532 lines
75 KiB
XML
532 lines
75 KiB
XML
<document id="61F085871E4E0B43D14B67AF965EBE42" ID-CLB-Dataset="305150" ID-DOI="10.1002/tax.13176" ID-GBIF-Dataset="73548f05-40de-4973-a03f-cc7ca3fcce2a" ID-ISSN="1996-8175" ID-Zenodo-Dep="14047615" IM.bibliography_approvedBy="julia" IM.illustrations_approvedBy="julia" IM.materialsCitations_approvedBy="julia" IM.tables_approvedBy="julia" IM.taxonomicNames_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1730918502192" checkinUser="felipe" docAuthor="Khan, Gulzar, Mayland-Quellhorst, Eike, Kosachev, Petr A., Mandáková, Terezie, Lysak, Martin A. & Albach, Dirk C." docDate="2024" docId="03E0551FFFCE990A31F5F90AFDA8BBF0" docLanguage="en" docName="TAXON.73.2.530-546.pdf" docOrigin="TAXON 73 (2)" docSource="http://dx.doi.org/10.1002/tax.13176" docStyle="DocumentStyle:239A75409313D6488D9D82529296B06F.1:TAXON.2020-.journal_article.open" docStyleId="239A75409313D6488D9D82529296B06F" docStyleName="TAXON.2020-.journal_article.open" docStyleVersion="1" docTitle="Veronica (Pseudolysimachium)" docType="treatment" docVersion="3" lastPageNumber="539" masterDocId="FFD92D67FFC899033161FFD2FFCABF02" masterDocTitle="Altai Mountains - cradle of hybrids and introgressants: A case study in Veronica subg. Pseudolysimachium (Plantaginaceae)" masterLastPageNumber="546" masterPageNumber="530" pageNumber="536" updateTime="1731438523770" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-ND-4.0">
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<mods:title id="31819B65AF02C68DB67D6DFC272B71F3">Altai Mountains - cradle of hybrids and introgressants: A case study in Veronica subg. Pseudolysimachium (Plantaginaceae)</mods:title>
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<mods:namePart id="7A8C129D1FE60DD6A3741BC812592465">Khan, Gulzar</mods:namePart>
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<mods:affiliation id="626EFC4A2A57763A64B375FE99168218">Eike Mayland-Quellhorst, & Petr A. Kosachev, & Institute for Biology and Environmental Sciences, Carl von Ossietzky-University Oldenburg, Carl von Ossietzky-Str. & type. The GISH investigation in one case strongly supports homoploid hybridization (origin of V. × schmakovii from V. longifolia and V. porphyriana. Divergence times of Altai Veronica species are estimated to be within & -</mods:affiliation>
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<mods:namePart id="56ECE4DBFA1397CC86D38DC8D403794F">Mayland-Quellhorst, Eike</mods:namePart>
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<mods:affiliation id="0F85AB9142E9A4A40EB96B32655E4B26">Eike Mayland-Quellhorst, & Petr A. Kosachev, & Institute for Biology and Environmental Sciences, Carl von Ossietzky-University Oldenburg, Carl von Ossietzky-Str. & type. The GISH investigation in one case strongly supports homoploid hybridization (origin of V. × schmakovii from V. longifolia and V. porphyriana. Divergence times of Altai Veronica species are estimated to be within & -</mods:affiliation>
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<mods:namePart id="3BDD4CF63E836BB63C7CFD9FC9859CEF">Kosachev, Petr A.</mods:namePart>
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<mods:affiliation id="3D37ED90043550775EA59E5D11802676">Terezie Mandáková, & Faculty of Biology, Altai State University, Lenina 61, 656049 Barnaul, Russia & million years ago with high probability of gene flow over that time. Our results also demonstrate that the direction of gene flow is mainly from the locally endemic V. porphyriana. We hypothesize that the large Siberian plains and topographically diverse foreland of the Altai Mountains provide an ideal setting for hybridization with the potential for adaptive introgression of alleles conferring tolerance to cooler climates, to the lowland species migrating into the Altai Mountains.</mods:affiliation>
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<mods:namePart id="856BD22326E17E928DAA2604C062CC75">Mandáková, Terezie</mods:namePart>
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<mods:affiliation id="1725422608EC9B1E6A4EA9CEDEA6ACC6">& Dirk C. Albach & Central European Institute of Technology (CEITEC), Masaryk University, Brno, Czech Republic</mods:affiliation>
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<mods:namePart id="3447F00812FD779A90318A76860F4243">Lysak, Martin A.</mods:namePart>
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<mods:affiliation id="4B119696D290182AC661F34F0C6A1382">& Dirk C. Albach & Central European Institute of Technology (CEITEC), Masaryk University, Brno, Czech Republic</mods:affiliation>
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<mods:namePart id="299134D6FE147D56FD643F676673D75B">Albach, Dirk C.</mods:namePart>
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<mods:affiliation id="D3A4A84FF88E39F4C88D33C3EED9A42A">Eike Mayland-Quellhorst, & Petr A. Kosachev, & Institute for Biology and Environmental Sciences, Carl von Ossietzky-University Oldenburg, Carl von Ossietzky-Str. & type. The GISH investigation in one case strongly supports homoploid hybridization (origin of V. × schmakovii from V. longifolia and V. porphyriana. Divergence times of Altai Veronica species are estimated to be within & -</mods:affiliation>
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<mods:title id="7A52B07B6CB1E0561464B29527401DE8">TAXON</mods:title>
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<mods:date id="A76AFE14DA314B57E2DBE2C4E53FBCAF">2024</mods:date>
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<mods:number id="A7BEC7F3444480E3B4E37D0F203F12AA">2024-04-30</mods:number>
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<mods:number id="7728FCF44C8A1F988C5BA9A5E340BEAA">73</mods:number>
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<mods:url id="4E8E0D82514CB40836FE6CC2E498B414">http://dx.doi.org/10.1002/tax.13176</mods:url>
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<mods:classification id="4737E30F8625B3D130B85B749D4F03FB">journal article</mods:classification>
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<mods:identifier id="4750BD53D29CDEE682F77C5F2C789879" type="CLB-Dataset">305150</mods:identifier>
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<mods:identifier id="F5A1D48C565F9B8BA1D87658EAF11D75" type="DOI">10.1002/tax.13176</mods:identifier>
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<mods:identifier id="5D34260143B0A03ADFE802F76383CAF4" type="ISSN">1996-8175</mods:identifier>
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<mods:identifier id="B18C2F05D52C2230D9FA3A3914791A64" type="Zenodo-Dep">14047615</mods:identifier>
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<treatment id="03E0551FFFCE990A31F5F90AFDA8BBF0" ID-DOI="http://doi.org/10.5281/zenodo.14107803" ID-Zenodo-Dep="14107803" LSID="urn:lsid:plazi:treatment:03E0551FFFCE990A31F5F90AFDA8BBF0" httpUri="http://treatment.plazi.org/id/03E0551FFFCE990A31F5F90AFDA8BBF0" lastPageId="9" lastPageNumber="539" pageId="6" pageNumber="536">
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<subSubSection id="C353B782FFCE990531F5F90AFEE7B9F1" box="[148,301,1752,1779]" pageId="6" pageNumber="536" type="nomenclature">
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<paragraph id="8BF6E409FFCE990531F5F90AFEE7B9F1" blockId="6.[148,301,1752,1779]" box="[148,301,1752,1779]" pageId="6" pageNumber="536">
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<heading id="D0BE5365FFCE990531F5F90AFEE7B9F1" box="[148,301,1752,1779]" fontSize="36" level="1" pageId="6" pageNumber="536" reason="1">
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■
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<taxonomicName id="4C499F8AFFCE990531D1F90BFEE7B9F1" box="[176,301,1753,1779]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="subGenus" subGenus="Pseudolysimachium">
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<emphasis id="B93D381BFFCE990531D1F90BFEE7B9F1" bold="true" box="[176,301,1753,1779]" pageId="6" pageNumber="536">RESULTS</emphasis>
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="C353B782FFCE990A31F5F8C9FDA8BBF0" lastPageId="9" lastPageNumber="539" pageId="6" pageNumber="536" type="discussion">
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<paragraph id="8BF6E409FFCE990531F5F8C9FD25B8B0" blockId="6.[103,751,1819,1970]" pageId="6" pageNumber="536">
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<emphasis id="B93D381BFFCE990531F5F8C9FDF0B830" bold="true" box="[148,570,1819,1843]" pageId="6" pageNumber="536">
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Species boundaries and cohesion.
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<emphasis id="B93D381BFFCE9905337EF8CFFDF0B830" bold="true" box="[543,570,1821,1842]" italics="true" pageId="6" pageNumber="536">—</emphasis>
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</emphasis>
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||
The assignment test in STRUCTURE with the dataset including all 233 individuals recovered
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<emphasis id="B93D381BFFCE9905304AF889FEF7B870" box="[299,317,1883,1906]" italics="true" pageId="6" pageNumber="536">K</emphasis>
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= 5 as optimal both by Evanno Δ
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<emphasis id="B93D381BFFCE990533CFF889FD0AB870" box="[686,704,1883,1906]" italics="true" pageId="6" pageNumber="536">K</emphasis>
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and likelihood scores (suppl.
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||
<figureCitation id="1372F88CFFCE990530E2F8A9FE1DB890" box="[387,471,1915,1938]" captionStart="Fig" captionStartId="10.[103,135,1865,1886]" captionTargetBox="[278,1262,661,1831]" captionTargetId="figure-267@10.[278,1262,661,1831]" captionTargetPageId="10" captionText="Fig. 3. Chromosome localization of rDNA and genomic in situ hybridization (GISH) in Veronica. Mitotic chromosome complements of: A, V. porphyriana; B, V. ×schmakovii; C, V. spicata; D, V. pinnata; E, V. longifolia; and F, V. incana hybridized with 35S (red fluorescence) and 5S (purple) rDNA probes. G, Mitotic chromosomes of V. ×schmakovii hybridized with gDNA of V. longifolia (red) and V. porphyriana (green). — Chromosomes were counterstained with DAPI. Scale bars, 10 μm." figureDoi="http://doi.org/10.5281/zenodo.14047621" httpUri="https://zenodo.org/record/14047621/files/figure.png" pageId="6" pageNumber="536">Fig. S3</figureCitation>
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), which differs from the expected
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<emphasis id="B93D381BFFCE990531B3F849FF2EB8B0" box="[210,228,1947,1970]" italics="true" pageId="6" pageNumber="536">K</emphasis>
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= 10 based on species assignment (
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<tableCitation id="C6CBD1B2FFCE990533E9F849FD15B8B0" box="[648,735,1947,1970]" captionStart="Table 1" captionStartId="2.[103,156,1099,1120]" captionTargetId="graphics-522@2.[103,1436,1533,1573]" captionTargetPageId="2" captionText="Table 1. Details of sampled morphotypes and their geographical distribution." httpUri="http://table.plazi.org/id/DF36B481FFCA99013106FB99FCE4BB62" pageId="6" pageNumber="536" tableUuid="DF36B481FFCA99013106FB99FCE4BB62">Table 1</tableCitation>
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).
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</paragraph>
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<paragraph id="8BF6E409FFCE99053274FF69FCBBBD10" blockId="6.[789,1437,187,1970]" pageId="6" pageNumber="536">
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The result of three different groupings,
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<emphasis id="B93D381BFFCE990535DAFF69FB07BFD0" box="[1211,1229,187,210]" italics="true" pageId="6" pageNumber="536">K</emphasis>
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= 4, 5, 6 recovered more or less highly genetically cohesive groups including
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<taxonomicName id="4C499F8AFFCE9905341DFF09FC0DBE10" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="longifolia">
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<emphasis id="B93D381BFFCE9905341DFF09FC0DBE10" italics="true" pageId="6" pageNumber="536">Veronica longifolia</emphasis>
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</taxonomicName>
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,
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<taxonomicName id="4C499F8AFFCE990532B7FF29FB82BE10" box="[982,1096,251,274]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="pinnata">
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<emphasis id="B93D381BFFCE990532B7FF29FC2CBE10" box="[982,998,251,274]" italics="true" pageId="6" pageNumber="536">V</emphasis>
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.
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<emphasis id="B93D381BFFCE99053295FF29FB82BE10" box="[1012,1096,251,274]" italics="true" pageId="6" pageNumber="536">pinnata</emphasis>
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</taxonomicName>
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<emphasis id="B93D381BFFCE99053529FF29FB3FBE10" box="[1096,1269,251,274]" italics="true" pageId="6" pageNumber="536">
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,
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<taxonomicName id="4C499F8AFFCE99053535FF29FB3FBE10" box="[1108,1269,251,274]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">V. porphyriana</taxonomicName>
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</emphasis>
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and
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<taxonomicName id="4C499F8AFFCE9905344BFF29FA5FBE10" box="[1322,1429,251,274]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="spicata">
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<emphasis id="B93D381BFFCE9905344BFF29FAF0BE10" box="[1322,1338,251,274]" italics="true" pageId="6" pageNumber="536">V</emphasis>
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.
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<emphasis id="B93D381BFFCE99053428FF29FA5FBE10" box="[1353,1429,251,274]" italics="true" pageId="6" pageNumber="536">spicata</emphasis>
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</taxonomicName>
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; however, the individuals from
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<taxonomicName id="4C499F8AFFCE99053500FEC9FB00BE30" box="[1121,1226,283,306]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="incana">
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<emphasis id="B93D381BFFCE99053500FEC9FBBBBE30" box="[1121,1137,283,306]" italics="true" pageId="6" pageNumber="536">V</emphasis>
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||
.
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<emphasis id="B93D381BFFCE990535E3FEC9FB00BE30" box="[1154,1226,283,306]" italics="true" pageId="6" pageNumber="536">incana</emphasis>
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</taxonomicName>
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, in K4 showed admixture of
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<taxonomicName id="4C499F8AFFCE990532ECFEE9FBD8BE50" box="[909,1042,315,338]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="longifolia">
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<emphasis id="B93D381BFFCE990532ECFEE9FC57BE50" box="[909,925,315,338]" italics="true" pageId="6" pageNumber="536">V</emphasis>
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.
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<emphasis id="B93D381BFFCE990532CAFEE9FBD8BE50" box="[939,1042,315,338]" italics="true" pageId="6" pageNumber="536">longifolia</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="4C499F8AFFCE99053529FEE9FB22BE50" box="[1096,1256,315,338]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">
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<emphasis id="B93D381BFFCE99053529FEE9FB22BE50" box="[1096,1256,315,338]" italics="true" pageId="6" pageNumber="536">V. porphyriana</emphasis>
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</taxonomicName>
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, but were recovered as a separate group in K5 and K6 with high genetic portion from
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<taxonomicName id="4C499F8AFFCE990532C5FEA9FB8DBE90" box="[932,1095,379,402]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">
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<emphasis id="B93D381BFFCE990532C5FEA9FC7EBE90" box="[932,948,379,402]" italics="true" pageId="6" pageNumber="536">V</emphasis>
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||
.
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<emphasis id="B93D381BFFCE990532A3FEA9FB8DBE90" box="[962,1095,379,402]" italics="true" pageId="6" pageNumber="536">porphyriana</emphasis>
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</taxonomicName>
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(
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<figureCitation id="1372F88CFFCE99053537FEA9FB5FBE90" box="[1110,1173,379,402]" captionStart="Fig" captionStartId="3.[151,183,1781,1802]" captionTargetBox="[151,1484,678,1747]" captionTargetId="figure-252@3.[153,1484,697,1747]" captionTargetPageId="3" captionText="Fig. 1. A, Genetic composition based on K = 5 of first-level STRUCTURE results and number of individuals included from that particular locality. Genetic composition of the localities with more than one individual has been averaged. Key to the colors is given in a separate inset. B–D, Probability of ancestry of each individual (horizontal axis; total 233 individuals) to each of K = 4, 5, 6 populations (vertical axis). The five populations correspond mostly to the morphotypes hypothesized for species and putative hybrids. V. ×alt, V. ×altaica; V. ×gri, V. ×grisea; V. ×kol, V. ×kolyvanensis; V. ×sap, V. ×sapozhnikovii; V. ×sch, V. ×schmakovii; V. ×ses, V. ×sessiliflora; V. ×smi, V. ×smirnovii; V. are, V. arenosa; V. inca, V. incana; V. lon, V. longifolia; V. pinn, V. pinnata; V. porp, V. porphyriana; V. reve, V. reverdattoi; V. saj, V. sajanensis; V. spic, V. spicata; V. spur, V. spuria; V. ×taig, V. ×taigischensis; uniden, unidentified." figureDoi="http://doi.org/10.5281/zenodo.14047617" httpUri="https://zenodo.org/record/14047617/files/figure.png" pageId="6" pageNumber="536">Fig. 1</figureCitation>
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). Removing the putative hybrids and including only putative purebreds (174 individuals, ~75% of total dataset), the STRUCTURE result of
|
||
<emphasis id="B93D381BFFCE99053415FE69FA4CBED0" box="[1396,1414,443,466]" italics="true" pageId="6" pageNumber="536">K</emphasis>
|
||
= 5 again showed more or less the same results (
|
||
<figureCitation id="1372F88CFFCE9905359FFE09FA85BEF0" box="[1278,1359,475,498]" captionStart="Fig" captionStartId="7.[151,183,1809,1830]" captionTargetBox="[151,1484,392,1773]" captionTargetId="graphics-103@7.[179,937,440,1773]" captionTargetPageId="7" captionText="Fig. 2. Assignments of individuals to K clusters/populations based on Bayesian, multivariate and likelihood approaches as implemented in STRUC- TURE, DAPC (in the adegenet package) and IQ-TREE programs respectively. A, Maximum likelihood unrooted tree reconstructed in IQ-TREE, calibrated with 1000 ultrafast bootstrap replicates. The numbers near the nodes represent bootstrap confidence. This analysis was based on 174 putative pure individuals. B, Probability of ancestry of each individual (horizontal axis; total 174 individuals) to each of K = 5 populations (vertical axis). The five populations correspond mostly to the morphotypes hypothesized for putative pure species. C, Discriminant analysis of principal components using the same dataset as in (B). D, Corresponding morphotypes of K = 5. — Photos: P.A. Kosachev." figureDoi="http://doi.org/10.5281/zenodo.14047619" httpUri="https://zenodo.org/record/14047619/files/figure.png" pageId="6" pageNumber="536">Fig. 2B</figureCitation>
|
||
, suppl.
|
||
<figureCitation id="1372F88CFFCE99053274FE29FCAEBD10" box="[789,868,507,530]" captionStart="Fig" captionStartId="10.[103,135,1865,1886]" captionTargetBox="[278,1262,661,1831]" captionTargetId="figure-267@10.[278,1262,661,1831]" captionTargetPageId="10" captionText="Fig. 3. Chromosome localization of rDNA and genomic in situ hybridization (GISH) in Veronica. Mitotic chromosome complements of: A, V. porphyriana; B, V. ×schmakovii; C, V. spicata; D, V. pinnata; E, V. longifolia; and F, V. incana hybridized with 35S (red fluorescence) and 5S (purple) rDNA probes. G, Mitotic chromosomes of V. ×schmakovii hybridized with gDNA of V. longifolia (red) and V. porphyriana (green). — Chromosomes were counterstained with DAPI. Scale bars, 10 μm." figureDoi="http://doi.org/10.5281/zenodo.14047621" httpUri="https://zenodo.org/record/14047621/files/figure.png" pageId="6" pageNumber="536">Fig. S3</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BF6E409FFCE99053223FDC9FA53BC10" blockId="6.[789,1437,187,1970]" pageId="6" pageNumber="536">
|
||
The DAPC recovered 20 principal components as the best to group our data and clearly resolved all the individuals into
|
||
<emphasis id="B93D381BFFCE990534EAFDE9FA57BD50" box="[1419,1437,571,594]" italics="true" pageId="6" pageNumber="536">K</emphasis>
|
||
= 5 (
|
||
<figureCitation id="1372F88CFFCE9905322DFD89FC6ABD70" box="[844,928,603,626]" captionStart="Fig" captionStartId="7.[151,183,1809,1830]" captionTargetBox="[151,1484,392,1773]" captionTargetId="graphics-103@7.[179,937,440,1773]" captionTargetPageId="7" captionText="Fig. 2. Assignments of individuals to K clusters/populations based on Bayesian, multivariate and likelihood approaches as implemented in STRUC- TURE, DAPC (in the adegenet package) and IQ-TREE programs respectively. A, Maximum likelihood unrooted tree reconstructed in IQ-TREE, calibrated with 1000 ultrafast bootstrap replicates. The numbers near the nodes represent bootstrap confidence. This analysis was based on 174 putative pure individuals. B, Probability of ancestry of each individual (horizontal axis; total 174 individuals) to each of K = 5 populations (vertical axis). The five populations correspond mostly to the morphotypes hypothesized for putative pure species. C, Discriminant analysis of principal components using the same dataset as in (B). D, Corresponding morphotypes of K = 5. — Photos: P.A. Kosachev." figureDoi="http://doi.org/10.5281/zenodo.14047619" httpUri="https://zenodo.org/record/14047619/files/figure.png" pageId="6" pageNumber="536">Fig. 2C</figureCitation>
|
||
; suppl.
|
||
<figureCitation id="1372F88CFFCE9905329BFD89FB87BD70" box="[1018,1101,603,626]" captionStart="Fig" captionStartId="11.[151,183,1865,1886]" captionTargetBox="[276,1316,1153,1829]" captionTargetId="graphics-603@11.[421,1237,1446,1533]" captionTargetPageId="11" captionText="Fig. 4. STRUCTURE results showing the probability of ancestry of each individual (horizontal axis) to each of K = 2 populations (vertical axis) in all the five scenarios. A, Veronica spicata × V. pinnata; B, V. incana and V. longifolia; C, V. longifolia and V. porphyriana; D & E, V. pinnata and V. porphyriana involving putative hybrids of V. ×schmakovii and V. ×sessiliflora. Details of the exact posterior probabilities of each putative hybrid individual and their corresponding parents are given in suppl. Table S3." figureDoi="http://doi.org/10.5281/zenodo.14107798" httpUri="https://zenodo.org/record/14107798/files/figure.png" pageId="6" pageNumber="536">Fig. S4</figureCitation>
|
||
). The DAPC results were supported by the unrooted trees inferred by IQ-TREE (
|
||
<figureCitation id="1372F88CFFCE9905345BFDA9FA47BD90" box="[1338,1421,635,658]" captionStart="Fig" captionStartId="7.[151,183,1809,1830]" captionTargetBox="[151,1484,392,1773]" captionTargetId="graphics-103@7.[179,937,440,1773]" captionTargetPageId="7" captionText="Fig. 2. Assignments of individuals to K clusters/populations based on Bayesian, multivariate and likelihood approaches as implemented in STRUC- TURE, DAPC (in the adegenet package) and IQ-TREE programs respectively. A, Maximum likelihood unrooted tree reconstructed in IQ-TREE, calibrated with 1000 ultrafast bootstrap replicates. The numbers near the nodes represent bootstrap confidence. This analysis was based on 174 putative pure individuals. B, Probability of ancestry of each individual (horizontal axis; total 174 individuals) to each of K = 5 populations (vertical axis). The five populations correspond mostly to the morphotypes hypothesized for putative pure species. C, Discriminant analysis of principal components using the same dataset as in (B). D, Corresponding morphotypes of K = 5. — Photos: P.A. Kosachev." figureDoi="http://doi.org/10.5281/zenodo.14047619" httpUri="https://zenodo.org/record/14047619/files/figure.png" pageId="6" pageNumber="536">Fig. 2A</figureCitation>
|
||
). The morphometric analysis (suppl. Appendix S1.) also supported the results of the genetic analyses in delimiting five main groups similar to those in
|
||
<figureCitation id="1372F88CFFCE99053502FD09FB6BBDF0" box="[1123,1185,731,754]" captionStart="Fig" captionStartId="3.[151,183,1781,1802]" captionTargetBox="[151,1484,678,1747]" captionTargetId="figure-252@3.[153,1484,697,1747]" captionTargetPageId="3" captionText="Fig. 1. A, Genetic composition based on K = 5 of first-level STRUCTURE results and number of individuals included from that particular locality. Genetic composition of the localities with more than one individual has been averaged. Key to the colors is given in a separate inset. B–D, Probability of ancestry of each individual (horizontal axis; total 233 individuals) to each of K = 4, 5, 6 populations (vertical axis). The five populations correspond mostly to the morphotypes hypothesized for species and putative hybrids. V. ×alt, V. ×altaica; V. ×gri, V. ×grisea; V. ×kol, V. ×kolyvanensis; V. ×sap, V. ×sapozhnikovii; V. ×sch, V. ×schmakovii; V. ×ses, V. ×sessiliflora; V. ×smi, V. ×smirnovii; V. are, V. arenosa; V. inca, V. incana; V. lon, V. longifolia; V. pinn, V. pinnata; V. porp, V. porphyriana; V. reve, V. reverdattoi; V. saj, V. sajanensis; V. spic, V. spicata; V. spur, V. spuria; V. ×taig, V. ×taigischensis; uniden, unidentified." figureDoi="http://doi.org/10.5281/zenodo.14047617" httpUri="https://zenodo.org/record/14047617/files/figure.png" pageId="6" pageNumber="536">Fig. 1</figureCitation>
|
||
. Genetic differentiation among species based on AMOVA was
|
||
<emphasis id="B93D381BFFCE990535D6FD29FB0DBC10" box="[1207,1223,763,786]" italics="true" pageId="6" pageNumber="536">F</emphasis>
|
||
<subScript id="17CDE64CFFCE990535A6FCD7FB14BC17" attach="left" box="[1223,1246,773,789]" fontSize="7" pageId="6" pageNumber="536">ST</subScript>
|
||
= 0.16 (
|
||
<tableCitation id="C6CBD1B2FFCE99053459FD29FA43BC10" box="[1336,1417,763,786]" captionStart="Table 2" captionStartId="9.[151,204,1537,1558]" captionTargetId="graphics-775@9.[151,1484,1738,1741]" captionTargetPageId="9" captionText="Table 2. Details of AMOVA in each group with different combination without including the putative hybrid." httpUri="http://table.plazi.org/id/DF36B481FFC1990A31F6F9D3FB48B917" pageId="6" pageNumber="536" tableUuid="DF36B481FFC1990A31F6F9D3FB48B917">Table 2</tableCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BF6E409FFCE99053223FCC9FAB8BCD0" blockId="6.[789,1437,187,1970]" pageId="6" pageNumber="536">
|
||
The hierarchical STRUCTURE analysis retrieved five cohesive groups including
|
||
<taxonomicName id="4C499F8AFFCE99053574FCE9FB74BC50" box="[1045,1214,827,850]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="incana">
|
||
<emphasis id="B93D381BFFCE99053574FCE9FB74BC50" box="[1045,1214,827,850]" italics="true" pageId="6" pageNumber="536">Veronica incana</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFCE990535AAFCE9FA80BC50" box="[1227,1354,827,850]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="longifolia">
|
||
<emphasis id="B93D381BFFCE990535AAFCE9FA80BC50" box="[1227,1354,827,850]" italics="true" pageId="6" pageNumber="536">V. longifolia</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFCE99053439FCE9FC89BC70" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="pinnata">
|
||
<emphasis id="B93D381BFFCE99053439FCE9FC89BC70" italics="true" pageId="6" pageNumber="536">V. pinnata</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFCE99053232FC89FC70BC70" box="[851,954,859,882]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="spicata">
|
||
<emphasis id="B93D381BFFCE99053232FC89FC70BC70" box="[851,954,859,882]" italics="true" pageId="6" pageNumber="536">V. spicata</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFCE99053291FC89FB44BC70" box="[1008,1166,859,882]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFCE99053291FC89FB44BC70" box="[1008,1166,859,882]" italics="true" pageId="6" pageNumber="536">V. porphyriana</emphasis>
|
||
</taxonomicName>
|
||
. In addition to these five groups,
|
||
<taxonomicName id="4C499F8AFFCE99053209FCA9FC1EBC90" box="[872,980,891,914]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="arenosa">
|
||
<emphasis id="B93D381BFFCE99053209FCA9FC1EBC90" box="[872,980,891,914]" italics="true" pageId="6" pageNumber="536">V. arenosa</emphasis>
|
||
</taxonomicName>
|
||
merged with
|
||
<taxonomicName id="4C499F8AFFCE9905353EFCA9FB0DBC90" box="[1119,1223,891,914]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="pinnata">
|
||
<emphasis id="B93D381BFFCE9905353EFCA9FB0DBC90" box="[1119,1223,891,914]" italics="true" pageId="6" pageNumber="536">V. pinnata</emphasis>
|
||
</taxonomicName>
|
||
, while
|
||
<taxonomicName id="4C499F8AFFCE99053470FCA9FA5CBC90" box="[1297,1430,891,914]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="sajanensis">
|
||
<emphasis id="B93D381BFFCE99053470FCA9FA5CBC90" box="[1297,1430,891,914]" italics="true" pageId="6" pageNumber="536">V. sajanensis</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C499F8AFFCE99053221FC49FC51BCB0" box="[832,923,923,946]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="spuria">
|
||
<emphasis id="B93D381BFFCE99053221FC49FC51BCB0" box="[832,923,923,946]" italics="true" pageId="6" pageNumber="536">V. spuria</emphasis>
|
||
</taxonomicName>
|
||
with
|
||
<taxonomicName id="4C499F8AFFCE990532B3FC49FBFFBCB0" box="[978,1077,923,946]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="spicata">
|
||
<emphasis id="B93D381BFFCE990532B3FC49FBFFBCB0" box="[978,1077,923,946]" italics="true" pageId="6" pageNumber="536">V. spicata</emphasis>
|
||
</taxonomicName>
|
||
. However,
|
||
<taxonomicName id="4C499F8AFFCE990535C9FC49FAFBBCB0" box="[1192,1329,923,946]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="reverdattoi">
|
||
<emphasis id="B93D381BFFCE990535C9FC49FAFBBCB0" box="[1192,1329,923,946]" italics="true" pageId="6" pageNumber="536">V. reverdattoi</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFCE99053401FC49FCB5BCD0" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="taigischensis">
|
||
<emphasis id="B93D381BFFCE99053401FC49FCB5BCD0" italics="true" pageId="6" pageNumber="536">V. taigischensis</emphasis>
|
||
</taxonomicName>
|
||
did not fit to any clear group (suppl.
|
||
<tableCitation id="C6CBD1B2FFCE99053464FC69FAAFBCD0" box="[1285,1381,955,978]" captionStart="Table 2" captionStartId="9.[151,204,1537,1558]" captionTargetId="graphics-775@9.[151,1484,1738,1741]" captionTargetPageId="9" captionText="Table 2. Details of AMOVA in each group with different combination without including the putative hybrid." httpUri="http://table.plazi.org/id/DF36B481FFC1990A31F6F9D3FB48B917" pageId="6" pageNumber="536" tableUuid="DF36B481FFC1990A31F6F9D3FB48B917">Table S2</tableCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BF6E409FFCE99053223FC08FC42B9B0" blockId="6.[789,1437,187,1970]" pageId="6" pageNumber="536">
|
||
<emphasis id="B93D381BFFCE99053223FC08FC32BB10" bold="true" pageId="6" pageNumber="536">
|
||
Chromosome localization of rDNA loci and classes of putative hybrids.
|
||
<emphasis id="B93D381BFFCE990532BCFC2FFC32BB10" bold="true" box="[989,1016,1021,1042]" italics="true" pageId="6" pageNumber="536">—</emphasis>
|
||
</emphasis>
|
||
Chromosome numbers and rDNA loci were determined in six species.
|
||
<taxonomicName id="4C499F8AFFCE99053516FBC9FA89BB30" box="[1143,1347,1051,1074]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="longifolia">
|
||
<emphasis id="B93D381BFFCE99053516FBC9FA89BB30" box="[1143,1347,1051,1074]" italics="true" pageId="6" pageNumber="536">Veronica longifolia</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFCE99053435FBC9FC8EBB50" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="pinnata">
|
||
<emphasis id="B93D381BFFCE99053435FBC9FC8EBB50" italics="true" pageId="6" pageNumber="536">V. pinnata</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFCE99053237FBE9FC31BB50" box="[854,1019,1083,1106]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFCE99053237FBE9FC31BB50" box="[854,1019,1083,1106]" italics="true" pageId="6" pageNumber="536">V. porphyriana</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFCE9905356CFBE9FB7ABB50" baseAuthorityName="Pfanzelt" baseAuthorityYear="1181" box="[1037,1200,1083,1106]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" isHybrid="true" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="schmakovii">
|
||
<emphasis id="B93D381BFFCE9905356CFBE9FB7ABB50" box="[1037,1200,1083,1106]" italics="true" pageId="6" pageNumber="536">V. ×schmakovii</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C499F8AFFCE99053595FBE9FA95BB50" box="[1268,1375,1083,1106]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="spicata">
|
||
<emphasis id="B93D381BFFCE99053595FBE9FA95BB50" box="[1268,1375,1083,1106]" italics="true" pageId="6" pageNumber="536">V. spicata</emphasis>
|
||
</taxonomicName>
|
||
were found to have 2
|
||
<emphasis id="B93D381BFFCE990532B1FB89FC17BB70" box="[976,989,1115,1138]" italics="true" pageId="6" pageNumber="536">n</emphasis>
|
||
= 2
|
||
<emphasis id="B93D381BFFCE99053577FB89FBE8BB70" box="[1046,1058,1115,1138]" italics="true" pageId="6" pageNumber="536">x</emphasis>
|
||
= 34 chromosomes and one 5S rDNA-bearing chromosome pair, whereas
|
||
<taxonomicName id="4C499F8AFFCE990535B5FBA9FAFCBB90" box="[1236,1334,1147,1170]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="incana">
|
||
<emphasis id="B93D381BFFCE990535B5FBA9FAFCBB90" box="[1236,1334,1147,1170]" italics="true" pageId="6" pageNumber="536">V. incana</emphasis>
|
||
</taxonomicName>
|
||
had a tetraploid chromosome number (2
|
||
<emphasis id="B93D381BFFCE99053511FB49FBB7BBB0" box="[1136,1149,1179,1202]" italics="true" pageId="6" pageNumber="536">n</emphasis>
|
||
= 4
|
||
<emphasis id="B93D381BFFCE990535CEFB49FB71BBB0" box="[1199,1211,1179,1202]" italics="true" pageId="6" pageNumber="536">x</emphasis>
|
||
= 68) with a corresponding double number of 5S loci, i.e., two 5S-bearing chromosome pairs. In contrast to the conserved number of 5S rDNA, we found a large interspecific variation in the number of 35S rDNA: four NOR-bearing chromosomes in
|
||
<taxonomicName id="4C499F8AFFCE99053434FAC9FCBFBA50" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFCE99053434FAC9FCBFBA50" italics="true" pageId="6" pageNumber="536">V. porphyriana</emphasis>
|
||
</taxonomicName>
|
||
, six in
|
||
<taxonomicName id="4C499F8AFFCE990532B9FAE9FBB1BA50" baseAuthorityName="Pfanzelt" baseAuthorityYear="1181" box="[984,1147,1339,1362]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" isHybrid="true" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="schmakovii">
|
||
<emphasis id="B93D381BFFCE990532B9FAE9FBB1BA50" box="[984,1147,1339,1362]" italics="true" pageId="6" pageNumber="536">V. ×schmakovii</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFCE990535DDFAE9FAE0BA50" box="[1212,1322,1339,1362]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="spicata">
|
||
<emphasis id="B93D381BFFCE990535DDFAE9FAE0BA50" box="[1212,1322,1339,1362]" italics="true" pageId="6" pageNumber="536">V. spicata</emphasis>
|
||
</taxonomicName>
|
||
, seven in
|
||
<taxonomicName id="4C499F8AFFCE99053274FA89FC4ABA70" box="[789,896,1371,1394]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="pinnata">
|
||
<emphasis id="B93D381BFFCE99053274FA89FC4ABA70" box="[789,896,1371,1394]" italics="true" pageId="6" pageNumber="536">V. pinnata</emphasis>
|
||
</taxonomicName>
|
||
, eight in
|
||
<taxonomicName id="4C499F8AFFCE99053285FA89FBAFBA70" box="[996,1125,1371,1394]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="longifolia">
|
||
<emphasis id="B93D381BFFCE99053285FA89FBAFBA70" box="[996,1125,1371,1394]" italics="true" pageId="6" pageNumber="536">V. longifolia</emphasis>
|
||
</taxonomicName>
|
||
, and at least nine in the tetraploid
|
||
<taxonomicName id="4C499F8AFFCE99053237FAA9FC76BA90" box="[854,956,1403,1426]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="incana">
|
||
<emphasis id="B93D381BFFCE99053237FAA9FC76BA90" box="[854,956,1403,1426]" italics="true" pageId="6" pageNumber="536">V. incana</emphasis>
|
||
</taxonomicName>
|
||
. In
|
||
<taxonomicName id="4C499F8AFFCE9905328CFAA9FB99BA90" box="[1005,1107,1403,1426]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="incana">
|
||
<emphasis id="B93D381BFFCE9905328CFAA9FB99BA90" box="[1005,1107,1403,1426]" italics="true" pageId="6" pageNumber="536">V. incana</emphasis>
|
||
</taxonomicName>
|
||
, additional very weak signals were observed (
|
||
<figureCitation id="1372F88CFFCE990532A8FA49FBC7BAB0" box="[969,1037,1435,1458]" captionStart="Fig" captionStartId="10.[103,135,1865,1886]" captionTargetBox="[278,1262,661,1831]" captionTargetId="figure-267@10.[278,1262,661,1831]" captionTargetPageId="10" captionText="Fig. 3. Chromosome localization of rDNA and genomic in situ hybridization (GISH) in Veronica. Mitotic chromosome complements of: A, V. porphyriana; B, V. ×schmakovii; C, V. spicata; D, V. pinnata; E, V. longifolia; and F, V. incana hybridized with 35S (red fluorescence) and 5S (purple) rDNA probes. G, Mitotic chromosomes of V. ×schmakovii hybridized with gDNA of V. longifolia (red) and V. porphyriana (green). — Chromosomes were counterstained with DAPI. Scale bars, 10 μm." figureDoi="http://doi.org/10.5281/zenodo.14047621" httpUri="https://zenodo.org/record/14047621/files/figure.png" pageId="6" pageNumber="536">Fig. 3</figureCitation>
|
||
). Each gDNA of
|
||
<taxonomicName id="4C499F8AFFCE99053583FA49FAA3BAB0" box="[1250,1385,1435,1458]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="longifolia">
|
||
<emphasis id="B93D381BFFCE99053583FA49FAA3BAB0" box="[1250,1385,1435,1458]" italics="true" pageId="6" pageNumber="536">V. longifolia</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFCE99053274FA69FC73BAD0" box="[789,953,1467,1490]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFCE99053274FA69FC73BAD0" box="[789,953,1467,1490]" italics="true" pageId="6" pageNumber="536">V. porphyriana</emphasis>
|
||
</taxonomicName>
|
||
hybridized to one half of the chromosome complement of
|
||
<taxonomicName id="4C499F8AFFCE990532A9FA09FBA1BAF0" baseAuthorityName="Pfanzelt" baseAuthorityYear="1181" box="[968,1131,1499,1522]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" isHybrid="true" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="schmakovii">
|
||
<emphasis id="B93D381BFFCE990532A9FA09FBA1BAF0" box="[968,1131,1499,1522]" italics="true" pageId="6" pageNumber="536">V. ×schmakovii</emphasis>
|
||
</taxonomicName>
|
||
. Therefore, GISH strongly suggests a homoploid hybrid origin of
|
||
<taxonomicName id="4C499F8AFFCE9905359BFA29FA56B910" baseAuthorityName="Pfanzelt" baseAuthorityYear="1181" box="[1274,1436,1531,1554]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" isHybrid="true" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="schmakovii">
|
||
<emphasis id="B93D381BFFCE9905359BFA29FA56B910" box="[1274,1436,1531,1554]" italics="true" pageId="6" pageNumber="536">V. ×schmakovii</emphasis>
|
||
</taxonomicName>
|
||
from parental genomes closely related to
|
||
<taxonomicName id="4C499F8AFFCE99053584F9C9FAA1B930" box="[1253,1387,1563,1586]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="longifolia">
|
||
<emphasis id="B93D381BFFCE99053584F9C9FAA1B930" box="[1253,1387,1563,1586]" italics="true" pageId="6" pageNumber="536">V. longifolia</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFCE99053274F9E9FC7FB950" box="[789,949,1595,1618]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFCE99053274F9E9FC7FB950" box="[789,949,1595,1618]" italics="true" pageId="6" pageNumber="536">V. porphyriana</emphasis>
|
||
</taxonomicName>
|
||
. This is also supported by the number of 35S rDNA-bearing chromosomes in
|
||
<taxonomicName id="4C499F8AFFCE99053513F989FADFB970" baseAuthorityName="Pfanzelt" baseAuthorityYear="1181" box="[1138,1301,1627,1650]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" isHybrid="true" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="schmakovii">
|
||
<emphasis id="B93D381BFFCE99053513F989FADFB970" box="[1138,1301,1627,1650]" italics="true" pageId="6" pageNumber="536">V. ×schmakovii</emphasis>
|
||
</taxonomicName>
|
||
(six), an intermediate value between
|
||
<taxonomicName id="4C499F8AFFCE99053542F9A9FB69B990" box="[1059,1187,1659,1682]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="longifolia">
|
||
<emphasis id="B93D381BFFCE99053542F9A9FB69B990" box="[1059,1187,1659,1682]" italics="true" pageId="6" pageNumber="536">V. longifolia</emphasis>
|
||
</taxonomicName>
|
||
(eight) and
|
||
<taxonomicName id="4C499F8AFFCE9905347FF9A9FCF7B9B0" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFCE9905347FF9A9FCF7B9B0" italics="true" pageId="6" pageNumber="536">V. porphyriana</emphasis>
|
||
</taxonomicName>
|
||
(four).
|
||
</paragraph>
|
||
<paragraph id="8BF6E409FFCE990A3223F969FEC2BE70" blockId="6.[789,1437,187,1970]" lastBlockId="9.[151,799,187,1266]" lastPageId="9" lastPageNumber="539" pageId="6" pageNumber="536">
|
||
The hypothesis of hybridization between and among species of
|
||
<taxonomicName id="4C499F8AFFCE99053204F909FC0BB9F0" box="[869,961,1755,1778]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="6" pageNumber="536" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="B93D381BFFCE99053204F909FC0BB9F0" box="[869,961,1755,1778]" italics="true" pageId="6" pageNumber="536">Veronica</emphasis>
|
||
</taxonomicName>
|
||
in the
|
||
<collectingRegion id="498D2AEBFFCE99053570F909FB8FB9F0" box="[1041,1093,1755,1778]" country="Russia" name="Altay" pageId="6" pageNumber="536">Altai</collectingRegion>
|
||
Mountains based on their morphotypes was mostly supported by STRUCTURE, and NewHybrids (
|
||
<figureCitation id="1372F88CFFCE990532D7F8C9FBCBB830" box="[950,1025,1819,1842]" captionStart="Fig" captionStartId="11.[151,183,1865,1886]" captionTargetBox="[276,1316,1153,1829]" captionTargetId="graphics-603@11.[421,1237,1446,1533]" captionTargetPageId="11" captionText="Fig. 4. STRUCTURE results showing the probability of ancestry of each individual (horizontal axis) to each of K = 2 populations (vertical axis) in all the five scenarios. A, Veronica spicata × V. pinnata; B, V. incana and V. longifolia; C, V. longifolia and V. porphyriana; D & E, V. pinnata and V. porphyriana involving putative hybrids of V. ×schmakovii and V. ×sessiliflora. Details of the exact posterior probabilities of each putative hybrid individual and their corresponding parents are given in suppl. Table S3." figureDoi="http://doi.org/10.5281/zenodo.14107798" httpUri="https://zenodo.org/record/14107798/files/figure.png" pageId="6" pageNumber="536">Fig. 4</figureCitation>
|
||
; suppl. Table S3). The first-level STRUCTURE analysis is a bit more inconclusive in that respect showing admixture of various levels involving both the biparental hybrids and potential triparental hybrid (
|
||
<figureCitation id="1372F88CFFCE9905342BF8A9FA47B890" box="[1354,1421,1915,1938]" captionStart="Fig" captionStartId="7.[151,183,1809,1830]" captionTargetBox="[151,1484,392,1773]" captionTargetId="graphics-103@7.[179,937,440,1773]" captionTargetPageId="7" captionText="Fig. 2. Assignments of individuals to K clusters/populations based on Bayesian, multivariate and likelihood approaches as implemented in STRUC- TURE, DAPC (in the adegenet package) and IQ-TREE programs respectively. A, Maximum likelihood unrooted tree reconstructed in IQ-TREE, calibrated with 1000 ultrafast bootstrap replicates. The numbers near the nodes represent bootstrap confidence. This analysis was based on 174 putative pure individuals. B, Probability of ancestry of each individual (horizontal axis; total 174 individuals) to each of K = 5 populations (vertical axis). The five populations correspond mostly to the morphotypes hypothesized for putative pure species. C, Discriminant analysis of principal components using the same dataset as in (B). D, Corresponding morphotypes of K = 5. — Photos: P.A. Kosachev." figureDoi="http://doi.org/10.5281/zenodo.14047619" httpUri="https://zenodo.org/record/14047619/files/figure.png" pageId="6" pageNumber="536">Fig. 2</figureCitation>
|
||
). The maximum likelihood tree based on all individuals also revealed that the putative hybrid individuals are mostly sister to one of the suggested parents (suppl.
|
||
<figureCitation id="1372F88CFFCF9904335EFF09FD45BFF0" box="[575,655,219,242]" captionStart="Fig" captionStartId="12.[103,135,1697,1718]" captionTargetBox="[255,1368,842,1661]" captionTargetId="graphics-396@12.[278,1111,857,1635]" captionTargetPageId="12" captionText="Fig. 5. Results of the G-PhoCS analysis for effective population sizes, and gene flow using only pure individuals (no admixture). The inferred current effective population size (Ne) of each species are given for all the five species. The direction of the arrows represents the probability of migration among the species both in forward and reverse directions. The width of the bars represents the effective population size of each species. For population size estimation, we used the equations Ne (effective population size) = θ / 4μg; and T (divergence time) = τ · g / μ; where substitution rate/site/year (μ) = 2.44E-9, generation time for population (g) = 10 years. Migration rates are based on per generation parameter (Msx = msx · θx / 4), which is the proportion of individuals in population x arrived by migration from another population per generation. Gene flow has been calculated using the total migration rate, cases where the total rate is low, it approximates the probability of gene flow between the two species. However, for higher rates, we adjusted probabilities into rates with the equation P = 1 − e−m (where P = the probability of gene flow, e = exponent, and m = total migration rate; following vonHoldt & al., 2016). The phylogenetic tree on which the G-PhoCS analysis has been based is given in suppl. Fig. S2. For complete details, see in Materials and Methods as well as suppl. Tables S4 and S5 for migration rates (msx), τ and θ values, and divergence times." figureDoi="http://doi.org/10.5281/zenodo.14047623" httpUri="https://zenodo.org/record/14047623/files/figure.png" pageId="7" pageNumber="537">Fig. S5</figureCitation>
|
||
). The genetically admixed individuals were mostly the ones identified as hybrids a priori by morphological characteristics (suppl.
|
||
<figureCitation id="1372F88CFFCF99043395FEC9FF7EBE50" captionStart="Fig" captionStartId="3.[151,183,1781,1802]" captionTargetBox="[151,1484,678,1747]" captionTargetId="figure-252@3.[153,1484,697,1747]" captionTargetPageId="3" captionText="Fig. 1. A, Genetic composition based on K = 5 of first-level STRUCTURE results and number of individuals included from that particular locality. Genetic composition of the localities with more than one individual has been averaged. Key to the colors is given in a separate inset. B–D, Probability of ancestry of each individual (horizontal axis; total 233 individuals) to each of K = 4, 5, 6 populations (vertical axis). The five populations correspond mostly to the morphotypes hypothesized for species and putative hybrids. V. ×alt, V. ×altaica; V. ×gri, V. ×grisea; V. ×kol, V. ×kolyvanensis; V. ×sap, V. ×sapozhnikovii; V. ×sch, V. ×schmakovii; V. ×ses, V. ×sessiliflora; V. ×smi, V. ×smirnovii; V. are, V. arenosa; V. inca, V. incana; V. lon, V. longifolia; V. pinn, V. pinnata; V. porp, V. porphyriana; V. reve, V. reverdattoi; V. saj, V. sajanensis; V. spic, V. spicata; V. spur, V. spuria; V. ×taig, V. ×taigischensis; uniden, unidentified." figureDoi="http://doi.org/10.5281/zenodo.14047617" httpUri="https://zenodo.org/record/14047617/files/figure.png" pageId="7" pageNumber="537">Fig. S1</figureCitation>
|
||
); however, we also recovered admixture in individuals considered to be pure species, e.g.,
|
||
<taxonomicName id="4C499F8AFFCF990435A7FF69FA9CBFD0" box="[1222,1366,187,210]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="7" pageNumber="537" phylum="Tracheophyta" rank="species" species="sajanensis">
|
||
<emphasis id="B93D381BFFCF990435A7FF69FB1CBFD0" box="[1222,1238,187,210]" italics="true" pageId="7" pageNumber="537">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFCF99043584FF69FA9CBFD0" box="[1253,1366,187,210]" italics="true" pageId="7" pageNumber="537">sajanensis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B93D381BFFCF99043437FF69FA90BFD0" box="[1366,1370,187,210]" italics="true" pageId="7" pageNumber="537">,</emphasis>
|
||
<taxonomicName id="4C499F8AFFCF99043402FF69FA0CBFD0" box="[1379,1478,187,210]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="7" pageNumber="537" phylum="Tracheophyta" rank="species" species="spuria">
|
||
<emphasis id="B93D381BFFCF99043402FF69FAB9BFD0" box="[1379,1395,187,210]" italics="true" pageId="7" pageNumber="537">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFCF990434E3FF69FA0CBFD0" box="[1410,1478,187,210]" italics="true" pageId="7" pageNumber="537">spuria</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFCF99043224FF09FC23BFF0" box="[837,1001,219,242]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="7" pageNumber="537" phylum="Tracheophyta" rank="species" species="taigischensis">
|
||
<emphasis id="B93D381BFFCF99043224FF09FC9FBFF0" box="[837,853,219,242]" italics="true" pageId="7" pageNumber="537">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFCF99043200FF09FC23BFF0" box="[865,1001,219,242]" italics="true" pageId="7" pageNumber="537">taigischensis</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C499F8AFFCF99043541FF09FB64BFF0" box="[1056,1198,219,242]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="7" pageNumber="537" phylum="Tracheophyta" rank="species" species="reverdattoi">
|
||
<emphasis id="B93D381BFFCF99043541FF09FBFABFF0" box="[1056,1072,219,242]" italics="true" pageId="7" pageNumber="537">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFCF9904355AFF09FB64BFF0" box="[1083,1198,219,242]" italics="true" pageId="7" pageNumber="537">reverdattoi</emphasis>
|
||
</taxonomicName>
|
||
. The STRUCTURE results showed that in most cases the putative hybrid individuals in all five scenarios had posterior probabilities ranging between 0.15 <Q <0.50 or 0.50 <Q <0.85, which suggests a hybrid swarm from F1 to later-generation hybrids (
|
||
<figureCitation id="1372F88CFFC1990A33E6FF69FD00BFD0" box="[647,714,187,210]" captionStart="Fig" captionStartId="11.[151,183,1865,1886]" captionTargetBox="[276,1316,1153,1829]" captionTargetId="graphics-603@11.[421,1237,1446,1533]" captionTargetPageId="11" captionText="Fig. 4. STRUCTURE results showing the probability of ancestry of each individual (horizontal axis) to each of K = 2 populations (vertical axis) in all the five scenarios. A, Veronica spicata × V. pinnata; B, V. incana and V. longifolia; C, V. longifolia and V. porphyriana; D & E, V. pinnata and V. porphyriana involving putative hybrids of V. ×schmakovii and V. ×sessiliflora. Details of the exact posterior probabilities of each putative hybrid individual and their corresponding parents are given in suppl. Table S3." figureDoi="http://doi.org/10.5281/zenodo.14107798" httpUri="https://zenodo.org/record/14107798/files/figure.png" pageId="9" pageNumber="539">Fig. 4</figureCitation>
|
||
; suppl. Table S3). The analysis implemented in NewHybrids to group the hybrid individuals supported the results of STRUCTURE with the exception of scenario 5 and recovered hybrids of different classes but dominated by F1 (suppl. Table S3).
|
||
</paragraph>
|
||
<caption id="DF36B481FFCF990431F6F8C3FB7CB8B3" ID-DOI="http://doi.org/10.5281/zenodo.14047619" ID-Zenodo-Dep="14047619" httpUri="https://zenodo.org/record/14047619/files/figure.png" pageId="7" pageNumber="537" startId="7.[151,183,1809,1830]" targetBox="[151,1484,392,1773]" targetPageId="7" targetType="figure">
|
||
<paragraph id="8BF6E409FFCF990431F6F8C3FB7CB8B3" blockId="7.[151,1485,1809,1970]" pageId="7" pageNumber="537">
|
||
<emphasis id="B93D381BFFCF990431F6F8C3FF1AB824" bold="true" box="[151,208,1809,1830]" pageId="7" pageNumber="537">Fig. 2.</emphasis>
|
||
Assignments of individuals to
|
||
<emphasis id="B93D381BFFCF99043088F8C3FE32B827" box="[489,504,1809,1829]" italics="true" pageId="7" pageNumber="537">K</emphasis>
|
||
clusters/populations based on Bayesian, multivariate and likelihood approaches as implemented in STRUCTURE, DAPC (in the adegenet package) and IQ-TREE programs respectively.
|
||
<emphasis id="B93D381BFFCF9904320FF8FFFCB5B843" bold="true" box="[878,895,1837,1857]" pageId="7" pageNumber="537">A</emphasis>
|
||
, Maximum likelihood unrooted tree reconstructed in IQ-TREE, calibrated with 1000 ultrafast bootstrap replicates. The numbers near the nodes represent bootstrap confidence. This analysis was based on 174 putative pure individuals.
|
||
<emphasis id="B93D381BFFCF9904300EF8B7FEB4B87B" bold="true" box="[367,382,1893,1913]" pageId="7" pageNumber="537">B</emphasis>
|
||
, Probability of ancestry of each individual (horizontal axis; total 174 individuals) to each of
|
||
<emphasis id="B93D381BFFCF990435B4F8B7FB2EB87B" box="[1237,1252,1893,1913]" italics="true" pageId="7" pageNumber="537">K</emphasis>
|
||
= 5 populations (vertical axis). The five populations correspond mostly to the morphotypes hypothesized for putative pure species.
|
||
<emphasis id="B93D381BFFCF99043514F853FB4CB897" bold="true" box="[1141,1158,1921,1941]" pageId="7" pageNumber="537">C</emphasis>
|
||
, Discriminant analysis of principal components using the same dataset as in (B).
|
||
<emphasis id="B93D381BFFCF9904335DF84FFD87B8B3" bold="true" box="[572,589,1949,1969]" pageId="7" pageNumber="537">D</emphasis>
|
||
, Corresponding morphotypes of
|
||
<emphasis id="B93D381BFFCF990432E1F84FFC45B8B3" box="[896,911,1949,1969]" italics="true" pageId="7" pageNumber="537">K</emphasis>
|
||
= 5. — Photos: P.A. Kosachev.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF36B481FFC0990B33CFF8A8FC9CB88D" box="[686,854,1914,1935]" isContinuationCaption="true" pageId="8" pageNumber="538" startId="8.[686,718,1914,1935]" targetBox="[103,1436,175,1882]" targetPageId="8" targetType="figure">
|
||
<paragraph id="8BF6E409FFC0990B33CFF8A8FC9CB88D" blockId="8.[686,854,1914,1935]" box="[686,854,1914,1935]" pageId="8" pageNumber="538">
|
||
<emphasis id="B93D381BFFC0990B33CFF8A8FD23B88D" bold="true" box="[686,745,1914,1935]" pageId="8" pageNumber="538">Fig. 2.</emphasis>
|
||
Continued.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BF6E409FFC1990A31A5FEA9FDA8BBF0" blockId="9.[151,799,187,1266]" pageId="9" pageNumber="539">
|
||
<emphasis id="B93D381BFFC1990A31A5FEA9FD00BE90" bold="true" box="[196,714,379,403]" pageId="9" pageNumber="539">
|
||
Demographic history of pure species.
|
||
<emphasis id="B93D381BFFC1990A33CEFEAFFD00BE90" bold="true" box="[687,714,381,402]" italics="true" pageId="9" pageNumber="539">—</emphasis>
|
||
</emphasis>
|
||
Using G-PhoCS to model the associated demographic history in the species of
|
||
<taxonomicName id="4C499F8AFFC1990A304BFE69FE4CBED0" box="[298,390,443,466]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="genus">
|
||
<emphasis id="B93D381BFFC1990A304BFE69FE4CBED0" box="[298,390,443,466]" italics="true" pageId="9" pageNumber="539">Veronica</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="1372F88CFFC1990A30F5FE69FE18BED0" box="[404,466,443,466]" captionStart="Fig" captionStartId="12.[103,135,1697,1718]" captionTargetBox="[255,1368,842,1661]" captionTargetId="graphics-396@12.[278,1111,857,1635]" captionTargetPageId="12" captionText="Fig. 5. Results of the G-PhoCS analysis for effective population sizes, and gene flow using only pure individuals (no admixture). The inferred current effective population size (Ne) of each species are given for all the five species. The direction of the arrows represents the probability of migration among the species both in forward and reverse directions. The width of the bars represents the effective population size of each species. For population size estimation, we used the equations Ne (effective population size) = θ / 4μg; and T (divergence time) = τ · g / μ; where substitution rate/site/year (μ) = 2.44E-9, generation time for population (g) = 10 years. Migration rates are based on per generation parameter (Msx = msx · θx / 4), which is the proportion of individuals in population x arrived by migration from another population per generation. Gene flow has been calculated using the total migration rate, cases where the total rate is low, it approximates the probability of gene flow between the two species. However, for higher rates, we adjusted probabilities into rates with the equation P = 1 − e−m (where P = the probability of gene flow, e = exponent, and m = total migration rate; following vonHoldt & al., 2016). The phylogenetic tree on which the G-PhoCS analysis has been based is given in suppl. Fig. S2. For complete details, see in Materials and Methods as well as suppl. Tables S4 and S5 for migration rates (msx), τ and θ values, and divergence times." figureDoi="http://doi.org/10.5281/zenodo.14047623" httpUri="https://zenodo.org/record/14047623/files/figure.png" pageId="9" pageNumber="539">Fig. 5</figureCitation>
|
||
), we retrieved divergence times between the major clades and branches to be recent (1.14 Mya to 1.57 Mya) at 95% HPD (highest posterior density; see suppl. Table S4 for complete information). Even with the upper HPD at 95%, the divergence time was in the range of 1.35 Mya to 1.84 Mya. Among the five species, the earliest recently diverged species are
|
||
<taxonomicName id="4C499F8AFFC1990A30AFFDA9FDBCBD90" box="[462,630,635,658]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFC1990A30AFFDA9FE14BD90" box="[462,478,635,658]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A3090FDA9FDBCBD90" box="[497,630,635,658]" italics="true" pageId="9" pageNumber="539">porphyriana</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFC1990A33D2FDA9FCD4BD90" box="[691,798,635,658]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="incana">
|
||
<emphasis id="B93D381BFFC1990A33D2FDA9FD09BD90" box="[691,707,635,658]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A33B7FDA9FCD4BD90" box="[726,798,635,658]" italics="true" pageId="9" pageNumber="539">incana</emphasis>
|
||
</taxonomicName>
|
||
(1.14 Mya), while the most divergent species is
|
||
<taxonomicName id="4C499F8AFFC1990A33CDFD49FCD4BDB0" box="[684,798,667,690]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="pinnata">
|
||
<emphasis id="B93D381BFFC1990A33CDFD49FD76BDB0" box="[684,700,667,690]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A33ACFD49FCD4BDB0" box="[717,798,667,690]" italics="true" pageId="9" pageNumber="539">pinnata</emphasis>
|
||
</taxonomicName>
|
||
(1.84 Mya) at upper 95% HPD. Similarly, the lowest effective population size is that of
|
||
<taxonomicName id="4C499F8AFFC1990A30CEFD09FDEABDF0" box="[431,544,731,754]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="pinnata">
|
||
<emphasis id="B93D381BFFC1990A30CEFD09FE75BDF0" box="[431,447,731,754]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A30AEFD09FDEABDF0" box="[463,544,731,754]" italics="true" pageId="9" pageNumber="539">pinnata</emphasis>
|
||
</taxonomicName>
|
||
(4280 individuals) followed by
|
||
<taxonomicName id="4C499F8AFFC1990A319DFD29FE54BC10" box="[252,414,763,786]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFC1990A319DFD29FEC6BC10" box="[252,268,763,786]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A3078FD29FE54BC10" box="[281,414,763,786]" italics="true" pageId="9" pageNumber="539">porphyriana</emphasis>
|
||
</taxonomicName>
|
||
(6150 individuals), while the largest one belongs to
|
||
<taxonomicName id="4C499F8AFFC1990A3057FCC9FE56BC30" box="[310,412,795,818]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="spicata">
|
||
<emphasis id="B93D381BFFC1990A3057FCC9FE56BC30" box="[310,412,795,818]" italics="true" pageId="9" pageNumber="539">V. spicata</emphasis>
|
||
</taxonomicName>
|
||
(21,000 individuals) at average 95% HPD. The effective population sizes of their ancestors were also different in comparison with the extant effective population sizes of all the five species, e.g., the smallest one of
|
||
<taxonomicName id="4C499F8AFFC1990A31F6FC49FEF6BCB0" box="[151,316,923,946]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFC1990A31F6FC49FF6DBCB0" box="[151,167,923,946]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A31D6FC49FEF6BCB0" box="[183,316,923,946]" italics="true" pageId="9" pageNumber="539">porphyriana</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFC1990A3012FC49FE15BCB0" box="[371,479,923,946]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="spicata">
|
||
<emphasis id="B93D381BFFC1990A3012FC49FE49BCB0" box="[371,387,923,946]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A30F2FC49FE15BCB0" box="[403,479,923,946]" italics="true" pageId="9" pageNumber="539">spicata</emphasis>
|
||
</taxonomicName>
|
||
ancestors (1160 individuals), whereas the largest one for the ancestor population of all species (16,100 individuals) (
|
||
<figureCitation id="1372F88CFFC1990A30C7FC09FE29BCF0" box="[422,483,987,1010]" captionStart="Fig" captionStartId="12.[103,135,1697,1718]" captionTargetBox="[255,1368,842,1661]" captionTargetId="graphics-396@12.[278,1111,857,1635]" captionTargetPageId="12" captionText="Fig. 5. Results of the G-PhoCS analysis for effective population sizes, and gene flow using only pure individuals (no admixture). The inferred current effective population size (Ne) of each species are given for all the five species. The direction of the arrows represents the probability of migration among the species both in forward and reverse directions. The width of the bars represents the effective population size of each species. For population size estimation, we used the equations Ne (effective population size) = θ / 4μg; and T (divergence time) = τ · g / μ; where substitution rate/site/year (μ) = 2.44E-9, generation time for population (g) = 10 years. Migration rates are based on per generation parameter (Msx = msx · θx / 4), which is the proportion of individuals in population x arrived by migration from another population per generation. Gene flow has been calculated using the total migration rate, cases where the total rate is low, it approximates the probability of gene flow between the two species. However, for higher rates, we adjusted probabilities into rates with the equation P = 1 − e−m (where P = the probability of gene flow, e = exponent, and m = total migration rate; following vonHoldt & al., 2016). The phylogenetic tree on which the G-PhoCS analysis has been based is given in suppl. Fig. S2. For complete details, see in Materials and Methods as well as suppl. Tables S4 and S5 for migration rates (msx), τ and θ values, and divergence times." figureDoi="http://doi.org/10.5281/zenodo.14047623" httpUri="https://zenodo.org/record/14047623/files/figure.png" pageId="9" pageNumber="539">Fig. 5</figureCitation>
|
||
; suppl. Table S4). Our modeling of gene flow in both forward and reverse directions showed high probability of frequent gene flow from
|
||
<taxonomicName id="4C499F8AFFC1990A33A3FBC9FF0DBB50" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="longifolia">
|
||
<emphasis id="B93D381BFFC1990A33A3FBC9FD18BB30" box="[706,722,1051,1074]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A33BEFBC9FF0DBB50" italics="true" pageId="9" pageNumber="539">longifolia</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFC1990A319DFBE9FEA6BB50" box="[252,364,1083,1106]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="pinnata">
|
||
<emphasis id="B93D381BFFC1990A319DFBE9FEC6BB50" box="[252,268,1083,1106]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A307AFBE9FEA6BB50" box="[283,364,1083,1106]" italics="true" pageId="9" pageNumber="539">pinnata</emphasis>
|
||
</taxonomicName>
|
||
to
|
||
<taxonomicName id="4C499F8AFFC1990A30EEFBE9FE3FBB50" box="[399,501,1083,1106]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="incana">
|
||
<emphasis id="B93D381BFFC1990A30EEFBE9FE55BB50" box="[399,415,1083,1106]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A30CCFBE9FE3FBB50" box="[429,501,1083,1106]" italics="true" pageId="9" pageNumber="539">incana</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFC1990A3365FBE9FD6EBB50" box="[516,676,1083,1106]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFC1990A3365FBE9FD6EBB50" box="[516,676,1083,1106]" italics="true" pageId="9" pageNumber="539">V. porphyriana</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFC1990A33B8FBE9FF0FBB70" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="spicata">
|
||
<emphasis id="B93D381BFFC1990A33B8FBE9FD23BB50" box="[729,745,1083,1106]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A3396FBE9FF0FBB70" italics="true" pageId="9" pageNumber="539">spicata</emphasis>
|
||
</taxonomicName>
|
||
. Nevertheless, from
|
||
<taxonomicName id="4C499F8AFFC1990A30CEFB89FDD0BB70" box="[431,538,1115,1138]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="incana">
|
||
<emphasis id="B93D381BFFC1990A30CEFB89FE75BB70" box="[431,447,1115,1138]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A30B3FB89FDD0BB70" box="[466,538,1115,1138]" italics="true" pageId="9" pageNumber="539">incana</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFC1990A3336FB89FD35BB70" box="[599,767,1115,1138]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFC1990A3336FB89FDADBB70" box="[599,615,1115,1138]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A331BFB89FD35BB70" box="[634,767,1115,1138]" italics="true" pageId="9" pageNumber="539">porphyriana</emphasis>
|
||
</taxonomicName>
|
||
to the other three species, the probability of gene flow was also comparatively high. Similarly, the gene flow among
|
||
<taxonomicName id="4C499F8AFFC1990A33D5FB49FCDDBBB0" box="[692,791,1179,1202]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="incana">
|
||
<emphasis id="B93D381BFFC1990A33D5FB49FD0EBBB0" box="[692,708,1179,1202]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A33AEFB49FCDDBBB0" box="[719,791,1179,1202]" italics="true" pageId="9" pageNumber="539">incana</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C499F8AFFC1990A31F6FB69FEF1BBD0" box="[151,315,1211,1234]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="porphyriana">
|
||
<emphasis id="B93D381BFFC1990A31F6FB69FF6DBBD0" box="[151,167,1211,1234]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A31D7FB69FEF1BBD0" box="[182,315,1211,1234]" italics="true" pageId="9" pageNumber="539">porphyriana</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C499F8AFFC1990A3011FB69FE11BBD0" box="[368,475,1211,1234]" class="Magnoliopsida" family="Plantaginaceae" genus="Veronica" kingdom="Plantae" order="Lamiales" pageId="9" pageNumber="539" phylum="Tracheophyta" rank="species" species="spicata">
|
||
<emphasis id="B93D381BFFC1990A3011FB69FE4ABBD0" box="[368,384,1211,1234]" italics="true" pageId="9" pageNumber="539">V</emphasis>
|
||
.
|
||
<emphasis id="B93D381BFFC1990A30EEFB69FE11BBD0" box="[399,475,1211,1234]" italics="true" pageId="9" pageNumber="539">spicata</emphasis>
|
||
</taxonomicName>
|
||
was high in both forward and reverse directions (
|
||
<figureCitation id="1372F88CFFC1990A3001FB09FE55BBF0" box="[352,415,1243,1266]" captionStart="Fig" captionStartId="12.[103,135,1697,1718]" captionTargetBox="[255,1368,842,1661]" captionTargetId="graphics-396@12.[278,1111,857,1635]" captionTargetPageId="12" captionText="Fig. 5. Results of the G-PhoCS analysis for effective population sizes, and gene flow using only pure individuals (no admixture). The inferred current effective population size (Ne) of each species are given for all the five species. The direction of the arrows represents the probability of migration among the species both in forward and reverse directions. The width of the bars represents the effective population size of each species. For population size estimation, we used the equations Ne (effective population size) = θ / 4μg; and T (divergence time) = τ · g / μ; where substitution rate/site/year (μ) = 2.44E-9, generation time for population (g) = 10 years. Migration rates are based on per generation parameter (Msx = msx · θx / 4), which is the proportion of individuals in population x arrived by migration from another population per generation. Gene flow has been calculated using the total migration rate, cases where the total rate is low, it approximates the probability of gene flow between the two species. However, for higher rates, we adjusted probabilities into rates with the equation P = 1 − e−m (where P = the probability of gene flow, e = exponent, and m = total migration rate; following vonHoldt & al., 2016). The phylogenetic tree on which the G-PhoCS analysis has been based is given in suppl. Fig. S2. For complete details, see in Materials and Methods as well as suppl. Tables S4 and S5 for migration rates (msx), τ and θ values, and divergence times." figureDoi="http://doi.org/10.5281/zenodo.14047623" httpUri="https://zenodo.org/record/14047623/files/figure.png" pageId="9" pageNumber="539">Fig. 5</figureCitation>
|
||
; suppl. Table S4).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |