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<document id="D4F14B2F5AF2D598D3A9090582DCDC49" ID-DOI="10.5852/ejt.2024.926.2469" ID-ISSN="2118-9773" ID-Zenodo-Dep="10847806" ID-ZooBank="A00D7669-2151-46AE-A066-9AF4D0F2BEEE" IM.bibliography_approvedBy="guilherme" IM.illustrations_approvedBy="guilherme" IM.materialsCitations_approvedBy="guilherme" IM.metadata_approvedBy="felipe" 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="03B0879DFF88FFF4FD3DD336FC2EFB85" 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 undefined-2" docType="treatment" docVersion="8" lastPageNumber="53" 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="51" updateTime="1711368481034" updateUser="guilherme" 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="0A2A6AF1EA8B2DEED697936A61A90239">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="7B37519E6BAFED360B7C0FEBCCF9CBA7">Jochum, Adrienne</mods:namePart>
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<mods:namePart id="EB26F8739184325D4DD6C9055CF788B4">Inäbnit, Thomas</mods:namePart>
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<mods:affiliation id="790A6F73D424A338293DDBD9FC5F7ED7">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="ED826A1796FC47AEF7DB2E93575FDF7B">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="B96DEA99FF88FFF2FF42D39BFDD5FB60" box="[189,513,1178,1204]" italics="true" pageId="50" pageNumber="51">Zospeum troglobalcanicum</emphasis>
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– 
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<bibRefCitation id="EF884B7AFF88FFF2FDE0D39AFCF1FB60" author="Inabnit T. &amp; Jochum A. &amp; Kampschulte M. &amp; Martels G. &amp; Ruthensteiner B. &amp; Slapnik R. &amp; Nesselhauf C. &amp; Neubert E." box="[543,805,1178,1204]" pageId="50" pageNumber="51" pagination="135 - 177" refId="ref31276" refString="Inabnit T., Jochum A., Kampschulte M., Martels G., Ruthensteiner B., Slapnik R., Nesselhauf C. &amp; Neubert E. 2019. An integrative taxonomic study reveals carychiid microsnails of the troglobitic genus Zospeum in the Eastern and Dinaric Alps (Gastropoda, Ellobioidea, Carychiinae). Organisms Diversity &amp; Evolution 19: (2): 135 - 177. https: // doi. org / 10.1007 / s 13127 - 019 - 00400 - 8" type="journal article" year="2019">
Inäbnit 
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2019: 19
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, fig. 7u.
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<emphasis id="B96DEA99FF88FFF2FF42D3DBFEE2FB21" bold="true" box="[189,310,1243,1269]" pageId="50" pageNumber="51">Diagnosis</emphasis>
(from 
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Inäbnit 
<emphasis id="B96DEA99FF88FFF2FE1ED3DCFDC9FB21" box="[481,541,1243,1269]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
2019
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)
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Shell ca 
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, transparent, with a globose conical form, peristome with well-defined parietal shield, totally smooth columella, lamellae missing completely.
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leg.; 
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(ex RSC 1981)
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.
</paragraph>
</subSubSection>
<subSubSection id="C3036500FF88FFF3FF42D102FB45FE59" lastPageId="51" lastPageNumber="52" pageId="50" pageNumber="51" type="description">
<paragraph id="8BA6368BFF88FFF2FF42D102FD79F9C8" blockId="50.[189,685,1538,1564]" box="[189,685,1538,1564]" pageId="50" pageNumber="51">
<emphasis id="B96DEA99FF88FFF2FF42D102FE9AF9C8" bold="true" box="[189,334,1538,1564]" pageId="50" pageNumber="51">Description</emphasis>
(using Micro-CT image data)
</paragraph>
<paragraph id="8BA6368BFF88FFF2FF42D131FEE3F9B9" blockId="50.[188,1399,1584,1645]" pageId="50" pageNumber="51">
<smallCapsWord id="8D40A057FF88FFF2FF42D131FEBAF99D" baselines="1604,1604" box="[189,366,1585,1611]" lowerCaseFontSize="8" mainFontSize="11" normCase="title" normString="Measurements" pageId="50" pageNumber="51">MEASUREMENTS</smallCapsWord>
(Micro-CT). sh: 
<quantity id="4CE19B6EFF88FFF2FDC8D130FD76F99F" box="[567,674,1584,1611]" metricMagnitude="-3" metricUnit="m" metricValue="1.3" pageId="50" pageNumber="51" unit="mm" value="1.3">1.30 mm</quantity>
; sw: 1.00 mm; ah: 
<quantity id="4CE19B6EFF88FFF2FC83D130FC33F99F" box="[892,999,1584,1611]" metricMagnitude="-4" metricUnit="m" metricValue="5.8" pageId="50" pageNumber="51" unit="mm" value="0.58">0.58 mm</quantity>
; aw: 
<quantity id="4CE19B6EFF88FFF2FBDCD130FB59F99F" box="[1059,1165,1584,1611]" metricMagnitude="-4" metricUnit="m" metricValue="6.0" pageId="50" pageNumber="51" unit="mm" value="0.6">0.60 mm</quantity>
; hlw: 
<quantity id="4CE19B6EFF88FFF2FB2CD130FAEAF99F" box="[1235,1342,1584,1611]" metricMagnitude="-4" metricUnit="m" metricValue="8.299999999999999" pageId="50" pageNumber="51" unit="mm" value="0.83">0.83 mm</quantity>
; SA: 71.60 deg.
</paragraph>
<paragraph id="8BA6368BFF88FFF3FF42D195FB45FE59" blockId="50.[189,1400,1684,1993]" lastBlockId="51.[189,1400,265,397]" lastPageId="51" lastPageNumber="52" pageId="50" pageNumber="51">
Shell ca 
<quantity id="4CE19B6EFF88FFF2FEDED195FEA9F964" box="[289,381,1685,1712]" metricMagnitude="-3" metricUnit="m" metricValue="1.3" pageId="50" pageNumber="51" unit="mm" value="1.3">1.3 mm</quantity>
, conical, with 5¼ regularly coiled, flatly convex whorls; transparent when fresh; suture uneven and shallow; height of last whorl greater than half of shell height; aperture reniform, narrow, slightly wider than high; teleoconch sculpture with some irregular blunt growth lines; suture zone of penultimate and final whorl with indentations from which growth lines extend irregularly; peristome thin and flattened at upper palatal side, with thickened callused zone starting after ca ⅓ the length of the palatal side; columellar side of peristome slightly oblique and callused, palatal and basal palatal edge narrowly reflected; parietal shield well-defined, thickly callused, long and straight with notch at upper parietal and palatal junction; upper rim of peristome recedes ca 1/5 the width of the penultimate whorl (aperture facing left); columella centrally aligned and totally smooth, lower part of internal penultimate whorl shows a low ridge-like fold independent of and above the formation of the columella but not leading onto it; ventral side with a shallow umbilical depression with some puckering of blunt ridges leading into it behind the columellar side of the peristome; columellar side of peristome positioned directly above umbilical depression; alignment of last 
<superScript id="7C6C9BC3FF89FFF3FCC4D672FC90FE54" attach="right" box="[827,836,370,384]" fontSize="6" pageId="51" pageNumber="52">1</superScript>
/ 
<subScript id="179D34CEFF89FFF3FCB3D67DFC81FE5F" attach="left" box="[844,853,381,395]" fontSize="6" pageId="51" pageNumber="52">5</subScript>
whorl somewhat compact.
</paragraph>
</subSubSection>
<subSubSection id="C3036500FF89FFF3FF42D6B4FE31FDF4" pageId="51" pageNumber="52" type="distribution">
<paragraph id="8BA6368BFF89FFF3FF42D6B4FE82FE1A" blockId="51.[189,342,436,462]" box="[189,342,436,462]" pageId="51" pageNumber="52">
<heading id="D0EE81E7FF89FFF3FF42D6B4FE82FE1A" bold="true" box="[189,342,436,462]" fontSize="11" level="3" pageId="51" pageNumber="52" reason="3">
<emphasis id="B96DEA99FF89FFF3FF42D6B4FE82FE1A" bold="true" box="[189,342,436,462]" pageId="51" pageNumber="52">Distribution</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFF89FFF3FF42D6E2FE31FDF4" blockId="51.[189,1398,482,544]" pageId="51" pageNumber="52">
Taleža pećina is located in the Trebinjska šuma region, a 
<quantity id="4CE19B6EFF89FFF3FC85D6E3FC13FE28" box="[890,967,482,509]" metricMagnitude="4" metricUnit="m" metricValue="2.3" pageId="51" pageNumber="52" unit="km" value="23.0">23 km</quantity>
long karst rocky plateau of eastern 
<collectingCountry id="F30E761BFF89FFF3FF42D506FE35FDF4" box="[189,481,518,544]" name="Bosnia and Herzegovina" pageId="51" pageNumber="52">Bosnia and Herzegovina</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="C3036500FF89FFF4FF42D547FC2EFB85" lastPageId="52" lastPageNumber="53" pageId="51" pageNumber="52" type="discussion">
<paragraph id="8BA6368BFF89FFF3FF42D547FEFBFDB5" blockId="51.[189,303,583,609]" box="[189,303,583,609]" pageId="51" pageNumber="52">
<heading id="D0EE81E7FF89FFF3FF42D547FEFBFDB5" bold="true" box="[189,303,583,609]" fontSize="11" level="3" pageId="51" pageNumber="52" reason="3">
<emphasis id="B96DEA99FF89FFF3FF42D547FEFBFDB5" bold="true" box="[189,303,583,609]" pageId="51" pageNumber="52">Remarks</emphasis>
</heading>
</paragraph>
<paragraph id="8BA6368BFF89FFF3FF42D576FDEAFC18" blockId="51.[188,1399,630,973]" pageId="51" pageNumber="52">
<bibRefCitation id="EF884B7AFF89FFF3FF42D576FE60FD44" author="Inabnit T. &amp; Jochum A. &amp; Kampschulte M. &amp; Martels G. &amp; Ruthensteiner B. &amp; Slapnik R. &amp; Nesselhauf C. &amp; Neubert E." box="[189,436,630,656]" pageId="51" pageNumber="52" pagination="135 - 177" refId="ref31276" refString="Inabnit T., Jochum A., Kampschulte M., Martels G., Ruthensteiner B., Slapnik R., Nesselhauf C. &amp; Neubert E. 2019. An integrative taxonomic study reveals carychiid microsnails of the troglobitic genus Zospeum in the Eastern and Dinaric Alps (Gastropoda, Ellobioidea, Carychiinae). Organisms Diversity &amp; Evolution 19: (2): 135 - 177. https: // doi. org / 10.1007 / s 13127 - 019 - 00400 - 8" type="journal article" year="2019">
Inäbnit 
<emphasis id="B96DEA99FF89FFF3FEE4D576FE8EFD44" box="[283,346,630,656]" italics="true" pageId="51" pageNumber="52">et al.</emphasis>
(2019)
</bibRefCitation>
compared this shell to the Western Balkan congeners in their molecular study of the radiation of Dinaride 
<taxonomicName id="4C194D08FF89FFF3FDEDD59AFDAFFD67" authorityName="Bourguignat" authorityYear="1856" box="[530,635,666,691]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="51" pageNumber="52" phylum="Mollusca" rank="genus">
<emphasis id="B96DEA99FF89FFF3FDEDD59AFDAFFD67" box="[530,635,666,691]" italics="true" pageId="51" pageNumber="52">Zospeum</emphasis>
</taxonomicName>
, concluding that it mostly shared an overall resemblance to the photograph of 
<taxonomicName id="4C194D08FF89FFF3FE8FD5BDFDB0FD02" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[368,612,700,726]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="51" pageNumber="52" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FF89FFF3FE8FD5BDFDB0FD02" box="[368,612,700,726]" italics="true" pageId="51" pageNumber="52">Z. troglobalcanicum</emphasis>
</taxonomicName>
supplied by 
<bibRefCitation id="EF884B7AFF89FFF3FCFDD5BCFC05FD02" author="Absolon K." box="[770,977,700,726]" pageId="51" pageNumber="52" pagination="574 - 576" refId="ref30332" refString="Absolon K. 1916 b. Z vyzkumnych cest po krasech Balkanu. O balkanske temnostni zvirene. Zlata Praha 33 (48): 574 - 576; (49): 586 - 588; (50): 597 - 600; (51): 609 - 612; (52): 622 - 624." type="journal article" year="1916">Absolon (1916b)</bibRefCitation>
. This conclusion was based on the globose conical form of the shell, the peristome with a well-defined parietal shield and that lamellae were lacking completely. Though 
<bibRefCitation id="EF884B7AFF89FFF3FDF1D402FD2EFCC9" author="Inabnit T. &amp; Jochum A. &amp; Kampschulte M. &amp; Martels G. &amp; Ruthensteiner B. &amp; Slapnik R. &amp; Nesselhauf C. &amp; Neubert E." box="[526,762,770,797]" pageId="51" pageNumber="52" pagination="135 - 177" refId="ref31276" refString="Inabnit T., Jochum A., Kampschulte M., Martels G., Ruthensteiner B., Slapnik R., Nesselhauf C. &amp; Neubert E. 2019. An integrative taxonomic study reveals carychiid microsnails of the troglobitic genus Zospeum in the Eastern and Dinaric Alps (Gastropoda, Ellobioidea, Carychiinae). Organisms Diversity &amp; Evolution 19: (2): 135 - 177. https: // doi. org / 10.1007 / s 13127 - 019 - 00400 - 8" type="journal article" year="2019">
Inäbnit 
<emphasis id="B96DEA99FF89FFF3FD95D403FD71FCC8" box="[618,677,770,796]" italics="true" pageId="51" pageNumber="52">et al.</emphasis>
(2019)
</bibRefCitation>
considered this morph 
<taxonomicName id="4C194D08FF89FFF3FBEED403FAD7FCC8" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[1041,1283,770,796]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="51" pageNumber="52" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FF89FFF3FBEED403FAD7FCC8" box="[1041,1283,770,796]" italics="true" pageId="51" pageNumber="52">Z. troglobalcanicum</emphasis>
</taxonomicName>
, it differs from the 
<typeStatus id="54A28829FF89FFF3FED4D426FE43FC94" box="[299,407,806,832]" pageId="51" pageNumber="52" type="lectotype">lectotype</typeStatus>
by the smooth columella and the configuration of the umbilical zone, including the shallow umbilical depression and the relatively tight alignment of the columellar side of the peristome in relation to it. It is also much smaller than 
<taxonomicName id="4C194D08FF89FFF3FD57D46DFC4DFC52" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[680,921,876,903]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="51" pageNumber="52" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FF89FFF3FD57D46DFD6CFC52" box="[680,696,877,902]" italics="true" pageId="51" pageNumber="52">Z</emphasis>
. 
<emphasis id="B96DEA99FF89FFF3FD38D46CFC4DFC52" box="[711,921,876,902]" italics="true" pageId="51" pageNumber="52">troglobalcanicum</emphasis>
</taxonomicName>
, constituting one of the smallest morphs in this study, except for 
<taxonomicName id="4C194D08FF89FFF3FE14D490FB8EFC7D" authority="Jochum, Schilthuizen &amp; Ruthensteiner" authorityName="Jochum, Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[491,1114,911,937]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="51" pageNumber="52" phylum="Mollusca" rank="species" species="njunjicae" status="sp. nov.">
<emphasis id="B96DEA99FF89FFF3FE14D490FDA9FC7D" box="[491,637,912,937]" italics="true" pageId="51" pageNumber="52">Z. njunjicae</emphasis>
Jochum, Schilthuizen &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FF89FFF3FB99D490FB13FC7E" box="[1126,1223,912,938]" pageId="51" pageNumber="52" rank="species">sp. nov.</taxonomicNameLabel>
, which on the average, bears the smallest shell.
</paragraph>
<paragraph id="8BA6368BFF89FFF4FF42D4F4FA82FE92" blockId="51.[188,1399,1012,1109]" lastBlockId="52.[187,1398,265,327]" lastPageId="52" lastPageNumber="53" pageId="51" pageNumber="52">
This shell is very similar to 
<taxonomicName id="4C194D08FF89FFF3FE02D4F5FC73FBDA" authority="Jochum &amp; Ruthensteiner" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[509,935,1012,1038]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="51" pageNumber="52" phylum="Mollusca" rank="species" species="neuberti" status="sp. nov.">
<emphasis id="B96DEA99FF89FFF3FE02D4F5FDAAFBDA" box="[509,638,1012,1038]" italics="true" pageId="51" pageNumber="52">Z. neuberti</emphasis>
Jochum &amp; Ruthensteiner
</taxonomicName>
<taxonomicNameLabel id="A25E57E2FF89FFF3FC52D4F4FBD2FBDA" box="[941,1030,1012,1038]" pageId="51" pageNumber="52" rank="species">sp. nov.</taxonomicNameLabel>
in that the columella is smooth, albeit somewhat attenuate, with the umbilical zone bearing similar morphology. The spire is wider, the aperture width is broader and there are fewer low ribs behind the peristome compared to the three individuals of 
<taxonomicName id="4C194D08FF8EFFF4FE9ED60AFE36FEF7" authorityName="Jochum &amp; Michalik &amp; Inäbnit &amp; Kneubühler &amp; Slapnik &amp; Vrabec &amp; Schilthuizen &amp; Ruthensteiner" authorityYear="2024" box="[353,482,265,291]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="52" pageNumber="53" phylum="Mollusca" rank="species" species="neuberti">
<emphasis id="B96DEA99FF8EFFF4FE9ED60AFE36FEF7" box="[353,482,265,291]" italics="true" pageId="52" pageNumber="53">Z. neuberti</emphasis>
</taxonomicName>
from the 
<typeStatus id="54A28829FF8EFFF4FDAED60AFD50FEF0" box="[593,644,266,292]" pageId="52" pageNumber="53">type</typeStatus>
locality, Grabovica cave. Due to the lack of comparative material from Taleža pećina, no conclusive conchological determination can be made based on this one shell.
</paragraph>
<caption id="DF666603FF89FFF3FF41D040FC84F833" ID-DOI="http://doi.org/10.5281/zenodo.10847864" ID-Zenodo-Dep="10847864" httpUri="https://zenodo.org/record/10847864/files/figure.png" pageId="51" pageNumber="52" startId="51.[190,233,1856,1882]" targetBox="[189,1398,1191,1817]" targetPageId="51">
<paragraph id="8BA6368BFF89FFF3FF41D040FC84F833" blockId="51.[188,1398,1856,2024]" pageId="51" pageNumber="52">
<emphasis id="B96DEA99FF89FFF3FF41D040FECDF88E" bold="true" box="[190,281,1856,1882]" pageId="51" pageNumber="52">Fig. 20.</emphasis>
3D visualizations of Micro-CT data of 
<taxonomicName id="4C194D08FF89FFF3FD2AD041FCAAF88F" box="[725,894,1857,1883]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="51" pageNumber="52" phylum="Mollusca" rank="species" species="undefined-2">
<emphasis id="B96DEA99FF89FFF3FD2AD041FC94F88E" box="[725,832,1857,1882]" italics="true" pageId="51" pageNumber="52">Zospeum</emphasis>
sp. 2
</taxonomicName>
, Taleža pećina, bei Trebinje, BiH, presented as 
<taxonomicName id="4C194D08FF89FFF3FF1FD064FE02F8AA" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[224,470,1892,1918]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="51" pageNumber="52" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FF89FFF3FF1FD064FE02F8AA" box="[224,470,1892,1918]" italics="true" pageId="51" pageNumber="52">Z. troglobalcanicum</emphasis>
</taxonomicName>
in 
<bibRefCitation id="EF884B7AFF89FFF3FDFCD064FD3CF8AA" author="Inabnit T. &amp; Jochum A. &amp; Slapnik R. &amp; Neubert E." box="[515,744,1892,1918]" pageId="51" pageNumber="52" pagination="175 - 193" refId="ref31366" refString="Inabnit T., Jochum A., Slapnik R. &amp; Neubert E. 2021. New genetic data reveals a new species of Zospeum in Bosnia (Gastropoda, Ellobioidea, Carychiinae). ZooKeys 1071: 175 - 193. https: // doi. org / 10.3897 / zookeys. 1071.66417" type="journal article" year="2021">
Inäbnit 
<emphasis id="B96DEA99FF89FFF3FD9DD064FD75F8A9" box="[610,673,1892,1918]" italics="true" pageId="51" pageNumber="52">et al.</emphasis>
2021
</bibRefCitation>
(NMBE 553414/1 (ex RSC 1981). 
<emphasis id="B96DEA99FF89FFF3FB5CD063FB6CF8A9" bold="true" box="[1187,1208,1891,1917]" pageId="51" pageNumber="52">A</emphasis>
. Aperture view showing low, ridge-like fold on inner penultimate whorl. 
<emphasis id="B96DEA99FF89FFF3FCA7D087FCB8F875" bold="true" box="[856,876,1927,1953]" pageId="51" pageNumber="52">B</emphasis>
. Aperture facing right view showing smooth columella. 
<emphasis id="B96DEA99FF89FFF3FEBDD0AAFE83F810" bold="true" box="[322,343,1962,1988]" pageId="51" pageNumber="52">C</emphasis>
. Apical view. 
<emphasis id="B96DEA99FF89FFF3FDF3D0AAFDCBF810" bold="true" box="[524,543,1962,1988]" pageId="51" pageNumber="52">D</emphasis>
. Dorsal view. 
<emphasis id="B96DEA99FF89FFF3FD2BD0AAFD3CF810" bold="true" box="[724,744,1962,1988]" pageId="51" pageNumber="52">E</emphasis>
. Aperture facing right view. 
<emphasis id="B96DEA99FF89FFF3FBB4D0AAFB89F810" bold="true" box="[1099,1117,1962,1988]" pageId="51" pageNumber="52">F</emphasis>
. Ventral view showing peristome edge directly parallel to umbilical depression.
</paragraph>
</caption>
<paragraph id="8BA6368BFF8EFFF4FF42D66EFE6FFE5F" blockId="52.[189,443,366,395]" box="[189,443,366,395]" pageId="52" pageNumber="53">
<emphasis id="B96DEA99FF8EFFF4FF42D66EFE6FFE5F" bold="true" box="[189,443,366,395]" italics="true" pageId="52" pageNumber="53">Molecular analysis</emphasis>
</paragraph>
<paragraph id="8BA6368BFF8EFFF4FF42D6A4FD15FDB9" blockId="52.[189,1399,419,621]" pageId="52" pageNumber="53">
<taxonomicName id="4C194D08FF8EFFF4FF42D6A4FDD3FE69" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[189,519,419,445]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="52" pageNumber="53" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FF8EFFF4FF42D6A4FDD3FE69" box="[189,519,419,445]" italics="true" pageId="52" pageNumber="53">Zospeum troglobalcanicum</emphasis>
</taxonomicName>
is revealed based on a single 
<typeStatus id="54A28829FF8EFFF4FC75D6A4FC31FE6A" box="[906,997,420,446]" pageId="52" pageNumber="53" type="syntype">syntype</typeStatus>
shell discovered in the NHMW collection and morphologically assessed along with shells deriving from 15 populations of southern Balkan 
<taxonomicName id="4C194D08FF8EFFF4FEE7D6EAFE57FDD7" authorityName="Bourguignat" authorityYear="1856" box="[280,387,490,515]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="52" pageNumber="53" phylum="Mollusca" rank="genus">
<emphasis id="B96DEA99FF8EFFF4FEE7D6EAFE57FDD7" box="[280,387,490,515]" italics="true" pageId="52" pageNumber="53">Zospeum</emphasis>
</taxonomicName>
from both museum collections and recent sampling efforts (
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A. Measurements include shell height (sh), shell width (sw), aperture height (ah), aperture width (aw), height of last whorl (hlw) and spire angle (SA). B. Internal view showing columella (co), lamella (la), peristome (pe) and penultimate whorl (pw). C. Apical view showing protoconch (pr), teleoconch (te) and peristome (pe). D. Ventral view showing peristome configuration (pe) parietal shield (ps) and umbilicus (um)." captionText-1="Fig. 3. Light microscopic images of shells from the caves Benetina pećina (SE Bosnia and Herzegovina) and Cetinska pećina (Montenegro) with specimen labels. A. Zospeum troglobalcanicum Absolon, 1916, lectotype (NHMW Moll.Coll.Edlauer 32.749) from Benetina pećina. B–C. Zospeum tortuosum Jochum &amp; Ruthensteiner sp. nov., paratypes (MCSMNH-PMSL-Moll.-FVelkovrh 34099[spm2-3]) from Cetinska pećina. D. Zospeum tortuosum Jochum &amp; Ruthensteiner sp. nov., holotype (MCSMNH- PMSL-Moll.-FVelkovrh 34099[spm1]) from Cetinska pećina. E. Deformed shell (MCSMNH-PMSL- Moll.-FVelkovrh 34099[spm4]) from Cetinska pećina." captionText-10="Fig. 12. Zospeum neuberti Jochum &amp; Ruthensteiner sp. nov. (NHMW Mol.Coll.Edlauer 32006). A–B. Light microscopic images of paratypes showing aperture and dorsal views and collection labels. C. Light microscopic images of holotype showing aperture and dorsal views. D–I. 3D visualizations of Micro-CT data of holotype. D. Aperture view. E. Aperture facing right view showing smooth columella. F. Apical view.G. Dorsal view showing smooth columella. H. Aperture facing left view. I. Ventral view." captionText-11="Fig. 13. Light microscopic images of Zospeum constrictum Jochum &amp; Ruthensteiner sp. nov. (NHMW Mol.Coll.Edlauer 16.693) and collection labels.A. Holotype, aperture, and dorsal views.B–D. Paratypes, aperture and dorsal views." captionText-12="Fig. 14. 3D visualizations of Micro-CT data of Zospeum constrictum Jochum &amp; Ruthensteiner sp. nov. (NHMW Mol.Coll.Edlauer 16.693). A–F. Holotype. A. Aperture and dorsal views. B. Aperture facing right view. C. Apical view showing upper right side of final whorl tucked underneath spire. D. Dorsal view. E. Aperture facing left view. F. Ventral view. G–L. Paratype (Fig. 13B). G. Aperture view. H. Aperture facing right view. I. Apical view showing upper right side of final whorl tucked underneath spire. J. Dorsal view. K. Aperture facing left view. L. Ventral view." captionText-13="Fig. 15. Zospeum biokovoense Jochum &amp; Ruthensteiner sp. nov. (NHMW Mol.Coll.Edlauer 16.390). A–B. Light microscopic images showing aperture and dorsal views and collection labels. A. Juvenile shell. B. Holotype. C–H. 3D visualizations of Micro-CT data of holotype. C. Aperture view showing gnarled deformation of the columella. D. Aperture facing right view showing fat columella and lamella. E. Apical view. F. Dorsal view showing lopsided spire and incomplete lamella on top of fat columella. G. Aperture facing left view. H. Ventral view showing puckered umbilical depression." captionText-14="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." captionText-15="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." captionText-16="Fig. 18. 3D visualizations of Micro-CT data of Zospeum simplex Inäbnit, Jochum &amp; Neubert, 2021, (NMHW-MO-113642 ex RSC 3760) from Dobravljevac jama, Gornji Brišnik, Bosnia and Herzegovina. A–F. Specimen 1 (spm1). A. Aperture view. B. Aperture facing right view. C. Apical view. D. Dorsal view. E. Aperture facing left view. F. Ventral view. G–L. Specimen 2 (spm2). Same perspectives as Specimen 1." captionText-17="Fig. 19. Zospeum simplex Inäbnit, Jochum &amp; Neubert, 2021, (SMF 349425) Scanning Electron Microscopy images. A. Protoconch and upper teleoconch showing microstructure of superficial pitting. B. Close up view of pitting microstructure. C. Last whorl with axial ribbing extending beyond peristome lip. D. Close up view of second whorl showing rows of interrupted dashes of radial pitting. E. Growth lines and radial banding on teleoconch. F. Close up view of growth lines and shell microstructure." captionText-2="Fig. 4. 3D visualizations using Micro-CT data of shells from the caves Benetina pećina (SE Bosnia and Herzegovina) and Cetinska pećina (Montenegro). A–F. Zospeum troglobalcanicum Absolon, 1916, lectotype (NHMW Moll.Coll.Edlauer 32.749). A. Aperture view. B. Aperture facing right view. C. Apical view. D. Dorsal view. E. Aperture facing left view. F. Ventral view showing umbilical depression. G–L. Zospeum tortuosum Jochum &amp; Ruthensteiner sp. nov., holotype (MCSMNH-PMSL- Moll.-FVelkovrh 34099[spm1]). G. Aperture view. H. Aperture facing right view. I. Apical view. J. Dorsal view. K. Aperture facing left view. L. Ventral view showing callused peristome positioned mostly on top of umbilicus." captionText-3="Fig. 5. 3D visualizations of Micro-CT data of the deformed shell, (MCSMNH-PMSL-Moll.-FVelkovrh 34099[spm4]) from Cetinska pećina. A. Aperture view. B. Aperture facing right view. C. Apical view. D. Dorsal view. E. Aperture facing left view. F. Ventral view with umbilicus covered by last quarter of shell whorl." captionText-4="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." captionText-5="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])." captionText-6="Fig. 8. 3D visualizations of Micro-CT data of shells from the caves Lipska pećina and Duboki do (Montenegro). A–F. 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. G–L. 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." captionText-7="Fig. 9. Zospeum amplioscutum Jochum &amp; Ruthensteiner sp. nov., (NHMW Mol.Coll.Edlauer 19.069) with sample labels. A. Light microscopic images of paratype (NHMW Mol.Coll.Edlauer 19.069), apertural and dorsal views.B. Light microscopic images of holotype (NHMW Mol.Coll.Edlauer 19.069), aperture and dorsal views. C–H. 3D visualizations of X-ray Micro-CT data of holotype (NHMW Mol. Coll.Edlauer 19.069). C. Aperture view. D. Aperture facing right view. E. Apical view. F. Dorsal view. G. Aperture facing left view. H. Ventral view showing coiling projecting high up on shell and peristome oblique to umbilicus." captionText-8="Fig. 10. Zospeum tumidum Jochum, Schilthuizen &amp; Ruthensteiner sp. nov., holotype (NMBE 572615). A. Light microscopic images, showing aperture and dorsal views and TxEx Collection label. B. 3D visualizations of Micro-CT data, apical view. C. Ventral view showing deep umbilical depression. D. Aperture view.E. Aperture facing right view showing tapered columella and lamellar bulge. F. Dorsal view. G. Aperture facing left view." captionText-9="Fig. 11. Zospeum njunjicae Jochum, Schilthuizen &amp; Ruthensteiner sp. nov. A–B. Light microscopic images of paratypes (NMBE 572616) showing aperture and dorsal views. C. Light microscopic images of holotype (NMBE 572617) showing aperture and dorsal views. D–I. 3D visualizations of X-ray micro- CT data of holotype (NMBE 572617). D. Aperture view. E. Aperture facing right view showing well defined lamella. F. Apical view. G. Dorsal view. H. Aperture facing left view. I. Ventral view." figureDoi-0="http://doi.org/10.5281/zenodo.10847810" figureDoi-1="http://doi.org/10.5281/zenodo.10847812" figureDoi-10="http://doi.org/10.5281/zenodo.10847838" figureDoi-11="http://doi.org/10.5281/zenodo.10847842" figureDoi-12="http://doi.org/10.5281/zenodo.10847846" figureDoi-13="http://doi.org/10.5281/zenodo.10847848" figureDoi-14="http://doi.org/10.5281/zenodo.10847850" figureDoi-15="http://doi.org/10.5281/zenodo.10847854" figureDoi-16="http://doi.org/10.5281/zenodo.10847858" figureDoi-17="http://doi.org/10.5281/zenodo.10847860" figureDoi-2="http://doi.org/10.5281/zenodo.10847814" figureDoi-3="http://doi.org/10.5281/zenodo.10847816" figureDoi-4="http://doi.org/10.5281/zenodo.10847820" figureDoi-5="http://doi.org/10.5281/zenodo.10847822" figureDoi-6="http://doi.org/10.5281/zenodo.10847826" figureDoi-7="http://doi.org/10.5281/zenodo.10847828" figureDoi-8="http://doi.org/10.5281/zenodo.10847832" figureDoi-9="http://doi.org/10.5281/zenodo.10847834" httpUri-0="https://zenodo.org/record/10847810/files/figure.png" httpUri-1="https://zenodo.org/record/10847812/files/figure.png" httpUri-10="https://zenodo.org/record/10847838/files/figure.png" httpUri-11="https://zenodo.org/record/10847842/files/figure.png" httpUri-12="https://zenodo.org/record/10847846/files/figure.png" httpUri-13="https://zenodo.org/record/10847848/files/figure.png" httpUri-14="https://zenodo.org/record/10847850/files/figure.png" httpUri-15="https://zenodo.org/record/10847854/files/figure.png" httpUri-16="https://zenodo.org/record/10847858/files/figure.png" httpUri-17="https://zenodo.org/record/10847860/files/figure.png" httpUri-2="https://zenodo.org/record/10847814/files/figure.png" httpUri-3="https://zenodo.org/record/10847816/files/figure.png" httpUri-4="https://zenodo.org/record/10847820/files/figure.png" httpUri-5="https://zenodo.org/record/10847822/files/figure.png" httpUri-6="https://zenodo.org/record/10847826/files/figure.png" httpUri-7="https://zenodo.org/record/10847828/files/figure.png" httpUri-8="https://zenodo.org/record/10847832/files/figure.png" httpUri-9="https://zenodo.org/record/10847834/files/figure.png" pageId="52" pageNumber="53">Figs 2–19</figureCitation>
). Two recently collected individuals from one population from Njeguši, 
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were additionally assessed by DNA sequencing in accordance with the most recent integrative study by 
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Inäbnit 
<emphasis id="B96DEA99FF8EFFF4FB5FD531FB03FD9E" box="[1184,1239,560,586]" italics="true" pageId="52" pageNumber="53">et al</emphasis>
. (2021)
</bibRefCitation>
. The phylogeny is updated in this work (
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).
</paragraph>
<paragraph id="8BA6368BFF8EFFF4FF42D595FB41FCCC" blockId="52.[189,1399,660,795]" pageId="52" pageNumber="53">
Tree topologies in both the ML and the BI tree were identical to each other and largely congruent to the ones published in 
<bibRefCitation id="EF884B7AFF8EFFF4FE6AD5B8FD53FD06" author="Inabnit T. &amp; Jochum A. &amp; Slapnik R. &amp; Neubert E." box="[405,647,696,722]" pageId="52" pageNumber="53" pagination="175 - 193" refId="ref31366" refString="Inabnit T., Jochum A., Slapnik R. &amp; Neubert E. 2021. New genetic data reveals a new species of Zospeum in Bosnia (Gastropoda, Ellobioidea, Carychiinae). ZooKeys 1071: 175 - 193. https: // doi. org / 10.3897 / zookeys. 1071.66417" type="journal article" year="2021">
Inäbnit 
<emphasis id="B96DEA99FF8EFFF4FE0ED5B9FDF2FD06" box="[497,550,696,722]" italics="true" pageId="52" pageNumber="53">et al</emphasis>
. (2021)
</bibRefCitation>
. The two new specimens from Njeguši, 
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(
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<emphasis id="B96DEA99FF8EFFF4FAF8D5B9FAA3FD06" box="[1287,1399,696,722]" italics="true" pageId="52" pageNumber="53">Z. kolbae</emphasis>
Jochum, Inäbnit, Kneubühler &amp; Ruthensteiner
</taxonomicName>
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NMBE 571122–571123), were recovered as the sister group of 
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<emphasis id="B96DEA99FF8EFFF4FE91D5FFFE52FCCC" box="[366,390,767,792]" italics="true" pageId="52" pageNumber="53">Z.</emphasis>
aff. 
<emphasis id="B96DEA99FF8EFFF4FE43D5FEFD5AFCCC" box="[444,654,766,792]" italics="true" pageId="52" pageNumber="53">troglobalcanicum</emphasis>
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from Špilja Jezero in southernmost 
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.
</paragraph>
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The ABGD analysis for the 
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<emphasis id="B96DEA99FF8EFFF4FE02D440FDADFC8D" box="[509,633,832,857]" italics="true" pageId="52" pageNumber="53">Z. pretneri</emphasis>
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-group yielded two different topologies, one with six clades (Prior Maximal Distance P = 
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3.59 
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02; barcode gap distance: 0.042) and one with eight clades (Prior Maximal Distance P = 
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4.64 
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03; barcode gap distance: 0.004). Both topologies regard the species from Njeguši, 
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, as a distinct species. The topology with six clades corresponds to the one that was ultimately used for species delimitation in 
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Inäbnit 
<emphasis id="B96DEA99FF8EFFF4FD65D4CDFD01FC32" box="[666,725,972,998]" italics="true" pageId="52" pageNumber="53">et al.</emphasis>
(2021)
</bibRefCitation>
and which accepts all currently accepted species within the group as independent lineages, while the topology with eight clades subdivides 
<taxonomicName id="4C194D08FF8EFFF4FB02D4F0FAACFBDE" authorityName="Inabnit, Jochum &amp; Neubert" authorityYear="2021" box="[1277,1400,1008,1034]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" kingdom="Animalia" order="Ellobiida" pageId="52" pageNumber="53" phylum="Mollusca" rank="species" species="simplex">
<emphasis id="B96DEA99FF8EFFF4FB02D4F0FAACFBDE" box="[1277,1400,1008,1034]" italics="true" pageId="52" pageNumber="53">Z. simplex</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C194D08FF8EFFF4FF10D314FDC4FBF9" authorityName="Absolon, 1916 Absolon" authorityYear="1916" box="[239,528,1043,1069]" class="Gastropoda" family="Ellobiidae" genus="Zospeum" isUncertain="true" kingdom="Animalia" order="Ellobiida" pageId="52" pageNumber="53" phylum="Mollusca" rank="species" species="troglobalcanicum">
<emphasis id="B96DEA99FF8EFFF4FF10D314FED3FBF9" box="[239,263,1044,1069]" italics="true" pageId="52" pageNumber="53">Z.</emphasis>
aff. 
<emphasis id="B96DEA99FF8EFFF4FEC1D313FDC4FBF9" box="[318,528,1043,1069]" italics="true" pageId="52" pageNumber="53">troglobalcanicum</emphasis>
</taxonomicName>
into two lineages each (with the two lineages in the latter co-occurring in the same cave). We will use the topology with six clades from now on.
</paragraph>
</subSubSection>
</treatment>
</document>