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<mods:title id="4A51CCA4C051680A27A8F25251EF71DB">Phylogeny, species delimitation and population structure of the steppe-inhabiting land snail genus Helicopsis in Eastern Europe</mods:title>
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<subSubSection id="C3156553FFC00E72FF7F88D1FD06F975" pageId="12" pageNumber="1120" type="nomenclature">
<paragraph id="8BB036D8FFC00E72FF7F88D1FD06F975" blockId="12.[163,779,1690,1744]" pageId="12" pageNumber="1120">
PHYLOGEOGRAPHYOF
<taxonomicName id="4C0F4D5BFFC00E72FE4B88D0FDA0F917" ID-CoL="635L7" ID-ENA="265987" authority="ASACHARACTERISTIC REPRESENTATIVE OF THE" authorityName="ASACHARACTERISTIC REPRESENTATIVE OF THE" box="[407,543,1691,1714]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC00E72FE4B88D0FDA0F917" box="[407,543,1691,1714]" italics="true" pageId="12" pageNumber="1120">HELICOPSIS</emphasis>
</taxonomicName>
ASACHARACTERISTIC REPRESENTATIVE OF THE STEPPE FAUNA
</paragraph>
</subSubSection>
<subSubSection id="C3156553FFC00E70FF7F88AAFDBBFDCF" lastPageId="14" lastPageNumber="1122" pageId="12" pageNumber="1120" type="distribution">
<paragraph id="8BB036D8FFC00E72FF7F88AAFABBFEF0" blockId="12.[163,779,1761,1906]" lastBlockId="12.[827,1444,197,1905]" pageId="12" pageNumber="1120">
The geographical origin of
<taxonomicName id="4C0F4D5BFFC00E72FE0F88AAFDF9F953" box="[467,582,1761,1782]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC00E72FE0F88AAFDF9F953" box="[467,582,1761,1782]" italics="true" pageId="12" pageNumber="1120">Helicopsis</emphasis>
</taxonomicName>
is unclear. Given that the clade including the Central and Eastern European species is sister to a clade comprising species from
<collectingCountry id="F3187648FFC00E72FEE28976FE31F8F7" box="[318,398,1853,1874]" name="Greece" pageId="12" pageNumber="1120">Greece</collectingCountry>
and
<collectingCountry id="F3187648FFC00E72FE178975FDBFF8F6" box="[459,512,1854,1875]" name="Iran" pageId="12" pageNumber="1120">Iran</collectingCountry>
, and the relationships with other genera from the eastern Mediterranean and the Caucasus (
<figureCitation id="13342A5DFFC00E72FBCE8E8EFBEEFF7F" box="[1042,1105,197,219]" 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="12" pageNumber="1120">Fig. 2</figureCitation>
; Supporting Information,
<figureCitation id="13342A5DFFC00E72FAA48E8EFCE7FF5C" 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="12" pageNumber="1120">Fig. S1</figureCitation>
), an origin on the Balkan Peninsula is most likely. The nested position of the Central European clade among the Pontic species indicates an eastern origin of the Central European species (
<figureCitation id="13342A5DFFC00E72FB6A8F0BFB4AFEF0" box="[1206,1269,320,342]" 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="12" pageNumber="1120">Fig. 3</figureCitation>
).
</paragraph>
<paragraph id="8BB036D8FFC00E72FC8F8F14FB0DFCE5" blockId="12.[827,1444,197,1905]" pageId="12" pageNumber="1120">
As in many other steppe-inhabiting organisms (
<bibRefCitation id="EF9E4B29FFC00E72FC9F8F36FB92FE36" author="Kajtoch L &amp; Cieslak E &amp; Varga Z &amp; Paul W &amp; Mazur MA &amp; Sramko G &amp; Kubisz D" box="[835,1069,381,403]" pageId="12" pageNumber="1120" pagination="2309 - 2339" refId="ref12624" refString="Kajtoch L, Cieslak E, Varga Z, Paul W, Mazur MA, Sramko G, Kubisz D. 2016. Phylogeographic patterns of steppe species in eastern Central Europe: a review and the implications for conservation. Biodiversity and Conservation 25: 2309 - 2339." type="journal article" year="2016">
Kajtoch
<emphasis id="B97BEACAFFC00E72FC748F35FC5BFE37" box="[936,996,381,403]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
, 2016
</bibRefCitation>
), the differentiation pattern of the Central and Eastern European
<taxonomicName id="4C0F4D5BFFC00E72FB098FD7FA1DFE17" box="[1237,1442,412,434]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="undetermined">
<emphasis id="B97BEACAFFC00E72FB098FD7FAF5FE14" box="[1237,1354,412,433]" italics="true" pageId="12" pageNumber="1120">Helicopsis</emphasis>
species
</taxonomicName>
reflects the geographic structuring of the steppe belt in Europe in Northern steppes in Central Europe, steppes of the Pannonian Basin and steppes in the Pontic region. Whereas the geographic structure of the steppe belt is reflected in most vagile species at the level of intraspecific lineages (
<bibRefCitation id="EF9E4B29FFC00E72FB4D8C1FFAD6FDCF" author="Kajtoch L &amp; Cieslak E &amp; Varga Z &amp; Paul W &amp; Mazur MA &amp; Sramko G &amp; Kubisz D" box="[1169,1385,596,618]" pageId="12" pageNumber="1120" pagination="2309 - 2339" refId="ref12624" refString="Kajtoch L, Cieslak E, Varga Z, Paul W, Mazur MA, Sramko G, Kubisz D. 2016. Phylogeographic patterns of steppe species in eastern Central Europe: a review and the implications for conservation. Biodiversity and Conservation 25: 2309 - 2339." type="journal article" year="2016">
Kajtoch
<emphasis id="B97BEACAFFC00E72FB338C1FFA99FDCC" box="[1263,1318,596,617]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
, 2016
</bibRefCitation>
), the separation of the regions resulted in a differentiation of species in different regions in
<taxonomicName id="4C0F4D5BFFC00E72FB068CDAFAE8FD03" box="[1242,1367,657,678]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC00E72FB068CDAFAE8FD03" box="[1242,1367,657,678]" italics="true" pageId="12" pageNumber="1120">Helicopsis</emphasis>
</taxonomicName>
. With regard to specific differentiation in subregions of the steppe belt,
<taxonomicName id="4C0F4D5BFFC00E72FC1E8C85FB8AFD46" box="[962,1077,718,739]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC00E72FC1E8C85FB8AFD46" box="[962,1077,718,739]" italics="true" pageId="12" pageNumber="1120">Helicopsis</emphasis>
</taxonomicName>
is more similar to subterranean mole rats (
<bibRefCitation id="EF9E4B29FFC00E72FC628CA6FB14FCA7" author="Nemeth A &amp; Homonnay ZG &amp; Krizsik V &amp; Csorba M &amp; Pavlicek T &amp; Hegyeli Z &amp; Hadid Y &amp; Sugar S &amp; Farkas J &amp; Csorba G" box="[958,1195,749,771]" pageId="12" pageNumber="1120" pagination="903 - 914" refId="ref13035" refString="Nemeth A, Homonnay ZG, Krizsik V, Csorba M, Pavlicek T, Hegyeli Z, Hadid Y, Sugar S, Farkas J, Csorba G. 2013. Old views and new insights: taxonomic revision of the Bukovina blind mole rat, Spalax graecus (Rodentia: Spalacinae). Zoological Journal of the Linnean Society 169: 903 - 914." type="journal article" year="2013">
Németh
<emphasis id="B97BEACAFFC00E72FBFA8CA5FBDDFCA7" box="[1062,1122,749,771]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
, 2013
</bibRefCitation>
) and southern birch mice (
<bibRefCitation id="EF9E4B29FFC00E72FC5B8D47FB32FC87" author="Cserkesz T &amp; Rusin M &amp; Sramko G" box="[903,1165,780,802]" pageId="12" pageNumber="1120" pagination="114 - 130" refId="ref11497" refString="Cserkesz T, Rusin M, Sramko G. 2016. An integrative systematic revision of the European southern birch mice (Rodentia: Sminthidae, Sicista subtilis group). Mammal Review 46: 114 - 130." type="journal article" year="2016">
Cserkész
<emphasis id="B97BEACAFFC00E72FBDD8D47FBFFFC84" box="[1025,1088,780,801]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
, 2016
</bibRefCitation>
) than to other steppe invertebrates investigated so far.
</paragraph>
<paragraph id="8BB036D8FFC00E73FC8F8D02FB50FB3E" blockId="12.[827,1444,197,1905]" lastBlockId="13.[809,1425,1065,1884]" lastPageId="13" lastPageNumber="1121" pageId="12" pageNumber="1120">
The Northern steppes in Central Europe and the steppes of the Pannonian Basin are inhabited by the Central European clade including
<taxonomicName id="4C0F4D5BFFC00E72FADD8DCCFA23FC3E" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[1281,1436,902,924]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC00E72FADD8DCCFA23FC3E" box="[1281,1436,902,924]" italics="true" pageId="12" pageNumber="1120">H. hungarica</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D5BFFC00E72FCE78DEDFC75FC1E" authorityName="E. Gittenberger" authorityYear="1969" box="[827,970,933,955]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="austriaca">
<emphasis id="B97BEACAFFC00E72FCE78DEDFCEFFC1E" box="[827,848,934,955]" italics="true" pageId="12" pageNumber="1120">H</emphasis>
.
<emphasis id="B97BEACAFFC00E72FC818DEDFC75FC1E" box="[861,970,934,955]" italics="true" pageId="12" pageNumber="1120">austriaca</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC00E72FBD88DEDFBCCFC1E" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[1028,1139,934,955]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC00E72FBD88DEDFBCCFC1E" box="[1028,1139,934,955]" italics="true" pageId="12" pageNumber="1120">H. striata</emphasis>
</taxonomicName>
(
<figureCitation id="13342A5DFFC00E72FB5F8DEEFB7CFC1F" box="[1155,1219,933,955]" captionStart="Figure 8" captionStartId="13.[145,223,919,941]" captionTargetBox="[146,1425,197,878]" captionTargetId="figure-477@13.[145,1425,195,879]" captionTargetPageId="13" captionText="Figure 8. Distribution of Helicopsis species in Central and Eastern Europe and localities from which 16S rDNA sequences are available (symbols). The geographical boundaries of H. striata, H. hungarica and H. lunulata are not yet clear because of the lack of reliable morphological characteristics." figureDoi="http://doi.org/10.5281/zenodo.5636881" httpUri="https://zenodo.org/record/5636881/files/figure.png" pageId="12" pageNumber="1120">Fig. 8</figureCitation>
).
<taxonomicName id="4C0F4D5BFFC00E72FB058DEEFA1CFC1E" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[1241,1443,933,955]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC00E72FB058DEEFA1CFC1E" box="[1241,1443,933,955]" italics="true" pageId="12" pageNumber="1120">Helicopsis striata</emphasis>
</taxonomicName>
is widespread in the Northern steppes, but overlaps with
<taxonomicName id="4C0F4D5BFFC00E72FCA58DA8FBA8FC52" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[889,1047,994,1016]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC00E72FCA58DA8FBA8FC52" box="[889,1047,994,1016]" italics="true" pageId="12" pageNumber="1120">H. hungarica</emphasis>
</taxonomicName>
in the north-western Pannonian Basin (
<bibRefCitation id="EF9E4B29FFC00E72FC4F8A4AFB95FBB2" author="Hudec V" box="[915,1066,1025,1047]" pageId="12" pageNumber="1120" pagination="161 - 176" refId="ref12432" refString="Hudec V. 1966. Zur Problematik des Vorkommens der Schnecken Helicopsis striata (Mull.) und H. hungarica (Soos H. Wagner) im Karpatenbecken. Biologia 3: 161 - 176." type="journal article" year="1966">Hudec, 1966</bibRefCitation>
;
<bibRefCitation id="EF9E4B29FFC00E72FBE78A4AFAB2FBB3" author="Duda M &amp; Haring E &amp; Bieringer G &amp; Eschner A &amp; Mrkvicka A &amp; Mason K" box="[1083,1293,1025,1047]" pageId="12" pageNumber="1120" 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="B97BEACAFFC00E72FB598A49FB7CFBB3" box="[1157,1219,1025,1047]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
, 2018
</bibRefCitation>
).
<taxonomicName id="4C0F4D5BFFC00E72FAFB8A4AFC17FB90" authorityName="E. Gittenberger" authorityYear="1969" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="austriaca">
<emphasis id="B97BEACAFFC00E72FAFB8A4AFC17FB90" italics="true" pageId="12" pageNumber="1120">Helicopsis austriaca</emphasis>
</taxonomicName>
is endemic to the eastern fringe of the Alps at the margin of the Pannonian Basin (
<bibRefCitation id="EF9E4B29FFC00E72FACB8A75FCCDFBD7" author="Duda M &amp; Haring E &amp; Bieringer G &amp; Eschner A &amp; Mrkvicka A &amp; Mason K" pageId="12" pageNumber="1120" 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="B97BEACAFFC00E72FABC8A74FA23FBF6" box="[1376,1436,1086,1108]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
, 2018
</bibRefCitation>
). Until recently,
<taxonomicName id="4C0F4D5BFFC00E72FBE98A15FB72FBD7" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[1077,1229,1117,1139]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC00E72FBE98A15FB72FBD7" box="[1077,1229,1117,1139]" italics="true" pageId="12" pageNumber="1120">H. hungarica</emphasis>
</taxonomicName>
was thought to be endemic to the Pannonian Basin. However,
<bibRefCitation id="EF9E4B29FFC00E72FAE98A37FC63FB15" author="Sychev AA &amp; Snegin EA" pageId="12" pageNumber="1120" 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>
published a mitochondrial sequence of a specimen identified as
<taxonomicName id="4C0F4D5BFFC00E72FB888AF1FB7EFB6A" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[1108,1217,1210,1231]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC00E72FB888AF1FB7EFB6A" box="[1108,1217,1210,1231]" italics="true" pageId="12" pageNumber="1120">H. striata</emphasis>
</taxonomicName>
from Gubkin in the Belgorod region in the Central Russian Upland, which was identified as
<taxonomicName id="4C0F4D5BFFC00E72FBD48ABCFB1EFAAE" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[1032,1185,1270,1292]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC00E72FBD48ABCFB1EFAAE" box="[1032,1185,1270,1292]" italics="true" pageId="12" pageNumber="1120">H. hungarica</emphasis>
</taxonomicName>
by
<bibRefCitation id="EF9E4B29FFC00E72FB108ABDFA22FAA9" author="Duda M &amp; Haring E &amp; Bieringer G &amp; Eschner A &amp; Mrkvicka A &amp; Mason K" box="[1228,1437,1270,1292]" pageId="12" pageNumber="1120" 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="B97BEACAFFC00E72FACF8ABCFAF3FAAE" box="[1299,1356,1270,1292]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
(2018)
</bibRefCitation>
. We here report an additional record of this species from Teleshovka in the Belgorod region.
<bibRefCitation id="EF9E4B29FFC00E72FB088B7FFA1CFAEF" author="Duda M &amp; Haring E &amp; Bieringer G &amp; Eschner A &amp; Mrkvicka A &amp; Mason K" box="[1236,1443,1332,1354]" pageId="12" pageNumber="1120" 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="B97BEACAFFC00E72FAC68B7FFAECFAEC" box="[1306,1363,1332,1353]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
(2018)
</bibRefCitation>
discussed three scenarios for the origin of
<taxonomicName id="4C0F4D5BFFC00E72FAD58B18FA22FAC2" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[1289,1437,1362,1384]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC00E72FAD58B18FA22FAC2" box="[1289,1437,1362,1384]" italics="true" pageId="12" pageNumber="1120">H. hungarica</emphasis>
</taxonomicName>
. We consider an origin of the species in the Pannonian Basin more likely than in the Central Russian Upland because it forms a well-supported clade with the Central European
<taxonomicName id="4C0F4D5BFFC00E72FBCC8B85FBC1FA46" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[1040,1150,1486,1507]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC00E72FBCC8B85FBC1FA46" box="[1040,1150,1486,1507]" italics="true" pageId="12" pageNumber="1120">H. striata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC00E72FB6B8B85FAFBFA46" authorityName="E. Gittenberger" authorityYear="1969" box="[1207,1348,1486,1507]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="austriaca">
<emphasis id="B97BEACAFFC00E72FB6B8B85FAFBFA46" box="[1207,1348,1486,1507]" italics="true" pageId="12" pageNumber="1120">H. austriaca</emphasis>
</taxonomicName>
and not with
<taxonomicName id="4C0F4D5BFFC00E72FCAA8BA7FBBCF9A4" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[886,1027,1516,1537]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC00E72FCAA8BA7FBBCF9A4" box="[886,1027,1516,1537]" italics="true" pageId="12" pageNumber="1120">H. filimargo</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC00E72FBE28BA7FB7BF9A4" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1086,1220,1516,1537]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC00E72FBE28BA7FBECF9A4" box="[1086,1107,1516,1537]" italics="true" pageId="12" pageNumber="1120">H</emphasis>
.
<emphasis id="B97BEACAFFC00E72FBBC8BA7FB7BF9A4" box="[1120,1220,1516,1537]" italics="true" pageId="12" pageNumber="1120">lunulata</emphasis>
</taxonomicName>
(
<figureCitation id="13342A5DFFC00E72FB0F8BA7FAABF9A4" box="[1235,1300,1516,1538]" 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="12" pageNumber="1120">Fig. 2</figureCitation>
; Supporting Information,
<figureCitation id="13342A5DFFC00E72FC038841FB89F9BA" box="[991,1078,1546,1568]" 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="12" pageNumber="1120">Fig. S1</figureCitation>
), which currently occupy the eastern part of the range of
<taxonomicName id="4C0F4D5BFFC00E72FB4A8862FAAFF99B" box="[1174,1296,1577,1598]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC00E72FB4A8862FAAFF99B" box="[1174,1296,1577,1598]" italics="true" pageId="12" pageNumber="1120">Helicopsis</emphasis>
</taxonomicName>
(
<figureCitation id="13342A5DFFC00E72FAFF8862FAD8F99B" box="[1315,1383,1577,1599]" captionStart="Figure 8" captionStartId="13.[145,223,919,941]" captionTargetBox="[146,1425,197,878]" captionTargetId="figure-477@13.[145,1425,195,879]" captionTargetPageId="13" captionText="Figure 8. Distribution of Helicopsis species in Central and Eastern Europe and localities from which 16S rDNA sequences are available (symbols). The geographical boundaries of H. striata, H. hungarica and H. lunulata are not yet clear because of the lack of reliable morphological characteristics." figureDoi="http://doi.org/10.5281/zenodo.5636881" httpUri="https://zenodo.org/record/5636881/files/figure.png" pageId="12" pageNumber="1120">Fig. 8</figureCitation>
). We have not found
<taxonomicName id="4C0F4D5BFFC00E72FC338803FB37F9F8" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[1007,1160,1608,1629]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC00E72FC338803FB37F9F8" box="[1007,1160,1608,1629]" italics="true" pageId="12" pageNumber="1120">H. hungarica</emphasis>
</taxonomicName>
populations in
<collectingCountry id="F3187648FFC00E72FAE18803FA20F9FB" box="[1341,1439,1608,1630]" name="Ukraine" pageId="12" pageNumber="1120">Ukraine</collectingCountry>
. Thus, the distributional gap between the Pannonian Basin and the Central Russian Upland seems to be real. However,
<taxonomicName id="4C0F4D5BFFC00E72FC3088E8FBDCF91D" box="[1004,1123,1699,1720]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC00E72FC3088E8FBDCF91D" box="[1004,1123,1699,1720]" italics="true" pageId="12" pageNumber="1120">Helicopsis</emphasis>
</taxonomicName>
was widespread in central and northern
<collectingCountry id="F3187648FFC00E72FC388889FBF8F97D" box="[996,1095,1730,1752]" name="Ukraine" pageId="12" pageNumber="1120">Ukraine</collectingCountry>
in the Pleistocene (
<bibRefCitation id="EF9E4B29FFC00E72FAE08889FCCCF953" author="Kunitsa NA" pageId="12" pageNumber="1120" refId="ref12862" refString="Kunitsa NA. 2007. Priroda Ukrainy v Plejstocene (po dannym malakofaunisticheskogo analiza). Chernovtsi: Ruta." type="book" year="2007">Kunitsa, 2007</bibRefCitation>
), where it is currently absent (
<bibRefCitation id="EF9E4B29FFC00E72FB3788AAFA23F952" author="Balashov IA" box="[1259,1436,1761,1783]" 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>
;
<bibRefCitation id="EF9E4B29FFC00E72FCE788B4FB83F8B0" author="Balashov I &amp; Vasyliuk O &amp; Shyriaieva D &amp; Shvydka Z &amp; Oskyrko O &amp; Marushchak O &amp; Stetsun H &amp; Bezsmertna O &amp; Babytskij A &amp; Kostiushyn V" box="[827,1084,1791,1813]" pageId="12" pageNumber="1120" pagination="3 - 12" refId="ref11275" refString="Balashov I, Vasyliuk O, Shyriaieva D, Shvydka Z, Oskyrko O, Marushchak O, Stetsun H, Bezsmertna O, Babytskij A, Kostiushyn V. 2018 b. Terrestrial molluscs in the dry grasslands of the Dnipro Upland (central Ukraine): new records, rare species and conservation potential. Vestnik Zoologii 52: 3 - 12." type="journal article" year="2018">
Balashov
<emphasis id="B97BEACAFFC00E72FC72894BFC57F8B1" box="[942,1000,1791,1813]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
, 2018b
</bibRefCitation>
).
<bibRefCitation id="EF9E4B29FFC00E72FB8F88B4FA87F8B0" author="Sychev AA &amp; Snegin EA &amp; Shapovalov AS &amp; Ponomarenko EV &amp; Chendev YG" box="[1107,1336,1791,1813]" pageId="12" pageNumber="1120" pagination="146 - 164" refId="ref14103" refString="Sychev AA, Snegin EA, Shapovalov AS, Ponomarenko EV, Chendev YG. 2015. K voprosu o strukture fauny nazemnykh molljuskov zapovednogo uchastka &quot; Yamskaya step' &quot; v pozdnem golotsene. Vestnik Tomskogo gosudarstvennogo universiteta, Biologija 2: 146 - 164." type="journal article" year="2015">
Sychev
<emphasis id="B97BEACAFFC00E72FB71894BFB58F8B1" box="[1197,1255,1791,1813]" italics="true" pageId="12" pageNumber="1120">et al.</emphasis>
(2015)
</bibRefCitation>
reported
<taxonomicName id="4C0F4D5BFFC00E72FCE78955FC0DF896" box="[827,946,1822,1843]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC00E72FCE78955FC0DF896" box="[827,946,1822,1843]" italics="true" pageId="12" pageNumber="1120">Helicopsis</emphasis>
</taxonomicName>
from an almost 3000-year-old soil horizon near Gubkin in the Belgorod region. Thus, it is possible that
<taxonomicName id="4C0F4D5BFFC00E72FCAB8917FBA8F8D5" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[887,1047,1883,1905]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="12" pageNumber="1120" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC00E72FCAB8917FBA8F8D5" box="[887,1047,1883,1905]" italics="true" pageId="12" pageNumber="1120">H. hungarica</emphasis>
</taxonomicName>
expanded eastwards during the Pleistocene and later went extinct in the large area between the Pannonian Basin and the Central Russian Upland. However, the fossil shells cannot be reliably identified at the species level and it is also possible that the Pleistocene
<taxonomicName id="4C0F4D5BFFC10E73FE328AEFFDD9FB1C" box="[494,614,1188,1209]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC10E73FE328AEFFDD9FB1C" box="[494,614,1188,1209]" italics="true" pageId="13" pageNumber="1121">Helicopsis</emphasis>
</taxonomicName>
populations from central and northern
<collectingCountry id="F3187648FFC10E73FE008A88FD80FB7C" box="[476,575,1219,1241]" name="Ukraine" pageId="13" pageNumber="1121">Ukraine</collectingCountry>
belonged to
<taxonomicName id="4C0F4D5BFFC10E73FD028A88FF4BFB53" baseAuthorityName="Krynicki" baseAuthorityYear="1833" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC10E73FD028A88FD4CFB7D" box="[734,755,1219,1240]" italics="true" pageId="13" pageNumber="1121">H</emphasis>
.
<emphasis id="B97BEACAFFC10E73FF4D8AAAFF4BFB53" box="[145,244,1249,1270]" italics="true" pageId="13" pageNumber="1121">lunulata</emphasis>
</taxonomicName>
and formed the connection between its main range in western and southern
<collectingCountry id="F3187648FFC10E73FDFC8B4BFD3AFAB3" box="[544,645,1280,1302]" name="Ukraine" pageId="13" pageNumber="1121">Ukraine</collectingCountry>
with the nowadays isolated occurrences in the
<collectingRegion id="49CBF83AFFC10E73FDE28B54FD22FA90" box="[574,669,1311,1333]" country="Ukraine" name="Donets'k" pageId="13" pageNumber="1121">Donetsk</collectingRegion>
Upland and the Central Russian Upland. An alternative explanation for the isolated occurrence of
<taxonomicName id="4C0F4D5BFFC10E73FDB88B16FD46FAD4" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[612,761,1372,1394]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC10E73FDB88B16FD46FAD4" box="[612,761,1372,1394]" italics="true" pageId="13" pageNumber="1121">H. hungarica</emphasis>
</taxonomicName>
haplotypes in the Central Russian Upland might be an introduction with mining equipment for the extensive iron-ore quarries near Gubkin. However, this hypothesis does not explain the presence of fossil
<taxonomicName id="4C0F4D5BFFC10E73FF4D8BBEFEB4F9AF" box="[145,267,1525,1546]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="genus">
<emphasis id="B97BEACAFFC10E73FF4D8BBEFEB4F9AF" box="[145,267,1525,1546]" italics="true" pageId="13" pageNumber="1121">Helicopsis</emphasis>
</taxonomicName>
in that region and the occurrence of the species in natural steppe communities. The specific identity of the specimens from the Central Russian Upland with the Pannonian
<taxonomicName id="4C0F4D5BFFC10E73FE3F8819FDC0F9C3" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[483,639,1617,1639]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC10E73FE3F8819FDC0F9C3" box="[483,639,1617,1639]" italics="true" pageId="13" pageNumber="1121">H. hungarica</emphasis>
</taxonomicName>
should be confirmed with non-mitochondrial genetic markers to ensure that the
<taxonomicName id="4C0F4D5BFFC10E73FE9288C4FE57F901" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[334,488,1679,1700]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC10E73FE9288C4FE57F901" box="[334,488,1679,1700]" italics="true" pageId="13" pageNumber="1121">H. hungarica</emphasis>
</taxonomicName>
haplotypes are not the result of an introgression. There is no evidence for the final scenario of
<bibRefCitation id="EF9E4B29FFC10E73FE848887FD90F947" author="Duda M &amp; Haring E &amp; Bieringer G &amp; Eschner A &amp; Mrkvicka A &amp; Mason K" box="[344,559,1740,1762]" pageId="13" pageNumber="1121" 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="B97BEACAFFC10E73FE7D8886FE63F944" box="[417,476,1740,1762]" italics="true" pageId="13" pageNumber="1121">et al.</emphasis>
(2018)
</bibRefCitation>
that the present distribution of
<taxonomicName id="4C0F4D5BFFC10E73FEE288A0FE69F8A5" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[318,470,1771,1792]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC10E73FEE288A0FE69F8A5" box="[318,470,1771,1792]" italics="true" pageId="13" pageNumber="1121">H. hungarica</emphasis>
</taxonomicName>
in the Carpathian Basin derives from a hitherto unknown southern refuge. The southern range border of
<taxonomicName id="4C0F4D5BFFC10E73FE1F8962FDDFF898" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[451,608,1832,1854]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC10E73FE1F8962FDDFF898" box="[451,608,1832,1854]" italics="true" pageId="13" pageNumber="1121">H. hungarica</emphasis>
</taxonomicName>
is unknown. It is possible that the species extends southwards into the western Balkan Peninsula. However, there is no necessity to postulate a southern refugium for the survival of the species, given that
<taxonomicName id="4C0F4D5BFFC10E73FB278A2CFA2EFBD9" authorityName="E. Gittenberger" authorityYear="1969" box="[1275,1425,1127,1148]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="austriaca">
<emphasis id="B97BEACAFFC10E73FB278A2CFA2EFBD9" box="[1275,1425,1127,1148]" italics="true" pageId="13" pageNumber="1121">H. austriaca</emphasis>
</taxonomicName>
obviously survived at asimilar latitude.
</paragraph>
<caption id="DF706650FFC10E73FF4D8DDCFD13FC4D" ID-DOI="http://doi.org/10.5281/zenodo.5636881" ID-Zenodo-Dep="5636881" httpUri="https://zenodo.org/record/5636881/files/figure.png" pageId="13" pageNumber="1121" startId="13.[145,223,919,941]" targetBox="[146,1425,197,878]" targetPageId="13">
<paragraph id="8BB036D8FFC10E73FF4D8DDCFD13FC4D" blockId="13.[145,1424,919,1000]" pageId="13" pageNumber="1121">
<emphasis id="B97BEACAFFC10E73FF4D8DDCFF47FC09" bold="true" box="[145,248,919,941]" pageId="13" pageNumber="1121">Figure 8.</emphasis>
Distribution of
<taxonomicName id="4C0F4D5BFFC10E73FE7F8DDCFDE3FC08" box="[419,604,919,941]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="undetermined">
<emphasis id="B97BEACAFFC10E73FE7F8DDCFDB2FC08" box="[419,525,919,941]" italics="true" pageId="13" pageNumber="1121">Helicopsis</emphasis>
species
</taxonomicName>
in Central and Eastern Europe and localities from which 16S rDNA sequences are available (symbols). The geographical boundaries of
<taxonomicName id="4C0F4D5BFFC10E73FD018DFEFCFDFC6E" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[733,834,949,971]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC10E73FD018DFEFCFDFC6E" box="[733,834,949,971]" italics="true" pageId="13" pageNumber="1121">H. striata</emphasis>
</taxonomicName>
,
<taxonomicName id="4C0F4D5BFFC10E73FC928DFEFC67FC6E" baseAuthorityName="Duda" baseAuthorityYear="2019" box="[846,984,949,971]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="hungarica">
<emphasis id="B97BEACAFFC10E73FC928DFEFC67FC6E" box="[846,984,949,971]" italics="true" pageId="13" pageNumber="1121">H. hungarica</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC10E73FBD18DFEFB38FC6E" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1037,1159,949,971]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC10E73FBD18DFEFB9FFC6F" box="[1037,1056,949,970]" italics="true" pageId="13" pageNumber="1121">H</emphasis>
.
<emphasis id="B97BEACAFFC10E73FBF08DFEFB38FC6E" box="[1068,1159,949,971]" italics="true" pageId="13" pageNumber="1121">lunulata</emphasis>
</taxonomicName>
are not yet clear because of the lack of reliable morphological characteristics.
</paragraph>
</caption>
<paragraph id="8BB036D8FFC10E73FC9D8AEFFB25F901" blockId="13.[809,1425,1065,1884]" pageId="13" pageNumber="1121">
The Pontic region is characterized by
<taxonomicName id="4C0F4D5BFFC10E73FADD8AEEFA2EFB1C" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1281,1425,1188,1210]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC10E73FADD8AEEFA2EFB1C" box="[1281,1425,1188,1210]" italics="true" pageId="13" pageNumber="1121">H. filimargo</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC10E73FC838A88FC53FB7D" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[863,1004,1219,1240]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC10E73FC838A88FCCBFB7D" box="[863,884,1219,1240]" italics="true" pageId="13" pageNumber="1121">H</emphasis>
.
<emphasis id="B97BEACAFFC10E73FC588A88FC53FB7D" box="[900,1004,1219,1240]" italics="true" pageId="13" pageNumber="1121">lunulata</emphasis>
</taxonomicName>
(
<figureCitation id="13342A5DFFC10E73FC228A88FBF6FB7D" box="[1022,1097,1219,1241]" 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="13" pageNumber="1121">Figs 1</figureCitation>
,
<figureCitation id="13342A5DFFC10E73FB858A88FBD8FB7D" box="[1113,1127,1219,1240]" captionStart="Figure 8" captionStartId="13.[145,223,919,941]" captionTargetBox="[146,1425,197,878]" captionTargetId="figure-477@13.[145,1425,195,879]" captionTargetPageId="13" captionText="Figure 8. Distribution of Helicopsis species in Central and Eastern Europe and localities from which 16S rDNA sequences are available (symbols). The geographical boundaries of H. striata, H. hungarica and H. lunulata are not yet clear because of the lack of reliable morphological characteristics." figureDoi="http://doi.org/10.5281/zenodo.5636881" httpUri="https://zenodo.org/record/5636881/files/figure.png" pageId="13" pageNumber="1121">8</figureCitation>
).
<taxonomicName id="4C0F4D5BFFC10E73FB5C8A88FACBFB7D" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1152,1396,1219,1240]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC10E73FB5C8A88FACBFB7D" box="[1152,1396,1219,1240]" italics="true" pageId="13" pageNumber="1121">Helicopsis filimargo</emphasis>
</taxonomicName>
is found east of the Dnieper River and especially in Crimea, whereas
<taxonomicName id="4C0F4D5BFFC10E73FC248B4AFB3FFAB0" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1016,1152,1280,1302]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC10E73FC248B4AFBB2FAB3" box="[1016,1037,1281,1302]" italics="true" pageId="13" pageNumber="1121">H</emphasis>
.
<emphasis id="B97BEACAFFC10E73FBC78B4BFB3FFAB0" box="[1051,1152,1280,1301]" italics="true" pageId="13" pageNumber="1121">lunulata</emphasis>
</taxonomicName>
occupies the part west of the Dnieper. It is not clear whether the isolated populations of
<taxonomicName id="4C0F4D5BFFC10E73FC0C8B75FBEAFAF7" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[976,1109,1341,1363]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC10E73FC0C8B75FBEAFAF7" box="[976,1109,1341,1363]" italics="true" pageId="13" pageNumber="1121">H. lunulata</emphasis>
</taxonomicName>
in the Donetsk Upland and the adjacent Central Russian Upland are the remains of a formerly continuous range (see above) or whether they are the result of transport by humans. The former hypothesis is supported by the fact that
<taxonomicName id="4C0F4D5BFFC10E73FAD48BF2FA2FFA68" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1288,1424,1464,1486]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC10E73FAD48BF2FA2FFA68" box="[1288,1424,1464,1486]" italics="true" pageId="13" pageNumber="1121">H. lunulata</emphasis>
</taxonomicName>
occupies only undisturbed natural steppes in the Donetsk Upland, whereas
<taxonomicName id="4C0F4D5BFFC10E73FBB28BBDFABCF9AF" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1134,1283,1525,1547]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC10E73FBB28BBDFABCF9AF" box="[1134,1283,1525,1547]" italics="true" pageId="13" pageNumber="1121">H. filimargo</emphasis>
</taxonomicName>
can occupy transformed habitats. The geographical boundaries of
<taxonomicName id="4C0F4D5BFFC10E73FC998878FC0AF9ED" baseAuthorityName="Muller" baseAuthorityYear="1774" box="[837,949,1587,1608]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC10E73FC998878FC0AF9ED" box="[837,949,1587,1608]" italics="true" pageId="13" pageNumber="1121">H. striata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C0F4D5BFFC10E73FC338878FBC9F9E2" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1007,1142,1586,1608]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC10E73FC338878FBC9F9E2" box="[1007,1142,1586,1608]" italics="true" pageId="13" pageNumber="1121">H. lunulata</emphasis>
</taxonomicName>
are unclear. It has to be checked whether the populations from south-eastern
<collectingCountry id="F3187648FFC10E73FCF5883BFCC6F923" box="[809,889,1648,1670]" name="Poland" pageId="13" pageNumber="1121">Poland</collectingCountry>
, classified as
<taxonomicName id="4C0F4D5BFFC10E73FBCD883AFA83F920" authority="(Stepczak, 2004)" baseAuthorityName="Stepczak" baseAuthorityYear="2004" box="[1041,1340,1648,1670]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC10E73FBCD883AFBC2F920" box="[1041,1149,1648,1670]" italics="true" pageId="13" pageNumber="1121">H. striata</emphasis>
(
<bibRefCitation id="EF9E4B29FFC10E73FB57883BFA8CF920" author="Stepczak K" box="[1163,1331,1648,1670]" pageId="13" pageNumber="1121" pagination="1774" refId="ref13905" refString="Stepczak K. 2004. Helicopsis striata (O. F. Muller, 1774). In: Glowacinski Z, Nowacki J, eds. Polska czerwona ksiega zwierzat. Bezkregowce. Krakow: Instytut Ochrony Przyrody Polskiej Akademii Nauk." type="book chapter" year="2004">Stępczak, 2004</bibRefCitation>
)
</taxonomicName>
, belong to that species or to
<taxonomicName id="4C0F4D5BFFC10E73FBD388C4FB2CF906" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[1039,1171,1678,1700]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC10E73FBD388C4FB2CF906" box="[1039,1171,1678,1700]" italics="true" pageId="13" pageNumber="1121">H. lunulata</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BB036D8FFC10E70FC9E88E6FDBBFDCF" blockId="13.[809,1425,1065,1884]" lastBlockId="14.[163,779,197,618]" lastPageId="14" lastPageNumber="1122" pageId="13" pageNumber="1121">
<bibRefCitation id="EF9E4B29FFC10E73FC9E88E6FBFDF966" author="Stewart JR &amp; Lister AM &amp; Barnes I &amp; Dalen L" box="[834,1090,1709,1731]" pageId="13" pageNumber="1121" pagination="661 - 671" refId="ref13978" refString="Stewart JR, Lister AM, Barnes I, Dalen L. 2010. Refugia revisited: individualistic responses of species in space and time. Proceedings of the Royal Society B: Biological Sciences 277: 661 - 671." type="journal article" year="2010">
Stewart
<emphasis id="B97BEACAFFC10E73FC7188E5FC54F967" box="[941,1003,1709,1731]" italics="true" pageId="13" pageNumber="1121">et al.</emphasis>
(2010)
</bibRefCitation>
suggested that the ranges of some steppe species are currently contracted in interglacial refugia. This is the case for
<taxonomicName id="4C0F4D5BFFC10E73FB3188A0FAE5F8A5" baseAuthorityName="Stepczak" baseAuthorityYear="2004" box="[1261,1370,1771,1792]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="13" pageNumber="1121" phylum="Ascomycota" rank="species" species="striata">
<emphasis id="B97BEACAFFC10E73FB3188A0FAE5F8A5" box="[1261,1370,1771,1792]" italics="true" pageId="13" pageNumber="1121">H. striata</emphasis>
</taxonomicName>
that was more widespread in Central Europe westwards to
<collectingCountry id="F3187648FFC10E73FC9B8963FC16F89B" box="[839,937,1832,1854]" name="United Kingdom" pageId="13" pageNumber="1121">England</collectingCountry>
during the glacials (
<bibRefCitation id="EF9E4B29FFC10E73FB408963FA87F89B" author="Jaeckel SGA" box="[1180,1336,1832,1854]" pageId="13" pageNumber="1121" pagination="25 - 294" refId="ref12495" refString="Jaeckel SGA. 1962. Erganzungen und Berichtigungen zum rezenten und quartaren Vorkommen der mitteleuropaischen Mollusken. In: Brohmer P, Ehrmann P, Ulmer G, eds. Die Tierwelt Mitteleuropas 2 (1, Erg.). Leipzig: Quelle and Meyer, 25 - 294." type="book chapter" year="1962">Jaeckel, 1962</bibRefCitation>
;
<bibRefCitation id="EF9E4B29FFC10E73FA9A8963FCDBF8F9" author="Lozek V" pageId="13" pageNumber="1121" pagination="1 - 374" refId="ref12927" refString="Lozek V. 1964. Quartarmollusken der Tschechoslowakei. Rozpravy Ustredniho Ustavu Geologickeho 31: 1 - 374, 32 pls." type="journal article" year="1964">Ložek, 1964</bibRefCitation>
), but is currently restricted to some of the Northern steppe fragments and the north-western part of the Pannonian Basin. However, not all steppe species are contracting in the interglacials. This is shown by significantly negative Fus
<emphasis id="B97BEACAFFC20E70FD548F69FD20FE98" box="[648,671,290,317]" italics="true" pageId="14" pageNumber="1122">F s</emphasis>
statistic indicating population expansion in
<taxonomicName id="4C0F4D5BFFC20E70FD9E8F0AFD6EFEF0" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[578,721,320,342]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="14" pageNumber="1122" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC20E70FD9E8F0AFDE8FEF3" box="[578,599,321,342]" italics="true" pageId="14" pageNumber="1122">H</emphasis>
.
<emphasis id="B97BEACAFFC20E70FDB98F0BFD6EFEF0" box="[613,721,320,341]" italics="true" pageId="14" pageNumber="1122">filimargo</emphasis>
</taxonomicName>
. The highest mitochondrial haplotype diversity, and a high diversity with regard to nuclear AFLP markers, were found in
<collectingRegion id="49CBF83AFFC20E70FE908FD7FE1AFE17" box="[332,421,412,434]" country="Ukraine" name="Crimea" pageId="14" pageNumber="1122">Crimea</collectingRegion>
.
<taxonomicName id="4C0F4D5BFFC20E70FE6D8FD7FD23FE14" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[433,668,412,433]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="14" pageNumber="1122" phylum="Ascomycota" rank="species" species="filimargo">
<emphasis id="B97BEACAFFC20E70FE6D8FD7FD23FE14" box="[433,668,412,433]" italics="true" pageId="14" pageNumber="1122">Helicopsis filimargo</emphasis>
</taxonomicName>
probably originated in
<collectingRegion id="49CBF83AFFC20E70FE9A8FF1FE22FE75" box="[326,413,442,464]" country="Ukraine" name="Crimea" pageId="14" pageNumber="1122">Crimea</collectingRegion>
and expanded from there into the Black Sea Lowland and the
<collectingRegion id="49CBF83AFFC20E70FDCE8F92FDCEFE4A" box="[530,625,473,495]" country="Ukraine" name="Donets'k" pageId="14" pageNumber="1122">Donetsk</collectingRegion>
Upland. Fus
<emphasis id="B97BEACAFFC20E70FF7F8FB3FF06FDB1" box="[163,185,504,532]" italics="true" pageId="14" pageNumber="1122">F s</emphasis>
statistic was not significant in
<taxonomicName id="4C0F4D5BFFC20E70FDF38FB2FD08FDA8" baseAuthorityName="Krynicki" baseAuthorityYear="1833" box="[559,695,504,526]" class="Dothideomycetes" family="Tubeufiaceae" genus="Helicopsis" kingdom="Fungi" order="Tubeufiales" pageId="14" pageNumber="1122" phylum="Ascomycota" rank="species" species="lunulata">
<emphasis id="B97BEACAFFC20E70FDF38FB2FDFBFDAB" box="[559,580,505,526]" italics="true" pageId="14" pageNumber="1122">H</emphasis>
.
<emphasis id="B97BEACAFFC20E70FD8E8FB3FD08FDA8" box="[594,695,504,525]" italics="true" pageId="14" pageNumber="1122">lunulata</emphasis>
</taxonomicName>
, which is patchily distributed across
<collectingCountry id="F3187648FFC20E70FDD98C5DFDD6FD89" box="[517,617,534,556]" name="Ukraine" pageId="14" pageNumber="1122">Ukraine</collectingCountry>
. However, we cannot exclude that the lack of significance was the result of the small sample size.
</paragraph>
</subSubSection>
</treatment>
</document>