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<document id="B6688835719D6F6D34C2388F08DC0F45" ID-CLB-Dataset="3658" ID-DOI="10.1093/zoolinnean/zlaa156" ID-GBIF-Dataset="c61c4597-0700-4f1d-bf39-ff3059c60d42" ID-ISSN="0024-4082" ID-Zenodo-Dep="5636863" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1635756554962" checkinUser="plazi" docAuthor="Balashov, Igor A., Neiber, Marco T. &amp; Hausdorf, Bernhard" docDate="2020" docId="03A687CEFFCB0E72FC6488C7FDACF9ED" docLanguage="en" docName="zlaa156.pdf" docOrigin="Zoological Journal of the Linnean Society 193" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-2023.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-2023.journal_article" docStyleVersion="7" docTitle="Helicopsis Fitzinger 1833" docType="treatment" docVersion="6" lastPageNumber="1119" masterDocId="FF9FFFB6FFCC0E7EFFDC8E4BFFBFFFA5" masterDocTitle="Phylogeny, species delimitation and population structure of the steppe-inhabiting land snail genus Helicopsis in Eastern Europe" masterLastPageNumber="1125" masterPageNumber="1108" pageNumber="1115" updateTime="1699467228909" updateUser="ExternalLinkService">
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<mods:title id="52703F82C5F1B63F98326FFE3A725998">Phylogeny, species delimitation and population structure of the steppe-inhabiting land snail genus Helicopsis in Eastern Europe</mods:title>
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<mods:namePart id="5F58EC2B0031AE442D03B35E00263232">Balashov, Igor A.</mods:namePart>
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<mods:namePart id="4E1FF99DCFE5EB78BE86428C477F1C90">Neiber, Marco T.</mods:namePart>
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<subSubSection id="C3156553FFCB0E79FC6488C7FABEF901" box="[952,1281,1676,1701]" pageId="7" pageNumber="1115" type="nomenclature">
<paragraph id="8BB036D8FFCB0E79FC6488C7FABEF901" blockId="7.[952,1281,1676,1701]" box="[952,1281,1676,1701]" pageId="7" pageNumber="1115">
<heading id="D0F881B4FFCB0E79FC6488C7FABEF901" box="[952,1281,1676,1701]" centered="true" fontSize="9" level="2" pageId="7" pageNumber="1115" reason="2">
SYSTEMATICS OF
<taxonomicName id="4C0F4D5BFFCB0E79FBA188DBFABEF901" ID-CoL="635L7" ID-ENA="265987" authorityName="Fitzinger" authorityYear="1833" box="[1149,1281,1680,1700]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="7" pageNumber="1115" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFCB0E79FBA188DBFABEF901" box="[1149,1281,1680,1700]" italics="true" pageId="7" pageNumber="1115">HELICOPSIS</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C3156553FFCB0E72FCF588FEFDACF9ED" lastPageId="12" lastPageNumber="1120" pageId="7" pageNumber="1115" type="description">
<paragraph id="8BB036D8FFCB0E76FCF588FEFAECF9CC" blockId="7.[809,1426,1717,1892]" lastBlockId="8.[827,1443,1527,1856]" lastPageId="8" lastPageNumber="1116" pageId="7" pageNumber="1115">
Our phylogenetic analysis of 16S rDNA sequences (
<figureCitation id="13342A5DFFCB0E79FCED8898FCCFF94D" box="[817,880,1747,1769]" captionStart="Figure 2" captionStartId="5.[145,227,1817,1839]" captionTargetBox="[272,1296,410,1747]" captionTargetId="graphics-142@5.[272,1273,411,1748]" captionTargetPageId="5" captionText="Figure 2. Bayesian 50% majority-rule consensus tree of Helicopsis and related groups based on sequences of the mitochondrial 16S rDNA. Nodes supported by posterior probabilities ≥ 0.95 are marked by a dot. For information about sequenced specimens and detailed support values, see Supporting Information, Figure S1 and Table S1." figureDoi="http://doi.org/10.5281/zenodo.5636867" httpUri="https://zenodo.org/record/5636867/files/figure.png" pageId="7" pageNumber="1115">Fig. 2</figureCitation>
; Supporting Information,
<figureCitation id="13342A5DFFCB0E79FB458898FB57F94D" box="[1177,1256,1747,1769]" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="7" pageNumber="1115">Fig. S1</figureCitation>
) confirms that
<taxonomicName id="4C0F4D5BFFCB0E79FCF588B9FC20F8A2" authorityName="Fitzinger" authorityYear="1833" box="[809,927,1778,1799]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="7" pageNumber="1115" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFCB0E79FCF588B9FC20F8A2" box="[809,927,1778,1799]" italics="true" pageId="7" pageNumber="1115">Helicopsis</emphasis>
</taxonomicName>
forms a clade with
<taxonomicName id="4C0F4D5BFFCB0E79FB5A88B9FB4AF8A2" authorityName="Monterosato" authorityYear="1892" box="[1158,1269,1778,1799]" class="Gastropoda" family="Geomitridae" genus="Xerolenta" kingdom="Animalia" order="Stylommatophora" pageId="7" pageNumber="1115" phylum="Mollusca" rank="genus">
<emphasis id="B97BEACAFFCB0E79FB5A88B9FB4AF8A2" box="[1158,1269,1778,1799]" italics="true" pageId="7" pageNumber="1115">Xerolenta</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D5BFFCB0E79FADF88B9FA35F8A2" authorityName="Monterosato" authorityYear="1892" box="[1283,1418,1778,1799]" class="Gastropoda" family="Geomitridae" genus="Xeromunda" kingdom="Animalia" order="Stylommatophora" pageId="7" pageNumber="1115" phylum="Mollusca" rank="genus">
<emphasis id="B97BEACAFFCB0E79FADF88B9FA35F8A2" box="[1283,1418,1778,1799]" italics="true" pageId="7" pageNumber="1115">Xeromunda</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D5BFFCB0E79FCF5895AFC2AF883" authorityName="Monterosato" authorityYear="1892" box="[809,917,1809,1830]" class="Gastropoda" family="Geomitridae" genus="Xeropicta" kingdom="Animalia" order="Stylommatophora" pageId="7" pageNumber="1115" phylum="Mollusca" rank="genus">
<emphasis id="B97BEACAFFCB0E79FCF5895AFC2AF883" box="[809,917,1809,1830]" italics="true" pageId="7" pageNumber="1115">Xeropicta</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFCB0E79FC12895AFB35F883" authorityName="Westerlund" authorityYear="1879" box="[974,1162,1809,1830]" class="Gastropoda" family="Geomitridae" genus="Pseudoxerophila" kingdom="Animalia" order="Stylommatophora" pageId="7" pageNumber="1115" phylum="Mollusca" rank="genus">
<emphasis id="B97BEACAFFCB0E79FC12895AFB35F883" box="[974,1162,1809,1830]" italics="true" pageId="7" pageNumber="1115">Pseudoxerophila</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9E4B29FFCB0E79FB45895AFCDAF8E0" author="Hausdorf B &amp; Bossneck U" pageId="7" pageNumber="1115" pagination="194 - 200" refId="ref12265" refString="Hausdorf B, Bossneck U. 2016. Helicopsis persica n. sp. from northern Iran (Gastropoda: Geomitridae). Zootaxa 4066: 194 - 200." type="journal article" year="2016">Hausdorf &amp; Bössneck 2016</bibRefCitation>
;
<bibRefCitation id="EF9E4B29FFCB0E79FCA58964FBCFF8E1" author="Neiber MT &amp; Razkin O &amp; Hausdorf B" box="[889,1136,1839,1861]" pageId="7" pageNumber="1115" pagination="169 - 184" refId="ref12949" refString="Neiber MT, Razkin O, Hausdorf B. 2017. Molecular phylogeny and biogeography of the land snail family Hygromiidae (Gastropoda: Helicoidea). Molecular Phylogenetics and Evolution 111: 169 - 184." type="journal article" year="2017">
Neiber
<emphasis id="B97BEACAFFCB0E79FC00897BFBA0F8E1" box="[988,1055,1839,1861]" italics="true" pageId="7" pageNumber="1115">et al.</emphasis>
, 2017
</bibRefCitation>
) and shows that also
<taxonomicName id="4C0F4D5BFFCB0E79FCF58905FB08F8C1" authority="Hudec &amp; Lezhawa, 1967" authorityName="Hudec &amp; Lezhawa" authorityYear="1967" box="[809,1207,1870,1892]" class="Gastropoda" family="Hygromiidae" genus="Kalitinaia" kingdom="Animalia" order="Stylommatophora" pageId="7" pageNumber="1115" phylum="Mollusca" rank="genus">
<emphasis id="B97BEACAFFCB0E79FCF58905FC21F8C6" box="[809,926,1870,1891]" italics="true" pageId="7" pageNumber="1115">Kalitinaia</emphasis>
Hudec &amp; Lezhawa, 1967
</taxonomicName>
from the Caucasus region belongs in this clade named Helicopsini. The delimitation of
<taxonomicName id="4C0F4D5BFFC40E76FE91885DFE7FF98E" authorityName="Fitzinger" authorityYear="1833" box="[333,448,1558,1579]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC40E76FE91885DFE7FF98E" box="[333,448,1558,1579]" italics="true" pageId="8" pageNumber="1116">Helicopsis</emphasis>
</taxonomicName>
was considered questionable, because it is characterized only by plesiomorphic character states (
<bibRefCitation id="EF9E4B29FFC40E76FE548818FDF4F9CC" author="Hausdorf B" box="[392,587,1619,1641]" pageId="8" pageNumber="1116" pagination="65 - 71" refId="ref12176" refString="Hausdorf B. 1996. Helicopsis aelleni n. sp. from northern Iran, with remarks on Helicopsis Fitzinger 1833 (Gastropoda: Pulmonata: Hygromiidae). Archiv fur Molluskenkunde 126: 65 - 71." type="journal article" year="1996">Hausdorf, 1996</bibRefCitation>
). Actually, the molecular phylogeny based on mitochondrial 16S rDNA sequences (
<figureCitation id="13342A5DFFC40E76FEAE88DAFE0DF903" box="[370,434,1681,1703]" captionStart="Figure 2" captionStartId="5.[145,227,1817,1839]" captionTargetBox="[272,1296,410,1747]" captionTargetId="graphics-142@5.[272,1273,411,1748]" captionTargetPageId="5" captionText="Figure 2. Bayesian 50% majority-rule consensus tree of Helicopsis and related groups based on sequences of the mitochondrial 16S rDNA. Nodes supported by posterior probabilities ≥ 0.95 are marked by a dot. For information about sequenced specimens and detailed support values, see Supporting Information, Figure S1 and Table S1." figureDoi="http://doi.org/10.5281/zenodo.5636867" httpUri="https://zenodo.org/record/5636867/files/figure.png" pageId="8" pageNumber="1116">Fig. 2</figureCitation>
; Supporting Information,
<figureCitation id="13342A5DFFC40E76FD3C88DAFF7EF960" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="8" pageNumber="1116">Fig. S1</figureCitation>
) revealed that
<taxonomicName id="4C0F4D5BFFC40E76FEA788FBFD87F961" baseAuthorityName="Nagele" baseAuthorityYear="1903" box="[379,568,1711,1733]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="subcalcarata">
<emphasis id="B97BEACAFFC40E76FEA788FBFE2FF960" box="[379,400,1712,1733]" italics="true" pageId="8" pageNumber="1116">H</emphasis>
.
<emphasis id="B97BEACAFFC40E76FE4388E4FD87F961" box="[415,568,1711,1732]" italics="true" pageId="8" pageNumber="1116">subcalcarata</emphasis>
</taxonomicName>
from
<collectingCountry id="F3187648FFC40E76FD5C88E4FD69F960" box="[640,726,1711,1733]" name="Turkey" pageId="8" pageNumber="1116">Turkey</collectingCountry>
and
<taxonomicName id="4C0F4D5BFFC40E76FF7F8885FEFAF941" box="[163,325,1742,1764]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="undetermined">
<emphasis id="B97BEACAFFC40E76FF7F8885FEA3F946" box="[163,284,1742,1763]" italics="true" pageId="8" pageNumber="1116">Helicopsis</emphasis>
sp.
</taxonomicName>
from
<collectingCountry id="F3187648FFC40E76FE528885FE5AF946" box="[398,485,1742,1763]" name="Cyprus" pageId="8" pageNumber="1116">Cyprus</collectingCountry>
, which is closely related to
<taxonomicName id="4C0F4D5BFFC40E76FF1F88A6FD86F8A6" authority="(Westerlund, 1889)" baseAuthorityName="Westerlund" baseAuthorityYear="1889" box="[195,569,1773,1795]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="cypriola">
<emphasis id="B97BEACAFFC40E76FF1F88A6FF67F8A7" box="[195,216,1773,1794]" italics="true" pageId="8" pageNumber="1116">H</emphasis>
.
<emphasis id="B97BEACAFFC40E76FF3488A6FEF7F8A7" box="[232,328,1773,1794]" italics="true" pageId="8" pageNumber="1116">cypriola</emphasis>
(Westerlund, 1889)
</taxonomicName>
, do not belong to
<taxonomicName id="4C0F4D5BFFC40E76FF7F8940FEA1F885" authorityName="Fitzinger" authorityYear="1833" box="[163,286,1803,1824]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC40E76FF7F8940FEA1F885" box="[163,286,1803,1824]" italics="true" pageId="8" pageNumber="1116">Helicopsis</emphasis>
</taxonomicName>
.
<taxonomicName id="4C0F4D5BFFC40E76FEF28940FDF1F885" baseAuthorityName="Nagele" baseAuthorityYear="1903" box="[302,590,1803,1824]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="subcalcarata">
<emphasis id="B97BEACAFFC40E76FEF28940FDF1F885" box="[302,590,1803,1824]" italics="true" pageId="8" pageNumber="1116">Helicopsis subcalcarata</emphasis>
</taxonomicName>
is sister to the remaining Helicopsini and, thus, a separate genus is required for this species.
<taxonomicName id="4C0F4D5BFFC40E76FBBB8BBCFAB9F9A8" box="[1127,1286,1527,1549]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="undetermined">
<emphasis id="B97BEACAFFC40E76FBBB8BBCFB61F9A9" box="[1127,1246,1527,1548]" italics="true" pageId="8" pageNumber="1116">Helicopsis</emphasis>
sp.
</taxonomicName>
from
<collectingCountry id="F3187648FFC40E76FA928BB3FA1DF9A8" box="[1358,1442,1528,1549]" name="Cyprus" pageId="8" pageNumber="1116">Cyprus</collectingCountry>
is closely related to the eastern Mediterranean genus
<taxonomicName id="4C0F4D5BFFC40E76FCE7887EFBB9F9EF" authorityName="Westerlund" authorityYear="1879" box="[827,1030,1589,1610]" class="Gastropoda" family="Geomitridae" genus="Pseudoxerophila" kingdom="Animalia" order="Stylommatophora" pageId="8" pageNumber="1116" phylum="Mollusca" rank="genus">
<emphasis id="B97BEACAFFC40E76FCE7887EFBB9F9EF" box="[827,1030,1589,1610]" italics="true" pageId="8" pageNumber="1116">Pseudoxerophila</emphasis>
</taxonomicName>
. Amore detailed analysis of the relationships of these species is in preparation.
</paragraph>
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<paragraph id="8BB036D8FFC40E76FF7F8B63FE9BFA74" blockId="8.[163,1443,1320,1489]" pageId="8" pageNumber="1116">
<emphasis id="B97BEACAFFC40E76FF7F8B63FEAFFA9B" bold="true" box="[163,272,1320,1342]" pageId="8" pageNumber="1116">Figure 4.</emphasis>
Neighbour-net network of
<taxonomicName id="4C0F4D5BFFC40E76FDE38B62FD11FA9A" authorityName="Fitzinger" authorityYear="1833" box="[575,686,1321,1343]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC40E76FDE38B62FD11FA9A" box="[575,686,1321,1343]" italics="true" pageId="8" pageNumber="1116">Helicopsis</emphasis>
</taxonomicName>
from Ukraine based on Jaccard distances obtained from AFLP data. Coloration of clusters corresponds to the STRUCTURE solution for
<emphasis id="B97BEACAFFC40E76FCA78B0CFC33FAF9" box="[891,908,1351,1372]" italics="true" pageId="8" pageNumber="1116">K</emphasis>
= 2. Red corresponds to
<taxonomicName id="4C0F4D5BFFC40E76FB4F8B0CFAA9FAF9" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1171,1302,1350,1372]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC40E76FB4F8B0CFB19FAF9" box="[1171,1190,1351,1372]" italics="true" pageId="8" pageNumber="1116">H</emphasis>
.
<emphasis id="B97BEACAFFC40E76FB6F8B0DFAA9FAF9" box="[1203,1302,1350,1372]" italics="true" pageId="8" pageNumber="1116">filimargo</emphasis>
</taxonomicName>
, yellow to
<taxonomicName id="4C0F4D5BFFC40E76FA568B0CFF41FADC" baseAuthorityName="Krynicki" baseAuthorityYear="1833" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC40E76FA568B0CFA22FAF9" box="[1418,1437,1351,1372]" italics="true" pageId="8" pageNumber="1116">H</emphasis>
.
<emphasis id="B97BEACAFFC40E76FF7F8B28FF41FADC" box="[163,254,1379,1401]" italics="true" pageId="8" pageNumber="1116">lunulata</emphasis>
</taxonomicName>
. DNA voucher numbers for specimens are given at the tips of the network. Coloured dots at the tips of the network correspond to cluster assignments (on majority-rule basis) of specimens in the STRUCTURE solutions for
<emphasis id="B97BEACAFFC40E76FAC78BCAFA93FA33" box="[1307,1324,1409,1430]" italics="true" pageId="8" pageNumber="1116">K</emphasis>
= 5 (inner) and
<emphasis id="B97BEACAFFC40E76FF0C8BD4FF5EFA11" box="[208,225,1439,1460]" italics="true" pageId="8" pageNumber="1116">K</emphasis>
= 7 (outer). For locality data and the distribution of clusters, see also Supporting Information, Table S1 and Figure 6, respectively.
</paragraph>
</caption>
<paragraph id="8BB036D8FFC40E77FC8F8839FDA8F9DE" blockId="8.[827,1443,1527,1856]" lastBlockId="9.[145,762,1270,1874]" lastPageId="9" lastPageNumber="1117" pageId="8" pageNumber="1116">
<taxonomicName id="4C0F4D5BFFC40E76FC8F8839FB32F922" authorityName="Hausdorf" authorityYear="1990" box="[851,1165,1650,1671]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="gittenbergeri">
<emphasis id="B97BEACAFFC40E76FC8F8839FB32F922" box="[851,1165,1650,1671]" italics="true" pageId="8" pageNumber="1116">Helicopsis gittenbergeri</emphasis>
</taxonomicName>
from
<collectingCountry id="F3187648FFC40E76FB388839FA83F922" box="[1252,1340,1650,1671]" name="Greece" pageId="8" pageNumber="1116">Greece</collectingCountry>
formed together with
<taxonomicName id="4C0F4D5BFFC40E76FC3088DAFBDBF903" authorityName="Hausdorf" authorityYear="2006" box="[1004,1124,1681,1702]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="aelleni">
<emphasis id="B97BEACAFFC40E76FC3088DAFBBEF903" box="[1004,1025,1681,1702]" italics="true" pageId="8" pageNumber="1116">H</emphasis>
.
<emphasis id="B97BEACAFFC40E76FBCE88DAFBDBF903" box="[1042,1124,1681,1702]" italics="true" pageId="8" pageNumber="1116">aelleni</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC40E76FB7988DAFA9EF903" authorityName="Hausdorf &amp; Bossneck" authorityYear="2016" box="[1189,1313,1681,1702]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="persica">
<emphasis id="B97BEACAFFC40E76FB7988DAFB05F903" box="[1189,1210,1681,1702]" italics="true" pageId="8" pageNumber="1116">H</emphasis>
.
<emphasis id="B97BEACAFFC40E76FB1788DAFA9EF903" box="[1227,1313,1681,1702]" italics="true" pageId="8" pageNumber="1116">persica</emphasis>
</taxonomicName>
from
<collectingCountry id="F3187648FFC40E76FAB288DAFA1DF903" box="[1390,1442,1681,1702]" name="Iran" pageId="8" pageNumber="1116">Iran</collectingCountry>
the well-supported sister-group of the
<taxonomicName id="4C0F4D5BFFC40E76FAF888E4FC31F941" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="undetermined">
<emphasis id="B97BEACAFFC40E76FAF888E4FA1DF961" box="[1316,1442,1711,1732]" italics="true" pageId="8" pageNumber="1116">Helicopsis</emphasis>
species
</taxonomicName>
from Eastern and Central Europe [as in the phylogenetic analysis of
<bibRefCitation id="EF9E4B29FFC40E76FB8D88A6FA2BF8A6" author="Hausdorf B &amp; Bossneck U" box="[1105,1428,1773,1795]" pageId="8" pageNumber="1116" pagination="194 - 200" refId="ref12265" refString="Hausdorf B, Bossneck U. 2016. Helicopsis persica n. sp. from northern Iran (Gastropoda: Geomitridae). Zootaxa 4066: 194 - 200." type="journal article" year="2016">Hausdorf &amp; Bössneck (2016)</bibRefCitation>
]. The monophyly of the Eastern and Central European
<taxonomicName id="4C0F4D5BFFC40E76FCE78961FBABF8E5" box="[827,1044,1834,1856]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="8" pageNumber="1116" phylum="Ascomycota" rank="species" species="undetermined">
<emphasis id="B97BEACAFFC40E76FCE78961FC0AF89A" box="[827,949,1834,1855]" italics="true" pageId="8" pageNumber="1116">Helicopsis</emphasis>
species
</taxonomicName>
, as well as the monophyly of the Central European
<taxonomicName id="4C0F4D5BFFC50E77FE828ABDFE6EFAAE" authorityName="Fitzinger" authorityYear="1833" box="[350,465,1270,1291]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC50E77FE828ABDFE6EFAAE" box="[350,465,1270,1291]" italics="true" pageId="9" pageNumber="1117">Helicopsis</emphasis>
</taxonomicName>
group including
<taxonomicName id="4C0F4D5BFFC50E77FD558ABCFD4CFAA9" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[649,755,1271,1292]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC50E77FD558ABCFD4CFAA9" box="[649,755,1271,1292]" italics="true" pageId="9" pageNumber="1117">H.striata</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D5BFFC50E77FF4D8B5EFE9FFA8F" authorityName="E. Gittenberger" authorityYear="1969" box="[145,288,1301,1322]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="austriaca">
<emphasis id="B97BEACAFFC50E77FF4D8B5EFE9FFA8F" box="[145,288,1301,1322]" italics="true" pageId="9" pageNumber="1117">H. austriaca</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC50E77FE878B5EFE4BFA8F" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[347,500,1301,1322]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC50E77FE878B5EFE4BFA8F" box="[347,500,1301,1322]" italics="true" pageId="9" pageNumber="1117">H. hungarica</emphasis>
</taxonomicName>
, are well supported in the 16S rDNA tree (
<figureCitation id="13342A5DFFC50E77FE588B78FE78FAED" box="[388,455,1331,1353]" captionStart="Figure 2" captionStartId="5.[145,227,1817,1839]" captionTargetBox="[272,1296,410,1747]" captionTargetId="graphics-142@5.[272,1273,411,1748]" captionTargetPageId="5" captionText="Figure 2. Bayesian 50% majority-rule consensus tree of Helicopsis and related groups based on sequences of the mitochondrial 16S rDNA. Nodes supported by posterior probabilities ≥ 0.95 are marked by a dot. For information about sequenced specimens and detailed support values, see Supporting Information, Figure S1 and Table S1." figureDoi="http://doi.org/10.5281/zenodo.5636867" httpUri="https://zenodo.org/record/5636867/files/figure.png" pageId="9" pageNumber="1117">Fig. 2</figureCitation>
; Supporting Information,
<figureCitation id="13342A5DFFC50E77FF4D8B19FF5EFAC2" box="[145,225,1362,1384]" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="9" pageNumber="1117">Fig. S1</figureCitation>
) and the tree based on the concatenated
<emphasis id="B97BEACAFFC50E77FD1F8B19FD4CFAC2" box="[707,755,1362,1383]" italics="true" pageId="9" pageNumber="1117">COI</emphasis>
, 12S rDNA and 16S rDNA sequences (
<figureCitation id="13342A5DFFC50E77FDB68B3AFD0EFA23" box="[618,689,1393,1415]" captionStart="Figure 3" captionStartId="6.[827,907,1649,1671]" captionTargetBox="[836,1433,1245,1607]" captionTargetId="graphics-830@6.[835,1247,1251,1596]" captionTargetPageId="6" captionText="Figure 3. Bayesian 50% majority-rule consensus tree of European Helicopsis based on concatenated sequences of the mitochondrial COI, 12S rDNA and 16S rDNA. Support values at the nodes correspond to Bayesian posterior probabilities (left), maximum likelihood (middle) and maximum parsimony (right) bootstrap values. Extraction voucher numbers are given at the tips of the tree. For information about sequenced specimens, see Supporting Information, Table S1." figureDoi="http://doi.org/10.5281/zenodo.5636869" httpUri="https://zenodo.org/record/5636869/files/figure.png" pageId="9" pageNumber="1117">Fig. 3</figureCitation>
). The populations from Eastern Europe, which are the focus of this study, form two highly supported clades in these trees, for which the oldest names are
<taxonomicName id="4C0F4D5BFFC50E77FDAA8B86FEEBF9A4" authority="(Krynicki, 1833)" baseAuthorityName="Krynicki" baseAuthorityYear="1833" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC50E77FDAA8B86FD46FA44" box="[630,761,1484,1506]" italics="true" pageId="9" pageNumber="1117">H. lunulata</emphasis>
(Krynicki, 1833)
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC50E77FE4F8BA7FD4BF9A4" authority="(Krynicki, 1833)" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[403,756,1515,1537]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC50E77FE4F8BA7FE17F9A4" box="[403,424,1516,1537]" italics="true" pageId="9" pageNumber="1117">H</emphasis>
.
<emphasis id="B97BEACAFFC50E77FE6B8BA0FD99F9A5" box="[439,550,1515,1536]" italics="true" pageId="9" pageNumber="1117">filimargo</emphasis>
(Krynicki, 1833)
</taxonomicName>
. The analyses of the concatenated sequences show that
<taxonomicName id="4C0F4D5BFFC50E77FF4D8862FE9FF998" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[145,288,1576,1598]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC50E77FF4D8862FE9FF998" box="[145,288,1576,1598]" italics="true" pageId="9" pageNumber="1117">H. filimargo</emphasis>
</taxonomicName>
is sister to the Central European clade, while
<taxonomicName id="4C0F4D5BFFC50E77FF078803FED9F9F9" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[219,358,1607,1629]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC50E77FF078803FF4FF9F8" box="[219,240,1608,1629]" italics="true" pageId="9" pageNumber="1117">H</emphasis>
.
<emphasis id="B97BEACAFFC50E77FF23880CFED9F9F9" box="[255,358,1607,1628]" italics="true" pageId="9" pageNumber="1117">lunulata</emphasis>
</taxonomicName>
is sister to all other Central and Eastern European species (
<figureCitation id="13342A5DFFC50E77FE14882DFDB7F9DE" box="[456,520,1638,1660]" captionStart="Figure 3" captionStartId="6.[827,907,1649,1671]" captionTargetBox="[836,1433,1245,1607]" captionTargetId="graphics-830@6.[835,1247,1251,1596]" captionTargetPageId="6" captionText="Figure 3. Bayesian 50% majority-rule consensus tree of European Helicopsis based on concatenated sequences of the mitochondrial COI, 12S rDNA and 16S rDNA. Support values at the nodes correspond to Bayesian posterior probabilities (left), maximum likelihood (middle) and maximum parsimony (right) bootstrap values. Extraction voucher numbers are given at the tips of the tree. For information about sequenced specimens, see Supporting Information, Table S1." figureDoi="http://doi.org/10.5281/zenodo.5636869" httpUri="https://zenodo.org/record/5636869/files/figure.png" pageId="9" pageNumber="1117">Fig. 3</figureCitation>
).
</paragraph>
<caption id="DF706650FFC50E77FF4D8A22FD49FB7D" ID-DOI="http://doi.org/10.5281/zenodo.5636873" ID-Zenodo-Dep="5636873" httpUri="https://zenodo.org/record/5636873/files/figure.png" pageId="9" pageNumber="1117" startId="9.[145,223,1129,1151]" targetBox="[145,1426,199,1090]" targetPageId="9">
<paragraph id="8BB036D8FFC50E77FF4D8A22FD49FB7D" blockId="9.[145,1425,1129,1240]" pageId="9" pageNumber="1117">
<emphasis id="B97BEACAFFC50E77FF4D8A22FF46FBDA" bold="true" box="[145,249,1129,1151]" pageId="9" pageNumber="1117">Figure 5.</emphasis>
Assignment of specimens of
<taxonomicName id="4C0F4D5BFFC50E77FDF08A21FD29FB25" authorityName="Fitzinger" authorityYear="1833" box="[556,662,1130,1152]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC50E77FDF08A21FD29FB25" box="[556,662,1130,1152]" italics="true" pageId="9" pageNumber="1117">Helicopsis</emphasis>
</taxonomicName>
from Ukraine to clusters resulting from the admixture analysis of AFLP data with STRUCTURE. A, solution for
<emphasis id="B97BEACAFFC50E77FD9C8AC3FDEEFB38" box="[576,593,1160,1181]" italics="true" pageId="9" pageNumber="1117">K</emphasis>
= 2. B, solution for
<emphasis id="B97BEACAFFC50E77FCFF8AC3FC8BFB38" box="[803,820,1160,1181]" italics="true" pageId="9" pageNumber="1117">K</emphasis>
= 5. C, solution for
<emphasis id="B97BEACAFFC50E77FBDA8AC3FBA8FB38" box="[1030,1047,1160,1181]" italics="true" pageId="9" pageNumber="1117">K</emphasis>
= 7. Numbers below the plots refer to populations (see Fig. 6; Supporting Information, Table S1). Letters above the plots refer to mitochondrial clades: a,
<taxonomicName id="4C0F4D5BFFC50E77FAA58AEEFF4DFB7D" baseAuthorityName="Krynicki" baseAuthorityYear="1833" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC50E77FAA58AEEFA33FB1F" box="[1401,1420,1189,1210]" italics="true" pageId="9" pageNumber="1117">H</emphasis>
.
<emphasis id="B97BEACAFFC50E77FF4D8A89FF4DFB7D" box="[145,242,1218,1240]" italics="true" pageId="9" pageNumber="1117">filimargo</emphasis>
</taxonomicName>
(clade A); b,
<taxonomicName id="4C0F4D5BFFC50E77FEAB8A89FE49FB7D" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[375,502,1218,1240]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC50E77FEAB8A89FE35FB72" box="[375,394,1218,1239]" italics="true" pageId="9" pageNumber="1117">H</emphasis>
.
<emphasis id="B97BEACAFFC50E77FE498A89FE49FB7D" box="[405,502,1218,1240]" italics="true" pageId="9" pageNumber="1117">filimargo</emphasis>
</taxonomicName>
(clade B); l,
<taxonomicName id="4C0F4D5BFFC50E77FDAB8A89FD4EFB7D" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[631,753,1218,1240]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC50E77FDAB8A89FD35FB72" box="[631,650,1218,1239]" italics="true" pageId="9" pageNumber="1117">H</emphasis>
.
<emphasis id="B97BEACAFFC50E77FD4A8A89FD4EFB7D" box="[662,753,1218,1240]" italics="true" pageId="9" pageNumber="1117">lunulata</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BB036D8FFC50E77FF7588CFFA9AF8F7" blockId="9.[145,762,1270,1874]" lastBlockId="9.[809,1426,1270,1874]" pageId="9" pageNumber="1117">
The
<taxonomicName id="4C0F4D5BFFC50E77FF0188CEFEDBF93C" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[221,356,1668,1690]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC50E77FF0188CEFF4DF93F" box="[221,242,1669,1690]" italics="true" pageId="9" pageNumber="1117">H</emphasis>
.
<emphasis id="B97BEACAFFC50E77FF2388CFFEDBF93C" box="[255,356,1668,1689]" italics="true" pageId="9" pageNumber="1117">lunulata</emphasis>
</taxonomicName>
clade includes mainly populations with depressed conical shells previously identified as
<taxonomicName id="4C0F4D5BFFC50E77FF4D8889FEBEF972" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[145,257,1730,1751]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC50E77FF4D8889FEBEF972" box="[145,257,1730,1751]" italics="true" pageId="9" pageNumber="1117">H. striata</emphasis>
</taxonomicName>
. However, individuals with disc-like shells, identified as
<taxonomicName id="4C0F4D5BFFC50E77FEF288AAFE7DF950" baseAuthorityName="Rossmassler" baseAuthorityYear="1838" box="[302,450,1760,1782]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="instabilis">
<emphasis id="B97BEACAFFC50E77FEF288AAFE7DF950" box="[302,450,1760,1782]" italics="true" pageId="9" pageNumber="1117">H. instabilis</emphasis>
</taxonomicName>
based on shell characters and originating from the Lviv region, the
<typeStatus id="54B4887AFFC50E77FD1E894BFD48F8B0" box="[706,759,1792,1813]" pageId="9" pageNumber="1117">type</typeStatus>
locality of this nominal species, also belonged to this clade. The same is true for the disc-like specimens from
<collectingCountry id="F3187648FFC50E77FCB18ABDFC61FAA9" box="[877,990,1270,1292]" name="Romania" pageId="9" pageNumber="1117">Romania</collectingCountry>
. Finally, specimens also identified as
<emphasis id="B97BEACAFFC50E77FC8C8B5EFCD2FA8F" box="[848,877,1301,1322]" italics="true" pageId="9" pageNumber="1117">H.</emphasis>
(
<taxonomicName id="4C0F4D5BFFC50E77FC5F8B5FFC46FA8C" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[899,1017,1300,1321]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC50E77FC5F8B5FFC46FA8C" box="[899,1017,1300,1321]" italics="true" pageId="9" pageNumber="1117">filimargo</emphasis>
</taxonomicName>
)
<taxonomicName id="4C0F4D5BFFC50E77FBD38B5EFBCEFA8F" baseAuthorityName="Krynicki" baseAuthorityYear="1836" box="[1039,1137,1301,1322]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="arenosa">
<emphasis id="B97BEACAFFC50E77FBD38B5EFBCEFA8F" box="[1039,1137,1301,1322]" italics="true" pageId="9" pageNumber="1117">arenosa</emphasis>
</taxonomicName>
from Yelanets Steppe Nature Reserve in the Mykolaiv region belong to this clade.
<bibRefCitation id="EF9E4B29FFC50E77FCA58B19FBC9FAC2" author="Balashov IA" box="[889,1142,1362,1384]" pageId="9" pageNumber="1117" refId="ref11096" refString="Balashov IA. 2016. Fauna Ukrainy. Tom 29. Mollyuski. Vypusk 5. Stebelchatoglazye (Stylommatophora). Kiev: Naukova Dumka." type="book" year="2016">Balashov (2016: 471</bibRefCitation>
, fig. 264) thought that the specimens at this locality represent sympatric populations of
<taxonomicName id="4C0F4D5BFFC50E77FC018BDBFBEEFA00" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[989,1105,1424,1445]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC50E77FC018BDBFBEEFA00" box="[989,1105,1424,1445]" italics="true" pageId="9" pageNumber="1117">H. striata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC50E77FB4C8BDBFA35FA00" baseAuthorityName="Krynicki" baseAuthorityYear="1836" box="[1168,1418,1423,1445]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="subSpecies" species="filimargo" subSpecies="arenosa">
<emphasis id="B97BEACAFFC50E77FB4C8BDBFA35FA00" box="[1168,1418,1423,1445]" italics="true" pageId="9" pageNumber="1117">H. filimargo arenosa</emphasis>
</taxonomicName>
. However, both, specimens identified as
<taxonomicName id="4C0F4D5BFFC50E77FB2C8BE5FAE0FA66" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[1264,1375,1454,1475]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC50E77FB2C8BE5FAE0FA66" box="[1264,1375,1454,1475]" italics="true" pageId="9" pageNumber="1117">H. striata</emphasis>
</taxonomicName>
and specimens identified as
<taxonomicName id="4C0F4D5BFFC50E77FB9B8B86FA83FA47" baseAuthorityName="Krynicki" baseAuthorityYear="1836" box="[1095,1340,1484,1506]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="subSpecies" species="filimargo" subSpecies="arenosa">
<emphasis id="B97BEACAFFC50E77FB9B8B86FA83FA47" box="[1095,1340,1484,1506]" italics="true" pageId="9" pageNumber="1117">H. filimargo arenosa</emphasis>
</taxonomicName>
belong to the same clade. Beside specimens from
<collectingCountry id="F3187648FFC50E77FAFD8BA0FA32F9A4" box="[1313,1421,1515,1537]" name="Romania" pageId="9" pageNumber="1117">Romania</collectingCountry>
,
<collectingCountry id="F3187648FFC50E77FCF58841FC31F985" box="[809,910,1546,1568]" name="Ukraine" pageId="9" pageNumber="1117">Ukraine</collectingCountry>
and adjacent
<collectingCountry id="F3187648FFC50E77FB9D8841FB29F985" box="[1089,1174,1546,1568]" name="Russia" pageId="9" pageNumber="1117">Russia</collectingCountry>
, an individual of an introduced population from north-eastern
<collectingCountry id="F3187648FFC50E77FAFF8862FA2FF99B" box="[1315,1424,1577,1598]" name="Germany" pageId="9" pageNumber="1117">Germany</collectingCountry>
(see
<bibRefCitation id="EF9E4B29FFC50E77FCBD880CFBFCF9F8" author="Zettler ML &amp; Jueg U &amp; Menzel-Harloff H &amp; Gollnitz U &amp; Petrick S &amp; Weber E &amp; Seemann R" box="[865,1091,1607,1629]" pageId="9" pageNumber="1117" refId="ref14280" refString="Zettler ML, Jueg U, Menzel-Harloff H, Gollnitz U, Petrick S, Weber E, Seemann R. 2006. Die Land- und Susswassermollusken Mecklenburg-Vorpommerns. Schwerin: Obotritendruck." type="book" year="2006">
Zettler
<emphasis id="B97BEACAFFC50E77FC618803FC45F9F9" box="[957,1018,1607,1629]" italics="true" pageId="9" pageNumber="1117">et al.</emphasis>
, 2006
</bibRefCitation>
) also belongs in this clade. The disc-like shells of this population agree with shells described as
<taxonomicName id="4C0F4D5BFFC50E77FBF688CFFB24F93C" authorityName="A. Ferussac" authorityYear="1821" box="[1066,1179,1668,1689]" class="Gastropoda" family="Geomitridae" genus="Helicella" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="1117" phylum="Mollusca" rank="genus">
<emphasis id="B97BEACAFFC50E77FBF688CFFB24F93C" box="[1066,1179,1668,1689]" italics="true" pageId="9" pageNumber="1117">Helicella</emphasis>
</taxonomicName>
?
<taxonomicName id="4C0F4D5BFFC50E77FB6888CFFAA9F93C" baseAuthorityName="Pinter" baseAuthorityYear="1969" box="[1204,1302,1668,1689]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="depulsa">
<emphasis id="B97BEACAFFC50E77FB6888CFFAA9F93C" box="[1204,1302,1668,1689]" italics="true" pageId="9" pageNumber="1117">depulsa</emphasis>
</taxonomicName>
from the surroundings of
<collectingRegion id="49CBF83AFFC50E77FC3788E8FB8DF91D" box="[1003,1074,1699,1720]" country="Bulgaria" name="Varna" pageId="9" pageNumber="1117">Varna</collectingRegion>
in
<collectingCountry id="F3187648FFC50E77FB8688E8FB7DF91C" box="[1114,1218,1699,1721]" name="Bulgaria" pageId="9" pageNumber="1117">Bulgaria</collectingCountry>
by
<bibRefCitation id="EF9E4B29FFC50E77FB3288E8FA35F91C" author="Pinter L" box="[1262,1418,1699,1721]" pageId="9" pageNumber="1117" pagination="91 - 96" refId="ref13155" refString="Pinter L. 1969. Neue Mollusken aus Bulgarien (Gastropoda: Helicidae). Acta Zoologica Academiae Scientiarum Hungaricae 15: 91 - 96." type="journal article" year="1969">Pintér (1969)</bibRefCitation>
, which is similar to
<taxonomicName id="4C0F4D5BFFC50E77FBCE8889FB18F972" baseAuthorityName="Rossmassler" baseAuthorityYear="1838" box="[1042,1191,1730,1751]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="instabilis">
<emphasis id="B97BEACAFFC50E77FBCE8889FB18F972" box="[1042,1191,1730,1751]" italics="true" pageId="9" pageNumber="1117">H. instabilis</emphasis>
</taxonomicName>
.
<taxonomicName id="4C0F4D5BFFC50E77FB6A8889FA9FF972" authorityName="A. Ferussac" authorityYear="1821" box="[1206,1312,1730,1751]" class="Gastropoda" family="Geomitridae" genus="Helicella" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="1117" phylum="Mollusca" rank="genus">
<emphasis id="B97BEACAFFC50E77FB6A8889FA9FF972" box="[1206,1312,1730,1751]" italics="true" pageId="9" pageNumber="1117">Helicella</emphasis>
</taxonomicName>
?
<taxonomicName id="4C0F4D5BFFC50E77FAE88889FA2EF972" baseAuthorityName="Pinter" baseAuthorityYear="1969" box="[1332,1425,1730,1751]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="9" pageNumber="1117" phylum="Ascomycota" rank="species" species="depulsa">
<emphasis id="B97BEACAFFC50E77FAE88889FA2EF972" box="[1332,1425,1730,1751]" italics="true" pageId="9" pageNumber="1117">depulsa</emphasis>
</taxonomicName>
has been classified as a subspecies or a synonym of
<taxonomicName id="4C0F4D5BFFC50E77FCF588B4FABAF8B1" authority="(Welter-Schultes, 2012)" baseAuthorityName="Welter-Schultes" baseAuthorityYear="2012" box="[809,1285,1791,1813]" class="Gastropoda" family="Geomitridae" genus="Xerolenta" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="1117" phylum="Mollusca" rank="species" species="obvia">
<emphasis id="B97BEACAFFC50E77FCF588B4FC5CF8B1" box="[809,995,1791,1812]" italics="true" pageId="9" pageNumber="1117">Xerolenta obvia</emphasis>
(
<bibRefCitation id="EF9E4B29FFC50E77FC2988B4FB45F8B1" author="Welter-Schultes F" box="[1013,1274,1791,1813]" pageId="9" pageNumber="1117" refId="ref14259" refString="Welter-Schultes F. 2012. European non-marine molluscs, a guide for species identification. Gottingen: Planet Poster Editions." type="book" year="2012">Welter-Schultes, 2012</bibRefCitation>
)
</taxonomicName>
. Specimens from the surroundings of
<collectingRegion id="49CBF83AFFC50E77FB8F8955FB25F896" box="[1107,1178,1822,1843]" country="Bulgaria" name="Varna" pageId="9" pageNumber="1117">Varna</collectingRegion>
have to be examined to clarify the identity of the nominal taxon.
</paragraph>
<caption id="DF706650FFC60E74FF788818FEA5F967" ID-DOI="http://doi.org/10.5281/zenodo.5636877" ID-Zenodo-Dep="5636877" httpUri="https://zenodo.org/record/5636877/files/figure.png" pageId="10" pageNumber="1118" startId="10.[164,244,1619,1641]" targetBox="[356,1248,198,1577]" targetPageId="10">
<paragraph id="8BB036D8FFC60E74FF788818FEA5F967" blockId="10.[163,1443,1619,1730]" pageId="10" pageNumber="1118">
<emphasis id="B97BEACAFFC60E74FF788818FEB0F9CD" bold="true" box="[164,271,1619,1641]" pageId="10" pageNumber="1118">Figure 6.</emphasis>
Distribution of clusters obtained from the admixture analysis of AFLP data of
<taxonomicName id="4C0F4D5BFFC60E74FBBA881FFB6DF9CF" authorityName="Fitzinger" authorityYear="1833" box="[1126,1234,1620,1642]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="10" pageNumber="1118" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC60E74FBBA881FFB6DF9CF" box="[1126,1234,1620,1642]" italics="true" pageId="10" pageNumber="1118">Helicopsis</emphasis>
</taxonomicName>
from Ukraine with STRUCTURE. A, solution for
<emphasis id="B97BEACAFFC60E74FE048839FE56F922" box="[472,489,1650,1671]" italics="true" pageId="10" pageNumber="1118">K</emphasis>
= 2. B, solution for
<emphasis id="B97BEACAFFC60E74FD6C8839FD7EF922" box="[688,705,1650,1671]" italics="true" pageId="10" pageNumber="1118">K</emphasis>
= 5. C, solution for
<emphasis id="B97BEACAFFC60E74FC558839FC25F922" box="[905,922,1650,1671]" italics="true" pageId="10" pageNumber="1118">K</emphasis>
= 7. Population numbers are given next to the bar plots (Supporting Information, Table S1). The bar plots show the proportional assignment of an individual to a cluster (see also Fig. 5).
</paragraph>
</caption>
<paragraph id="8BB036D8FFC60E75FF6788BEFD13FED1" blockId="10.[163,778,1781,1895]" lastBlockId="11.[145,762,197,1905]" lastPageId="11" lastPageNumber="1119" pageId="10" pageNumber="1118">
The clustering of the various forms that share the discussed mitochondrial haplotype clade in the network based on the nuclear AFLP markers (
<figureCitation id="13342A5DFFC60E74FF77891AFF53F8C3" box="[171,236,1873,1895]" captionStart="Figure 4" captionStartId="8.[163,244,1320,1342]" captionTargetBox="[200,1421,195,1281]" captionTargetId="graphics-203@8.[199,1421,195,1281]" captionTargetPageId="8" captionText="Figure 4. Neighbour-net network of Helicopsis from Ukraine based on Jaccard distances obtained from AFLP data. Coloration of clusters corresponds to the STRUCTURE solution for K = 2. Red corresponds to H. filimargo, yellow to H. lunulata. DNA voucher numbers for specimens are given at the tips of the network. Coloured dots at the tips of the network correspond to cluster assignments (on majority-rule basis) of specimens in the STRUCTURE solutions for K = 5 (inner) and K = 7 (outer). For locality data and the distribution of clusters, see also Supporting Information, Table S1 and Figure 6, respectively." figureDoi="http://doi.org/10.5281/zenodo.5636871" httpUri="https://zenodo.org/record/5636871/files/figure.png" pageId="10" pageNumber="1118">Fig. 4</figureCitation>
) and in a STRUCTURE analysis of these data (
<figureCitation id="13342A5DFFC60E74FC9888BEFC37F8AE" box="[836,904,1781,1803]" captionStart="Figure 5" captionStartId="9.[145,223,1129,1151]" captionTargetBox="[145,1426,199,1090]" captionTargetId="graphics-677@9.[259,1424,524,787]" captionTargetPageId="9" captionText="Figure 5. Assignment of specimens of Helicopsis from Ukraine to clusters resulting from the admixture analysis of AFLP data with STRUCTURE. A, solution for K = 2. B, solution for K = 5. C, solution for K = 7. Numbers below the plots refer to populations (see Fig. 6; Supporting Information, Table S1). Letters above the plots refer to mitochondrial clades: a, H. filimargo (clade A); b, H. filimargo (clade B); l, H. lunulata." figureDoi="http://doi.org/10.5281/zenodo.5636873" httpUri="https://zenodo.org/record/5636873/files/figure.png" pageId="10" pageNumber="1118">Fig. 5</figureCitation>
) demonstrated that they actually represent a single species. This species is distinct from
<taxonomicName id="4C0F4D5BFFC60E74FCE78978FC12F8ED" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[827,941,1843,1864]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="10" pageNumber="1118" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC60E74FCE78978FC12F8ED" box="[827,941,1843,1864]" italics="true" pageId="10" pageNumber="1118">H. striata</emphasis>
</taxonomicName>
, with which most populations of this clade were identified.
<taxonomicName id="4C0F4D5BFFC60E74FBDB891AFB5DF8C2" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[1031,1250,1873,1895]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="10" pageNumber="1118" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC60E74FBDB891AFB5DF8C2" box="[1031,1250,1873,1895]" italics="true" pageId="10" pageNumber="1118">Helicopsis striata</emphasis>
</taxonomicName>
is more closely related with the Eastern Alpine
<taxonomicName id="4C0F4D5BFFC70E75FDF48E8DFD7DFF7E" authorityName="E. Gittenberger" authorityYear="1969" box="[552,706,198,219]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="austriaca">
<emphasis id="B97BEACAFFC70E75FDF48E8DFD7DFF7E" box="[552,706,198,219]" italics="true" pageId="11" pageNumber="1119">H. austriaca</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC70E75FF4D8EAEFE91FF5C" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[145,302,228,250]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC70E75FF4D8EAEFE91FF5C" box="[145,302,228,250]" italics="true" pageId="11" pageNumber="1119">H. hungarica</emphasis>
</taxonomicName>
from the Pannonian Basin, than with the
<taxonomicName id="4C0F4D5BFFC70E75FF1C8F49FE85FEB2" authority="Fitzinger, 1833" box="[192,314,258,279]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="undetermined">
<emphasis id="B97BEACAFFC70E75FF1C8F49FE85FEB2" box="[192,314,258,279]" italics="true" pageId="11" pageNumber="1119">Helicopsis</emphasis>
</taxonomicName>
species from Eastern Europe (
<figureCitation id="13342A5DFFC70E75FD738F49FD4DFEBD" box="[687,754,258,280]" captionStart="Figure 2" captionStartId="5.[145,227,1817,1839]" captionTargetBox="[272,1296,410,1747]" captionTargetId="graphics-142@5.[272,1273,411,1748]" captionTargetPageId="5" captionText="Figure 2. Bayesian 50% majority-rule consensus tree of Helicopsis and related groups based on sequences of the mitochondrial 16S rDNA. Nodes supported by posterior probabilities ≥ 0.95 are marked by a dot. For information about sequenced specimens and detailed support values, see Supporting Information, Figure S1 and Table S1." figureDoi="http://doi.org/10.5281/zenodo.5636867" httpUri="https://zenodo.org/record/5636867/files/figure.png" pageId="11" pageNumber="1119">Fig. 2</figureCitation>
; Supporting Information,
<figureCitation id="13342A5DFFC70E75FE638F6AFDACFE93" box="[447,531,289,311]" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="11" pageNumber="1119">Fig. S1</figureCitation>
) and is apparently restricted to the Northern steppes and the northwestern Pannonian Basin in Central Europe.
</paragraph>
<paragraph id="8BB036D8FFC70E75FF758F36FD3AFA61" blockId="11.[145,762,197,1905]" pageId="11" pageNumber="1119">
There are several names for forms that belong to the discussed species. The most often used name is
<taxonomicName id="4C0F4D5BFFC70E75FF4D8FF0FDB4FE74" authority="(Rossmassler, 1838)" baseAuthorityName="Rossmassler" baseAuthorityYear="1838" box="[145,523,442,465]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="instabilis">
<emphasis id="B97BEACAFFC70E75FF4D8FF0FEA0FE6A" box="[145,287,442,464]" italics="true" pageId="11" pageNumber="1119">H. instabilis</emphasis>
(Rossmässler, 1838)
</taxonomicName>
, given for a disc-like form from
<collectingRegion id="49CBF83AFFC70E75FED08F92FE80FE4A" box="[268,319,473,495]" country="Ukraine" name="L'viv" pageId="11" pageNumber="1119">Lviv</collectingRegion>
in Western
<collectingCountry id="F3187648FFC70E75FE178F92FD93FE4A" box="[459,556,473,495]" name="Ukraine" pageId="11" pageNumber="1119">Ukraine</collectingCountry>
, but also used for disc-like forms from
<collectingCountry id="F3187648FFC70E75FEAA8FB3FE60FDAB" box="[374,479,504,526]" name="Romania" pageId="11" pageNumber="1119">Romania</collectingCountry>
. However, the oldest and, thus, valid name of the species is
<taxonomicName id="4C0F4D5BFFC70E75FDDC8C5CFF6DFDEE" authority="(Krynicki, 1833)" baseAuthorityName="Krynicki" baseAuthorityYear="1833" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FDDC8C5CFD3CFD8E" box="[512,643,534,556]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
(Krynicki, 1833)
</taxonomicName>
, given for a depressed conical form from Odessa [
<typeStatus id="54B4887AFFC70E75FF4B8C1FFF40FDCF" box="[151,255,596,618]" pageId="11" pageNumber="1119" type="lectotype">lectotype</typeStatus>
see
<bibRefCitation id="EF9E4B29FFC70E75FEF08C1FFE65FDCC" author="Hausdorf B" box="[300,474,596,618]" pageId="11" pageNumber="1119" pagination="57 - 71" refId="ref12143" refString="Hausdorf B. 1990. Zur Kenntnis einiger Arten der Gattung Helicopsis Fitzinger aus Griechenland und der Turkei (Gastropoda: Hygromiidae). Archiv fur Molluskenkunde 120: 57 - 71." type="journal article" year="1990">Hausdorf (1990</bibRefCitation>
: pl. 2 fig. 4);
<typeStatus id="54B4887AFFC70E75FD828C1FFD46FDCF" box="[606,761,596,618]" pageId="11" pageNumber="1119" type="paralectotype">paralectotype</typeStatus>
see
<bibRefCitation id="EF9E4B29FFC70E75FF608C38FE5EFD2D" author="Sysoev AV &amp; Schileyko AA" box="[188,481,626,648]" pageId="11" pageNumber="1119" refId="ref14149" refString="Sysoev AV, Schileyko AA. 2009. Land snails and slugs of Russia and adjacent countries. Sofia &amp; Moscow: Pensoft." type="book" year="2009">Sysoev &amp; Schileyko (2009</bibRefCitation>
: fig. 113A)]. Our records show that
<taxonomicName id="4C0F4D5BFFC70E75FED08CD9FE2BFD03" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[268,404,657,679]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FED08CD9FE2BFD03" box="[268,404,657,679]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
occurs in Transylvania, in the Podolian Upland and the Black Sea Lowland and in the Donetsk Upland and the adjacent Central Russian Upland (
<figureCitation id="13342A5DFFC70E75FF208CA6FEFFFCA7" box="[252,320,749,771]" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="11" pageNumber="1119">Fig. 1</figureCitation>
; Supporting Information,
<tableCitation id="C68D0363FFC70E75FD5C8CA6FD55FCA7" box="[640,746,749,771]" captionStart="Table 1" captionStartId="7.[144,208,196,217]" captionTargetPageId="7" captionText="Table 1. Within group (WG) and between group mean p-distances of Helicopsis taxa based on partial 16S rDNA sequences" httpUri="http://table.plazi.org/id/DF706650FFCB0E79FF4C8E8FFF45FF52" pageId="11" pageNumber="1119" tableUuid="DF706650FFCB0E79FF4C8E8FFF45FF52">Table S1</tableCitation>
), from where a sequence belonging to the
<taxonomicName id="4C0F4D5BFFC70E75FDAC8D47FD46FC84" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[624,761,780,801]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FDAC8D47FD46FC84" box="[624,761,780,801]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
clade has already been reported by
<bibRefCitation id="EF9E4B29FFC70E75FDEE8D61FF5EFCFA" author="Sychev AA &amp; Snegin EA" pageId="11" pageNumber="1119" pagination="175 - 189" refId="ref14071" refString="Sychev AA, Snegin EA. 2016. K probleme sistematiki roda Helicopsis (Gastropoda: Pulmonata: Hygromiidae) na territorii Vostochnoy Evropy. Ruthenica 26: 175 - 189." type="journal article" year="2016">Sychev &amp; Snegin (2016)</bibRefCitation>
from Zasosna. Unfortunately, no AFLP data were available from these eastern specimens. Thus, we cannot completely exclude the possibility that these specimens belong to
<taxonomicName id="4C0F4D5BFFC70E75FDCC8DEDFD1CFC1F" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[528,675,933,955]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FDCC8DEDFD1CFC1F" box="[528,675,933,955]" italics="true" pageId="11" pageNumber="1119">H. filimargo</emphasis>
</taxonomicName>
with a mitochondrial introgression, although there is no evidence for this. There are records of
<taxonomicName id="4C0F4D5BFFC70E75FDB88DA8FD46FC52" baseAuthorityName="Rossmassler" baseAuthorityYear="1838" box="[612,761,994,1016]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="instabilis">
<emphasis id="B97BEACAFFC70E75FDB88DA8FD46FC52" box="[612,761,994,1016]" italics="true" pageId="11" pageNumber="1119">H. instabilis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC70E75FF188A49FE8AFBB2" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[196,309,1026,1047]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC70E75FF188A49FE8AFBB2" box="[196,309,1026,1047]" italics="true" pageId="11" pageNumber="1119">H. striata</emphasis>
</taxonomicName>
from
<collectingCountry id="F3187648FFC70E75FEA78A4AFE61FBB2" box="[379,478,1025,1047]" name="Moldova" pageId="11" pageNumber="1119">Moldova</collectingCountry>
(
<bibRefCitation id="EF9E4B29FFC70E75FE338A4AFD4FFBB2" author="Balashov IA &amp; Son MO &amp; Coada V &amp; Welter-Schultes F" box="[495,752,1025,1047]" pageId="11" pageNumber="1119" pagination="175 - 181" refId="ref11189" refString="Balashov IA, Son MO, Coada V, Welter-Schultes F. 2013 b. An updated annotated checklist of the molluscs of the Republic of Moldova. Folia Malacologica 21: 175 - 181." type="journal article" year="2013">
Balashov
<emphasis id="B97BEACAFFC70E75FDBE8A49FD23FBB3" box="[610,668,1025,1047]" italics="true" pageId="11" pageNumber="1119">et al.</emphasis>
, 2013b
</bibRefCitation>
) and
<collectingCountry id="F3187648FFC70E75FF1E8A6BFE99FB93" box="[194,294,1056,1078]" name="Bulgaria" pageId="11" pageNumber="1119">Bulgaria</collectingCountry>
(
<bibRefCitation id="EF9E4B29FFC70E75FEE88A6BFDCEFB93" author="Damjanov SG &amp; Likharev IM" box="[308,625,1056,1078]" pageId="11" pageNumber="1119" refId="ref11536" refString="Damjanov SG, Likharev IM. 1975. Fauna na Bulgariya. 4. Sukhozemni okhlyuvi (Gastropoda terrestria). Sofia: Bulgarska Akademiya na Naukite." type="book" year="1975">Damjanov &amp; Likharev, 1975</bibRefCitation>
), indicating that
<taxonomicName id="4C0F4D5BFFC70E75FF158A74FEF0FBF6" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[201,335,1086,1108]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FF158A74FEF0FBF6" box="[201,335,1086,1108]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
is probably also distributed in these countries. However, it has to be checked whether all populations identified as
<taxonomicName id="4C0F4D5BFFC70E75FDDA8A37FD1BFB34" baseAuthorityName="Rossmassler" baseAuthorityYear="1838" box="[518,676,1148,1169]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="instabilis">
<emphasis id="B97BEACAFFC70E75FDDA8A37FD1BFB34" box="[518,676,1148,1169]" italics="true" pageId="11" pageNumber="1119">H. instabilis</emphasis>
</taxonomicName>
so far actually belong to
<taxonomicName id="4C0F4D5BFFC70E75FEA88AD0FDBEFB0A" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[372,513,1178,1200]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FEA88AD0FDBEFB0A" box="[372,513,1178,1200]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
. Beside populations with disc-like shells corresponding to
<taxonomicName id="4C0F4D5BFFC70E75FD828AF1FD4DFB6B" baseAuthorityName="Rossmassler" baseAuthorityYear="1838" box="[606,754,1209,1231]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="instabilis">
<emphasis id="B97BEACAFFC70E75FD828AF1FD4DFB6B" box="[606,754,1209,1231]" italics="true" pageId="11" pageNumber="1119">H. instabilis</emphasis>
</taxonomicName>
, there are also populations with depressed conical shells in Transylvania that are often classified as an endemic species,
<taxonomicName id="4C0F4D5BFFC70E75FE848B5DFE50FA8F" baseAuthorityName="Bielz" baseAuthorityYear="1851" box="[344,495,1301,1323]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="cereoflava">
<emphasis id="B97BEACAFFC70E75FE848B5DFE50FA8F" box="[344,495,1301,1323]" italics="true" pageId="11" pageNumber="1119">H. cereoflava</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF9E4B29FFC70E75FE238B5EFD28FA8F" author="Grossu AV" box="[511,663,1301,1322]" pageId="11" pageNumber="1119" refId="ref12061" refString="Grossu AV. 1983. Gastropoda Romaniae 4. Suprafam.: Arionacea, Zonitacea, Ariophantacea si Helicacea. Bucharest: Editura Litera." type="book" year="1983">Grossu, 1983</bibRefCitation>
; Welter-Schultes, 2012). Considering the large intraspecific variability observed in
<taxonomicName id="4C0F4D5BFFC70E75FE418B18FD9CFAC2" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[413,547,1362,1384]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FE418B18FD9CFAC2" box="[413,547,1362,1384]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
, it is possible that all Transylvanian populations of
<taxonomicName id="4C0F4D5BFFC70E75FDCE8B3AFD38FA23" authorityName="Fitzinger" authorityYear="1833" box="[530,647,1393,1414]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC70E75FDCE8B3AFD38FA23" box="[530,647,1393,1414]" italics="true" pageId="11" pageNumber="1119">Helicopsis</emphasis>
</taxonomicName>
belong to this species. However, typical
<taxonomicName id="4C0F4D5BFFC70E75FE308BDBFD3DFA00" baseAuthorityName="Bielz" baseAuthorityYear="1851" box="[492,642,1424,1445]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="cereoflava">
<emphasis id="B97BEACAFFC70E75FE308BDBFD3DFA00" box="[492,642,1424,1445]" italics="true" pageId="11" pageNumber="1119">H. cereoflava</emphasis>
</taxonomicName>
should be studied genetically to prove this hypothesis.
</paragraph>
<paragraph id="8BB036D8FFC70E75FF758B86FDBCF891" blockId="11.[145,762,197,1905]" pageId="11" pageNumber="1119">
The
<taxonomicName id="4C0F4D5BFFC70E75FF3D8B85FEC7FA47" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[225,376,1485,1507]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FF3D8B85FEC7FA47" box="[225,376,1485,1507]" italics="true" pageId="11" pageNumber="1119">H. filimargo</emphasis>
</taxonomicName>
clade is found in
<collectingCountry id="F3187648FFC70E75FD858B86FD02FA46" box="[601,701,1485,1507]" name="Ukraine" pageId="11" pageNumber="1119">Ukraine</collectingCountry>
east of the Dnieper River, from
<collectingRegion id="49CBF83AFFC70E75FE0D8BA7FD97F9A7" box="[465,552,1516,1538]" country="Ukraine" name="Crimea" pageId="11" pageNumber="1119">Crimea</collectingRegion>
and the adjacent eastern part of the Black Sea Lowland to the
<collectingRegion id="49CBF83AFFC70E75FD468841FD46F985" box="[666,761,1546,1568]" country="Ukraine" name="Donets'k" pageId="11" pageNumber="1119">Donetsk</collectingRegion>
Upland and the Central Russian Upland (
<figureCitation id="13342A5DFFC70E75FDAE8862FD0DF99B" box="[626,690,1577,1599]" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="11" pageNumber="1119">Fig. 1</figureCitation>
). This group comprises even more diverse morphological forms than
<taxonomicName id="4C0F4D5BFFC70E75FEF7882CFE7DF9DE" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[299,450,1638,1660]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FEF7882CFE7DF9DE" box="[299,450,1638,1660]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
. It includes, beside the nominal species listed from
<collectingRegion id="49CBF83AFFC70E75FE1388CEFD9BF93E" box="[463,548,1669,1691]" country="Ukraine" name="Crimea" pageId="11" pageNumber="1119">Crimea</collectingRegion>
and the Black Sea Lowland,
<taxonomicName id="4C0F4D5BFFC70E75FEDB88EFFE26F91C" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[263,409,1700,1721]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FEDB88EFFE26F91C" box="[263,409,1700,1721]" italics="true" pageId="11" pageNumber="1119">H. filimargo</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D5BFFC70E75FE7588EFFD95F91C" baseAuthorityName="Krynicki" baseAuthorityYear="1836" box="[425,554,1700,1721]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="arenosa">
<emphasis id="B97BEACAFFC70E75FE7588EFFD95F91C" box="[425,554,1700,1721]" italics="true" pageId="11" pageNumber="1119">H. arenosa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC70E75FDB488EFFD46F91C" baseAuthorityName="Clessin" baseAuthorityYear="1883" box="[616,761,1700,1721]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="retowskii">
<emphasis id="B97BEACAFFC70E75FDB488EFFD46F91C" box="[616,761,1700,1721]" italics="true" pageId="11" pageNumber="1119">H. retowskii</emphasis>
</taxonomicName>
and the recently described supposed endemics from the
<collectingRegion id="49CBF83AFFC70E75FF6388AAFE9FF952" box="[191,288,1761,1783]" country="Ukraine" name="Donets'k" pageId="11" pageNumber="1119">Donetsk</collectingRegion>
Upland,
<taxonomicName id="4C0F4D5BFFC70E75FE4C88A9FDA6F953" authorityName="Gural-Sverlova" authorityYear="2010" box="[400,537,1761,1783]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="luganica">
<emphasis id="B97BEACAFFC70E75FE4C88A9FDA6F953" box="[400,537,1761,1783]" italics="true" pageId="11" pageNumber="1119">H. luganica</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D5BFFC70E75FDF488A9FD7BF953" authorityName="Gural-Sverlova" authorityYear="2010" box="[552,708,1761,1783]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="martynovi">
<emphasis id="B97BEACAFFC70E75FDF488A9FD7BF953" box="[552,708,1761,1783]" italics="true" pageId="11" pageNumber="1119">H. martynovi</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC70E75FF4D894BFEF5F8B1" authorityName="Gural-Sverlova" authorityYear="2010" box="[145,330,1791,1813]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="subfilimargo">
<emphasis id="B97BEACAFFC70E75FF4D894BFEF5F8B1" box="[145,330,1791,1813]" italics="true" pageId="11" pageNumber="1119">H. subfilimargo</emphasis>
</taxonomicName>
, plus some populations classified as
<taxonomicName id="4C0F4D5BFFC70E75FF4D8954FF41F891" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[145,254,1823,1844]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC70E75FF4D8954FF41F891" box="[145,254,1823,1844]" italics="true" pageId="11" pageNumber="1119">H. striata</emphasis>
</taxonomicName>
from eastern
<collectingCountry id="F3187648FFC70E75FE418955FE40F891" box="[413,511,1822,1844]" name="Ukraine" pageId="11" pageNumber="1119">Ukraine</collectingCountry>
.
</paragraph>
<paragraph id="8BB036D8FFC70E75FF758976FB7BFE37" blockId="11.[145,762,197,1905]" lastBlockId="11.[809,1426,197,1905]" pageId="11" pageNumber="1119">
<taxonomicName id="4C0F4D5BFFC70E75FF758976FE2EF8F7" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[169,401,1853,1874]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FF758976FE2EF8F7" box="[169,401,1853,1874]" italics="true" pageId="11" pageNumber="1119">Helicopsis filimargo</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC70E75FE108975FDF7F8F6" baseAuthorityName="Krynicki" baseAuthorityYear="1836" box="[460,584,1854,1875]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="arenosa">
<emphasis id="B97BEACAFFC70E75FE108975FDF7F8F6" box="[460,584,1854,1875]" italics="true" pageId="11" pageNumber="1119">H. arenosa</emphasis>
</taxonomicName>
have also been recorded from Dobrogea in
<collectingCountry id="F3187648FFC70E75FE058910FDFCF8D4" box="[473,579,1883,1905]" name="Romania" pageId="11" pageNumber="1119">Romania</collectingCountry>
(
<bibRefCitation id="EF9E4B29FFC70E75FD898917FD4FF8D4" author="Grossu AV" box="[597,752,1884,1905]" pageId="11" pageNumber="1119" refId="ref12061" refString="Grossu AV. 1983. Gastropoda Romaniae 4. Suprafam.: Arionacea, Zonitacea, Ariophantacea si Helicacea. Bucharest: Editura Litera." type="book" year="1983">Grossu, 1983</bibRefCitation>
) and the latter species was also found in
<collectingCountry id="F3187648FFC70E75FAFA8E8EFA2FFF7E" box="[1318,1424,197,219]" name="Bulgaria" pageId="11" pageNumber="1119">Bulgaria</collectingCountry>
(
<bibRefCitation id="EF9E4B29FFC70E75FCED8EA8FBD0FF5C" author="Damjanov SG &amp; Likharev IM" box="[817,1135,227,249]" pageId="11" pageNumber="1119" refId="ref11536" refString="Damjanov SG, Likharev IM. 1975. Fauna na Bulgariya. 4. Sukhozemni okhlyuvi (Gastropoda terrestria). Sofia: Bulgarska Akademiya na Naukite." type="book" year="1975">Damjanov &amp; Likharev, 1975</bibRefCitation>
). However, given the lack of diagnostic morphological characters distinguishing
<taxonomicName id="4C0F4D5BFFC70E75FCF58F69FC01FE93" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[809,958,289,311]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FCF58F69FC01FE93" box="[809,958,289,311]" italics="true" pageId="11" pageNumber="1119">H. filimargo</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC70E75FC228F69FB33FE93" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1022,1164,289,311]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FC228F69FB33FE93" box="[1022,1164,289,311]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
, it has to be checked genetically whether the populations from
<collectingCountry id="F3187648FFC70E75FB278F74FADEFEF0" box="[1275,1377,319,341]" name="Romania" pageId="11" pageNumber="1119">Romania</collectingCountry>
and
<collectingCountry id="F3187648FFC70E75FCF58F15FC2EFED1" box="[809,913,350,372]" name="Bulgaria" pageId="11" pageNumber="1119">Bulgaria</collectingCountry>
actually belong to
<taxonomicName id="4C0F4D5BFFC70E75FBA98F14FAB9FED6" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1141,1286,350,372]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FBA98F14FAB9FED6" box="[1141,1286,350,372]" italics="true" pageId="11" pageNumber="1119">H. filimargo</emphasis>
</taxonomicName>
or whether they represent forms of
<taxonomicName id="4C0F4D5BFFC70E75FBE58F36FB02FE37" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1081,1213,381,402]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FBE58F36FB02FE37" box="[1081,1213,381,402]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BB036D8FFC70E75FC9D8FD0FBDAFCD8" blockId="11.[809,1426,197,1905]" pageId="11" pageNumber="1119">
The
<taxonomicName id="4C0F4D5BFFC70E75FCA08FD7FBA6FE15" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[892,1049,411,433]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FCA08FD7FBA6FE15" box="[892,1049,411,433]" italics="true" pageId="11" pageNumber="1119">H. filimargo</emphasis>
</taxonomicName>
clade is subdivided into two geographically restricted clades in the mitochondrial tree (
<figureCitation id="13342A5DFFC70E75FCB78F92FC0FFE4B" box="[875,944,473,495]" captionStart="Figure 2" captionStartId="5.[145,227,1817,1839]" captionTargetBox="[272,1296,410,1747]" captionTargetId="graphics-142@5.[272,1273,411,1748]" captionTargetPageId="5" captionText="Figure 2. Bayesian 50% majority-rule consensus tree of Helicopsis and related groups based on sequences of the mitochondrial 16S rDNA. Nodes supported by posterior probabilities ≥ 0.95 are marked by a dot. For information about sequenced specimens and detailed support values, see Supporting Information, Figure S1 and Table S1." figureDoi="http://doi.org/10.5281/zenodo.5636867" httpUri="https://zenodo.org/record/5636867/files/figure.png" pageId="11" pageNumber="1119">Fig. 2</figureCitation>
; Supporting Information,
<figureCitation id="13342A5DFFC70E75FB2E8F92FAF8FE4B" box="[1266,1351,473,495]" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="11" pageNumber="1119">Fig. S1</figureCitation>
). One occupies most of the range, whereas the other is restricted to southernmost
<collectingRegion id="49CBF83AFFC70E75FBB98C5DFB05FD89" box="[1125,1210,534,556]" country="Ukraine" name="Crimea" pageId="11" pageNumber="1119">Crimea</collectingRegion>
(
<figureCitation id="13342A5DFFC70E75FB168C5DFAB4FD8E" box="[1226,1291,534,556]" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="11" pageNumber="1119">Fig. 1</figureCitation>
). There are a few populations along the mountains where both haplotypes occurred together. The specimens from southernmost
<collectingRegion id="49CBF83AFFC70E75FC138C39FB9BFD2D" box="[975,1060,626,648]" country="Ukraine" name="Crimea" pageId="11" pageNumber="1119">Crimea</collectingRegion>
are concentrated in one cluster of the network based on nuclear AFLP markers (
<figureCitation id="13342A5DFFC70E75FA998CDAFA3CFD03" box="[1349,1411,657,679]" captionStart="Figure 4" captionStartId="8.[163,244,1320,1342]" captionTargetBox="[200,1421,195,1281]" captionTargetId="graphics-203@8.[199,1421,195,1281]" captionTargetPageId="8" captionText="Figure 4. Neighbour-net network of Helicopsis from Ukraine based on Jaccard distances obtained from AFLP data. Coloration of clusters corresponds to the STRUCTURE solution for K = 2. Red corresponds to H. filimargo, yellow to H. lunulata. DNA voucher numbers for specimens are given at the tips of the network. Coloured dots at the tips of the network correspond to cluster assignments (on majority-rule basis) of specimens in the STRUCTURE solutions for K = 5 (inner) and K = 7 (outer). For locality data and the distribution of clusters, see also Supporting Information, Table S1 and Figure 6, respectively." figureDoi="http://doi.org/10.5281/zenodo.5636871" httpUri="https://zenodo.org/record/5636871/files/figure.png" pageId="11" pageNumber="1119">Fig. 4</figureCitation>
). However, there are also specimens from the rest of the range with the other mitochondrial subclade among them. This indicates that differentiation between the two subgroups is also evolving in the nuclear genome, but is not yet advanced. This is also corroborated by the STRUCTURE analyses (
<figureCitation id="13342A5DFFC70E75FBA78D02FB01FCFA" box="[1147,1214,841,863]" captionStart="Figure 5" captionStartId="9.[145,223,1129,1151]" captionTargetBox="[145,1426,199,1090]" captionTargetId="graphics-677@9.[259,1424,524,787]" captionTargetPageId="9" captionText="Figure 5. Assignment of specimens of Helicopsis from Ukraine to clusters resulting from the admixture analysis of AFLP data with STRUCTURE. A, solution for K = 2. B, solution for K = 5. C, solution for K = 7. Numbers below the plots refer to populations (see Fig. 6; Supporting Information, Table S1). Letters above the plots refer to mitochondrial clades: a, H. filimargo (clade A); b, H. filimargo (clade B); l, H. lunulata." figureDoi="http://doi.org/10.5281/zenodo.5636873" httpUri="https://zenodo.org/record/5636873/files/figure.png" pageId="11" pageNumber="1119">Fig. 5</figureCitation>
). Thus, we do not classify them as subspecies.
</paragraph>
<paragraph id="8BB036D8FFC70E75FC9D8DCDFBAAFAEC" blockId="11.[809,1426,197,1905]" pageId="11" pageNumber="1119">
The STRUCTURE analyses shows little admixture between
<taxonomicName id="4C0F4D5BFFC70E75FC428DEEFB81FC1F" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[926,1086,933,954]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FC428DEEFB81FC1F" box="[926,1086,933,954]" italics="true" pageId="11" pageNumber="1119">H. filimargo</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC70E75FB5B8DEEFAA1FC1F" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1159,1310,933,954]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FB5B8DEEFAA1FC1F" box="[1159,1310,933,954]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
(
<figureCitation id="13342A5DFFC70E75FAE98DEEFA3FFC1E" box="[1333,1408,933,955]" captionStart="Figure 5" captionStartId="9.[145,223,1129,1151]" captionTargetBox="[145,1426,199,1090]" captionTargetId="graphics-677@9.[259,1424,524,787]" captionTargetPageId="9" captionText="Figure 5. Assignment of specimens of Helicopsis from Ukraine to clusters resulting from the admixture analysis of AFLP data with STRUCTURE. A, solution for K = 2. B, solution for K = 5. C, solution for K = 7. Numbers below the plots refer to populations (see Fig. 6; Supporting Information, Table S1). Letters above the plots refer to mitochondrial clades: a, H. filimargo (clade A); b, H. filimargo (clade B); l, H. lunulata." figureDoi="http://doi.org/10.5281/zenodo.5636873" httpUri="https://zenodo.org/record/5636873/files/figure.png" pageId="11" pageNumber="1119">Fig. 5</figureCitation>
), supporting the species status of the two taxa. The only exception was the specimen from the introduced population in north-eastern
<collectingCountry id="F3187648FFC70E75FBA48A4AFB5AFBB3" box="[1144,1253,1025,1046]" name="Germany" pageId="11" pageNumber="1119">Germany</collectingCountry>
. However, this specimen is older and not preserved in 100% alcohol, so that the slightly different AFLP pattern might be an artefact resulting from DNA degradation. The specific distinctness of
<taxonomicName id="4C0F4D5BFFC70E75FBF28A37FB06FB35" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1070,1209,1147,1169]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC70E75FBF28A37FB06FB35" box="[1070,1209,1147,1169]" italics="true" pageId="11" pageNumber="1119">H. filimargo</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC70E75FB2E8A37FAC8FB35" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1266,1399,1147,1169]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC70E75FB2E8A37FAC8FB35" box="[1266,1399,1147,1169]" italics="true" pageId="11" pageNumber="1119">H. lunulata</emphasis>
</taxonomicName>
is also corroborated by the isolation by distance analysis that shows that the distances between individuals of the two taxa are larger than expected based on their geographical distances and the relationship of genetic distances with geographical distances found within the two taxa (
<figureCitation id="13342A5DFFC70E75FC1A8B78FBBAFAEC" box="[966,1029,1331,1353]" captionStart="Figure 7" captionStartId="12.[163,248,649,671]" captionTargetBox="[175,770,195,569]" captionTargetId="graphics-743@12.[234,770,202,538]" captionTargetPageId="12" captionText="Figure 7. Relationships between Jaccard distances between individuals of Helicopsis filimargo and H. lunulata based on AFLP data and logarithmized geographical distances. Black circles and red triangles: distances between individuals belonging to H. filimargo and H. lunulata, respectively; green crosses: distances between individuals belonging to different species; black and red broken lines: regression lines fitted within species; green broken line: regression line fitted on the within-group distances only (i.e. the black circles and red triangles taken together); green solid line: regression line fitted on all distances together; blue lines: centres of the betweengroups geographical distances." figureDoi="http://doi.org/10.5281/zenodo.5636879" httpUri="https://zenodo.org/record/5636879/files/figure.png" pageId="11" pageNumber="1119">Fig. 7</figureCitation>
).
</paragraph>
<paragraph id="8BB036D8FFC70E75FC9D8B19FC30F952" blockId="11.[809,1426,197,1905]" pageId="11" pageNumber="1119">
An individual from Teleshovka in the Belgorod region in the Central Russian Upland represented another clade in the mitochondrial tree (
<figureCitation id="13342A5DFFC70E75FA9C8BC4FA35FA01" box="[1344,1418,1423,1445]" captionStart="Figure 2" captionStartId="5.[145,227,1817,1839]" captionTargetBox="[272,1296,410,1747]" captionTargetId="graphics-142@5.[272,1273,411,1748]" captionTargetPageId="5" captionText="Figure 2. Bayesian 50% majority-rule consensus tree of Helicopsis and related groups based on sequences of the mitochondrial 16S rDNA. Nodes supported by posterior probabilities ≥ 0.95 are marked by a dot. For information about sequenced specimens and detailed support values, see Supporting Information, Figure S1 and Table S1." figureDoi="http://doi.org/10.5281/zenodo.5636867" httpUri="https://zenodo.org/record/5636867/files/figure.png" pageId="11" pageNumber="1119">Fig. 2</figureCitation>
; Supporting Information,
<figureCitation id="13342A5DFFC70E75FBB38BE5FB75FA66" box="[1135,1226,1454,1476]" captionStart="Figure 1" captionStartId="2.[163,241,1817,1839]" captionTargetBox="[164,1444,1087,1776]" captionTargetId="figure-476@2.[163,1444,1085,1777]" captionTargetPageId="2" captionText="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." figureDoi="http://doi.org/10.5281/zenodo.5636865" httpUri="https://zenodo.org/record/5636865/files/figure.png" pageId="11" pageNumber="1119">Fig. S1</figureCitation>
), which can be identified as
<taxonomicName id="4C0F4D5BFFC70E75FC1E8B86FBE4FA47" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[962,1115,1485,1506]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC70E75FC1E8B86FBE4FA47" box="[962,1115,1485,1506]" italics="true" pageId="11" pageNumber="1119">H. hungarica</emphasis>
</taxonomicName>
based on sequences of the species from
<collectingCountry id="F3187648FFC70E75FC618BA7FB9CF9A4" box="[957,1059,1516,1537]" name="Hungary" pageId="11" pageNumber="1119">Hungary</collectingCountry>
and
<collectingCountry id="F3187648FFC70E75FB878BA0FB0EF9A4" box="[1115,1201,1515,1537]" name="Austria" pageId="11" pageNumber="1119">Austria</collectingCountry>
(
<bibRefCitation id="EF9E4B29FFC70E75FB1D8BA0FA3CF9A5" author="Duda M &amp; Haring E &amp; Bieringer G &amp; Eschner A &amp; Mrkvicka A &amp; Mason K" box="[1217,1411,1515,1537]" pageId="11" pageNumber="1119" pagination="432 - 450" refId="ref11688" refString="Duda M, Haring E, Bieringer G, Eschner A, Mrkvicka A, Mason K. 2018. Taxonomic reassessment of Helicopsis austriaca Gittenberger, 1969 and its relationships to H. striata (OF Muller, 1774) and H. hungarica (Soos &amp; H. Wagner, 1935) (Eupulmonata: Helicoidea). Journal of Molluscan Studies 84: 432 - 450." type="journal article" year="2018">
Duda
<emphasis id="B97BEACAFFC70E75FADA8BA7FA80F9A5" box="[1286,1343,1515,1537]" italics="true" pageId="11" pageNumber="1119">et al.</emphasis>
, 2018
</bibRefCitation>
). Actually, a sequence of the same haplotype group has already been reported as
<taxonomicName id="4C0F4D5BFFC70E75FB878862FB70F99B" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[1115,1231,1577,1598]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC70E75FB878862FB70F99B" box="[1115,1231,1577,1598]" italics="true" pageId="11" pageNumber="1119">H. striata</emphasis>
</taxonomicName>
from Gubkin in the Belgorod Region by
<bibRefCitation id="EF9E4B29FFC70E75FB9F880CFAE2F9F8" author="Sychev AA &amp; Snegin EA" box="[1091,1373,1607,1629]" pageId="11" pageNumber="1119" pagination="175 - 189" refId="ref14071" refString="Sychev AA, Snegin EA. 2016. K probleme sistematiki roda Helicopsis (Gastropoda: Pulmonata: Hygromiidae) na territorii Vostochnoy Evropy. Ruthenica 26: 175 - 189." type="journal article" year="2016">Sychev &amp; Snegin (2016)</bibRefCitation>
and identified as
<taxonomicName id="4C0F4D5BFFC70E75FC1D882CFBE4F9DE" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[961,1115,1638,1660]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC70E75FC1D882CFBE4F9DE" box="[961,1115,1638,1660]" italics="true" pageId="11" pageNumber="1119">H. hungarica</emphasis>
</taxonomicName>
by
<bibRefCitation id="EF9E4B29FFC70E75FB5A882DFAE8F9D9" author="Duda M &amp; Haring E &amp; Bieringer G &amp; Eschner A &amp; Mrkvicka A &amp; Mason K" box="[1158,1367,1638,1660]" pageId="11" pageNumber="1119" pagination="432 - 450" refId="ref11688" refString="Duda M, Haring E, Bieringer G, Eschner A, Mrkvicka A, Mason K. 2018. Taxonomic reassessment of Helicopsis austriaca Gittenberger, 1969 and its relationships to H. striata (OF Muller, 1774) and H. hungarica (Soos &amp; H. Wagner, 1935) (Eupulmonata: Helicoidea). Journal of Molluscan Studies 84: 432 - 450." type="journal article" year="2018">
Duda
<emphasis id="B97BEACAFFC70E75FB11882CFAB8F9DE" box="[1229,1287,1638,1660]" italics="true" pageId="11" pageNumber="1119">et al.</emphasis>
(2018)
</bibRefCitation>
. The specific identity of the specimens from the Central Russian Upland with the Pannonian
<taxonomicName id="4C0F4D5BFFC70E75FB2D88EFFA2EF91D" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[1265,1425,1699,1721]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="11" pageNumber="1119" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC70E75FB2D88EFFA2EF91D" box="[1265,1425,1699,1721]" italics="true" pageId="11" pageNumber="1119">H. hungarica</emphasis>
</taxonomicName>
should be confirmed with non-mitochondrial genetic markers.
</paragraph>
<paragraph id="8BB036D8FFC70E72FC9D88B4FDACF9ED" blockId="11.[809,1426,197,1905]" lastBlockId="12.[163,780,1065,1608]" lastPageId="12" lastPageNumber="1120" pageId="11" pageNumber="1119">
Our results, based on mitochondrial DNA sequences (
<figureCitation id="13342A5DFFC70E75FCEE8955FC24F896" box="[818,923,1822,1844]" captionStart-0="Figure 1" captionStart-1="Figure 2" captionStart-2="Figure 3" captionStartId-0="2.[163,241,1817,1839]" captionStartId-1="5.[145,227,1817,1839]" captionStartId-2="6.[827,907,1649,1671]" captionTargetBox-0="[164,1444,1087,1776]" captionTargetBox-1="[272,1296,410,1747]" captionTargetBox-2="[836,1433,1245,1607]" captionTargetId-0="figure-476@2.[163,1444,1085,1777]" captionTargetId-1="graphics-142@5.[272,1273,411,1748]" captionTargetId-2="graphics-830@6.[835,1247,1251,1596]" captionTargetPageId-0="2" captionTargetPageId-1="5" captionTargetPageId-2="6" captionText-0="Figure 1. Sampled Helicopsis populations in Ukraine, the adjacent Central Russian Upland and Romania and distribution of mitochondrial haplotype clades: H. lunulata (yellow dots), H. hungarica (magenta triangles) and H. filimargo (clade A, red diamonds, clade B, green diamonds and mixed population of clades A and B, blue diamonds)." captionText-1="Figure 2. Bayesian 50% majority-rule consensus tree of Helicopsis and related groups based on sequences of the mitochondrial 16S rDNA. Nodes supported by posterior probabilities ≥ 0.95 are marked by a dot. For information about sequenced specimens and detailed support values, see Supporting Information, Figure S1 and Table S1." captionText-2="Figure 3. Bayesian 50% majority-rule consensus tree of European Helicopsis based on concatenated sequences of the mitochondrial COI, 12S rDNA and 16S rDNA. Support values at the nodes correspond to Bayesian posterior probabilities (left), maximum likelihood (middle) and maximum parsimony (right) bootstrap values. Extraction voucher numbers are given at the tips of the tree. For information about sequenced specimens, see Supporting Information, Table S1." figureDoi-0="http://doi.org/10.5281/zenodo.5636865" figureDoi-1="http://doi.org/10.5281/zenodo.5636867" figureDoi-2="http://doi.org/10.5281/zenodo.5636869" httpUri-0="https://zenodo.org/record/5636865/files/figure.png" httpUri-1="https://zenodo.org/record/5636867/files/figure.png" httpUri-2="https://zenodo.org/record/5636869/files/figure.png" pageId="11" pageNumber="1119">Figs 13</figureCitation>
) and nuclear AFLP markers (
<figureCitation id="13342A5DFFC70E75FACB8955FA3FF896" box="[1303,1408,1822,1844]" captionStart-0="Figure 4" captionStart-1="Figure 5" captionStart-2="Figure 6" captionStart-3="Figure 7" captionStartId-0="8.[163,244,1320,1342]" captionStartId-1="9.[145,223,1129,1151]" captionStartId-2="10.[164,244,1619,1641]" captionStartId-3="12.[163,248,649,671]" captionTargetBox-0="[200,1421,195,1281]" captionTargetBox-1="[145,1426,199,1090]" captionTargetBox-2="[356,1248,198,1577]" captionTargetBox-3="[175,770,195,569]" captionTargetId-0="graphics-203@8.[199,1421,195,1281]" captionTargetId-1="graphics-677@9.[259,1424,524,787]" captionTargetId-2="figure-127@10.[353,1252,195,1580]" captionTargetId-3="graphics-743@12.[234,770,202,538]" captionTargetPageId-0="8" captionTargetPageId-1="9" captionTargetPageId-2="10" captionTargetPageId-3="12" captionText-0="Figure 4. Neighbour-net network of Helicopsis from Ukraine based on Jaccard distances obtained from AFLP data. Coloration of clusters corresponds to the STRUCTURE solution for K = 2. Red corresponds to H. filimargo, yellow to H. lunulata. DNA voucher numbers for specimens are given at the tips of the network. Coloured dots at the tips of the network correspond to cluster assignments (on majority-rule basis) of specimens in the STRUCTURE solutions for K = 5 (inner) and K = 7 (outer). For locality data and the distribution of clusters, see also Supporting Information, Table S1 and Figure 6, respectively." captionText-1="Figure 5. Assignment of specimens of Helicopsis from Ukraine to clusters resulting from the admixture analysis of AFLP data with STRUCTURE. A, solution for K = 2. B, solution for K = 5. C, solution for K = 7. Numbers below the plots refer to populations (see Fig. 6; Supporting Information, Table S1). Letters above the plots refer to mitochondrial clades: a, H. filimargo (clade A); b, H. filimargo (clade B); l, H. lunulata." captionText-2="Figure 6. Distribution of clusters obtained from the admixture analysis of AFLP data of Helicopsis from Ukraine with STRUCTURE. A, solution for K = 2. B, solution for K = 5. C, solution for K = 7. Population numbers are given next to the bar plots (Supporting Information, Table S1). The bar plots show the proportional assignment of an individual to a cluster (see also Fig. 5)." captionText-3="Figure 7. Relationships between Jaccard distances between individuals of Helicopsis filimargo and H. lunulata based on AFLP data and logarithmized geographical distances. Black circles and red triangles: distances between individuals belonging to H. filimargo and H. lunulata, respectively; green crosses: distances between individuals belonging to different species; black and red broken lines: regression lines fitted within species; green broken line: regression line fitted on the within-group distances only (i.e. the black circles and red triangles taken together); green solid line: regression line fitted on all distances together; blue lines: centres of the betweengroups geographical distances." figureDoi-0="http://doi.org/10.5281/zenodo.5636871" figureDoi-1="http://doi.org/10.5281/zenodo.5636873" figureDoi-2="http://doi.org/10.5281/zenodo.5636877" figureDoi-3="http://doi.org/10.5281/zenodo.5636879" httpUri-0="https://zenodo.org/record/5636871/files/figure.png" httpUri-1="https://zenodo.org/record/5636873/files/figure.png" httpUri-2="https://zenodo.org/record/5636877/files/figure.png" httpUri-3="https://zenodo.org/record/5636879/files/figure.png" pageId="11" pageNumber="1119">Figs 47</figureCitation>
), demonstrate that the native populations from
<collectingCountry id="F3187648FFC70E75FAEE8977FA2EF8F7" box="[1330,1425,1852,1874]" name="Ukraine" pageId="11" pageNumber="1119">Ukraine</collectingCountry>
and adjacent regions can be classified into two species,
<taxonomicName id="4C0F4D5BFFC00E72FF7F8A61FE9AFB9B" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[163,293,1065,1087]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC00E72FF7F8A61FE9AFB9B" box="[163,293,1065,1087]" italics="true" pageId="12" pageNumber="1120">H. lunulata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC00E72FE868A61FE5CFB9B" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[346,483,1065,1087]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC00E72FE868A61FE5CFB9B" box="[346,483,1065,1087]" italics="true" pageId="12" pageNumber="1120">H. filimargo</emphasis>
</taxonomicName>
. This result resembles the suggestion of
<bibRefCitation id="EF9E4B29FFC00E72FEE78A03FE4FFBFB" author="Balashov IA" box="[315,496,1096,1118]" pageId="12" pageNumber="1120" refId="ref11096" refString="Balashov IA. 2016. Fauna Ukrainy. Tom 29. Mollyuski. Vypusk 5. Stebelchatoglazye (Stylommatophora). Kiev: Naukova Dumka." type="book" year="2016">Balashov (2016)</bibRefCitation>
to classify the Ukrainian
<taxonomicName id="4C0F4D5BFFC00E72FF7F8A2CFEA1FBD9" authorityName="Fitzinger" authorityYear="1833" box="[163,286,1127,1148]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC00E72FF7F8A2CFEA1FBD9" box="[163,286,1127,1148]" italics="true" pageId="12" pageNumber="1120">Helicopsis</emphasis>
</taxonomicName>
populations into two species,
<taxonomicName id="4C0F4D5BFFC00E72FD488A2CFCB4FBD9" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[660,779,1127,1148]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC00E72FD488A2CFCB4FBD9" box="[660,779,1127,1148]" italics="true" pageId="12" pageNumber="1120">H. striata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC00E72FF0B8ACDFED8FB3F" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[215,359,1157,1179]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC00E72FF0B8ACDFED8FB3F" box="[215,359,1157,1179]" italics="true" pageId="12" pageNumber="1120">H. filimargo</emphasis>
</taxonomicName>
. However, the delimitation and the distribution of the species in the two classifications are different. Although
<taxonomicName id="4C0F4D5BFFC00E72FE608A88FDF9FB7D" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[444,582,1219,1240]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC00E72FE608A88FDF9FB7D" box="[444,582,1219,1240]" italics="true" pageId="12" pageNumber="1120">H. lunulata</emphasis>
</taxonomicName>
includes mainly populations with depressed conical shells formerly assigned to
<taxonomicName id="4C0F4D5BFFC00E72FEED8B4AFE1CFAB3" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[305,419,1281,1302]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC00E72FEED8B4AFE1CFAB3" box="[305,419,1281,1302]" italics="true" pageId="12" pageNumber="1120">H. striata</emphasis>
</taxonomicName>
, it also comprises populations with disc-like shells formerly assigned to
<taxonomicName id="4C0F4D5BFFC00E72FDA58B54FCBBFA96" baseAuthorityName="Rossmassler" baseAuthorityYear="1838" box="[633,772,1310,1332]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="instabilis">
<emphasis id="B97BEACAFFC00E72FDA58B54FCBBFA96" box="[633,772,1310,1332]" italics="true" pageId="12" pageNumber="1120">H. instabilis</emphasis>
</taxonomicName>
, which was included in
<taxonomicName id="4C0F4D5BFFC00E72FE7C8B75FD96FAF7" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[416,553,1341,1363]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC00E72FE7C8B75FD96FAF7" box="[416,553,1341,1363]" italics="true" pageId="12" pageNumber="1120">H. filimargo</emphasis>
</taxonomicName>
by
<bibRefCitation id="EF9E4B29FFC00E72FD938B76FCBBFAF6" author="Balashov IA" box="[591,772,1341,1363]" pageId="12" pageNumber="1120" refId="ref11096" refString="Balashov IA. 2016. Fauna Ukrainy. Tom 29. Mollyuski. Vypusk 5. Stebelchatoglazye (Stylommatophora). Kiev: Naukova Dumka." type="book" year="2016">Balashov (2016)</bibRefCitation>
. Moreover,
<bibRefCitation id="EF9E4B29FFC00E72FEC58B17FE70FAD7" author="Balashov IA" box="[281,463,1372,1394]" pageId="12" pageNumber="1120" refId="ref11096" refString="Balashov IA. 2016. Fauna Ukrainy. Tom 29. Mollyuski. Vypusk 5. Stebelchatoglazye (Stylommatophora). Kiev: Naukova Dumka." type="book" year="2016">Balashov (2016)</bibRefCitation>
classified some
<taxonomicName id="4C0F4D5BFFC00E72FD5B8B16FCB4FAD4" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[647,779,1372,1394]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC00E72FD5B8B16FCB4FAD4" box="[647,779,1372,1394]" italics="true" pageId="12" pageNumber="1120">H. lunulata</emphasis>
</taxonomicName>
as
<taxonomicName id="4C0F4D5BFFC00E72FF188B30FE08FA35" baseAuthorityName="Krynicki" baseAuthorityYear="1836" box="[196,439,1402,1424]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="subSpecies" species="filimargo" subSpecies="arenosa">
<emphasis id="B97BEACAFFC00E72FF188B30FE08FA35" box="[196,439,1402,1424]" italics="true" pageId="12" pageNumber="1120">H. filimargo arenosa</emphasis>
</taxonomicName>
and supposed that there are syntopic co-occurrences of
<taxonomicName id="4C0F4D5BFFC00E72FE0D8BD1FD80FA0A" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[465,575,1434,1455]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC00E72FE0D8BD1FD80FA0A" box="[465,575,1434,1455]" italics="true" pageId="12" pageNumber="1120">H. striata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC00E72FDA58BD1FCBAFA0B" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[633,773,1433,1455]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC00E72FDA58BD1FCBAFA0B" box="[633,773,1433,1455]" italics="true" pageId="12" pageNumber="1120">H. filimargo</emphasis>
</taxonomicName>
. We have not found syntopic co-occurrences of
<taxonomicName id="4C0F4D5BFFC00E72FF7F8B9CFE98FA4E" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[163,295,1494,1516]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC00E72FF7F8B9CFE98FA4E" box="[163,295,1494,1516]" italics="true" pageId="12" pageNumber="1120">H. lunulata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC00E72FE838B9CFE56FA4E" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[351,489,1494,1516]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC00E72FE838B9CFE56FA4E" box="[351,489,1494,1516]" italics="true" pageId="12" pageNumber="1120">H. filimargo</emphasis>
</taxonomicName>
, although we found some overlap of the ranges of the two species in the
<collectingRegion id="49CBF83AFFC00E72FD708BBEFCB4F9AE" box="[684,779,1525,1547]" country="Ukraine" name="Donets'k" pageId="12" pageNumber="1120">Donetsk</collectingRegion>
Upland and the adjacent Central Russian Upland so that co-occurrences are possible.
</paragraph>
</subSubSection>
</treatment>
</document>