation>
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<emphasis id="B96DEA99FF91FFEBFF42D1A2FEE2F968" bold="true" box="[189,310,1698,1724]" pageId="43" pageNumber="44">Diagnosis</emphasis>
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Shell 
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, conical with 4.5–5 flatly convex, regular, tightly coiled whorls; aperture elliptical-ovate to subquadrate-oblique; columellar side of peristome thickly callused; teleoconch sculpture with conspicuous irregular scratch-like growth lines and milky, horizontal banding on the penultimate and body whorls.
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<emphasis id="B96DEA99FF91FFEBFF42D07CFE90F842" bold="true" box="[189,324,1916,1942]" pageId="43" pageNumber="44">Etymology</emphasis>
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This species is named in honour of Prof. Dr.Annette Klussmann-Kolb for providing valuable mentorship and introducing AJ to the 
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.
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<emphasis id="B96DEA99FF96FFECFF42D609FEB8FEF6" bold="true" box="[189,364,264,291]" pageId="44" pageNumber="45">Type material</emphasis>
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</paragraph>
<paragraph id="8BA6368BFF96FFECFF42D65AFE26FDD6" blockId="44.[189,1399,311,514]" pageId="44" pageNumber="45">
<collectingCountry id="F30E761BFF96FFECFF42D65AFE5EFEA0" box="[189,394,346,372]" name="Serbia" pageId="44" pageNumber="45">MONTENEGRO</collectingCountry>
• [holotype and body destroyed for DNA extraction, photograph and DNA are the only remnants of the holotype]; 
<collectingCounty id="62C74E07FF96FFECFDCBD67DFD46FE43" box="[564,658,381,407]" pageId="44" pageNumber="45">Njeguši</collectingCounty>
, 
<location id="8EC66050FF96FFECFD61D67EFC9CFE4C" LSID="urn:lsid:plazi:treatment:03B0879DFF91FFEDFE8DD11BFB77FDD5:8EC66050FF96FFECFD61D67EFC9CFE4C" box="[670,840,381,408]" country="Serbia" county="Njegusi" latitude="42.4307" longLatPrecision="6" longitude="18.8115" municipality="St John's cave" name="St John's cave" pageId="44" pageNumber="45">St John’s cave</location>
; 
<geoCoordinate id="EE2D504CFF96FFECFCABD67EFC0AFE4C" box="[852,990,382,408]" degrees="42.4307" direction="north" orientation="latitude" pageId="44" pageNumber="45" precision="5" value="42.4307">42.4307° N</geoCoordinate>
, 
<geoCoordinate id="EE2D504CFF96FFECFC16D67DFBBAFE4C" box="[1001,1134,381,408]" degrees="18.8115" direction="east" orientation="longitude" pageId="44" pageNumber="45" precision="5" value="18.8115">18.8115° E</geoCoordinate>
, 
<elevation id="0034D1B8FF96FFECFB85D67DFAD0FE43" box="[1146,1284,381,408]" metricMagnitude="2" metricUnit="m" metricValue="9.15" pageId="44" pageNumber="45" unit="m" value="915.0">
<quantity id="4CE19B6EFF96FFECFB85D67DFB10FE4C" box="[1146,1220,381,408]" metricMagnitude="2" metricUnit="m" metricValue="9.15" pageId="44" pageNumber="45" unit="m" value="915.0">915 m</quantity>
a.s.l.
</elevation>
; ca - 
<quantity id="4CE19B6EFF96FFECFAC5D67DFAA3FE4C" box="[1338,1399,381,408]" metricMagnitude="1" metricUnit="m" metricValue="8.5" pageId="44" pageNumber="45" unit="m" value="85.0">85 m</quantity>
vertical depth below the cave entrance; 
<date id="FFA7104BFF96FFECFD6BD6A0FCFEFE6F" box="[660,810,416,443]" pageId="44" pageNumber="45" value="2021-08-05">
<collectingDate id="EFE3E9A3FF96FFECFD6BD6A0FCFEFE6F" box="[660,810,416,443]" pageId="44" pageNumber="45" value="2021-08-05">5 Aug. 2021</collectingDate>
</date>
; 
<collectorName id="26EC535DFF96FFECFCC8D6A1FC01FE6F" box="[823,981,417,443]" pageId="44" pageNumber="45">László Dányi</collectorName>
&amp; 
<collectorName id="26EC535DFF96FFECFC03D6A1FB10FE6F" box="[1020,1220,417,443]" pageId="44" pageNumber="45">Nikolett Ujhegyi</collectorName>
leg.; GenBank CO1 gene: ON037484; GenBank 16S gene: ON041449; GenBank H3 gene: ON088652; imaged shell of aliquot 
<specimenCode id="DBBF9EF0FF96FFECFEC8D6E8FE3AFDD6" box="[311,494,487,514]" collectionCode="NMBE" country="Switzerland" httpUri="http://grbio.org/cool/5hwi-0wgz" name="Naturhistorisches Museum der Burgergemeinde Bern" pageId="44" pageNumber="45">NMBE 571122</specimenCode>
.
</paragraph>
</materialsCitation>
<paragraph id="8BA6368BFF96FFECFF15D529FE88FD97" blockId="44.[189,1399,553,720]" box="[234,348,553,579]" pageId="44" pageNumber="45">
<emphasis id="B96DEA99FF96FFECFF15D529FE88FD97" bold="true" box="[234,348,553,579]" pageId="44" pageNumber="45">
<typeStatus id="54A28829FF96FFECFF15D529FE88FD97" box="[234,348,553,579]" pageId="44" pageNumber="45" type="paratype">Paratype</typeStatus>
</emphasis>
</paragraph>
<paragraph id="8BA6368BFF96FFECFF42D54CFEAEFD04" blockId="44.[189,1399,553,720]" pageId="44" pageNumber="45">
<materialsCitation id="3B713CD6FF96FFECFF42D54CFEAEFD04" ID-GBIF-Occurrence="4598261330" collectingDate="2021-08-05" collectionCode="NMBE" collectorName="Laszlo Danyi &amp; Nikolett Ujhegyi" country="Serbia" county="Njegusi" elevation="915" latitude="42.4307" location="MONTENEGRO" longLatPrecision="6" longitude="18.8115" municipality="St John's cave" pageId="44" pageNumber="45" specimenCode="NMBE 571123" specimenCount="1" typeStatus="paratype">
<collectingCountry id="F30E761BFF96FFECFF42D54CFE5EFDB2" box="[189,394,588,614]" name="Serbia" pageId="44" pageNumber="45">MONTENEGRO</collectingCountry>
• 
<specimenCount id="9D1FFD02FF96FFECFE56D54CFE2AFDB2" box="[425,510,588,614]" count="1" pageId="44" pageNumber="45" type="generic">1 spec.</specimenCount>
[
<typeStatus id="54A28829FF96FFECFDF0D54CFDA3FDB2" box="[527,631,588,614]" pageId="44" pageNumber="45">paratype</typeStatus>
and body destroyed for DNA extraction, photograph and DNA are the only remnants of the paratype]; same collection data as for holotype; GenBank CO1 gene: ON037485; GenBank 16S gene: ON041450; GenBank H3 gene: ON088653; imaged shell of aliquot 
<specimenCode id="DBBF9EF0FF96FFECFF40D5B6FEA2FD04" box="[191,374,693,720]" pageId="44" pageNumber="45">NMBE 571123</specimenCode>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="C3036500FF96FFECFF42D5F6FBA0FB69" pageId="44" pageNumber="45" type="description">
<paragraph id="8BA6368BFF96FFECFF42D5F6FE9AFCC4" blockId="44.[189,334,758,784]" box="[189,334,758,784]" pageId="44" pageNumber="45">
<heading id="D0EE81E7FF96FFECFF42D5F6FE9AFCC4" bold="true" box="[189,334,758,784]" fontSize="11" level="3" pageId="44" pageNumber="45" reason="3">
<emphasis id="B96DEA99FF96FFECFF42D5F6FE9AFCC4" bold="true" box="[189,334,758,784]" pageId="44" pageNumber="45">Description</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFF96FFECFF42D426FCB1FC7D" blockId="44.[189,1400,805,938]" pageId="44" pageNumber="45">
<smallCapsWord id="8D40A057FF96FFECFF42D426FEBAFCEA" baselines="825,825" box="[189,366,806,832]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Measurements" pageId="44" pageNumber="45">MEASUREMENTS</smallCapsWord>
. Snails processed for DNA sequence data and thus, the type series has been destroyed. 
<typeStatus id="54A28829FF96FFECFF41D448FEE4FCB6" box="[190,304,840,866]" pageId="44" pageNumber="45" type="holotype">Holotype</typeStatus>
: sh: 
<quantity id="4CE19B6EFF96FFECFE94D449FE0EFCB7" box="[363,474,841,867]" metricMagnitude="-3" metricUnit="m" metricValue="1.38" pageId="44" pageNumber="45" unit="mm" value="1.38">1.38 mm</quantity>
; sw: 1.00 mm; ah: 
<quantity id="4CE19B6EFF96FFECFD35D448FCEDFCB7" box="[714,825,840,867]" metricMagnitude="-4" metricUnit="m" metricValue="6.6" pageId="44" pageNumber="45" unit="mm" value="0.66">0.66 mm</quantity>
; aw: 
<quantity id="4CE19B6EFF96FFECFC82D448FC38FCB7" box="[893,1004,840,867]" metricMagnitude="-4" metricUnit="m" metricValue="5.8999999999999995" pageId="44" pageNumber="45" unit="mm" value="0.59">0.59 mm</quantity>
; hlw: 
<quantity id="4CE19B6EFF96FFECFBC5D448FB7DFCB7" box="[1082,1193,840,867]" metricMagnitude="-4" metricUnit="m" metricValue="9.6" pageId="44" pageNumber="45" unit="mm" value="0.96">0.96 mm</quantity>
; SA: 75.69 deg.; NMBE 571122 (
<figureCitation id="13222A0EFF96FFECFE7ED46CFE3BFC51" box="[385,495,875,902]" captionStart="Fig" captionStartId="45.[189,232,1787,1813]" captionTargetBox="[220,1366,561,1748]" captionTargetId="figure-212@45.[220,1366,561,1748]" captionTargetPageId="45" captionText="Fig. 16. Light microscopic images of full-bodied Zospeum Bourguignat, 1856 from Njeguši, St John’s cave. A–B. Zospeum kolbae Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner sp. nov. (NMBE 571122– 571123), individuals assessed by DNA sequencing with sigmoid intestine showing through shells (shells were destroyed post imaging for tissue extraction). A. Holotype (NMBE 571122), shell of aliquot, aperture, and aperture facing left view. B. Paratype (NMBE 571123), shell of aliquot, aperture, and aperture facing left view. C. Undescribed Zospeum sp. 1 (NMBE 577052) showing all perspectives. D–E. Undescribed Zospeum sp. 1 (NMBE 577053/2) showing all perspectives." figureDoi="http://doi.org/10.5281/zenodo.10847850" httpUri="https://zenodo.org/record/10847850/files/figure.png" pageId="44" pageNumber="45">Fig. 16A</figureCitation>
). 
<typeStatus id="54A28829FF96FFECFDFBD46CFDBBFC52" box="[516,623,876,902]" pageId="44" pageNumber="45" type="paratype">Paratype</typeStatus>
: (N = 1): sh: 
<quantity id="4CE19B6EFF96FFECFCF2D46BFCADFC52" box="[781,889,875,902]" metricMagnitude="-3" metricUnit="m" metricValue="1.59" pageId="44" pageNumber="45" unit="mm" value="1.59">1.59 mm</quantity>
; sw: 
<quantity id="4CE19B6EFF96FFECFC49D46BFBF7FC52" box="[950,1059,875,902]" metricMagnitude="-4" metricUnit="m" metricValue="9.5" pageId="44" pageNumber="45" unit="mm" value="0.95">0.95 mm</quantity>
; ah: 
<quantity id="4CE19B6EFF96FFECFBA4D46CFB13FC52" box="[1115,1223,876,902]" metricMagnitude="-4" metricUnit="m" metricValue="7.3" pageId="44" pageNumber="45" unit="mm" value="0.73">0.73 mm</quantity>
; aw: 
<quantity id="4CE19B6EFF96FFECFAF9D46BFAA6FC52" box="[1286,1394,875,902]" metricMagnitude="-4" metricUnit="m" metricValue="6.2" pageId="44" pageNumber="45" unit="mm" value="0.62">0.62 mm</quantity>
; hlw: 
<quantity id="4CE19B6EFF96FFECFF06D48FFE89FC7D" box="[249,349,911,937]" metricMagnitude="-4" metricUnit="m" metricValue="7.3" pageId="44" pageNumber="45" unit="mm" value="0.73">0.73mm</quantity>
; SA: 62.88 deg.; NMBE 571123 (
<figureCitation id="13222A0EFF96FFECFD13D48FFC83FC7D" box="[748,855,911,937]" captionStart="Fig" captionStartId="45.[189,232,1787,1813]" captionTargetBox="[220,1366,561,1748]" captionTargetId="figure-212@45.[220,1366,561,1748]" captionTargetPageId="45" captionText="Fig. 16. Light microscopic images of full-bodied Zospeum Bourguignat, 1856 from Njeguši, St John’s cave. A–B. Zospeum kolbae Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner sp. nov. (NMBE 571122– 571123), individuals assessed by DNA sequencing with sigmoid intestine showing through shells (shells were destroyed post imaging for tissue extraction). A. Holotype (NMBE 571122), shell of aliquot, aperture, and aperture facing left view. B. Paratype (NMBE 571123), shell of aliquot, aperture, and aperture facing left view. C. Undescribed Zospeum sp. 1 (NMBE 577052) showing all perspectives. D–E. Undescribed Zospeum sp. 1 (NMBE 577053/2) showing all perspectives." figureDoi="http://doi.org/10.5281/zenodo.10847850" httpUri="https://zenodo.org/record/10847850/files/figure.png" pageId="44" pageNumber="45">Fig. 16B</figureCitation>
).
</paragraph>
<paragraph id="8BA6368BFF96FFECFF42D4D0FBA0FB69" blockId="44.[189,1399,976,1214]" pageId="44" pageNumber="45">
Shell 
<quantity id="4CE19B6EFF96FFECFEF9D4D0FE6DFC3F" box="[262,441,976,1003]" metricMagnitude="-3" metricUnit="m" metricValue="1.4849999999999999" metricValueMax="1.59" metricValueMin="1.38" pageId="44" pageNumber="45" unit="mm" value="1.4849999999999999" valueMax="1.59" valueMin="1.38">1.38–1.59 mm</quantity>
, conical; transparent; with 4.5–5 flatly convex, regular, tightly coiled whorls; penultimate whorl equal or slightly over ½ height of body whorl (in lateral view); aperture elliptical-ovate to subquadrate; columellar side long; suture shallow; teleoconch sculpture with conspicuous irregular scratch-like growth lines and milky, horizontal banding on the penultimate and body whorls; some weak axial ribbing present immediately behind palatal lip; palatal lip narrowly reflected, palatal-basal side reflected; peristome thickly callused on basal columellar side; parietal shield long and straight with conspicuous narrow notch indentation at upper parietal and palatal junction.
</paragraph>
</subSubSection>
<subSubSection id="C3036500FF96FFECFF42D3E5FB36FAFA" box="[189,1250,1253,1326]" pageId="44" pageNumber="45" type="distribution">
<paragraph id="8BA6368BFF96FFECFF42D3E5FE82FB2B" blockId="44.[189,342,1253,1279]" box="[189,342,1253,1279]" pageId="44" pageNumber="45">
<heading id="D0EE81E7FF96FFECFF42D3E5FE82FB2B" bold="true" box="[189,342,1253,1279]" fontSize="11" level="3" pageId="44" pageNumber="45" reason="3">
<emphasis id="B96DEA99FF96FFECFF42D3E5FE82FB2B" bold="true" box="[189,342,1253,1279]" pageId="44" pageNumber="45">Distribution</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFF96FFECFF42D213FB36FAFA" blockId="44.[189,1250,1299,1326]" box="[189,1250,1299,1326]" pageId="44" pageNumber="45">
This species is only known from the 
<typeStatus id="54A28829FF96FFECFD91D214FD75FAFA" box="[622,673,1300,1326]" pageId="44" pageNumber="45">type</typeStatus>
locality, St John’s cave, in Njeguši, 
<collectingCountry id="F30E761BFF96FFECFBB4D214FB0AFAFA" box="[1099,1246,1300,1326]" name="Serbia" pageId="44" pageNumber="45">Montenegro</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="C3036500FF96FFEDFF42D255FB77FDD5" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45" type="discussion">
<paragraph id="8BA6368BFF96FFECFF42D255FEFBFABB" blockId="44.[189,303,1365,1391]" box="[189,303,1365,1391]" pageId="44" pageNumber="45">
<heading id="D0EE81E7FF96FFECFF42D255FEFBFABB" bold="true" box="[189,303,1365,1391]" fontSize="11" level="3" pageId="44" pageNumber="45" reason="3">
<emphasis id="B96DEA99FF96FFECFF42D255FEFBFABB" bold="true" box="[189,303,1365,1391]" pageId="44" pageNumber="45">Remarks</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFF96FFECFF42D283FAB7F805" blockId="44.[188,1399,1411,2001]" pageId="44" pageNumber="45">
The two shells demonstrate a significant range in shell height (
<quantity id="4CE19B6EFF96FFECFC29D283FB58FA4A" box="[982,1164,1411,1438]" metricMagnitude="-3" metricUnit="m" metricValue="1.49" metricValueMax="1.59" metricValueMin="1.39" pageId="44" pageNumber="45" unit="mm" value="1.49" valueMax="1.59" valueMin="1.39">1.39–1.59 mm</quantity>
), constituting what might be the ends of the size spectrum for 
<taxonomicName id="4C194D08FF96FFECFD31D2A7FAA3FA15" authority="Jochum, Inabnit, Kneubuhler &amp; Ruthensteiner" authorityName="Jochum, Inabnit, Kneubuhler &amp; Ruthensteiner" authorityYear="2024" box="[718,1399,1447,1473]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="kolbae" status="sp. nov.">
<emphasis id="B96DEA99FF96FFECFD31D2A7FC94FA15" box="[718,832,1447,1473]" italics="true" pageId="44" pageNumber="45">Z. kolbae</emphasis>
Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FF96FFECFF42D2CAFECEFA30" box="[189,282,1482,1508]" pageId="44" pageNumber="45" rank="species">sp. nov.</taxonomicNameLabel>
or that shell B is a potential hybrid with 
<taxonomicName id="4C194D08FF96FFECFCE8D2CBFADBFA30" authority="Jochum &amp; Ruthensteiner" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[791,1295,1482,1508]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="dubokidoense" status="sp. nov.">
<emphasis id="B96DEA99FF96FFECFCE8D2CBFC08FA30" box="[791,988,1482,1508]" italics="true" pageId="44" pageNumber="45">Z. dubokidoense</emphasis>
Jochum &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FF96FFECFAE6D2CAFAA3FA30" box="[1305,1399,1482,1508]" pageId="44" pageNumber="45" rank="species">sp. nov.</taxonomicNameLabel>
or a potential freak. Of all the shells in this study, shell B is by far the largest with that of one shell from 
<taxonomicName id="4C194D08FF96FFECFEFDD111FE1EF9FE" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[258,458,1552,1578]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="dubokidoense">
<emphasis id="B96DEA99FF96FFECFEFDD111FE1EF9FE" box="[258,458,1552,1578]" italics="true" pageId="44" pageNumber="45">Z. dubokidoense</emphasis>
</taxonomicName>
measuring a close second in shell height (
<quantity id="4CE19B6EFF96FFECFC1DD110FBB1F9FE" box="[994,1125,1552,1579]" metricMagnitude="-3" metricUnit="m" metricValue="1.546" pageId="44" pageNumber="45" unit="mm" value="1.546">1.546 mm</quantity>
). With its molecularly closest congener, 
<taxonomicName id="4C194D08FF96FFECFE6BD134FDDAF999" authorityName="Inabnit, Jochum &amp; Neubert" authorityYear="2021" box="[404,526,1587,1613]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="simplex">
<emphasis id="B96DEA99FF96FFECFE6BD134FDDAF999" box="[404,526,1587,1613]" italics="true" pageId="44" pageNumber="45">Z. simplex</emphasis>
</taxonomicName>
, (1.41– 
<quantity id="4CE19B6EFF96FFECFD95D134FD0CF99A" box="[618,728,1588,1614]" metricMagnitude="-3" metricUnit="m" metricValue="1.4" pageId="44" pageNumber="45" unit="mm" value="1.4">1.40 mm</quantity>
) it is comparable at the lower end of the size range but demonstrates a much broader spire angle than that of the narrower 
<taxonomicName id="4C194D08FF96FFECFC16D157FBB1F9A5" authorityName="Inabnit, Jochum &amp; Neubert" authorityYear="2021" box="[1001,1125,1623,1649]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="simplex">
<emphasis id="B96DEA99FF96FFECFC16D157FBB1F9A5" box="[1001,1125,1623,1649]" italics="true" pageId="44" pageNumber="45">Z. simplex</emphasis>
</taxonomicName>
and a markedly larger aperture (ah) compared to that of 
<taxonomicName id="4C194D08FF96FFECFDB3D17BFD12F940" authorityName="Inabnit, Jochum &amp; Neubert" authorityYear="2021" box="[588,710,1658,1684]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="simplex">
<emphasis id="B96DEA99FF96FFECFDB3D17BFD12F940" box="[588,710,1658,1684]" italics="true" pageId="44" pageNumber="45">Z. simplex</emphasis>
</taxonomicName>
. The elliptical-ovate to subquadrate shape of the aperture, the height of the penultimate whorl equaling or extending slightly over ½ the height of the body whorl (lateral view) are reminiscent of 
<taxonomicName id="4C194D08FF96FFECFDB2D1C1FCC7F90E" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[589,787,1728,1754]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="dubokidoense">
<emphasis id="B96DEA99FF96FFECFDB2D1C1FCC7F90E" box="[589,787,1728,1754]" italics="true" pageId="44" pageNumber="45">Z. dubokidoense</emphasis>
</taxonomicName>
from Duboki do cave (
<figureCitation id="13222A0EFF96FFECFBCBD1C0FB4DF90E" box="[1076,1177,1728,1754]" captionStart="Fig" captionStartId="28.[189,232,1857,1883]" captionTargetBox="[189,1398,887,1818]" captionTargetId="figure-222@28.[189,1398,887,1818]" captionTargetPageId="28" captionText="Fig. 6. Zospeum intermedium Jochum &amp; Ruthensteiner sp. nov. (RMNH.MOL.234132) from Gittenberger (1975). A. Light microscopic images of apertural and dorsal views. B. Sample labels. C–H. 3D visualizations of X-ray Micro-CT data. C. Aperture view. D. Aperture facing right view. E. Apical view. F. Dorsal view. G. Aperture facing right view. H. Umbilical view showing slight umbilical depression and lamella projecting from the columellar side." figureDoi="http://doi.org/10.5281/zenodo.10847820" httpUri="https://zenodo.org/record/10847820/files/figure.png" pageId="44" pageNumber="45">Figs 6D</figureCitation>
, 
<figureCitation id="13222A0EFF96FFECFB57D1C0FB1AF90E" box="[1192,1230,1728,1754]" captionStart="Fig" captionStartId="30.[188,231,1857,1883]" captionTargetBox="[190,1399,1183,1819]" captionTargetId="figure-365@30.[190,1399,1183,1819]" captionTargetPageId="30" captionText="Fig. 7. Lightmicroscopic images of shells from the caves Lipska pećina and Duboki do (Montenegro) with specimen labels. A–C. Zospeum troglobalcanicum Absolon, 1916 (MCSMNH-PMSL-Moll.-FVelkovrh 29603). D–F. Zospeum dubokidoense Jochum &amp; Ruthensteiner sp. nov. D. Holotype (MCSMNH- PMSL-Moll.-FVelkovrh 30360[spm1]). E–F. Paratypes, Duboki do, Montenegro (MCSMNH-PMSL- Moll.-FVelkovrh 30360[spm2-3])." figureDoi="http://doi.org/10.5281/zenodo.10847822" httpUri="https://zenodo.org/record/10847822/files/figure.png" pageId="44" pageNumber="45">7G</figureCitation>
) (MCSMNHPMSL-Moll.-FVelkovrh 30360[spm1]). Their affinity in conjunction with the recent discovery of a connection between St John’s cave and Duboki do cave (
<bibRefCitation id="EF884B7AFF96FFECFC99D007FB71F8F5" author="Csepreghy F. &amp; Lenkei P." box="[870,1189,1799,1825]" pageId="44" pageNumber="45" pagination="7" refId="ref30808" refString="Csepreghy F. &amp; Lenkei P. 2021. A Szent Janos-bg. osszekotese a Duboki Do-val, egyeb feltarasok, jovobeli lehetosegek. In: Kosztra B. (ed.) Book of Program and Abstracts 26. Barlangkutatok &quot; Szablyar Peter &quot; Szakmai Talalkozoja - 2021. November 19 - 21: 7. Tes, MKBT, Budapest." type="journal article" year="2021">Csepreghy &amp; Lenkei 2021</bibRefCitation>
) may be clarified by future sampling opportunities and DNA sequence analysis. Imaged with 
<taxonomicName id="4C194D08FF96FFECFBD0D02BFB49F890" authorityName="Jochum, Inabnit, Kneubuhler &amp; Ruthensteiner" authorityYear="2024" box="[1071,1181,1834,1860]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="kolbae">
<emphasis id="B96DEA99FF96FFECFBD0D02BFB49F890" box="[1071,1181,1834,1860]" italics="true" pageId="44" pageNumber="45">Z. kolbae</emphasis>
</taxonomicName>
, is an undescribed species (
<taxonomicName id="4C194D08FF96FFECFEDBD04EFE19F8B3" box="[292,461,1869,1896]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="undefined-1">
<emphasis id="B96DEA99FF96FFECFEDBD04EFE5BF8B3" box="[292,399,1870,1895]" italics="true" pageId="44" pageNumber="45">Zospeum</emphasis>
sp. 1
</taxonomicName>
) (
<figureCitation id="13222A0EFF96FFECFE16D04DFDA0F8B3" box="[489,628,1869,1895]" captionStart="Fig" captionStartId="45.[189,232,1787,1813]" captionTargetBox="[220,1366,561,1748]" captionTargetId="figure-212@45.[220,1366,561,1748]" captionTargetPageId="45" captionText="Fig. 16. Light microscopic images of full-bodied Zospeum Bourguignat, 1856 from Njeguši, St John’s cave. A–B. Zospeum kolbae Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner sp. nov. (NMBE 571122– 571123), individuals assessed by DNA sequencing with sigmoid intestine showing through shells (shells were destroyed post imaging for tissue extraction). A. Holotype (NMBE 571122), shell of aliquot, aperture, and aperture facing left view. B. Paratype (NMBE 571123), shell of aliquot, aperture, and aperture facing left view. C. Undescribed Zospeum sp. 1 (NMBE 577052) showing all perspectives. D–E. Undescribed Zospeum sp. 1 (NMBE 577053/2) showing all perspectives." figureDoi="http://doi.org/10.5281/zenodo.10847850" httpUri="https://zenodo.org/record/10847850/files/figure.png" pageId="44" pageNumber="45">Fig. 16C–E</figureCitation>
) (NMBE 577052–577053) found in the same cave chamber with 
<specimenCount id="9D1FFD02FF96FFECFF41D071FEE3F85E" box="[190,311,1904,1931]" count="2" pageId="44" pageNumber="45" type="adult">two adults</specimenCount>
showing a substantially smaller size (
<quantity id="4CE19B6EFF96FFECFD0DD070FC4DF85E" box="[754,921,1904,1930]" metricMagnitude="-2" metricUnit="m" metricValue="6.5645" metricValueMax="13.0" metricValueMin="0.129" pageId="44" pageNumber="45" unit="mm" value="65.645" valueMax="130.0" valueMin="1.29">1.29–130 mm</quantity>
) than that of 
<taxonomicName id="4C194D08FF96FFECFBCDD071FB75F85E" authorityName="Jochum, Inabnit, Kneubuhler &amp; Ruthensteiner" authorityYear="2024" box="[1074,1185,1904,1930]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="kolbae">
<emphasis id="B96DEA99FF96FFECFBCDD071FB75F85E" box="[1074,1185,1904,1930]" italics="true" pageId="44" pageNumber="45">Z. kolbae</emphasis>
</taxonomicName>
. A third, subfossil species, found in the dry part of the same cave, 
<taxonomicName id="4C194D08FF96FFECFD23D094FB79F879" authority="Jochum &amp; Ruthensteiner" authorityName="Jochum &amp; Ruthensteiner" authorityYear="2024" box="[732,1197,1939,1966]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="njegusiense" status="sp. nov.">
<emphasis id="B96DEA99FF96FFECFD23D094FC50F879" box="[732,900,1940,1965]" italics="true" pageId="44" pageNumber="45">Z. njegusiense</emphasis>
Jochum &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FF96FFECFB4DD094FAD8F87A" box="[1202,1292,1940,1966]" pageId="44" pageNumber="45" rank="species">sp. nov.</taxonomicNameLabel>
(
<figureCitation id="13222A0EFF96FFECFAE5D093FAB9F87A" box="[1306,1389,1939,1966]" captionStart="Fig" captionStartId="46.[189,232,1822,1848]" captionTargetBox="[189,1398,1058,1782]" captionTargetId="figure-216@46.[189,1398,1058,1782]" captionTargetPageId="46" captionText="Fig. 17. Z. njegusiense Jochum &amp; Ruthensteiner sp. nov., Scanning Electron Microscopy images. A. Holotype (NMBE 572639), aperture view showing subfossil shell with superficial erosion of shell layer. B. Paratype (NMBE 578378), aperture view showing close up of lamellar band on columella (Fig. C). C. Paratype (NMBE 578378), aperture view showing damaged shell and columella. D. Close up view of holotype (NMBE 572639), showing body whorl with low, irregular growth lines. E. Paratype (NMBE 578378), protoconch." figureDoi="http://doi.org/10.5281/zenodo.10847854" httpUri="https://zenodo.org/record/10847854/files/figure.png" pageId="44" pageNumber="45">Fig. 17</figureCitation>
) measures within the same size range as the two shells of 
<taxonomicName id="4C194D08FF96FFECFCA7D0B7FC13F805" authorityName="Jochum, Inabnit, Kneubuhler &amp; Ruthensteiner" authorityYear="2024" box="[856,967,1975,2001]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="44" pageNumber="45" phylum="Mollusca" rank="species" species="kolbae">
<emphasis id="B96DEA99FF96FFECFCA7D0B7FC13F805" box="[856,967,1975,2001]" italics="true" pageId="44" pageNumber="45">Z. kolbae</emphasis>
</taxonomicName>
but shows no other affinity with it.
</paragraph>
<paragraph id="8BA6368BFF97FFEDFF42D609FE89FEF6" blockId="45.[189,349,264,291]" box="[189,349,264,291]" pageId="45" pageNumber="46">
<heading id="D0EE81E7FF97FFEDFF42D609FE89FEF6" bold="true" box="[189,349,264,291]" fontSize="11" level="3" pageId="45" pageNumber="46" reason="3">
<emphasis id="B96DEA99FF97FFEDFF42D609FE89FEF6" bold="true" box="[189,349,264,291]" pageId="45" pageNumber="46">Type locality</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFF97FFEDFF42D637FB77FDD5" blockId="45.[189,1399,311,514]" pageId="45" pageNumber="46">
According to the collector, László Dányi, “The small chamber, where all the snails were collected, is really a rather dry part of the cave, a so called ‘fossil’ part which doesn’t have any active water-flow – even temporarily – since ages. However, on the rocks there is a thin layer of clay which is wet around micro fissures because of percolating water.” (
<figureCitation id="13222A0EFF97FFEDFD1DD6A1FC9BFE6F" box="[738,847,417,443]" captionStart="Fig" captionStartId="55.[189,232,1822,1848]" captionTargetBox="[191,1400,633,1782]" captionTargetId="figure-236@55.[191,1400,621,1782]" captionTargetPageId="55" captionText="Fig. 22. Sites of localities and collections. A. Site of type locality of Zospeum njunjicae Jochum, Schilthuizen &amp; Ruthensteiner sp. nov., Golubova pećina, Gornja Seoca, Montenegro. Credit: I. Njunjić. B. Collection site within St John’s cave, type locality of Z. kolbae Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner sp. nov. and Z. njegusiense Jochum &amp; Ruthensteiner sp. nov., Njeguši, Montenegro (42.4307° N, 18.8115° E) with speleologist, P. Kunisch. Credit: Péter Lenkei. C. Entrance to Golubova pećina (42.2093° N, 19.1306° E), Gornja Seoca, Montenegro. Credit: I. Njunjić." figureDoi="http://doi.org/10.5281/zenodo.10847870" httpUri="https://zenodo.org/record/10847870/files/figure.png" pageId="45" pageNumber="46">Fig. 22B</figureCitation>
). Maximum depth of St John’s cave is - 
<quantity id="4CE19B6EFF97FFEDFAD6D6A0FAA3FE6F" box="[1321,1399,416,443]" metricMagnitude="2" metricUnit="m" metricValue="5.37" pageId="45" pageNumber="46" unit="m" value="537.0">537 m</quantity>
(László Dányi pers. comm 
<date id="FFA7104BFF97FFEDFDFDD6C4FD55FE0A" box="[514,641,452,478]" pageId="45" pageNumber="46" value="2022-05">May 2022</date>
). Recent exploration by the Hungarian Speleo Camp detected a connection between St John’s cave and Duboki do cave (
<bibRefCitation id="EF884B7AFF97FFEDFCA6D6E7FB42FDD5" author="Csepreghy F. &amp; Lenkei P." box="[857,1174,487,513]" pageId="45" pageNumber="46" pagination="7" refId="ref30808" refString="Csepreghy F. &amp; Lenkei P. 2021. A Szent Janos-bg. osszekotese a Duboki Do-val, egyeb feltarasok, jovobeli lehetosegek. In: Kosztra B. (ed.) Book of Program and Abstracts 26. Barlangkutatok &quot; Szablyar Peter &quot; Szakmai Talalkozoja - 2021. November 19 - 21: 7. Tes, MKBT, Budapest." type="journal article" year="2021">Csepreghy &amp; Lenkei 2021</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>