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<mods:title id="6029B01192F2A89058A1A175BD45C81C">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="E5AA97B6A4EDA861669BA8E6A75F927B">Jochum, Adrienne</mods:namePart>
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<mods:namePart id="75E025D7CED259E42CB139220F85CD2B">Michalik, Peter</mods:namePart>
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<mods:affiliation id="BF9E2A96A0254E5A783EB19ABC39B2CB">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:namePart id="CB443733EF77843DC98B8F0F3C2B0959">Vrabec, Marko</mods:namePart>
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<mods:affiliation id="21C605C5312298F010972EB339579E05">Department of Geology, Faculty of Natural Sciences and Engineering, Aškerčeva 12, University of Ljubljana, 1000 Ljubljana, Slovenia.</mods:affiliation>
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<mods:affiliation id="7CFC3F3E8072C0878F52AA431479C8C5">Naturalis Biodiversity Center and Leiden University, Vondellaan 55, NL- 2332 AA Leiden, The Netherlands.</mods:affiliation>
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<mods:namePart id="EC416A0BAEBDFAA575D6E96788EC8F7B">Ruthensteiner, Bernhard</mods:namePart>
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<mods:affiliation id="2AD4CB21F5C12587197A7491B9343AF6">SNSB - Zoologische Staatssammlung München (ZSM), 81247 München, Germany.</mods:affiliation>
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<emphasis id="B96DEA99FFA1FFDBFE35D3A9FD09FB17" bold="true" box="[458,733,1192,1219]" italics="true" pageId="27" pageNumber="28">Zospeum intermedium</emphasis>
Jochum &amp; Ruthensteiner
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urn:lsid:zoobank.org:act:
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<paragraph id="8BA6368BFFA1FFDBFD08D3EFFCE8FADD" blockId="27.[361,1226,1192,1289]" box="[759,828,1263,1289]" pageId="27" pageNumber="28">
<figureCitation id="13222A0EFFA1FFDBFD08D3EFFCE8FADD" box="[759,828,1263,1289]" 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. CH. 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="27" pageNumber="28">Fig. 6</figureCitation>
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<taxonomicName id="4C194D08FFA1FFDBFF42D231FDD5FA9E" ID-CoL="7TPBF" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[189,513,1328,1354]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="27" pageNumber="28" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FFA1FFDBFF42D231FDD5FA9E" box="[189,513,1328,1354]" italics="true" pageId="27" pageNumber="28">Zospeum troglobalcanicum</emphasis>
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Maier in
<treatmentCitation id="0AB8109AFFA1FFDBFD75D230FC45FA9E" author="Gittenberger E." box="[650,913,1328,1355]" page="26" pageId="27" pageNumber="28" year="1975">
<bibRefCitation id="EF884B7AFFA1FFDBFD75D230FC45FA9E" author="Gittenberger E." box="[650,913,1328,1355]" pageId="27" pageNumber="28" pagination="21 - 37" refId="ref31037" refString="Gittenberger E. 1975. Cave snails found in southern Crna Gora. Glasnik Republickog zavoda za zastitutu prirode i Prirodnjackog muzeja 8: 21 - 37." type="journal article" year="1975">Gittenberger 1975: 26</bibRefCitation>
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: fig. 3.
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<emphasis id="B96DEA99FFA1FFDBFF42D271FEE2FA5F" bold="true" box="[189,310,1393,1419]" pageId="27" pageNumber="28">Diagnosis</emphasis>
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A tall shell ca
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, conical or elongate-conical with ca 5½ regularly coiled, slightly shouldered whorls; columella bears a well-formed lamella forming a conspicuous and short parietalis visible in umbilical view (
<figureCitation id="13222A0EFFA1FFDBFE83D2E7FE0EF9D5" box="[380,474,1511,1537]" 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. CH. 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="27" pageNumber="28">Fig. 6H</figureCitation>
); ventral side swollen with small umbilical depression.
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<emphasis id="B96DEA99FFA1FFDBFF42D128FE90F996" bold="true" box="[189,324,1576,1602]" pageId="27" pageNumber="28">Etymology</emphasis>
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This species is named after its mixture of significant shell characteristics in four species of
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<emphasis id="B96DEA99FFA1FFDBFAF3D157FAA3F9A4" box="[1292,1399,1623,1648]" italics="true" pageId="27" pageNumber="28">Zospeum</emphasis>
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from nearby caves described in this study, making for a composite, intermediary form of these species.
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<subSubSection id="C3036500FFA1FFDBFF42D1BBFC25F89E" pageId="27" pageNumber="28" type="materials_examined">
<paragraph id="8BA6368BFFA1FFDBFF42D1BBFEB8F901" blockId="27.[189,364,1723,1749]" box="[189,364,1723,1749]" pageId="27" pageNumber="28">
<heading id="D0EE81E7FFA1FFDBFF42D1BBFEB8F901" bold="true" box="[189,364,1723,1749]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="3">
<emphasis id="B96DEA99FFA1FFDBFF42D1BBFEB8F901" bold="true" box="[189,364,1723,1749]" pageId="27" pageNumber="28">Type material</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFFA1FFDBFF15D1EAFE88F8D0" blockId="27.[189,1399,1770,1867]" box="[234,348,1770,1796]" pageId="27" pageNumber="28">
<emphasis id="B96DEA99FFA1FFDBFF15D1EAFE88F8D0" bold="true" box="[234,348,1770,1796]" pageId="27" pageNumber="28">
<typeStatus id="54A28829FFA1FFDBFF15D1EAFE88F8D0" box="[234,348,1770,1796]" pageId="27" pageNumber="28" type="holotype">Holotype</typeStatus>
</emphasis>
</paragraph>
<paragraph id="8BA6368BFFA1FFDBFF42D00DFC25F89E" blockId="27.[189,1399,1770,1867]" pageId="27" pageNumber="28">
<materialsCitation id="3B713CD6FFA1FFDBFF42D00DFC25F89E" ID-GBIF-Occurrence="4598261322" collectingDate="1974-05-15" collectionCode="RMNH" collectorName="E. Gittenberger" country="Serbia" county="Crna Gora" location="Cetinjska pecina" municipality="Cetinje" pageId="27" pageNumber="28" specimenCode="RMNH.MOL.234132" specimenCount="1" typeStatus="holotype">
<collectingCountry id="F30E761BFFA1FFDBFF42D00DFE5EF8F3" box="[189,394,1805,1831]" name="Serbia" pageId="27" pageNumber="28">MONTENEGRO</collectingCountry>
<location id="8EC66050FFA1FFDBFE5FD00DFDB5F8F3" LSID="urn:lsid:plazi:treatment:03B0879DFFA1FFDDFE35D3A9FDDDFCB5:8EC66050FFA1FFDBFE5FD00DFDB5F8F3" box="[416,609,1805,1831]" country="Serbia" county="Crna Gora" municipality="Cetinje" name="Cetinjska pecina" pageId="27" pageNumber="28">Cetinjska pećina</location>
[= Cetinska pećina] near
<collectingMunicipality id="6BC2ACF1FFA1FFDBFC78D00DFC0BF8F3" box="[903,991,1805,1831]" pageId="27" pageNumber="28">Cetinje</collectingMunicipality>
, Crna Gora [= Montenegro] behind monastery;
<date id="FFA7104BFFA1FFDBFEB8D030FE32F89E" box="[327,486,1840,1867]" pageId="27" pageNumber="28" value="1974-05-15">
<collectingDate id="EFE3E9A3FFA1FFDBFEB8D030FE32F89E" box="[327,486,1840,1867]" pageId="27" pageNumber="28" value="1974-05-15">15 May 1974</collectingDate>
</date>
;
<collectorName id="26EC535DFFA1FFDBFE0CD031FD73F89E" box="[499,679,1840,1867]" pageId="27" pageNumber="28">E. Gittenberger</collectorName>
leg.;
<specimenCode id="DBBF9EF0FFA1FFDBFD16D030FC39F89E" box="[745,1005,1840,1866]" pageId="27" pageNumber="28">RMNH.MOL.234132</specimenCode>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="C3036500FFA1FFDCFF42D071FD86FD38" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28" type="description">
<paragraph id="8BA6368BFFA1FFDBFF42D071FE9AF85F" blockId="27.[189,334,1905,1931]" box="[189,334,1905,1931]" pageId="27" pageNumber="28">
<heading id="D0EE81E7FFA1FFDBFF42D071FE9AF85F" bold="true" box="[189,334,1905,1931]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="3">
<emphasis id="B96DEA99FFA1FFDBFF42D071FE9AF85F" bold="true" box="[189,334,1905,1931]" pageId="27" pageNumber="28">Description</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFFA1FFDBFF42D0A1FF3AF809" blockId="27.[188,1398,1952,2013]" pageId="27" pageNumber="28">
<smallCapsWord id="8D40A057FFA1FFDBFF42D0A1FEBAF86D" baselines="1972,1972" box="[189,366,1953,1979]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Measurements" pageId="27" pageNumber="28">MEASUREMENTS</smallCapsWord>
.
<typeStatus id="54A28829FFA1FFDBFE83D0A0FE3AF86E" box="[380,494,1952,1978]" pageId="27" pageNumber="28" type="holotype">Holotype</typeStatus>
: sh:
<quantity id="4CE19B6EFFA1FFDBFDDED0A0FD58F86F" box="[545,652,1952,1979]" metricMagnitude="-3" metricUnit="m" metricValue="1.57" pageId="27" pageNumber="28" unit="mm" value="1.57">1.57 mm</quantity>
; sw:
<quantity id="4CE19B6EFFA1FFDBFD39D0A0FCE5F86F" box="[710,817,1952,1979]" metricMagnitude="-3" metricUnit="m" metricValue="1.08" pageId="27" pageNumber="28" unit="mm" value="1.08">1.08 mm</quantity>
; ah:
<quantity id="4CE19B6EFFA1FFDBFC98D0A0FC05F86F" box="[871,977,1952,1979]" metricMagnitude="-4" metricUnit="m" metricValue="5.8" pageId="27" pageNumber="28" unit="mm" value="0.58">0.58 mm</quantity>
; aw: 0.66; hlw:
<quantity id="4CE19B6EFFA1FFDBFB75D0A0FB21F86F" box="[1162,1269,1952,1979]" metricMagnitude="-4" metricUnit="m" metricValue="9.5" pageId="27" pageNumber="28" unit="mm" value="0.95">0.95 mm</quantity>
; SA: 66.35 deg.
</paragraph>
<paragraph id="8BA6368BFFA6FFDCFF42D609FD86FD38" blockId="28.[188,1399,263,749]" pageId="28" pageNumber="29">
Shell conical or elongate conical, height ca
<quantity id="4CE19B6EFFA6FFDCFD38D609FCF6FEF7" box="[711,802,265,291]" metricMagnitude="-3" metricUnit="m" metricValue="1.6" pageId="28" pageNumber="29" unit="mm" value="1.6">1.6 mm</quantity>
with 5½ whorls; transparent. Whorls convex, and slightly shouldered, spire regularly and tightly coiled; suture moderately deep; aperture reniform, almost elliptical ovate; aperture wider than high, columellar side slightly rounded; slight notch at upper parietal and palatal junction; parietal callus (shield) well defined, straight, long and angular; palatal rim thin; palatal-basal lip narrowly reflected; teleoconch smooth with irregular growth lines; weak radial banding on body whorl; course irregular growth lines present immediately behind palatal-basal lip; height of last whorl less than half of shell height; apertural rim thin and receding
<superScript id="7C6C9BC3FFA6FFDCFBEDD6DCFBCEFE3E" attach="right" box="[1042,1050,476,490]" fontSize="6" pageId="28" pageNumber="29">1</superScript>
/
<subScript id="179D34CEFFA6FFDCFBDDD6E7FBFFFE21" attach="left" box="[1058,1067,487,501]" fontSize="6" pageId="28" pageNumber="29">5</subScript>
width of penultimate whorl (in aperture facing left view), exposing deeply set parietalis (
<figureCitation id="13222A0EFFA6FFDCFC53D6FFFBD8FDCD" box="[940,1036,511,537]" 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. CH. 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="28" pageNumber="29">Fig. 6G</figureCitation>
); columella centrally aligned, moderately thick, not swollen at base; low lamella under penultimate whorl forming a conspicuous, short parietalis reaching up as high as the umbilical depression in umbilical view; umbilical depression small; umbilicus callused at junction of peristome and base of columella, columellar side of peristome positioned above and to the right of the umbilicus, base swollen with little puckering; upper palatal rim not reflected (as seen from umbilical view); alignment of last ¼ whorl compact with a slightly rightward tendency (seen in umbilical view).
</paragraph>
</subSubSection>
<caption id="DF666603FFA6FFDCFF42D041FC39F83C" ID-DOI="http://doi.org/10.5281/zenodo.10847820" ID-Zenodo-Dep="10847820" httpUri="https://zenodo.org/record/10847820/files/figure.png" pageId="28" pageNumber="29" startId="28.[189,232,1857,1883]" targetBox="[189,1398,887,1818]" targetPageId="28">
<paragraph id="8BA6368BFFA6FFDCFF42D041FC39F83C" blockId="28.[189,1399,1857,2024]" pageId="28" pageNumber="29">
<emphasis id="B96DEA99FFA6FFDCFF42D041FEC3F88F" bold="true" box="[189,279,1857,1883]" pageId="28" pageNumber="29">Fig. 6.</emphasis>
<taxonomicName id="4C194D08FFA6FFDCFED6D042FC5EF88F" authority="Jochum &amp; Ruthensteiner" authorityName="Jochum &amp; Ruthensteiner" authorityYear="2024" box="[297,906,1857,1884]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="28" pageNumber="29" phylum="Mollusca" rank="species" species="intermedium" status="sp. nov.">
<emphasis id="B96DEA99FFA6FFDCFED6D042FDE9F88F" box="[297,573,1857,1883]" italics="true" pageId="28" pageNumber="29">Zospeum intermedium</emphasis>
Jochum &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FFA6FFDCFC62D042FBD0F88F" box="[925,1028,1858,1884]" pageId="28" pageNumber="29" rank="species">sp. nov.</taxonomicNameLabel>
(RMNH.MOL.234132) from
<bibRefCitation id="EF884B7AFFA6FFDCFF42D065FE7AF8AB" author="Gittenberger E." box="[189,430,1892,1919]" pageId="28" pageNumber="29" pagination="21 - 37" refId="ref31037" refString="Gittenberger E. 1975. Cave snails found in southern Crna Gora. Glasnik Republickog zavoda za zastitutu prirode i Prirodnjackog muzeja 8: 21 - 37." type="journal article" year="1975">Gittenberger (1975)</bibRefCitation>
.
<emphasis id="B96DEA99FFA6FFDCFE42D064FE06F8AA" bold="true" box="[445,466,1892,1918]" pageId="28" pageNumber="29">A</emphasis>
. Light microscopic images of apertural and dorsal views.
<emphasis id="B96DEA99FFA6FFDCFB5ED065FB61F8AB" bold="true" box="[1185,1205,1893,1919]" pageId="28" pageNumber="29">B</emphasis>
. Sample labels.
<emphasis id="B96DEA99FFA6FFDCFF42D088FF2CF876" bold="true" box="[189,248,1928,1954]" pageId="28" pageNumber="29">CH</emphasis>
. 3D visualizations of X-ray Micro-CT data.
<emphasis id="B96DEA99FFA6FFDCFCDED088FCE2F876" bold="true" box="[801,822,1928,1954]" pageId="28" pageNumber="29">C</emphasis>
. Aperture view.
<emphasis id="B96DEA99FFA6FFDCFBF9D088FBCFF876" bold="true" box="[1030,1051,1928,1954]" pageId="28" pageNumber="29">D</emphasis>
. Aperture facing right view.
<emphasis id="B96DEA99FFA6FFDCFF42D0ABFF05F811" bold="true" box="[189,209,1963,1989]" pageId="28" pageNumber="29">E</emphasis>
. Apical view.
<emphasis id="B96DEA99FFA6FFDCFE77D0ABFE4EF811" bold="true" box="[392,410,1963,1989]" pageId="28" pageNumber="29">F</emphasis>
. Dorsal view.
<emphasis id="B96DEA99FFA6FFDCFDACD0ABFDBEF811" bold="true" box="[595,618,1963,1989]" pageId="28" pageNumber="29">G</emphasis>
. Aperture facing right view.
<emphasis id="B96DEA99FFA6FFDCFC2BD0ABFC3FF811" bold="true" box="[980,1003,1963,1989]" pageId="28" pageNumber="29">H</emphasis>
. Umbilical view showing slight umbilical depression and lamella projecting from the columellar side.
</paragraph>
</caption>
<subSubSection id="C3036500FFA7FFDDFF42D609FDDDFCB5" pageId="29" pageNumber="30" type="discussion">
<paragraph id="8BA6368BFFA7FFDDFF42D609FEFBFEF7" blockId="29.[189,303,265,291]" box="[189,303,265,291]" pageId="29" pageNumber="30">
<heading id="D0EE81E7FFA7FFDDFF42D609FEFBFEF7" bold="true" box="[189,303,265,291]" fontSize="11" level="3" pageId="29" pageNumber="30" reason="3">
<emphasis id="B96DEA99FFA7FFDDFF42D609FEFBFEF7" bold="true" box="[189,303,265,291]" pageId="29" pageNumber="30">Remarks</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFFA7FFDDFF42D638FDDDFCB5" blockId="29.[187,1399,311,866]" pageId="29" pageNumber="30">
It is not known from which section of the cave this shell derived except that it was amidst a larger sample (RMNH.MOL.234134) comprising 64 shells from a part of the cave “with absolute darkness” (
<bibRefCitation id="EF884B7AFFA7FFDDFF39D67DFE7EFE4C" author="Gittenberger E." box="[198,426,381,408]" pageId="29" pageNumber="30" pagination="21 - 37" refId="ref31037" refString="Gittenberger E. 1975. Cave snails found in southern Crna Gora. Glasnik Republickog zavoda za zastitutu prirode i Prirodnjackog muzeja 8: 21 - 37." type="journal article" year="1975">Gittenberger 1975</bibRefCitation>
). The species represents an intermediary form between
<taxonomicName id="4C194D08FFA7FFDDFB89D67EFABBFE43" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[1142,1391,381,407]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FFA7FFDDFB89D67EFABBFE43" box="[1142,1391,381,407]" italics="true" pageId="29" pageNumber="30">Z. troglobalcanicum</emphasis>
</taxonomicName>
,
<taxonomicName id="4C194D08FFA7FFDDFF42D6A1FD56FE6F" authority="Jochum &amp; Ruthensteiner" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[189,642,417,443]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="tortuosum" status="sp. nov.">
<emphasis id="B96DEA99FFA7FFDDFF42D6A1FE81FE6E" box="[189,341,417,442]" italics="true" pageId="29" pageNumber="30">Z. tortuosum</emphasis>
Jochum &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FFA7FFDDFD75D6A1FD33FE6F" box="[650,743,417,443]" pageId="29" pageNumber="30" rank="species">sp. nov.</taxonomicNameLabel>
,
<taxonomicName id="4C194D08FFA7FFDDFD0CD6A1FB36FE6F" authority="Jochum &amp; Ruthensteiner" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[755,1250,417,443]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="dubokidoense" status="sp. nov.">
<emphasis id="B96DEA99FFA7FFDDFD0CD6A1FC62FE6F" box="[755,950,417,443]" italics="true" pageId="29" pageNumber="30">Z. dubokidoense</emphasis>
Jochum &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FFA7FFDDFB15D6A1FA91FE6F" box="[1258,1349,417,443]" pageId="29" pageNumber="30" rank="species">sp. nov.</taxonomicNameLabel>
and
<emphasis id="B96DEA99FFA7FFDDFF42D6C5FE97FE0A" box="[189,323,452,478]" italics="true" pageId="29" pageNumber="30">
<taxonomicName id="4C194D08FFA7FFDDFF42D6C5FEEBFE0A" authorityName="Inabnit, Jochum &amp; Neubert" authorityYear="2021" box="[189,319,452,478]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="simplex">Z. simplex</taxonomicName>
.
</emphasis>
It is significantly larger than these species in shell height, shell width, aperture height, aperture width, height of the last whorl and narrower in spire angle. The columella shows similarity with that of
<taxonomicName id="4C194D08FFA7FFDDFEF4D50BFE1AFDF0" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[267,462,522,549]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="57" phylum="Mollusca" rank="species" species="dubokidoense">
Z.
<emphasis id="B96DEA99FFA7FFDDFED4D50AFE1AFDF0" box="[299,462,522,548]" italics="true" pageId="29" pageNumber="30">dubokidoense</emphasis>
</taxonomicName>
in turning angle, its slightly off-centered alignment, and superficial irregularities. In dorsal and aperture facing left views, the columella is similar to that of
<taxonomicName id="4C194D08FFA7FFDDFBB2D52EFA96FD93" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[1101,1346,557,583]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FFA7FFDDFBB2D52EFA96FD93" box="[1101,1346,557,583]" italics="true" pageId="29" pageNumber="30">Z. troglobalcanicum</emphasis>
</taxonomicName>
and
<taxonomicName id="4C194D08FFA7FFDDFF42D551FEABFDBF" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[189,383,593,619]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="dubokidoense">
<emphasis id="B96DEA99FFA7FFDDFF42D551FEABFDBF" box="[189,383,593,619]" italics="true" pageId="29" pageNumber="30">Z. dubokidoense</emphasis>
</taxonomicName>
. However,
<taxonomicName id="4C194D08FFA7FFDDFDFAD551FC30FDBF" authority="Jochum &amp; Ruthensteiner" authorityName="Jochum &amp; Ruthensteiner" authorityYear="2024" box="[517,996,593,619]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="intermedium" status="sp. nov.">
<emphasis id="B96DEA99FFA7FFDDFDFAD551FD6EFDBF" box="[517,698,593,619]" italics="true" pageId="29" pageNumber="30">Z. intermedium</emphasis>
Jochum &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FFA7FFDDFC14D551FB91FDBF" box="[1003,1093,593,619]" pageId="29" pageNumber="30" rank="species">sp. nov.</taxonomicNameLabel>
differs from these species in that the umbilical depression grades from a pronounced umbilical depression in
<taxonomicName id="4C194D08FFA7FFDDFB6BD575FAA3FD5A" authorityName="Absolon" authorityYear="1916" box="[1172,1399,628,654]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="troglobalcnicum">
<emphasis id="B96DEA99FFA7FFDDFB6BD575FAA3FD5A" box="[1172,1399,628,654]" italics="true" pageId="29" pageNumber="30">Z. troglobalcnicum</emphasis>
</taxonomicName>
and
<taxonomicName id="4C194D08FFA7FFDDFF13D598FEB0FD65" authorityName="Inabnit, Jochum &amp; Neubert" authorityYear="2021" box="[236,356,663,689]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="simplex">
<emphasis id="B96DEA99FFA7FFDDFF13D598FEB0FD65" box="[236,356,663,689]" italics="true" pageId="29" pageNumber="30">Z. simplex</emphasis>
</taxonomicName>
to that of a slight umbilical depression in
<taxonomicName id="4C194D08FFA7FFDDFCC2D598FC24FD65" authorityName="Jochum &amp; Ruthensteiner" authorityYear="2024" box="[829,1008,663,689]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="intermedium">
<emphasis id="B96DEA99FFA7FFDDFCC2D598FC24FD65" box="[829,1008,663,689]" italics="true" pageId="29" pageNumber="30">Z. intermedium</emphasis>
</taxonomicName>
to that of no umbilical depression in
<taxonomicName id="4C194D08FFA7FFDDFF23D5BBFE4BFD00" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[220,415,698,724]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="dubokidoense">
<emphasis id="B96DEA99FFA7FFDDFF23D5BBFE4BFD00" box="[220,415,698,724]" italics="true" pageId="29" pageNumber="30">Z. dubokidoense</emphasis>
</taxonomicName>
, which is largely covered by the columellar side of the peristome directly above it.
<taxonomicName id="4C194D08FFA7FFDDFF41D5DEFEA3FD23" authorityName="Jochum &amp; Ruthensteiner" authorityYear="2024" box="[190,375,733,759]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="intermedium">
<emphasis id="B96DEA99FFA7FFDDFF41D5DEFEA3FD23" box="[190,375,733,759]" italics="true" pageId="29" pageNumber="30">Z. intermedium</emphasis>
</taxonomicName>
differs from the rest of the Southern Balkan species in this study in that it bears a pronounced lamella forming a deeply set, conspicuous parietalis. Only
<taxonomicName id="4C194D08FFA7FFDDFBF0D401FB05FCCF" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[1039,1233,769,795]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="dubokidoense">
<emphasis id="B96DEA99FFA7FFDDFBF0D401FB05FCCF" box="[1039,1233,769,795]" italics="true" pageId="29" pageNumber="30">Z. dubokidoense</emphasis>
</taxonomicName>
shows a very slight but visible parietalis (
<figureCitation id="13222A0EFFA7FFDDFDF2D424FDBEFCEA" box="[525,618,804,830]" captionStart="Fig" captionStartId="31.[189,232,1787,1813]" captionTargetBox="[187,1396,481,1747]" captionTargetId="figure-65@31.[187,1396,481,1747]" captionTargetPageId="31" captionText="Fig. 8. 3D visualizations of Micro-CT data of shells from the caves Lipska pećina and Duboki do (Montenegro). AF. Zospeum troglobalcanicum Absolon, 1916 (MCSMNH-PMSL-Moll.-FVelkovrh 29603) (Fig. 7A). A. Aperture view. B. Aperture facing right view. C. Apical view. D. Dorsal view. E. Aperture facing left view. F. Ventral view showing reduced umbilical depression. GL. Zospeum dubokidoense Jochum &amp; Ruthensteiner sp. nov., holotype (MCSMNH-PMSL-Moll.-FVelkovrh 30360[spm1]). G. Aperture view. H. Aperture facing right view. I. Apical view. J. Dorsal view. K. Aperture facing left view. L. Ventral view showing no umbilical depression." figureDoi="http://doi.org/10.5281/zenodo.10847826" httpUri="https://zenodo.org/record/10847826/files/figure.png" pageId="29" pageNumber="30">Fig. 8L</figureCitation>
).
<taxonomicName id="4C194D08FFA7FFDDFD7ED425FCA0FCEA" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[641,884,804,830]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="29" pageNumber="30" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FFA7FFDDFD7ED425FCA0FCEA" box="[641,884,804,830]" italics="true" pageId="29" pageNumber="30">Z. troglobalcanicum</emphasis>
</taxonomicName>
bears a threadlike partial lamella that does not form a visible parietalis.
</paragraph>
</subSubSection>
</treatment>
</document>